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TRANSACTIONS

OF

THE ZOOLOGICAL SOCIETY

OF LONDON.

VOLUME VI.

%j

LONDON:

PRINTED FOR THE SOCIETY :
SOLD AT THEIR HOUSE IN HANOVER-SQUARE;
AND BY MESSRS. LONGMANS, GREEN. READER, AND DYER, PATERNOSTER-ROW,

1869.

CONTENTS.

I. On the Characters and Affinities of Potamogale, a genus of Insectivorous Mammals.
By Guorce J. Auman, F.R.S., Corr. Mem. Zool. Soc. Lond., Regius Professor of
Natural History in the University of Edinburgh, and oes Keeper of the Edin-

burgh Museum of Natural History. . . . . See ER pACeML
Il. On some Indian Cetacea collected by Water Exuiot, Esg. By Professor Owen,
roe PE ILA Ee ON oat pe wes ge ee ee eme
Ill. On the Osteology of the Dodo ea ineptus, Linn. 2 By Professor Owsn, F.R.S.,
TELS EOE. a ee c MRR I nes UES LSS aR Re eek staan

IV. Description of the Skeleton ay Inia geoffrensis and of the Skull of Pontoporia
blainvillii, with Remarks on the Systematic Position of these Animals in the
Order Cuvacea. By Wiut1am Henry Fuowsr, /.BS., FRCS. FZS., &e.,
Conservator of the Museum of the Royal College of Surgeons of England . 87

V. On a Raptorial Bird transmitted by Mr. Anprrsson from Damara Land. By J. H.

GiRNEAELY ZS ee ey ee rr do een a Shag ep ae Aeen cotta a Lf,
VI. On some Fossil Birds from the Zebbug Cave, Malta. By W. K. Parker, ERS.,
IRSA UCR gen een ae SE Sig MS RES AE or on aL

VIL. Synopsis of the species of recent Crocodilians or Emydosaurians, chiefly founded on
the specimens in the British Museum and the Royal College f Surgeons. By

Dr. Jonn Epwarp Gray, F.B.S., V.P.Z.S., PDS. de... PEM tee AS)
VIII. Note to Memoir on the Indian Cetacea collected by SiR Waurer Exuior. By Pro-
fecacHOWINN HES: EN Zi Gs. sila: Geek en Se ey capaci aoe ie TE

IX. Contributions towards a more complete knowledge of the Skeleton of the Primates.
By Sv. Guorce Mrvart, F.L.S., Lecturer on Comparative Anatomy at St. Mary's
Hospital. Part 1. The Appendicular Skeleton of Simia. . . . . . - 175

X. Description of the Remains of three extinct Species of Elephant, collected by Captain
Spratt, O.B., R.N., in the Ossiferous Cavern of Zebbug, in the Island of Malta.
By Guorce Busx, F.R.S.; partly i the Notes eee the late H. Fauconer, M.D.,
BETIS ooh sco tek sp Ata: vse ge! ‘ : MLE tay cil ape eal)

iv CONTENTS.
XI. On «a Species of Dormouse (Myoxus) occurring in the Fossil state in Malta. By
AL Lame Anams, OB: FoGA8s 0. Ue a Ss beds Fs Deo

XII. On the Osteology of the Cachalot or Sperm-Whale (Physeter macrocephalus). By
Wim Henry Frowrer, 7.2.S., ARCS. PLS., F.GS., F.Z.S., Conservator

of the Museum of the Royal College of Surgeons of England . . . . . 309
XIII. On a Picture supposed to represent the Didine Bird of the Island of Bourbon
(Réunion). By Atrrep Newton, W.A., P.LS., F.ZS., de. . . . . . 378

XIV. An account of the Fishes of the States of Central America, based on collections
made by Capt. J. M. Dow, F. Gopmay, Esq., and O. Satvin, Esq. By ALBERT
Gunman i Al MD PhD ERS: ELS ees) ia oe naires et BT

XV. On Drvornis (Part XI.): containing a Description of the Inteqgument of the Sole,
and Tendons of a Toe, of the Foot a Dinornis robustus, Ow. By Professor
OWEN, BISA, Ge 3. be ep ee Weatn yi gs Se sag ag eee EOD

XVI. On Divornis (Part XII.): containing a Description of the Femur, Tibia, and Meta-
tarsus of Dinornis maximus, Owen. By Professor OwEn, F.R.S., F.Z.S., &e. 497

XVII. On the Osteology A the Batu CE ae dee ¥ W. K. Parker,
FBS PZ ae es ; gn, et ONE

TRANSACTIONS

OF

is 40:0 LOG LG. A,L,,S OCI ET Y.

I. On the Characters and Affinities of Potamogale, a genus of Insectivorous Mammals.
By Gzoren J. Auiman, F.R.S., Corr. Mem. Zool. Soc. Lond., Regius Professor of
Natural History in the University of Edinburgh, and Regius Keeper of the Edinburgh
Museum of Natural History.

Read June 13th, 1863.

{Puaves 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 interest from
the same quarter*.

A little examination was sufficient to show that the Old Calabar mammal belonged
to the order Jnsectivora, but that, with a well-marked insectivorous organization, it
possessed characters of a very peculiar kind, and such as separated it widely from every
genus hitherto referred 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 PROFESSOR 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. ‘his skin, however, is in a wretched condition; the
skull and teeth are altogether absent, and the 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 satisfactorily 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 Cynogale, under the name of Cynogale velox, Du Chaillu. Mr. Du
Chaillu, however, is not without doubt as to the correctness of assigning his new animal
to the genus Cynogale; and having in view the possibility of its being afterwards deemed
desirable to construct for it a separate genus, he suggests the name of Potamogale 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 Cynogale, but that it
probably did not even belong to the order Carnivora, while 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 Cynogale velox of Du Chaillu became the Mythomys velox
of Gray f.

Dr. Gray’s characterization of his new genus is much more correct than 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 his 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 should be brought to light.

* Du Chaillu, “On Animals from Equatorial Africa believed to be new,’ Proc. Bost. Soc. Nat. Hist.,
yol. vii. p. 353. + Proe. Zool. Soe., 1861, p. 275.

AND AFFINITIES OF POTAMOGALE. 3

It is exactly in this position that Mr. Du Chaillu’s name of Potamogale stands: it
has thus precedence over Gray’s name of Mythomys; and the laws of natural-history
nomenclature compel us to accept it. The synonomy of Mr. Du Chaillu’s animal will
accordingly stand as follows :—Potomogale (prov. gen.) velox, Du Chaillu,=Cynogale
veloa, Du Chaillu,= Wythomys 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 veloa.—A, one of the longer hairs, magnified about ten diameters; B, a por-

: : ; ora
tion of one of the shorter and ner hairs, magnified about 40 diameters to show its atructure 5 aya plete
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

1 PROFESSOR 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. 1B). 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 (a'). 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 (a°, 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. 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 POTAMOGALE. 9)

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; the tragus is indistinct, but the anti-

4
| Wil
\ he
|

~

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. 5a). The fingers are five in number, and are connected
at their bases by a very 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 furrow 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. 58). The toes are five in

Right ear, enlarged.

: Feet, plantar surface.—
number; the outer and inner toes are the shortest, the A. Right anterior.

inner being a little shorter than the outer; the second, B. Right posterior.
third, and fourth are nearly equal to one another in length ; the second and third are

6 PROFESSOR ALLMAN ON THE CHARACTERS

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 153;
inch ; from this point it gradually thins away, and, at an inch beyond it, its vertical
height is 15 inch, and its breadth 34, inch; while at three inches from the same
point its height is 1 inch, and its thickness 34; 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 (Pl. I. & 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 itis a canine, but in form it isa 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 certainly 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 :—
esis p 2th pe ea als
3-3’ "0-04" 3-3’ * 3-3?

In the upper jaw the first incisor resembles a canine ; it projects more than any other
tooth 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

>I

AND AFFINITIES OF POTAMOGALE.

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
slightly larger than that of the third incisor, but otherwise it resembles it. The second
premolar is also implanted by two fangs, and is otherwise 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-
gular 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 (Pl. 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 incisov.

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 front 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 vividly reminding us of the
teeth of certain sharks. In this respect they resemble the premolar teeth of the viver-

8 PROFESSOR ALLMAN ON THE CHARACTERS

ridan genus Cynogale—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 hzemal 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 IT.)

Cervical Region.—The transverse process of the atlas is broad and flattened horizon-
tally. ‘The neural spine is reduced toa 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 Region —There are sixteen, dorsal and five lumbar vertebee. 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 vertebrae the anapophysis

* The account which Mr. Du Chaillu (Joe. cit.) has given us of the habits of his Potamogale velo is entirely
in accordance with what the structure of the animal 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. Itis 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 iscaught. On account of the rapidity of its movements,
T 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 interval 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 with it still
remains distinct.

The neural 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
vertebre 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 first
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 vertebra, 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 vertebre,
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 vertebre; they then gradually diminish on the caudal
vertebree with the diminishing size of these, until towards the end of the tail they dis-
appear, and the vertebra become reduced to minute centra. All the caudal vertebra,
from the second to about the twenty-third, 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 deyelope a short hemai
spine, and send off at each side from their lower surface a broad horizontal plate.

VOL. VI.—PART I. C

10 PROFESSOR ALLMAN ON THE CHARACTERS

The Skull.—Viewed 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 Fig. 6.
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 bulle. 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, Skull, vertical as-
pect: nat. size.

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 anteriora: there are no

clinoid processes.
Fig. 7.

Skull, basal aspect: twice the nat. size.
The tympanic and petrosal bones unite to form tympano-petrosal bull 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
noite i 4 eeu . . és ;
cave in front, where it embraces the posterior margin of the frontal bones. These are

AND AFFINITIES OF POTAMOGALE. 11

nairow, 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 their 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, the
nasal, and the frontal, anteriorly with the premaxillary, and Fie. 8.
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 arch.
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 pterygoid ridges converge from before backwards,
and enclose a deep, narrow interpterygoid fossa, whcse 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 PROFESSOR ALLMAN ON THE CHARACTERS

edge 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 costee 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 143; 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 52;ths 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 33,ths 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 pollex 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 vertebr. 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 7gth of an inch wide, which 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 1,4; 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, forming 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 hallux 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 cecum, 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 ALLMAN 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; 6, margin of anus;
c, postanal chamber laid open from behind; d, vulva; «, orifice of uretha; f, anal glands; g, pouches into
which their ducts open; h, mucous pores; 7, uterus; &, cornu of uterus; J, ovary; m, oviduct; n, vagina ;
o, urinary bladder ; p, ureters. }

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

The position of the mamme 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 corpora
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-
tory 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 AFFINITIES OF POTAMOGALE. 15

.

PotamoGaLE, Du Chaillu.

Teeth, i.

3-3 0-0 3-3 3-
oa

3
3.3” Ce 0-0" Pp. 3-3 mM. 3 36.

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 hair 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, vulva, urethra, and ducts 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 Potamogalide may

be given.

The above paper had been already printed when I became acquainted with a descrip-
tion of Potamogale velox, contained im a communication presented to the Zoological
Society on the 25th of April, 1865, by Professor J. V. Barboza du Bocage, “ 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, however, accept the generic name of either Du Chaillu or Gray, but constructs

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

16 POFESSOR ALLMAN ON THE CHARACTERS AND AFFINITIES OF POTAMOGALE.

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

EXPLANATION OF PLATES I. & II,

Plate I. Potamogale velox, size of life.
Plate II. Skeleton of Potamogale velox, of the natural size,

YOUHA ZIVIONVLOd

JEM LS

W West imp

GH Ford

POTAMOGALE VELOX

a

II. On some Indian Cetacea collected by Wauter Exot, ig
By Professor Owen, F.B.S., F.Z.S., &e.

Read 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 DELPHINIDZ.

Genus DELPHINUs, Cuvier.
DELPHINUS (subgenus STENO, Gray) GADAMU, Owen.
The “ Gadamu” Dolphin. (Pl. 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

1 Now Sir Walter Elliot, K.C.S8,I.

VOL. VI.—PART I.

18 PROFESSOR OWEN ON INDIAN 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 (fig. 2, 6). 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 (7. ¢. 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 the 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 very 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 6 lines. The number of teeth, as noted by Mr. Elliot in one specimen,

27-27 —
was 37—5,—108; in a second specimen,;,—,, La == 96; in the skull transmitted, a8 =101.

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

PROFESSOR OWEN ON INDIAN CETACEA. 19

“elongate, about ¢ 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-
dibulz”’ of the restricted Delphini of Cuvier’s system.

In the skull, no. 423, of the “Gadamu” (PI. IV.), the symphysis mandibule (figs.
3 & 4,5,5) is more than 4th the length of the entire dental series, and about 4th 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 by a cross groove”.

In this section the present species differs from the Delphinus (Steno) malayanus in
colour, in number of teeth, and perhaps also in size. The D. malayanus is “ greyish
above and below;” the dental formula 22=°°=144. From the Delphinus (Steno)

36—36
frontatus of the Indian Ocean, with teeth 00 or = 84, D. gadamu differs

in the greater number of teeth. From Delphinus (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 D. gadamu in the length

. 40 —40
and slenderness of the snout and the more numerous teeth, the formula being 7-3,

= 160. :

In the skull of D. (Steno) gadamu (P1. 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 pterygoids, 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 in 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 (Pl. 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. 1844, p. 43. Not any of the
figures of the skulls of Steno, Gray, illustrate the symphysial character in question. In a specimen of Steno
frontatus in the British Museum the mandibular symphysis is about one-fourth of the entire length of the
skull. 2 Thid.

D2

20 PROFESSOR OWEN 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.

De.puinus (Steno?) 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 thé 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 (Pl. V. fig. 1), but is continued by an alteration of curve more directly into the
rostrum than it isin D. gadamu (Pl. III. fig. 1). The transverse groove, as indicated in
the drawing (PI. V. fig. 6, ¢), 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 forehead is nearly straight, very slightly undulated, not convexly
curved as in D. gadamu. ,

The specimen figured (Pl. 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, 33 inches in
depth at the base, and 5 inches there across; the skull shows better the pre-
dominance of the vertical over the transverse diameter of the rostral production of
the jaws. The “ictus oris,” 1 foot in length, bends gently upward from the base of
the snout to within 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
14} 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 curve, is
12 inches. The dorsal fin is relatively lower than in D. fusiformis, much more so than

PROFESSOR OWEN ON INDIAN CETACEA. 21

in D. 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
length 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. fusiformis, and is more obtusely terminated than in that species or in D. gadamu,
From the hind border of the caudal fin to the vent is 2 feet 5 inches: the vulva is
24 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
foregoing 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 D. 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

2—32

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 D. gadamu; the occipital condyles are larger, the superoccipital surface
is narrower, the temporal fosse more squared above; the premaxillaries do not rise to
form a distinct convexity at the upper part of the rostrum, as in D. 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 103 inches in D. gadamu, 11 inches in D. lentigi-
nosus. But the chief distinction is in the number of the teeth: in the skuil 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
D. lentiginosus is 9 inches 9 lines, but is not more than 8 inches 6 lines in D. gadamu.

The D. lentiginosus is known to the Waltair and Vizagapatam fishermen by the
Telugu name of “ Bolla Gadimi.”

DELPHINUS (STENO?) MACULIVENTER.
Spot-bellied Dolphin. (PI. VI. figs. 1 & 2.)
In the degree of convexity of the forehead the present species resembles the D.

Susiformis (Pl. V. fig. 1); but the head is relatively larger, and the body is deeper in ~
proportion to its length, than in either D. fusiformis or D. gadamu.

22 PROFESSOR 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 maculiventer.
The specimen from which figs. 1 and 2 were taken was a female, 6 feet 11 inches in
length, found at Waltair, 26th April, 1854. It is called by the fishermen “‘ Suvva.”

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 inches; 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 D. lentiginosus (Pl. 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 inches; 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 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 Pl. VI. figs. 1 & 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
beasmaller 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 oe 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 D. gadamu, 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 D. gadamu. 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

PROFESSOR OWEN ON INDIAN CETACEA. Z

bo

9
2)

with the eyes; crescentic, with the angles bent forward. The length of the snout is
6 inches, of the “ictus oris” 10 inches; from the end of the snout to the eye 1 foot;
from the same to the setting-on of the pectoral fin 1 foot 74 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 1] 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
in advance of the mid notch of the caudal fin; the vulva 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
(Pl. VII.), which presents the general characters of that section of Delphinide to which the
term “‘Lagenorhynchus” has been attached. It resembles in size and general characters the
skull of Lagenorhynchus electra, Gray ; but the occipital condyles are more approximate
below the foramen magnum, the presphenoid is narrower, the longitudinal channel
formed 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 (£); but the breadth
there is 54 inches in Lagenorhynchus electra and 5 inches in Lagenorhynchus fusiformis.
In this species a narrow slip of the vomer (fig. 4, 13), about an inch in length, appears
on the bony palate, 3 inches from the anterior end.

In Lagenorhynchus (Pl. VII.) the skull is broader in proportion to its length, and the
mandibular symphysis shorter, than in Sfeno (Pl. 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 POMEEGRA, Owen.

The Pomeegra Dolphin. (PI. VI. fig. 3.)

This species belongs to the same section of Delphinus as the Black Dolphin of the Cape
and Ceylon (Delphinus longirostris, Gray) and the Delphinus forsteri? of the Pacific.

1 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 colour, almost black, with a rather
lighter shade at the under part of the belly. Mr. Elliot, 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. = 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. forster?, the greatest circumference being at the fore part
of that fin. It is more slender in proportion to its length than any of the above-described
fusiform Dolphins belonging to the subsection Steno, Gray. The symphysis mandibule
(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. ga-
damu (Pl. 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 De/phini as restricted by Dr. Gray. In Delphinus euphrosyne, 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 writes, “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 Procana, Cuvier.
Puocana (Orca, Gray, Reinhardt) 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,

PROFESSOR OWEN ON INDIAN CEIACEA. 25

if even the proportions of the rostral part of the skull (Pl. IX. fig. 1, 21' 22) did not show
that it belongs to a different section of Delphinide!. The present part of a Cetacean
skeleton, as the skulls of those species, figured in Pls. 1V. 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.

Lari Ve (8 ee Rg ee Rw ey ee oe Cees Pom Pe oem rene |
Breadth, greatest, across zygomata . . . Ea Cee aN Bet
From the back of occipital condyle to rrtenetal process of malarbone 7 6

From the antorbital process of malar to anterior end of premaxillary 4 8
From the back part of nostrils to do. do. ge

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 Delphinide, including the
Grampuses and Porpoises, for which Cuvier proposed the subgeneric name Phocena?,
and which, in his ‘Ossemens Fossiles, tome y. part i. (1823), he distinguished as
“§ 2. Les Dauphins a téte obtuse” (p. 280), from “ ) 1. Les Dauphins @ bec” (p. 275)
(Delphinus, proper)’.

The number of Delphinide 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-
tebra, 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 quasi-generic
names is doubtful; and I shall refer the present skull, which appears to me to belong
to an undescribed species, to the Phocena brevirostris, as a member of the section of
Cuvier’s Phocene, characterized by conical teeth, in which its nearest alliance appears
to be with the Phocena globiceps, Cuv.*

1 The following is Mr. Elliot’s note respecting this specimen :—* August 1852. Got the skull of a porpoise
which one of the fishermen found dead at the mouth of the Vizagapatam river. He called it ‘Ganumu,’ and
described it as haying a rounded head, without beak, colour black or dark above, white below; perhaps a
Phocena or Globicephalus.”

2 Régne Anim. tome i. p. 290 (1829). 3 Thid. p. 287.

4 Tbid. p. 290; Annales du Muséum, tome xix. ; Ossem. Foss. tome y. part i. p. 290, tab. 21. pls. 1, 2, 3, figs:
11, 12, 13.

VOL. VI. PART I. E

26 PROFESSOR OWEN ON INDIAN CETACEA,

The elements of the occipital have coalesced. The basioccipital (Pl. IX. fig. 3,1)
forms the lower fifth of the foramen magnum, intervening for an extent, measured in a
straight line, of 103 lines between the lower ends of the occipital condyles (ib. 2’): it is
here thick and concave transversely: it becomes thinner vertically and expanded later-
ally as it advances to join the basisphenoid (ib. 5), 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. ov). ‘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 off 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. 3,4”); this border is divided by a notch from the
otocranial plate (5') of the basisphenoid, and just within the bottom of that notch’
opens the canal for the nervus vagus. The superoccipital (figs. 1,2, 3) rises and ex-
Pands, as in other Delphinide, into a broad and lofty convex plate reaching the vertex,
and there articulating with the parietals (7) and interparietal (7*); a low median
ridge (fig. 2, 3') divides vertically the upper half of the superoccipital. 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 grooved transversely by the lateral
sinus.

The alisphenoids (Pl. IX. figs. 1, 3,6) coalesce with the fore part of the lateral borders
of the basisphenoid, in advance of the otocrane (fig. 3, 07), 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 (7), and frontal (11), and to overlap the
process of the squamosal (fig. 3, 27’), continued, mesiad, from the glenoid cavity (7). The
suture between the interparietal(fig. 2, 7*) and superoccipital (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 (5), and extends in
the form of a compressed rostrum forward, contracting, to be enclosed by the pos-
terior sheath-shaped part of the vomer (13). The orbitosphenoids (ibid. 10) extend

PROFESSOR OWEN ON INDIAN CETACEA. 27

outward, overlapping the pterygoids (24), contract where they form the fore part of the
foramen lacerum anterius and the optic foramina, beyond which they expand to
support the orbital plate (fig. 5, 11’ ) of the frontal.’

The frontals (Pl. EX. figs. 1 & 2, 11, 11’), in great part overlapped, as in other Cetacea, by
the maxillaries ( 21 ), show 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 (7),
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, 1) articulates with
the parietal (7) and alisphenoid (6); 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 (26); 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 interparietal (7*), anteriorly with the nasals (15 ).

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, two inches behind the fore end of the vomer, the
rough thick anterior border of the coalesced prefrontals (fig. 2, 14), 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(14’ ). 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 (15 )—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 off
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
Phocena globiceps; the superorbital plate (fig. 1, 21*) 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. globiceps,
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. 5, 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 Delphinidw, consists of the antorbital (Pl. IX. fig. 1, 26’ ) and
styliform (26) portions. The former (26' ) is anarrow 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 under 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
autorbital fossa (7), external to which is the rough malomaxillary fossa (e).

PROFESSOR OWEN ON INDIAN CETACEA. 29

The squamosal consists chiefly of its articular or zygomatic part (Pl. IX. figs. 1 & 3, 27),
which is deep in proportion to its length, truncate, and three-sided; the outer side is
slightly convex and rather rough, 1” 5'” in depth posteriorly; the inner side is divided
between the articular cavity (fig. 3, g), rough for syndesmosis 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 (6) 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, 7),
I consider to be the “mastoid” confluent with the squamosal, together forming the bone
which should be termed ‘“‘ squamo-mastoid.” The mastoid part terminates below in a
rough, flattened, triangular surface (fig. 3, 8), 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, in the line 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, downward, and with a slight inclination forward. The mandible
offers no notable peculiarity, save that which relates to shortness in proportion
to the entire skull, concurrently with the same specific character of the upper jaw.
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 greatest vertical extent of the ramus being
2’ 6"; 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 common
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 36". 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-thirds of the irregular
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 incurvation; the length of crown is from 3’” to 4’”, the diameter of its
base 1’, that of the inserted root 2'”.

As in other Delphinidw, the bony palate is entire, save at the slight median divarica-
tion of the maxillaries and premaxillaries, and the major part of this median fissure is
closed by the yomer. 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 PROFESSOR OWEN ON INDIAN CETACEA.

The optic foramen communicates or is blended with a larger vacuity or fissure
between the orbitosphenoid, frontal and pterygoid, which might be termed the spheno-
frontal fissure. 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 “ intersphenal fissure ”?.

The removal of the loosely attached petrotympanic exposes the wide otocranial
vacuity (Pl. IX. fig. 3, or) in the basal walls of the cranium, which is a characteristic
feature of the Delphinoid as compared with the Physeteroid skull (Pl. 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
present species of Phocena it presents a subquadrate 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 Delphinide.

The departure from symmetry in the present Delphinoid skull is slight: it is seen
in the greater 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.

Family PHYSETERID (Cachalots or Sperm-Whales).

Genus Evpuysetes, Macleay.
PuysETER (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, Pl. XI. to scale) and female (side view, Pl. 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.” The male, measuring 6
feet 8 inches in length, was taken at Waltair, February 28, 1853. The female was taken
on the Ist of March, 1853, at the same part of the coast; she measured 6 feet in length.

‘ Tt is noticed as “le trou sphéno-orbitaire,”’ by Cuvier, ‘ Oss. Foss.’ tom. cit. p. 294.

PROFESSOR OWEN ON INDIAN CETACEA. 51

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 Delphinide.

The first and most important of these is the inferior position of the mouth, beyond
the small opening of which the blunt rostrum extends forward from 4 to 6 inches.
The blow-hole (Pl. X. fig. 2) is single, but is not medial in position or symmetrical in
shape; it is im 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
the angles or “cresses” directed 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 in the upper half of the
head, seen in profile, near the boundary dividing it from the lower half. From the
vertical line 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 semicircular 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 line, 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 imaccuracy
here. The length of the fin in both is 1 foot; its extreme breadth is 45 inches in the
male, 4 inches in the female: its line of attachment is in the lower third of the trunk,
as seen in profile. The dorsal fin is well developed, subfalcate in shape; its anterior
border is halfway between the snout and the base of the tail. The length of the base
of the fin is 10 inches in the male, 9 inches in the female: the height of the fin,
vertically at its back part, where the apex curves back a little beyond the basal attach-
ment, is 7 inches in both. The anterior border of the fin is slightly convex; its length,
in 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 tail, or tail-end of the
trunk, measures 34 to 4 inches in vertical and nearly 2 inches in transverse diameter.

The expansion of the trunk is pretty equal in every direction towards the dorsal fin, and
the upper surface gives the appearance of the fore part being subdepressed: the dunmu-
tion beyond the dorsal is more rapid from side to side than from above downward.
The greatest vertical diameter of the trunk is, in the male, | foot 64 inches, in the

32 PROFESSOR OWEN ON INDIAN CETACEA.

female 1 foot 4} inches: the greatest transverse diameter of the trunk in the female is
1 foot 2 inches.

The caudal fin, the shape of which is given in fig. 2, Pl. X., measures, in the female,
1 foot 7 inches in extreme breadth, and 7 inches 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 from
the same part in the female. It is 10 inches behind the vertical line dropped 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 inches 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 discoloured with blood.” The dentition is
=e=a20 (PIS XE fig sili yay ise).

9-9

The Physeteride (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 tegu-
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 truncate mass. The external blow-hole is reduced
by its operculum or flap toa single sigmoid fissure on the left side of the upper and fore
part of the head, 7. e. at or near to the summit of the truncate end of the 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
fossee 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 Cuvier, whether any large Sperm-Whale may exist, characterized
as above, but with a high 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 dentigerous
symphysis, still waits a reply from a direct 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”2, 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. 8. Wall, Curator. 8yo. Sydney, 1851. ealb psec

PROFESSOR OWEN ON INDIAN CETACEA. 53

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 observa
on the condition of the dorsal fin or hump was made.

Cuvier characterizes the ‘“Cachalot macrocéphale” (Catodon macrocephalus, Art.,
Physeter macrocephalus, Linn.) as having the back provided with a slightly raised
prominence, which some have called “ fin,” others “longitudinal ridge”, others “ hump”
or “tubercle” (doc. cit. p. 338): “Il a une dorsale trés-peu saillante vers I’ arriére
du dos, quelquefois réduite 4 une protubérance, ou 4 deux ou trois” (ib. p. 339). In the
*Regne Animal,’ Cuvier says, “Il n’a qu'une éminence 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? en existe-t-il dont l’évent soit percé prés du front sur le
milieu de la téte? en existe-t-il ou les branches de la machoire inférieure ne soient pas
réunies sur la plus grande partie de leur longueur en une symphyse cylindrique? Voila
ce qui reste a chercher, ce qui reste 4 prouver autrement que par des figures tracées par
des matelots. Ce nest quaprés que des hommes éclairés auront observé ces étres
avec soin, et en auront déposé les parties osseuses dans des collections ou elles puissent
étre vérifiées par des naturalistes, qu il sera possible a la critique de les admettre dans
le catalogue des animaux” (tom. 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 Physeteride—a Cachalot in fact—though small, in
which the dorsal is lofty, with the usual shape of such well-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 drawings by
the native artist, significantly indicated the family 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
transmission, with the drawings, of the skull of one of the individuals figured.

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 skull 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 skull first made known by De Blainville as of the Cachalot which
he called Physeter breviceps®, and in that subsequently described by Macleay * under

1 Op. cit. p. 9. ? Op. cit. p. 4.
3 Annales Francaises et Etrangéres d’Anatomie et de Physiologie, tom. ii. (Svo, 1838) p. 335: “Sur les
Cachalots.” 4 Op. cit.

VOL. VI.—PART I. F

54 PROFESSOR OWEN ON INDIAN CETACEA.

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

Description of the Skuil.
(Pls. XII., XIIL, & XIV. fig. 1.)

Short as is the upper jaw in proportion to the skull in Phocena brevirostris (Pl. IX.),
it is shorter in the subgenus or section of Physeteridw represented by the Physeter
breviceps, De Bl. (Pl. XIV. fig. 3), and shortest of all in the present species (ib. fig. 1).

In the following Table of admeasurements are given those of the Physeter (Euphysetes)
grayi, Macleay (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 breviceps, from the Cape of Good Hope.

|
|
|
}

P. simus. P. grayi. P. breviceps.

inches. lines. | inches. lines. | inches, lines.

| Length from the back of occipital condyles to end of snout....| 10 5 16 6 15 5
Breadth across postorbital processes ..............0000000- 14 0
Breadth across the beginning of malo-maxillary fissure ...... 9 6
From the back of occipital condyle to antorbital process of

LCA hs A OMOO COG OM ORTOU OO HORIIOD Ones SoCo, oer

| From the antorbital process of malar to end of snout ........
From the back of occipital condyles to posterior wall of left
MOSER oeayay ave craleve stares eens calcio ale CPS Pace cue pete Oe ears

| From the bottom of malo-maxillary fissure 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

GuinthallGia7s Shas See Oce Eno rOS Ane Sane oleae onan

| Breadth of snout at its extremity .......-...-ccseeese sess
| Breadth of premaxillaries at the malo-maxillary fissure ......
| Breadth between anterior ends of premanillaries............
| Antero-posterior diameter of left nostril
jstransyerse diameter/ot: left most: ovens « savers eavecto neon
| Antero-posterior diameter of right nostril
Transyerse diameter of right nostril. ..../...).s.ceeeeu es

| Length of interfrontal crest, straight line
| Width of occipital foramen
jeventicalsdiameten/of foramen... ts yosvoiis cuits eisai eerenee ee

Between outer edges of occipital condyles
Breadth between paroceipitals .. 22... shee edt ee dace nes

| From the lower border of basioccipital to vertex

| Length of mandible, in a straight line

/enethiofjalveolar: series)” 1.2/4.1. / uw ctide 4 acer. stern eee

Height of mandible at coronoid ridge

ES ive)
a
ou

Die

=
w

10

DW DNBNHHPWOSCHRFPONE ER BOF eal
HOADWODW AP WOT OH O poo oo 7
i
jen
AOOFURNHAPONNHE WHO Ont &
WWWOOSCKANORDRrOWDOSO ooo

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 Frangaises et Etrangéres d’Anatomie et de Physiologie, tom. ii. tab. x. (the admeasurements
are given in French inches), viz. :—‘ Longueur du erane 14 pouces et demi,’”=15" 5’”, Engl. “ Longueur de la
machoire inférieure 13 pouces,’=13" 10”, Engl. Ecartement de ses condyles 12 pouces,”=12" 9'” Engl.

PROFESSOR OWEN ON INDIAN CETACEA. 85

(Pls. XII., XTII., & XIV. fig. 2, 1) beneath the foramen magnum (ib. 0) 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
(Pl. XII. fig. 2, 2") are more sessile than in Phocena brevirostris, being raised only
by a linear border from the contiguous bone, except at their 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
Physeter macrocephalus (Pl. XIV. fig. 2, 0) 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 (Pl. XII. 2) extends outward and slightly downward, expanding a
little vertically, and thickening to form the paroccipital (4); this expanse is moderately
concave transversely, convex vertically. The border of the paroccipital is thick and
rugged: it is concave toward the otocrane (PI. XII. fig. 1, and Pl. 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 (Pls. XII. & XIII. fig. 2,7) from the otocranial plate of
the basioccipito-sphenoid (Pl. XII. fig. 1, and Pl. XIII. fig. 2,5’): this plate arches out-
ward and downward, with a slight obliquity backward, and is overlapped anteriorly by
the pterygoid (ib. 24’), which seems to form an anterior continuation thereof, converging
towards its fellow: but the free border of the basisphenoidal otocranial plate (5’) is
more obtuse and thicker than that of its pterygoid prolongation (24). A trace of the
suture between the exoccipital (Pl. XII. fig. 1, 2) and squamosal (ib. 27) remains. ‘The
ridge across the vertex (Pls. XII. & XIII. fig. 1, 7,11,3) is obtuse, but well marked :
the proportions contributed by the superoccipital (3), parietal (7), and interparietal (if
any) cannot be determined; and the frontal (11), as it ascends, contracting from the
superorbital roof, is also blended with those constituents of the ridge’. The instructive
harmonia between basi- (Pls. XIII. & XIV. fig. 1, 5) and presphenoid (ib. 9) remains.

The alisphenoid (Pl. 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

1 To afford a comparison with Physeter macrocephalus, I propose to append, in the present note, descrip-
tions of the homologous cranial bones of a foetus of that species described, in my ‘ Catalogue of the Osteological
Series in the Museum of the Royal College of Surgeons,’ 4to. 1853:—‘ The elements of the occipital neural
arch are unanchylosed. The lateral margins of the anterior half of the basioccipital are produced and bent
obliquely downward. The exoccipitals are much produced and expanded laterally: they are deeply notched
below. The superoccipital contributes the upper ends of both condyles: it is in the form of a vertical plate,
conyex from side to side: a strong internal yertical crest is produced forwards: it is overlapped at its lower
and lateral angles by the exoccipitals, 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). _

F2

56 PROFESSOR OWEN ON INDIAN CETACEA.

the temporal fossa (ib. ¢), 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 (¢r), from which
two channels lead toward the back part of the orbital roof (07), blending together
and widening as they grow shallow. The temporal fossa (Pl. XI. & Pl. XIII.
fig. 2, t) is 1” 1” in antero-posterior, and 2” in transverse extent, has its marginal
boundary almost completed by the approximation of the postfrontal (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 (Pl. XII. fig. 1, 27).

The presphenoid (Pls. 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
alisphenoids (6).

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 (Pl. XIV. fig. 2,2): 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 trigeminal one (Pl. 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 channel (ib. 10). There is no intraorbital fossa
answering to that in Phocena brevirostris. The roof of the orbit is unbroken, gently con-
cave from before backward, formed chiefly by the frontal (Pl. XII. fig. 1,11, 11'), which is
notched near the middle of the superorbital ridge: this is thick, obtuse, and produced
backward and downward into a postfrontal or postorbital process (ib. 12). Above the ridge,
the frontal (ib. 11’) contracts; its surface is here free from the maxillary (21’), is slightly
concave vertically, before it is reduced by the overlapping of the parietal (7) and superocci-
pital (3) behind, and of the maxillary ( 21’ ) in front, to the narrow strip (1), which rises,
bending convexly, to the vertex. The fore part of the superorbital ridge (11) is cb-
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 alisphenoids: 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 entocranial 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 alisphenoid in a groove anteriorly.” —Physeter macrocephalus, 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 ale; the under and anterior
part is overlapped by the vomer.”—Physeter macrocephalus, op. cit. p. 447.

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

PROFESSOR OWEN ON INDIAN CETACEA. 37

The vomer (Pl. XIII. figs. 1 & 2, 13, 13’) has partially coalesced with the presphenoid (ib.
fig. 2,9) and underlaps the prefrontals (Pl. XIV. fig. 1, 14): it appears upon the palate,
about an inch in advance of the posterior fissure (Pl. XIII. fig. 2, w), expands toa breadth
of 6 lines (13'), 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 premaxil-
laries (ib. 22), which cartilage terminates anteriorly the series of vertebral centrums, of
which the vomer is the inferior or cortical ossification. The fore margin of the confluent
prefrontals (ib. 14) isat 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. 07) and the corresponding wall of the small
right nostril (Pl. XIV. fig. 1, o/’). 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 (15'): this
ridge also sends off a kind of “spur” (15) from 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. y) 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 (13'),
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 direction from
withm, outward and backward, contracting to terminate mesiad of the fossa (d): they
develope no outer or free lamella in Euphysetes.

a long 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 sqnamosal. The straight median margins
of the frontals are thinned off and joined by a squamous frontal 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 broad
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 conyolutions.”
—Physeter macrocephalus, op. cit. p. 442.

1M. de Blainville figures, but makes no mention of this bony ridge bisecting the ‘ postnarial’’ cayity.
Dr. Gray, in appending the term Kogia to the Physeter breviceps, De Blainy. (Zoology of the Erebus and
Terror, ‘‘ Cetacea,” 4to. 1846, p. 22), is equally silent—indeed, adds nothing to De Blainyille’s meagre
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 Huphysetes from Physeter or Catodon,

2

38 PROFESSOR OWEN ON INDIAN CETACEA.

The maxillary (Pl. XII. fig. 1, Pl. XIII. figs. 1 and 2, 21) forms the major part of the
bony roof of the mouth: a small triangular strip of the premaxillary (Pl. XTIT. fig. 2, 22)
is wedged into the short anterior interspace between the maxillary (2) and vomer (13').
The palatal surface (21*) is moderately convex transversely, straight lengthwise, and
is impressed by an alveolar groove (a/) retaining one socket and tooth (Pl. XII.
fig. 1, x) at the fore end and continued in a straight line backward for 3 inches
(rather more on the left, rather less on the right side) without indications of
alveoli, and in a line not parallel with the outer margin of the bone, but receding
to a distance of 1 inch from it, posteriorly; so that the teeth, if developed there, would
be rather palatal than marginal in position. The outer border of the maxillary thickens
near the malo-maxillary fissure (21, 4), with a smooth convex exterior. That fissure dilates,
as it sinks obliquely backward and inward, to a breadth of from 5 lines to 4 lines, its
depth being | inch 6 lines (#4). These fissures mark off the rostral portion of the skull,
which is here an equilateral triangle, including above (Pl. XIII. fig. 1) parts of the
vomer (13), prefrontal (14), premaxillaries (22), and maxillaries (21): the surface so formed
is concaye transversely at its posterior three-fourths, almost straight longitudinally. The
maxillary, expanding backward beyond the rostrum, bends (at /, fig. 1) round the upper
and back part of the malo-maxillary fissure ; and in close conjunction (here partial con-
fluence) with the malar (26), it forms the large smooth tuberosity (21,26) 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 hollow as it sweeps inward to rise again upon the bifurcate sinuous ridge
(ib. 15,15”) which divides this singular 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 circumference, so bounded by the maxillaries and describing as much
of a circle, being a little produced backward, subangularly, at the hindmost part: the
open anterior fourth is continued upon the more shallow concavity of the triangular
rostrum. The right maxillary is vertically pierced by two foramina (Pl. XIII. fig. 1, a, d),
which converge to the common inferior outlet (ib. fig. 2,d). The upper fissure between
the maxillary and premaxillary widens and deepens as it extends backward, and terminates
in the canal (fig. 1, ¢), also conducting to the fossa (fig. 2, d), which, as it transmits
maxillary branches of the fifth pair from 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 maxillaries upon the forehead above and behind the
orbits.

The pterygoids (Pl. XIII. fig. 2, 24) meet at the midsurface of the roof of the mouth,
and extend the palatine suture (p/) 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 triedral process, representing

PROFESSOR OWEN ON INDIAN CETACEA. 39

the “hamular” one; the outer portion, partly marked off by a ridge from the palatine
plate of the inner 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. 24’) which appears to continue forward the otocranial
plate (s') 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
surface somewhat convex: the concave channel prolongs backward the nasal passage
(w) beyond the septum. A semicircular emargination divides the posterior subvertical
plate from the palatine portion (24) of the pterygoid. The total length of the pterygoid
is 4 inches 8 lines; the breadth of the pair of bones posteriorly is 5 inches; the sutural
union of the pterygoid with surrounding bones persists’.

The malar bone (PI. XII. fig. 1, Pl. XIII. figs. 1, 2, 26) is represented in the present skull
by the portion of that in Delphinidw (Pl. IX. figs. 1, 3,26") which is wedged like a
lacrymal? between the frontal (11’) and maxillary (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 inner
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 (#). The outer facet of the malar is
slightly concave vertically, convex transversely: the antero-internal facet is concave im
both directions, except where it curves anteriorly round the obtuse angle between it and
the outer surface: the internal or orbital surface is the narrowest, and is conxex trans-
versely, and straight vertically. The apex is subbifid, the outer part (Pl. XII. fig. 1, 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 Phocena brevirostris (Pl. 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 skull, which
interval I found occupied by a ligament (“sclerous” state of malar) in a young Cachalot?.

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

1 «The pterygoid, which is double the size of the palatine, extends backward to the basioccipital, articulating
in that course by its expanded upper border with the pre-, pbasi-, 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 by a deep notch, and cireumscribes the posterior bony aperture of the
nostril.” —Physeter macrocephalus, op. cit. p. 443.

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

3 «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 pone.” —Physeter macrocephalus, op. cit. p. 444.

40 PROFESSOR OWEN ON 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 circumference: the anterior boundary, which also forms the posterior
one of the lower outlet of the temporal fossa, is concave: the wall (Pl. XII. fig. 1, 27) which
the squamosal contributes to the posterior and internal part of the temporal fossa (¢)
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 (Pl. XIII. fig. 2, 8’), where it contributes, with the paroccipital (4),
to the otocranial cavity’.

In the interior of the cranium (Pl. 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, directed mesiad, which may be where the exoccipital suture ran: the outer
or lateral beginning of the tentorium receives a short angular ossification, which forms
the outer wall of the fossa (v), 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 (Pl. XII. fig. 1, Pl. 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, 7) between the otocranial plates of the basisphenoid (5) and paroccipitals
(4). At the fore part of the tentorial process (Pl. XIV. fig. 1,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 distinct foramen ovale, foramen rotundum, or foramen opticum: they all
seem here to be confounded in the intersphenal fissure (Pl. XIII. fig. 2, #7). From the
extreme shortness of the jaws, the nerves of sensation to the face must have been very
small. ‘The “sella” (Pl. XIV. figs. 1 & 3), scarcely impresses the basisphenoid : its best
antero-external boundaries are afforded by the superoptic processes of the orbitosphenoid
(ib. m). 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 ina 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, dentated sutural margins to
the frontal and exoccipital : its anterior border is grooved for the reception of the alisphenoid: the lower angle
is as it were truncated, and presents a rough surface for the attachment of the petro-tympanic: 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 produced
downward into a slender process.” —Physeter macrocephalus, op. cit. p. 444.

* In the Great Cachalot “a strong medial crest is produced forward from the inner surface of the super-
occipital” (loc. cit. p. 442),

PROFESSOR OWEN ON INDIAN CETACEA. 4]

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 (30), 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 with 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 third 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 teeth
(Pl. XII. fig. 1, and 4) 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 14 line, with a length of 24 lines, diminishing to a subrecurved apex.

The loss of symmetry in this skull is hardly observable im the general contour,
whether viewed from above (Pl. 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 skull (copied
in Pl. XIV. fig. 3), the occipital condyle is more prominent than in Huphysetes simus (P).
XII. fig. 1): the contour of the superoccipital is concave in Huphysetes 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 the
greater proportional length of the rostral part of the, skull—measured, viz., from the ma-
lomaxillary fissure (ib. & Pl. XIII. &) to the end of the upper jaw (2, ©): in Euphysetes
breviceps it forms about two-fifths of the entire length of the skull, in Luphysetes simus
about two-sevenths. The proportion of the maxillary, above the frontal and malar, on

1 «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 symphysial 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.”—Qp.
cit. p. 444, foetal Physeter macrocephalus.

VOL. VI —PART I. G

42 PROFESSOR OWEN ON INDIAN CETACEA.

the exterior of the skull is much greater in Huphysetes breviceps than in Euphysetes simus,
especially in vertical extent: in the upper view of the skull the porportion of the postnarial
cavity, especially in breadth, to the extent of the rostrum is less in Euphysetes breviceps
than in Huphysetes 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
sockets for as many, in each mandibular ramus: the entire tooth, figured by De Blain-
ville (copied in Pl. XIV. fig 2. 2), is 10 lines in length, and has a proportionally larger
and more curved crown than in Euphysetes simus. De Blainville writes, “Il me parait
& peu pres certain qu il n’y avait pas de dents 4 la machoire supérieure” (1. 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 their 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 Huphysetes breviceps, De
Blainy.: they are as wide apart as in Ewphysetes simus, but have crowns more slender
and recurved.

In the figures of the mandible given by De Blainville (oc. cit. pl. 10), and by
Macleay (Joc. cit. pl. 2. fig. 5), the breadth between the outer parts of the condyles
equals the length of the mandible in a straight line, that is, from the middle of the
chord drawn 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 Physeteride and the
Delphinide (see Phocena brevirostris, Pl. LX. fig. 1) is the non-production of the upper
or hinder expansion (naso-frontal plate) of the maxillary (Pl. XII. fig. 1, 21*, 21”) over
the orbital process of the frontal( 1,11’); which, therefore, in Euphysetes simus as in
Euphysetes breviceps, stands out free (Pl. XII. fig. 1,1’) 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 (Mr. Kreffts) of the Australian Museum, I have thought it might
be useful to add the following notes :—

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

PROFESSOR OWEN ON INDIAN CETACEA. 45

the twentieth (or forty-first from the skull), leaving ten, perhaps eleven, terminal
vertebre represented by depressed centrums, gradually diminishing to the last. The
seven cervicals are anchylosed: the diapophyses distinguish the atlas and axis, the former
of which vertebre 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 gradually, 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 pairs of ribs directly join the sternum,
which consists of three sternebers, each more or less completely divided at the middle
line into two bones. The first 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-
external angle of the manubrium. The second and six following ribs have both head
and tubercle, the former abutting against the interspace of their own 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 third, gaining length, losing breadth, and with more regular curvature, is arti-
culated by its short hemapophysis to the interspace between the second and third
sternebers. The fourth rib is joined to the end of the third sterneber. 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 previously recorded is, that they are all gradational,
and exemplify steps by which are gained the extreme modifications, especially in the

skull and dentition.
@ 2

{4 PROFESSOR OWEN ON INDIAN CETACEA.

Imperfect as may be the cetacean record, it yields several series of differential cha-
racters,—as, ¢.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 Balwna, Delphinus, Pho-
cena, Ziphius, Euphysetes, and Physeter, again culminating in Platanista—in the number
of teeth, from zero (Balena and old Delphinapteri), through Monodon, Ziphius, Huphy-
setes, to the multitude of teeth in Delphinus, 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 individuals, 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
(Salmo fario) 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
flow, and same nature of bed. Where 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 Hyperoodon, 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 observed steps in a gradation of such deyviational
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 being 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
grappling 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 attain2.
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 Balena, Balwnoptera,
Physeter, Hyperoodon, &c., 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, though 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 needed
for the Cetaceous Department of a Zoological Museum’.

1 Owen, ‘ On the Nature of Limbs,’ 1849, p. 86.

2 T 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.

4’ See Hansard, ‘ Debate on Museum of Natural History,’ May 19th, 1862, p. 1928.

46

Fig.

Fig.

Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.

Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.

PROFESSOR OWEN ON INDIAN CETACEA.

DESCRIPTION OF THE PLATES.

PLATE II.
Delphinus (Steno) gadamu: diminished to scale.

. Side view.
. Upper view: 6 blow-hole.

PLATE IV.
Delphinus gadamu.

. Side view of skull (wanting back part of cranium).
. Side view of mandible.

. Upper view of mandible: ss symphysis.

. Symphysial end, inner view of mandible.

. Bony palate.

All the figures are nearly half the natural size.

PLATE V.

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

PLATE VI.

. Delphinus maculiventer (to scale of Plate V.).
. The same, upper view.
. Delphinus pomeegra (id.).

PLATE VII.
Delphinus fusiformis.

. Side view of cranium and upper jaw.

. Side view of lower jaw.

. Upper view of cranium and upper jaw.
. Under view of ditto.

. Upper view of symphysis of lower jaw.

All the figures are nearly half the natural size.

PLATE VIII.

. Side view of cranium and upper jaw of Delphinus pomeegra.
. Under view of upper jaw of ditto.
. Side view of under jaw of ditto.

Fig.
Fig.
Fig.

Fig.
Fig.

Fig.
Fig.

Fig.
Fig.

Fig.
Fig.

Fig.
Fig.

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Fig.

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PROFESSOR OWEN ON INDIAN CETACEA.

. Upper view of symphysis of under jaw.
. Under view of upper jaw of Delphinus euphrosyne.

All the figures are half the natural size.

PLATE IX.

Phocena brevirostris.

. Side view of cranium and upper jaw.

. Upper view of ditto.

. Under view of ditto.

All the figures are nearly half the natural size.

PLATE X.

Euphysetes simus.

. Side view of female.
. Upper view of ditto (drawn to scale).

PLATE XI.
Euphysetes simus.

. Side view of male (to same scale as female, Pl. X.).
. Outline of ditto, with skeleton.

PLATE XII.

Euphysetes simus.

. Side view of skull.
. Back view of skull (rather more than half the natural size):

PLATE XIII.

Euphysetes simus.

. Upper view of skull.
. Under view of ditto (half the natural size).

PLATE XIV.

. Section of cranium of Euphysetes simus.
. Section of cranium of Physeter macrocephalus.
. Luphysetes breviceps.

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1 PHYSETER SIMUS.2.PHYSETER MACROCEPHALUS 3.PHYSETER

III. On the Osteology of the Dodo (Didus ineptus, Linn.).
By Professor Owrn, F.RS., F.Z.S., Le.

Read January 9th, 1866.
[Puates XV. To XXIV.]

§ 1. Introduction.

THE 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 admirable 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,
extinct bird which were known 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 under 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 Didun-
culus, directed the ornithologist and ornithotomist 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

1 ¢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.S8., F.R.G.8.,
President of the Ashmolean Society, &c., and A. G. Melville, M.D. Edin., M.R.C.S. 4to, London, 1848.

2 See Annals of Nat. Hist. ser. 2. vol. vi. p. 290 (1850).

3 «Bs war in 1843, dass ich auf den Gedanken kam, dass der Dodo eine Sancinall Taubenform sei; ich
iiberzeugte mich bald dass diese Auffassung die einzig richtige sei, und fing an eine Arbeit iiber diesen Gegen-
stand vorzubereiten. In 1845 wurde ich aber von meiner Regierung beauftragt eine Reise um die Welt mit
einem dinischen Kriegsschiff mitzumachen ; meine Arbeit musste also vorliiufig bei Seite gelegt werden. Schon
vor meine Abreise hat ich aber mehrere sowohl diinische wie fremde Naturforscher mit meiner Ansicht bekannt
gemacht, und der Beweis das es sich so verhiilt wird Owen finden kénnen :—

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

«2. in Sundeyall, Arsberiittelse om Framstegen i vertebrerade Djurens Naturalhistoria og Ethnographien,

1845-50, p. 254.”—Letter from Prof. J, Rurnmarpt to Dr. Arperr GUNTHER.

VOL. VI.—PART II. #

50 PROFESSOR 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 imeptus which has at length been
transmitted from the island of Mauritius to London, under the following circum-
stances.

In 1863, I 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 pear 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 natural 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 diligence, in his vacation-times (by no means extensive), which deserves much
credit and encouragement.

* Reinhardt, quoted by Strickland, op. cit. p. 41 (see also p. 70).
* This Collection was purchased by the Trustees of the 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.

‘7 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 Mahébourg, Island of Mauritius :—

“On the estate called ‘ Plaisance,’ about three miles from Mahébourg, in the island
of Mauritius, 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 tibiee, 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 tibiz 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 living—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 nearly thirty years that I have inhabited this colony, I have made frequent

inquiries of old people as to the finding of the bones of large birds, and have offered liberal
H 2

52 PROFESSOR 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-down. (Signed) “GrorGe Cuark. 1865.”
The above “Statement” was authenticated by the following testimony :—
“Having visited the place with Mr. Clark, I can vouch for the truth of the facts
herein mentioned. (Signed) “ WituiaM THomas BaANKs,
* Civil Chaplain, Mauritius.”
“The Rey. W. T. Banks, Civil Chaplain at Mahébourg, in this diocese, and Mr.
George Clark, Master of the Government School at Mahébourg, 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 (Didus 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 ..............eceeeeeees 14
Vertebres'‘and pelyis! <i... .cs-o-ssrs-ctses esse senneecsensncs 30
RDS ssisdietiss ocine'sectpneslcrecicinamoethciianice Mesaaviinciemep ete raes te 22
SternuUng cee seeanesere sate ccense somerset daszduaoraabcode 2

: NcapMlararchy any parts -epases ss acsp eae eects Seabascetye 7
Humerus, ulnatradius seeerccsseserteee sees pee aa en Ne 6
HEMOTA <isssmpwanantensatasaneeeeeeten cere senescence esestee 5
Tibias. i sievccnesanpedertaneeteetacaceetesgcaces snacemelshs ade 6
Fibula 52 cv 2 ctetencdaectetae toatrancuten Seslepepacieis ane aaese 4
Metatarsal, oct eues.-cee eee Gabo aguerneoudcdad uo abonoEsncac 4
Total number of parts of skeleton of the Dodo ............ 100

The known characters of the skull and metatarsus of the Didus ineptus served 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 was in type, as bones of the Dodo. Ihave 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. With 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. Vertebre.

The dorsal vertebree are chiefly represented, in this series of bones, by three which are
anchylosed together by their bodies and neural arches (Pl. XVII. figs. 1-5) : the posterior
articular surface of the body of the last of these vertebre (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 (Pl. 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 vertebra is the penultimate dorsal. The anterior arti-
cular surface of the foremost of the three (Pl. XVI. fig. 1, ¢) 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 vertebre are compressed
and wedged-shaped, slightly expanded at their coalesced. ends, produced below into
subquadrate hypapophyses in the first and second (Pl. 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 third (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.—R. O., June 1866.

1 Tn the quaint print, in folio 3, of the “ Narration Historique du Voiage faict par les huict Navires d’Am-
sterdam au mois de Mars l’An 1598. soubs la conduitte de l’admiral Jaques Corneille Necq,” &c., the first-
named object, No 1, “Sont Tortues qui se tiennent sur haut pays, frustez d aisles pour nage, de telle grandeur,
qwils chargent ung homme et rampent encore fort roidement, prennent aussi des Keriuisses de la grandeur d’un
pied qu’ils mengent. 2. Est ung oiseau, par nous nommé Oiseau de Nausée, 4 Yinstar dune Higne, ont le cul
rond, couvert de deux ou trois plumettes crespues, carent des aisles, mais en lieu dicelles ont ilz trois ou
quatre plumettes noires, des susdicts oiseaux ayons nous prins une certaine quantité, accompaigné d’aucunes
tourturelles et autres oiseaux, qui par noz compaignons furet prins, la premiere fois qu’il arrivoyent au pays,

" pour chercher la plus profonde et plus fraische Riviere, et si les navires y pourroyent estre sauvez, et retour-
nerent d’une grande joye, distribuant chasque navire, de leur Venoison prins, dont nous partismes le lendemain
vers le port, fournismes chasque navire d’un Pilote de ceux qui au parayant y avoyent esté, avons cuict cest
oiseau, estoit si coriace que ne le poyions asses boviller, mais avons mengé a demy cru. Si tost qu’arrivames
au port, envoya le Vice-Admiral nous, avecq une certaine troupe au pays, pour trouver aucun peuple, mais
nont trouvé personne, que des Tourturelles et autres en grande abondance, lesquels nous prismes et tuames,
car yeu qu’il n’y eust personne qui les effraia, n’avoient ilz de nous nulle crainte, tindret lieu, se laisserent
assomer. En some c’est un pays abodant en poissd et oiseaux, voire tellemet quwil excella tous les autres
audit voyage.”—Le Second Livre de la Navigation des Indes Orientales, fol., 1601.

54 PROFESSOR OWEN ON THE

‘The hypapophysis of the first of the three expands at its termination (Pl. 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
(Pl. XVII. figs. 1 & 5, hy). Each vertebra shows an elliptical articular cavity (ib. figs.
1 & 5, p, p2) 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 (Pl. XVI. fig. 1, d).

The neural arch circumscribes a canal the anterior outlet of which (ib. fig. 1, 2) is
oval with the small end downward, 5 lines in vertical, and 53 in transverse diameter:
the sides of the neural canal slightly project inward above the lower third: the posterior
outlet (Pl. XVII. fig. 4, x) 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 (PI.
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 expands
into the postzygapophyses: these have coalesced with the preezygapophyses in the suc-
ceeding vertebra (Pl. XVII. fig. 2, z), as has happened also between this and the third
vertebra. In the last of the three vertebre the postzygapophyses are entire (ib. z 3),
and show very slightly concave, oval articular surfaces, looking obliquely downward and
outward (ib. fig. 4, z). The conjugational foramina, continuously surrounded by bone,
are a full ellipse, and large, the anterior one (ib. figs. 1 & 5, f) being 5} lines in vertical
diameter; the second (ib. f") 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, ms) 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 (Pl. XVI. fig. 1, ms). A small foramen behind the base of each
of the coalesced zygapophyses (Pl. XVII. fig. 2, z z) leads to a canal descending to
the neural one, and indicates superiorly the limits of the otherwise continuously ossified
neural arches.

In the series of detached vertebra, one (Pl. 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. hy), 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 (d) has a larger costal surface: it sends for-
ward a convex ridge midway between the di- and zygapophysis (z). The neural canal
(fig. 7, 2) 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 (pl) existed, indicating the present to be the first of the dorsal
series, as shown in Pl. XV. ‘The 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 6 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. pl): this
projects backward 7 lines in length, terminating obtusely, and circumscribing a ver-
tebrarterial foramen (ib. v) of a full elliptic shape, 53 lines in long diameter. The
surfaces of the preezygapophyses (z) are larger, and look more upward and less inward,
than in the preceding and the dorsal vertebree: they are very slightly concave. ‘Those
of the postzygapophyses (fig. 8, 2’), with a downward and slightly outward aspect, are
in a similar degree convex. The neural canal, as usual in the cervical series, expands
at its outlets, most so posteriorly (fig. 9, 2); 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 vertebre (Pl. XV.). The third cervical has also the
hypapophysis (Pl. XXIII. fig. 3, hy).

In all the other cervicals of the present series the hypapophysis is wanting, but each
parapophysis developes a plate (Pl. XVII. figs. 10 & 11, Pl. XX. fig. 1, p) to form the
sides of the hemal canal through which the carotids ran; and the position of such ver-
tebre 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 vertebra, however, they are mutilated. They differ chiefly in
the position and shape of the anapophyses (fig. 10, @), which advance from above the

56 PROFESSOR OWEN ON THE

postzygapophyses (z’), converging towards the middle of the upper surface of the neural
arch, being arrested, save in one instance, at the sides of the ligamentous surface
occupying the common position of the base of the neural spine.

In the axis vertebra (Pl. 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, hy) descends below this surface: the anterior or odontoid surface
presents the usual form in birds; the odontoid process (ib. w) has a pit at its apex.
The prezygapophyses (fig. 12, z), of very small size, project from the outer and fore
border of the neural 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, 45 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. ms), 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 (Pl. 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
of the pleurapophyses and other processes.

§ 3. Rids. (Plates XV. & XVI.)

The specimens of ribs include both vertebral and sternal portions; that which appears
to be the second or third on the right side (Pl. XVI. figs. 7, 7 a) is 4 inches 4 lines
in length (following the outer curve), and expands to a breadth of 7 lines at its lower
part; the interval between the articular surfaces of the head and tubercle is 6 lines.
The appendage (ib. a) has coalesced with the middle of the hind margin of the shaft.
The neck is compressed, with a thin upper margin; the lower one is continued with
a curve upon a strong internal buttress-like ridge (ib. 4), 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 channel 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 the epipleural process. ‘The anterior border is thicker,
and is almost straight. Towards the sternal end the pleurapophysis contracts and thickens,
terminating in a rough syndesmotic elliptical surface, 3 lines by 2 (fig. 7,f), for the at-
tachment of the hemapophysis or sternal rib.

A vertebral rib (ib, fig, 2) which is entire, measures 9 inches in length (following the

it

oO

OSTEOLOGY OF THE DODO.

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 the 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 hemapophysis 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 margin indicative of the attachment of such process.

The body of a posterior vertebral rib (Pl. XVI. fig. 10) shows a fracture which has
been healed, with some irregular ossific deposit on the inner surface. All 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 (Pl. XV.) and the five right ones do not, either of them,
constitute a consecutive series. but have come from different individuals, of different
sizes, as exemplified in the third rib figured in Plates XV. and XVI.

The sternal ribs (Pl. XVI. figs. 5 & 12) are characterized by the two facets, nearly
or quite meeting at an open angle, into which their sternal end expands (ib. fig. 3, ¢).
One of these ribs, which is entire, shows the single, elliptic syndesmotic surface at the
opposite end (ib. 4); it is 3} inches in length, with a greatest breadth of 4 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 Pl. 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 hemal 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
yertebrie, with which the iliac bones are continuously confluent.

1 See, especially, Bontekoe’s figure, copied by Strickland, in the title-page and at p. 63 of the above-cited work.

VOL. VI.—PART II. I

58 PROFESSOR OWEN ON THE

The first sacral shows the transversely extended and concave articular surface of the
centrum (Pl. XIX. fig. 1,¢); the subcircular pit (ib. p) 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 subyertebral muscle
(longus colli?). The anterior outlet of the neural canal (ib. 2) is subcircular in one spe-
cimen, vertically elliptic in others, and 3 lines or less in transverse diameter. From the
sides of the neurapophyses stretch out the strong buttresses of bone which blend with
the under part of the ilia, giving off from the fore part of their base the preezygapophyses
(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. ms) is roughened by a syn-
desmotic surface ; it rises to a height of 14 lines, curving 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, ms) to the hindmost vertebra of the series, without any trace of
their primitive division; but this track rises, posteriorly, above the shallow channel on
each side, in which are the foramina (ib. 0), indicating most of the constituent vertebre.

The second sacral vertebra abuts against the ilium by a pleurapophysis (ib. fig. 1,
pl2), as well as a diapophysis (ib. d2); but the former is a slender, straight filament, or
narrow plate of bone, confluent at both ends.

In the next two vertebre the pleurapophysis (ib. p/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 first of these rib-buttresses inclines forward,
and is completely confluent with the ilium; the thicker one (ib. p/ 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 vertebra in which the pleurapophysis is not de-
veloped, the attachment to the ilia being by diapophyses only (ib. d d), which are short
slender lamelle, 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. p/ 8) 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 vertebra is traversed by a sharp median longitudinal
ridge, which is more feebly and interruptedly continued to near the end of the sacral
series.

Eight vertebre, 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, 0) 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, 7), extending obliquely backward to behind the acetabulum—the an-
terior division being narrower and concave, the posterior broader and convex but in 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. ¢ z), 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 5 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) 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. ¢ ¢) 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. 0) 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. 7’), 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. 7), 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 then
again diverge to the last sacral; they coalesce with the diapophyses (ib. fig. 2 d qd).

‘ Owen, ‘Anatomy of Vertebrates,’ 1866, vol. ii. p. 32.
12

60 PROFESSOR OWEN ON THE

The inner or under surface of the ilium is thickened into a kind of buttress (ib. fig. 1, e),
terminating behind the ischiadic foramen. The breadth of the iliac bones and inter-
vening 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, 64) have completely coalesced around the aceta-
bulum. The pubis, which in this part is 7 limes 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 concave 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, f), which is 9 lines in length, and is situated one half in
advance of, and the other half beneath, the ischiadic foramen (ib. m). This 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 pelvis are on the inner surface, above the 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. ¢) near its posterior junction with the ischium. The preerenal fossa (be-
tween pl 4 & pl 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 (Pls. 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 (Pl. XVI. 4) is 45 inches; from this diameter the bone contracts anteriorly to a
breadth of 33 inches at the costal processes (ib. d), and posteriorly it contracts more
rapidly to an obtuse, horizontally flattened apex (Pl. XVIII. fig. 3). The anterior
border of the sternum (Pl. XVI. fig. 4.) is widely and rather deeply emarginate at
the middle (¢), less deeply so on each side: the breadth of the mid notch (4 ¢ d) is
1 inch 9 lines, that of each side notch (6 d) is 1 inch 2 lines. The sternum is deeply
hollowed above (Pl. XXIII. fig. 4), correspondingly convex beneath (ib.); the keel (s)
is low and thick, commencing by a pair of broad obtuse ridges (Pls. XVI. fig. 4, &
XVIII. fig. 1, 7 7) from the mesial ends of the outer walls of the coracoid grooves

OSTEOLOGY OF THE DODO. 61

(ib. 6), 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 3 of an inch at the middle of the sternum, then
gradually sinks to the level of the bone, as it extends backward, at 1+ inch from the
hind end (Pl. XVIII. fig. 3), a little increasing in thickness as it subsides: its free
border describes a pretty regular convex curve (Pl. 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 (Pl. XXIII. fig. 4). Behind the costal surface (Pl. XVIII. ce),
on each side, extends a lamelliform process (Pls. XV. & XVIII. h), 4 an inch 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 (Pls. X. & XIV. fig. 3):
there is no trace of an entolateral process (ib. 7). The thin margin of the Dodo’s breast-
bone, behind the ectolateral process (Pls. XV. & XVIII. A), is entire and uninterrupted
to the obtuse apex, and the body of the sternum is imperforate: the notch (f) behind
the process (i) represents the ectolateral notch of the gallinaceous sternum (Pl. XXIV.
figs. 1 & 3, f'). The costal border (Pl. XVIII. fig. 2, ¢) 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 bilobed, the anterior one (¢ 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
(d)' 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 from the costal border; the inner side is smooth and convex: it is
not quite entire on either side. The coracoid grooves (Pl. XVI. fig. 4, 0 6’) are small
in proportion to the sternum, and are divided from each other by an interspace of about
an inch; the outer wall of the groove (’), 9 lines in extent, is moderately produced and
convex ; it appears to be a continuation of one of the initial ridges (7) of the keel: the
inner wall of the groove (4) is deeper, and is formed by the obtuse angle of the anterior
border of the sternum, between the medial and lateral emarginations. External to each
coracoid groove is a large elliptical pneumatic foramen (p) or depression. ‘There is no epi-
sternal process. On the convex outer surface of the body of the sternum the “ pectoral ”
ridge (Pl. XVIII. fig. 1, 4)’ 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

1 Called “ hyosternal ” in the Geoffroyan determination of parts of the bird’s sternum.
2 '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. 16-23.

62 PROFESSOR OWEN 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 (Pl. XXIII. fig. 4).

§ 6. Scapular Arch. (Plates XV. & XX.)

This consists of the scapula (Pl. XX. figs. 6, 7, 8 & 9, s1), coracoid (ib. figs. 4 & 5, sx), 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 articulation. The scapula (ib. figs. 6 & 7,8 & 9,
51), 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 short process (ib. 51); between this process and the articulation the breadth of the
bone is little more than 3 lines; the breadth of the articular end is 9 lines. Nearly
one-half of it is occupied by the almost flat, subcircular humeral surface (fig. 8, a), with
a diameter of 44 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, ¢) extends forward, curving toward the coracoid, and terminating obtusely.

The coracoid (ib. figs. 4, 5, 8 & 9, 52), 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 (7), and extends in advance
of the pneumatic foramen (Pl. XVI. fig. 4, p) 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 subsides,
at one-third of the extent of the bone, is sharp; the inner border is obtuse to near the .
inner angle (Pl. XX. figs. 4 & 5,7). 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, Pl. XX. A muscular ridge and rough surface (ib. fig. 9, 7) 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, first, 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 (/), from which the thick, obtuse, inner

“continuation of the scapular end projects inward, forward, with a slightly upward curve,

OSTEOLOGY OF THE DODO. 63

and shows the narrow oblong surface for the articulation and ultimate confluence of the
clavicle (58). The coracoid unites with the scapula at an angle of 100°.

The clavicle (ib. figs. 4 & 5, 5s), 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 scapule’” (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. (Pls. 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 (Pl. XX. figs. 12-14), but differing 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. @) 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. 5) is triangular, obtuse, short, and
bent, or directed toward the front side of the bone: the ulnar tuberosity (ib. ¢) is more
produced in that direction ; it is oblong, obtuse, with its base impressed by a large pit
both above (fig. 12, i) and below—the lower one (ib. g) 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, ¢) and a lower and internal
(ib. g), 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, 7),
half an inch in length; it gradually expands at the distal third to a breadth of 10 lines.
where the articulations offer 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 (Pl. 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 air-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 (Pls. 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 concaye ; the distal end expands to the same
extent, but is compressed, as usual.

64 PROFESSOR OWEN ON THE

The ulna (Pls. 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 length (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
slightly 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 pinion bone in the collection of remains submitted to me: this
part of the wing is conjecturally restored in dotted outline in Plate XV.

§ 8. Bones of the Leg. (Pls. XV., XXI., XXII. & XXIII).

Of the five femora in the above defined series of remains of the Dodo, two measure
6 inches 3 lines in length; one (Pl. X XI.) is 6 inches 44 lines; the shortest is a little
under 6 inches, with proportionate differences in the diameter of the shaft. All of them
show a small pneumatic foramen (Pl. XXI. figs. 1 & 2,p) 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, 6), adapted to the trochanterian prominence of the pelvis (Pl. XIX. 4), is
well-defined. The trochanter (Pl. X-XI. fig. 1, ¢) rises, ridge-like, 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, 7), inclining to the middle of
the fore part of the shaft, is continued from the trochanterian 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. 3, m) perforates the middle of the back part of the shaft; the
popliteal fossa (ib. fig. 3, 0) shows a few small pneumatic orifices; a triangular rough
flat surface divides the fossa from the outer condyle. Above the fibular depression
(ib. fig. 5, 7) there is a well-defined, slightly raised, rough surface (ib. &) for the head
of the ectogastrocnemius muscle. he ridge (ib. m) extending to the back part of the
inner condyle is not sharp; the rotular groove (ib. fig. 1, p) is deep and moderately wide,
with the inner boundary, formed by the narrow anterior part of the inner condyle (ib.
fig. 5, ¢), 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 Pl. XXI. fig. 5.

The head, neck, and great trochanter (Pl. 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 THE 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 tiie 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, p) 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
surface is well defined by an irregular inferior line or ridge (ib. fig. 2,2). The ectocnemial
process (ib. figs. 1, 3, 4, ¢) is thicker, shorter, and terminates roughly and obtusely.
There is a low, narrow ridge (ib. fig. 2, 7), about half an inch in length, on the inner
side of the proximal end of the shaft, beginning about 9 lines below the articular sur-
face at that end. ‘The fibular ridge (ib. figs. 1 & 3, h), beginning 1 inch 8 lines from
the proximal end, extends about 2 inches down the outer side of the shaft. The epi-
cnemial ridge (ib. figs. 1 & 4, &) is obtuse, and but little produced above the upper
articular surfaces or condyles (¢ 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, 7), 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. 4).
Their hind ends project very little beyond the level of that aspect of the shaft of the
tibia. An intermuscular ridge (ib. fig. 1, 7) strengthens into a tuberosity (r’) 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 (Pl. XXIII. fig. 8), and to about an inch anda
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 structure indicates the extent of the original
distal epiphysis of the tibia (fig. 8).

The fibula (Pl. 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 proximal
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 haye received every requisite
attention in this way in the excellent work on the Dodo and its kindred, already
quoted. A longitudinal section of the metctarsus, taken in the direction from side to side
(Pl. XXIII. fig. 10), shows the loose cancellous texture of the common epiphysis of

VOL. VI.—PART II. K

66 PROFESSOR OWEN ON 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 (77), 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 (7v) is wider and descends
lower before the breaking up of the inner surface into decussating lamelle or filaments,
than that of the inner metatarsal (i): 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
(Pl. XXIII. fig. 1, 32), confluent at the “ gonys,” are distinct from the hinder halves
of the rami formed by the confluent, or perhaps connate, articular, surangular and an-
gular elements (ib. 31): 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 characters of these important evidences of the specific
character of the remains from the Mahébourg morass’, they agree with those of Didus
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. 1) presents the same hemispheroid or reniform 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 superoccipital
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 Didunculus (Pl. XV. fig. 2) shows no trace of it; I have also failed to find it in the
skull of a Crown-pigeon (Gowra coronata). The superoccipital ridge is defined by the
subsidence of the surface beneath it being continued directly from the upper, almost
flat, smooth surface of the cranium: the middle part of the ridge is more produced than

1 «Ta Mare 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, Didus most
resembles Dinornis. The basioccipital curves downward, and unites with the basi-
sphenoid in developing the pair of larger tuberosities (Pl. XXIII. fig. 1, 5), which ter-
minate about 4 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 flattish 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. @) 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 fossze (P]. XV.), in the present specimens, show the same contraction in
proportion to their depth hy 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, )idwneulus shows a
closer resemblance to Didus than does Gowra, 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 cavity,
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 (Pl. XV.); in Didus the breadth
greatly exceeds the length of the interorbital frontal convexity, as compared with

K 2

68 PROFESSOR OWEN ON THE

Didunculus, and the convexity reigns in the transverse as well as the antero-posterior
direction ; in Didunculus, however, it is less concave transversely than in Gowra. In
the breadth or thickness of the interorbital septum Didus resembles Apteryx and Palap-
teryx and shows the same pneumatic cancellous structure. The posterior olfactory
chambers are partially divided, as in Dinornis, by an upper median septum; each
compartment, which is 7 lines across and an inch in length, is perforated posteriorly 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 (Pl. XXIII.
fig. 1, 22) 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.’ 2. ¢. 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 perforations and
grooves, with a sharp alveolar border, which describes a sigmoid curve lengthwise, and
with a deeper concavity of the palatal surface than in Dinornis or Didunculus. Moreover
the concavity is partially divided 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 slightly 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 with 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 (Pl. XXIII. fig, 1, 22*) is smooth and slightly
convex vertically ; both upper and lower borders are obtuse and thick.

Proc. Zool. Soe. lc. p. 5.

OSTEOLOGY OF THE DODO. 69

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 (Pl. XXIII. fig. 1, vc) is 1 inch 8 lines, 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, ¢.g. Microglossum aterrimum; but it is fully paralleled only by the Elephant
among air-breathing vertebrates, as may be seen by comparing the section Pl. 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 diploé 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 diploé above the cerebral cavity equals the vertical diameter of
that cavity in Didus: the diploé gradually decreases in thickness as it approaches the
foramen magnum. ‘The disposition of the osseous lamelle forming the cells or cavities
of the diploé is very different 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 cavity. The

1 Proe, Zool, Soc. 7. ¢. p. 6.

2 Zool. Trans. vol. iy. pl. 24. fig. 4.
3 Qdontography, pl. 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 skull, 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 the 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 (Pl. XXIII. fig. 1, 1 0) of the cranial cavity accords with
the 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, Casuarius, Rhea, Dromaius, Struthio, Aptornis,
Cnemiornis, Palapteryx, Atpyornis, Dinornis, &c. among terrestrial birds, of Notornis
among the lake-haunting Coots, and of Aptenodytes and Alca 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, Cnemiornis, Apyornis, Palapteryx, Dinornis, Didus, and Pezophaps, with the
largest of the Auks, have thus passed away, while Notornis and Apterya 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 Man 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 (Pl. XXIII. fig. 1, 0) 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. ¢7) 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. 2)
which appears to have received veins from the cranial diploé. 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 hemi-
spheres ; and this ridge shows a narrow groove as for a small longitudinal smus, A
transverse linear groove abruptly defines the fore part of the ridge.

The vertically expanded anterior part of the premaxillary (ib. fig. 1,22) has a large
pneumatic cavity communicating by a reticulate wall with the cells of a cancellous struc-
ture, larger than those of the cranial diploé. ‘The maxillary branch of the premaxillary
(ib. 22*) consists of a light open-work air-diploé, with a very thin outer case of bone.
‘The short symphysis mandibule shows a small cavity, surrounded 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 (¢.9. 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 PROFESSOR OWEN ON THE

wielding 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
might 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, more 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 vertebre: I have observed four to be so welded together by both centrums and
neural spines in Phenicopterus, 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 birds of prey five dorsal vertebre 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, Cranes, Psophia, Cariama, Palamedea, the Penguins, and
in all flightless land-birds save the Dodo, no such anchylosis takes place. The Colum-
bide are the species in which the dorsal vertebree, 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 develope, in Goura and Didunculus, hypapophyses closely corresponding
im 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 Columbide is in the greater number of the vertebre or segments which
are typically completed by bony hemapophyses articulating with pleurapophyses and
directly with their mass of coalesced and expanded hemal 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 hemapophysis terminating in a point before
reaching the sternum, and extensively connected with the antecedent hemapophysis or
sternal rib: in both genera two dorsal vertebra in advance of the typically complete one
have moveable pleurapophyses terminating freely in a point, with no hzemapophyses
other than the costal processes of the stemum may represent. In Goura, which has six
pairs of moveable or thoracic ribs, the second pair belong to the first of the three anchy-
losed dorsal vertebree: in Didwnculus, 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 Pl. 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-vertebree under my observation exhibit such
characters, the proportion of those with neural spines, or with hypapophyses, or both,
is the same as in the Colwmbidew. 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 vertebre of Didus, which I have
yet received, shows such circumscription of the hemal canal. The majority of the
cervicals in Didus (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 Didus has both the above processes, as in Columbide: 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 vertebre 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 regiom, encompassing a more capacious
thoracic-abdominal cavity. The ribs of the Vulture are more expanded than in Didus,
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 Didus 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 (Pl. XV. fig. 2,
Didumculus), 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 Colwm-
bide (Pl. XXIV. fig. 2, 4); consequently the inner or upper wall of the confluent grooves
forms a median prominence (ib. ¢) at the front margin of the sternum, contrasting with
the wide notch at that part of the bone in the Dodo (Pl. XVI. fig. 4). The next differ-
ence, as compared with Gouwra and most Pigeons, is the absence of the entolateral
processes (Pl. XXIV. fig. 3, 7) in the Dodo’s sternum: but Didunculus singularly exem-
plifies its nearer affinity to Didus by a like absence of those processes; only the sternal
margins behind the ectolateral processes (ib. fig. 1, h), instead of conyerging with a
slight convexity to an obtuse apex, as in Pl. XVIIL., describe a concavity, through an
expansion of the posterior truncate end of the breast-bone. The sternum of Didwnculus
may be said to show one pair of posterior notches (Pl. XXIV. fig. 1, f), that of other
Pigeons two pairs (ib. fig. 3, ff’); but the sternum of Didus, which is relatively broader,
shows no other trace of the anterior notch (Pl. XVIII. f) than is afforded by the
rounded angle at which the ectolateral process (/) rises from the bone. Although the

VOL. VI.—PART II. L

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 Didunculus as compared with Gowra, in which latter
Dove there are articular surfaces for three sternal ribs (Pl. XXIV. fig. 3, ¢ 1, 2,3), whilst
in Didunculus there are four (ib. fig. 1, ¢). Didunculus also exhibits, more strongly
than Goura, the obtuse ridges (ib. fig. 2,7) 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, Pl. XVI., which I do not find in the sternum of Gowra; 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 (Pl. XXIV. fig. 1).

In the direction of the ectolateral processes Gouwra (ib. fig. 3, h) 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 their backward course, not so convergent as in Didus. In Goura the subcostal ridge
is better marked than in Didunculus. In no Dove of flight is the body of the sternum
so broad and hollow as in Didus (Pl. 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 in the sternum of a Night-jar (Po-
dargus hwmeralis), The ectolateral processes (Pl. XXIV. fig. 4, h) rise behind the
moderately extended costal borders, ¢; and beyond them the body of the sternum con-
verges to an obtuse end, with 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, /’, of which one was present
on one side in the specimen figured.

Through the reduction of the coracoids in all flightless birds, there is an interval
between their sternal articulations: this is long and concaye in the Dodo, but is
longest and most deeply concave in Apteryax; it is long but almost straight in Rhea; 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, Bdicnemus, 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 power of flight, and expanded and excavated for the support of the larger gizzard
with its heavier grindstones’.

In comparing the pelvis of Didwnculus and Goura (Pl. XXIV. fig. 5) with that of
Didus (Pl. XTX. 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 vertebre ;
then follow, as in Didus, three vertebree without pleurapophyses, these reappearing in
the next two with their 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 Gowra 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 Dodlets; 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 Gowra. 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 Didus ; 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 Gowra those margins reach the
neural crest, but do not overtop it at any part. In Gowra the acetabula are more in
advance of a median position than in Didunculus, Columba magnifica, or Didus. Although
the ischiadic foramina are completed by terminal confluence of the ilium and ischium in

1 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 Dopo, 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 like 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 (Wilkin’s Edition, 4 vols.: London, 1836), vol. i.

p. 369; vol. ii. p. 173.
L2

76 PROFESSOR OWEN ON THE

Dromaius and Casuarius, yet the length of those foramina (which are unclosed) in
Struthio and Apteryx, concomitant with the greater relative length of the pelvis, shows
the difference of Didus 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 Didwneulus: 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 longer and narrower: the proportions of that in Didwnculus are more like
those in Didus. In Didunculus the pubis coalesces with the ischium behind the small
obturator foramen, but leaves a second or posterior elongate ischio-pubic vacuity. The
greatest amount of resemblances with the pelvis of the Dodo is found in that of different
members of the Dove-tribe.

In comparing the pelvis of the Dodo with that of the Vulture (Pl. XXIV. fig. 6), we
find in the latter that the first two confluent sacral vertebre supporting moveable ribs
are succeeded by several with short abutting ribs, the extent of this part of the sacrum
being nearly one-half of the whole, instead of one-fourth as in Didus and the Doves.
The reappearance of rib-abutments after four ribless sacrals is in the posterior third of
the sacrum, and they are continued to the end of that bone from the last four vertebre
of the series, constituting a very marked difference, both as to number and the character
of the vertebre in the sacral part of the pelvis.

With regard to the iliac bones, the anterior concave track occupies two-thirds of the
extent of the bone in Vuléur, not one-half as in Didus 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 Didus. In Ciconia the antacetabular part of the pelvis is relatively
longer, and the iliac bones are more expanded anteriorly. In Platalea the proportions
are more nearly those in Didus. In Ofis 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 ilium is longer, narrower, and deeper, in most
waders, than in Didus. In Eudyptes and Aptenodytes the ilia are more expanded ante-
riorly, but the whole pelvis is narrower and longer than in Didus. The Gar-fowl
(Alca impennis)', Uria, Podiceps, and Colymbus, 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 from 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 diploé 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. vy. pl. 51.

OSTEOLOGY OF THE DODO. T7

of the prosencephalic roof resembles that in Didus. 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 represents very feebly the tentorial tuberosity (Pl. XXIII.
fig. 1,0) in the Dodo: from it, in Gowra, is continued backward the arch of bone formed
by the superior semicircular canal, above which is the groove for the venous sinus, as
in Didus. The internal auditory fossa is less deep than in Didus: above it is a similarly
vertically oblong cerebellar pit. The nerve-foramina correspond with those in Didus:
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 diploé 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 internal 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 Didus; 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 5 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 Didus (Pl. XX. figs. 12 & 14) as in the
volant Doves (Pl. XXIV. figs. 8 & 9, Gowra); that, indeed, which is a ridge (77) on the
back part of the shaft in Didus, is a mere rough surface in Gowra, and does not show
in Didunculus. The pneumatic fossa, which varies in depth in the two humeri of the
Dodo, is in both relatively larger and shallower than in Gowra. ‘The pectoral process
is thinner, but relatively rather more produced, in Didunculus, The humerus in Gdi-
cnemus, Otis, and Charadrius has a more longitudinally extended, thinner, and more
produced pectoral ridge than in Didus and the Columbidw; 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 differences

' Trans, Zool. Soc. vol. iv. pl. 24. fig. 4.

78 PROFESSOR OWEN 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, pl. 66. figs. 7-10). In that extinct
species, although the humerus is 5} inches in length, the parts indicative 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 Gowra coronata (Pl. 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 Gidicnemus 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
detined 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. Soe. vol. y. pl. 65. fig. 3.

OSTEOLOGY OF THE DODO. 79

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

The femur in Cnemiornis’ and Dinornis* is much thicker, in proportion to its length,
than in either Aptornis or Didus. 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 axe 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 Gowra coronata is 4 inches 7 lines,
and its shaft is more slender, in proportion to its length (Pl. XXIV. fig. 13), than in
Didus (P\. 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 (Hdicnemus erepitans) the tibia, as in other Gralle, is longer in
proportion to its thickness than in Didus; the epicnemial process rises higher above and
projects further in front of the condylar surfaces 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, Ciconia, Mycteria, Porphyrio, opposes
similar or equivalent differences to those in Adicnemus, against the affinity of Didus to
any of those Gralle.

In the comparison of the tibia of this extinct flightless bird with that of the Cnemiornis,
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 Cneméiornis; 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.

1 Trans. Zool. Soc. vol. v. pl. 65. fig. 1. ? Thid, fig. 5,

80 PROFESSOR 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®. 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 continuance 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 Mauritian Doves (Columba nitidissima and C. meyer?) that
retained their powers of flight continue to exist there.

As I have no reason to offer 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 organic creation is equally perfect, the

1 Histoire Naturelle, &c., 4to, tom. xiv. “ Dégénération des Animaux: ” 1760,

? Philosophie Zoologique, 8vo, 1809, tom. i. chaps. 3, 6, & 7.

3 Avreeably with the principle of the “contest for existence” by which I explained the extinction of the
species of Dinornis, Trans. Zool. Soc. vol. 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 ineptus, L., through its degenerate or
imperfect structure, howsoever acquired, has perished. What have the stigmatizers of
Buffon 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 wings 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 grounds 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 suppose, 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 vitally 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

1 Strickland and Melville, ‘The Dodo and its Kindred,’ 4to, 1848, p. 34.

2 Op. cit. p. 34.

2 Sce letter in ‘The Times’ of May 21st, 1862, advocating the limitation of the National Museum of Natural
History to “six rooms,” signed Tuomas H. Huxxey, F.R.S.

4 Reply to the aboye 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 deyoid 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 PROFESSOR OWEN ON THE

as I seemed to do on this point in the array 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 Address 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 (Didus ineptus, L.), with an outline of
the bird as represented in the oil-painting presented to the British Museum
by Epwarps, 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.

Fig. 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.

. Front view of the fourth (or first of the three confluent) dorsal vertebre (centrum
and neural arch).

. Vertebral rib, or pleurapophysis, of the same vertebra, front view.

. Sternal rib, or hemapophysis, of the same vertebra: a, outer side; 0, upper or
pleural end; ¢, lower or sternal end; d, front margin ; ¢, inner surface.

. Front view of sternum, or connate mass of hemal 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.

I

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ae

—

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gai 08
wo po

Fig.

' «The doctrine of typical nuclei seems only a mode of evading the difficulty. Experience does not give us
the types of theory; and, after all, what are these types? It must be 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 similitudes 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, &., by W1zt1am R. Grove, Esq., Q.C., M.A. 8yvo, 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 Gould, ‘ Birds of Australia,’ part 22 (March, 1846).

OSTEOLOGY OF THE DODO. 83

Fig. 7. Anterior pleurapophysis with appendage, a, front view: c, capitular end; d,

Fig

Fig

Bch
ods

tubercular end; f, hemal end; 7 @, outer surface ; 7 6, inner surface.

An anterior pleurapophysis, front view.

Posterior surface of the upper end of a posterior pleurapophysis: 9 @, body and
lower end of ditto.

. 10. Part of a pleurapophysis which has been broken and healed.
. 11. Lower end of a posterior dorsal pleurapophysis, with connate rudiment of

appendage, @.

. 12. Heemapophysis.

PLATE XVII.

1. Fourth, fifth, and sixth dorsal vertebre, anchylosed, side view.
2. Ditto, ditto, upper view.

3. Ditto, ditto, under view.

4. Ditto, ditto, back view.

.O.
6
7
8
9

Ditto, ditto, mutilated, of another Dodo.

. Anterior dorsal vertebra, side view.

. Ditto, front view; p/, outline of heads of floating rib.
. Penultimate cervical vertebra, side view.

. Ditto, back view.

. 10. Middle cervical vertebra, upper view.

. 11. Ditto, under view.

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

PLATE XVIII.
. Under view of sternum.
. Upper or inner view.
. Back view.

PLATE XIX.
. Under or inner view of pelvis.

. Upper or outer view of pelvis.

PLATE XX.

. Middle cervical vertebra, upper view.
. Fifth cervical vertebra, upper view.

. Fourth cervical vertebra, under view.
. Right coracoid and clavicle.

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PROFESSOR OWEN ON THE

Left coracoid and clavicle.

Right scapula, outer view.

Right scapula, inner view.

Left moiety of scapular arch, outer view.
Ditto, inner view.

. Upper articular end of right coracoid.

. Lower ditto.

. Left humerus, anconal or back surface.
. Left ditto, ulnar or inner surface.

. Left ditto, palmar or front surface.

A. Ditto, proximal or upper end.
B. Ditto, radial side of upper half.
C. Ditto, distal end.

. Right radius.
5 AG.
slits

Right ulna, inner or radial side.
Right ulna, outer or ulnar side.

PLATE XXI.

. Left femur, front view.
. Ditto, inner view.

. Ditto, back view.

. Ditto, upper end.

. Ditto, lower end.

PLATE XXII.

. Left tibia, front view.

Ditto, inner view.

. Ditto, back view.

. Ditto, upper end.

. Ditto, lower end.

. Left fibula, outer view.
. Ditto, inner view.

. Ditto, upper view.

PLATE XXIII.

. Longitudinal vertical section of mutilated skull.
. Ditto of third cervical vertebra.
. Ditto of lower cervical vertebra.
. Transverse vertical section of sternum.

OSTEOLOGY OF THE DODO. 85

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

PLATE XXIV.

. Sternum of Didunculus, upper view.

. Ditto, front view.

. Sternum of Gouwra, upper view.

. Sternum of Podargus humeralis, under view.

. Pelvis of Gowra, under or inner view, half natural size.

. Pelvis of Gyps (Vulture), under or inner view, half natural size.
. Left moiety of scapular arch, Gowra.

. Left humerus of Gowra, anconal surface.

. Left humerus of Gouwra, palmar surface of upper end.

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

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

Fig. 12. Right femur of Gouwra, back view of upper end, and back view of lower end.
Fig. 13. Right tibia and fibula of Goura, front view.

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IV. Description of the Skeleton of Inia geoffrensis and of the Skull of Pontoporia
blainvillii, with Remarks on the Systematic Position of these Animals in the Order
Ceracea. By Witiiam Henry Fiower, F.RS., F.R.CS., F.Z.S., &c., Conservator
of the Museum of the Royal College of Surgeons of’ England.

Read November 22nd, 1866,

[Prares XXV. to XXVIII]
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 tributary 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 Jnia, 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,
narrow, 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 Delphinus 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 boliviensist.
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 :—‘t Toutes ces observations nous font
regarder cette espéce comme’ ayant des mceurs beaucoup plus terrestres qu’aucune des
espéces connues.”—-“ Tous ces caractéres réunis 4 uhe dorsale peu apparente, nous font
proposer la formation d’un nouveau genre, qui Sees le passage entre les sousous
[Platanista] et les stelléres” [Sirenia].

* Reise in Brasil. t. iii. pp. 1119 & 1133 (1831). Von Martius states that his Delphinus amazonicus agrees
very closely with Desmarest’s description of D. geoffroyi, and eyen suggests that it may possibly belong to the
game species. His description of the teeth is sufficient to determine 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 little figure he gives
(fig. 34) more resembles Delphinus fluviatilis (Gervais) of Castelnau’s Voyage than the Inia.

+ Nouv. Ann. Mus. Paris, tom, iii. p. 23 (1834).

VOL. V1.—PART III. oO

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

In the Zoology of d’Orbigny’s ‘ Voyage en Amérique méridionale,’ “‘ Mammiféres,”
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 (pl. 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 Musée
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 Delphinus geoffrensis, and subsequently by Desmarest*
as D. geoffroyi.

In a later notice by Professor Gervais, in the Zoology of Castelnau’s ‘ Expédition
dans les parties centrales de l’Amérique du Sud,’ “ Mammiféres,” p. 90 (1855), this sup-
sition is confirmed, and the name Jnia 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 the
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 vertebre 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
still retains its epiphysial condition. The total length of the living animal, judging
from the skull and vertebre, and allowing for the intervertebral spaces, would be but
little more than 9’, the skull being 16:4". The specimen obtained by dOrbigny 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 193"
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).
+ Catalogue of Seals and Whales in the British Museum, p. 227 (1866).

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLIL. 89

As Inia has always been supposed to have certain affinities with Platanista, I
have in the following description compared the different bones with those of that sin-
gularly modified Cetacean on the one hand, and of several of the ordinary Delphinide
on the 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 :—

Collected by

Mr. Bates, | Mr. Bates. Mr. Bartlett.
a. 4.

Brxtmenreslene timers pert ret crete: avers eiessiey aeteteriet-eeler vos etotetie Melee 20-7"! 19-4" 16-7"
Length of rostrum (from anterior end of premaxillary to bottom

of antorbital notch of maxillary) ....... ....2.-.-0e e--| 135 12-7 11:0
From anterior end of premaxillary to lower edge of nasal bones .| 16-7 15:7 136
From anterior end of premaxillary to hinder edge of palate.....)] 15:7
Greatest breadth, across zygomatic processes of squamosals ....
Breadth of foramen Magnum... ........ eeeees ee reece cece
Breadth of the occipital condyles .... 2... +. e+ seen eee eee
Breadth across antorbital processes of frontals............+.+-
Breadth of rostrum at base (bottom of antorbital notches of}

THAXUIATIOS) ©. 5 0:0 vv vin cies so ee oe sm sielen vies opesmasiolje sloe ene
Breadth of rostrum at middle .........+.- sin Pes Spaces Oo
Mandible, length ........ e022 cece eter ee tere eens cee
Length of symphysis ...... 0. eee e ners steer rere neers
Greatest breadth across the posterior ends of the rami..,... ....
Height of ramus at coronoid process.... -.++-+++++eees ee eeee |

ne

mb R e
nd woo

i
SAG 0 sale LS ae bl

ar
WDCC ier Ce

OWHw AN

H
COTO ATH OD ND Dp
Ropotwth womr
oOmdbmree

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 premaxillary bones, and in the greater elevation on
the same side of the postnarial prominence of the frontal bones. Both maxillary and
premaxillary 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-

* «Om Gangesdelphinen,” Trans, Roy. Dan. Acad, 1851, Translated in Ann, & Mag, Nat. Hist. for March

1852, + Broken,
02

90 MR. WOH. FLOWER ON THE OSTEOLOGY OF

nication (see Platés XXV. & XXYVI.), 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 channelled 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’Orbigny, is very much larger in proportion-to the
size of the.cranium than in any other Dolphin, except Platanista, 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 from 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. which enter into the formation of 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 ‘6" with the well-developed alisphenoid, thus completely

INIA GEOFFRENSIS AND PONTOPORIA. BLAINVILLII. 91

shutting off the.squamosal from the frontal; whereas in Platanista 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 Delphinus. 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 Delphinide. 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, Znia 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 Delphinide 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 maxillaries, which gives
the peculiar character to the upper surface of the skull of Jnia as compared with that of
Delphinus. The difference is only one of arrangement of the same parts; there is
nothing superadded like the extraordinary outgrowths upon the maxille ot 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 maxillary. 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. FLOWER 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 Delphinide, in which they are generally reduced to irregular, oval, unsymmetrical
nodules. Phocena, however, differs from its allies in this respect, and closely approxi-
mates to Inia. In Platanista also the nasal bones are well-developed flattened plates ;
but they partake of the great elongation, narrowness, 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 Phocena 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 Delphinide (e. g. Globiocephalus).

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 Zphiine.

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 13" 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 Eschricht, who pointed out that 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*. The easily separable
condition of the bones of the young Platanista skull in the Museum of the Royal
College of Surgeons has enabled me to confirm Eschricht’s view; for on removing this
plate the true palatine is seen, forming as usual the greater part of the anterior and

* Ossemens Fossiles, 4m édit, (1836) tome viii. p. 130.

INIA GEOFFRENSIS AND PONTOPORIA 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 Platanista stands alone
among Cetaceans; even Jnia 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
their 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 wall of the chamber which lodges the great post-
palatine air-sinus. ‘This plate is slightly developed and very thin, perforated by
numerous large lacune, and, owing to the non-development of the outer reflected portion
of the pterygoids, is completely free along its inferior edge.

The pterygoids 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 surface 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 Jnia, 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 me process of
cleaning the skull, had been broken away. It is certainly possible that such is the case ;

94 MR. W. H. FLOWER ON 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. their 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 irregular, 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
bones.

One circumstance in which the petro-tympanic bones of Jnia differ from those of
Platanista is their loose connexion with the rest of the cranium; for they are only
attached by ligament, as in Delphinus, and not locked in their place by a process of
the mastoid. In general form the tympanic bulle resemble those of Delphinus, 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’ 1-1".

The mandible presents a remarkable miniature resemblance to that of a Cachalot. It
differs from the mandible of all the true Delphinide 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 than 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 Gervais.
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 been destroyed.

+ After noticing that in certain Delphinoids the aperture left between the hinder edge of the alisphe-
noid, the exoccipital, basioccipital, and basisphenoid is exceedingly small, so that the tympano-periotic is
still more shut out from the cranial cavity than in Balena, Professor Huxley remarks that “in Platanista
the aperture is large, and the periotic appears in the interior of the cranial cavity 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 note
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 fully an inch in length. Whether 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 yary, even in most closely allied; forms, .

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLII. 95

marked rugosity of the surface 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 gradually
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 implanted by large and generally somewhat twisted 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 surface.

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

Eega has 28-28 109, and also two minute rudimentary teeth in the gum behind the
ig 26—27 yi &

last in the left maxilla. In the smaller skull from the same place there are
= 110. In the skull in the Paris Museum, brought by d’Orbigny, there are,

; =| 3888 : : :
according to Gervais, 3;—3,=132; but in the type specimen in the same museum, taken

from Lisbon, the number is given by de Blainville as —— 104. In the Berlin skull the

teeth are a =131 +. Von Martius in his diagnosis of the species gives S114.
aS

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

Antero-posterior diameter of the basihyal .............-.0..04 1-0
MitaSVeETSe CAMEL ese bila sorceress se sielseler smectite 1:3
Length of thyrohyal ............:seeeseeeeeeeeeeeee essen eeseeeees 2-0
Greatest DLEAMGI. 9. .cecece tice ries sete cin sels sirelniselnisleiele = else mieleciels 0:6
Distance between the outer extremities of the thyrohyals... 3-4
Length of stylohyal ...........:essseeeeseeeeenesereeeeeseaeeeeanees raf
Greatest thiGkMess) conse sactiest desi scecteaaselsleieclels|s\«\ss/s 0-4

The spinal column (Pl. XXV. figs. 1 & 2) appears complete to the end of the tail, and
-onsists of but 41 vertebra, the smallest number known in any Cetaceanf. 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 Orca and I'seudorca. + Peters, in a letter.
+ As the bones had been separated from each other adncleaned at the time that they came into my hands for

VOL. VI.—PART III. P

96 MR. 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 vertebre 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°53" long, or ;§35 of the whole column. In a common Por-
poise, measured for the purpose of comparison, it is but ~3G5. All the vertebre are
distinct, as in Platanista, Beluga, and Monodon alone among toothed Whales.

The atlas (Pl. X XVII. 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 the 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 Jnia, 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 which 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 which the
skeleton was prepared render this, at the least, extremely improbable. When it arrived in this country the
yertebre were all united by their natural ligaments. Unfortunately they were not counted when in this state.

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLIL. 97

The remaining five cervical 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 cervical vertebre 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 backwards. 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 own 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 have
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 Beluga similar rings are formed by the transverse processes of this vertebra, also in
the Platanista described by Eschricht, 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 them. 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 ct
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 (Halicore) agrees with Jnia in this respect, as well as in many other of
the characters of the neck-vertebre.

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 lamine 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 vertebre measure in length when placed in close contact 12:5",
Their bodies increase at first rapidly, then more gradually in length—the first mea-

P2

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

suring ‘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 laminze 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-
terior 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 vertebre
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 Delphinide the anterior ribs are
articulated by their tubercle to a well-developed transverse 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 point 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 vertebree. This arrangement, departing con-
siderably from that found in the ordinary mammal, occurs in Delphinus, Phocena, Orca,
Globiocephalus, Beluga, Monodon, and their immediate allies—in fact, in all the Del-
phinide 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 GEOFFRENSIS AND PONTOPORIA BLAINVILLII. 99

bady 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-
bre coexisting with it, as in the cervical region: or, to explain the case in other words,
the anterior ribs in both have an upper and a lower connexion with the vertebre ; 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 Jnia 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 vertebre have as usual a tolerably well-developed, thick and rounded transverse
process, springing from the arch at the junction of the pedicle with the lamina, 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 vertebrz, 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 vertebrie, 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 Delphinidw). Uereafter 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
yertebra 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 direction 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 from the
first thoracic vertebra with its doubly attached rib, to the last with its singly attached

100 MR. W. H. FLOWER 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 Delphinide 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 Del-
phinide 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 Jnia.

The remaining vertebre (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, vertebrae, 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 vertebre the first (always very small) pair of hema-
pophyses was attached. I think, however, that there can be little doubt that the
fourth of the vertebree 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 vertebre at only three, Jnia presents

* As a uniform system of nomenclature in enumerating the vertebrae of Cetacea is very desirable, it is to be
regretted that Eschricht 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 vertebra the one immediately preceding the first
chevron 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 vertebra in the Cetacea commonly called lumbar to the region of the tail,—a view further strengthened
by the fact that, in the ordinary mammals, the chevron bones, when present, begin generally not on the first, 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. Itis 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.

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

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLII. 101

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

The three lumbar vertebree 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 lamine 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-vertebre 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 chevron 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 lamine 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-
vertebrae of the Cetacea, occur first in 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 their characters at the point
where they enter the laterally expanded part of the tail and 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 vertebre 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 different 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 Jnia 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 (Pl. XXV. fig. 2). These bones agree in general characters
with the processes of the vertebree 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 (Pl. XXVII. fig. 2), the last being well developed
and articulating with the transverse processes of the corresponding vertebre. 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-

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

iret 2 ete cc. 37 ight higy scat om oee 6-9
SECOn Ae meen a dns o7 INN thgent ac ene 6:7
Mathis es econ nee eReme aes 6-9 ently cen tat coun 6:5
Blount Wiese acoso eens 108 Bileventhia-eeeseeesce 6:4
it Nees sete ecs ate track 7:3 Twelfth... sc: csaseesdeest 59
Sixteen scat esees oa 7-2 hinfeenthweesssseesses 5:3
Seventies tcc tisscicesens T1

The costal cartilages, as in Platanista and all the Physeteridz, 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 are to be seen
on this bone, which, if confirmed, would be most exceptional among Cetacea, and be
another feature of resemblance with the Sirenia.

The sternum (PI. X XVII. 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 differs
from this, however, in its greater solidity, especially towards the anterior part, and in
possessing two strong triangular processes (4) 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, with smooth, rounded edges. On each side of this are
two thick conical processes (a), projecting directly forwards, 7" apart at their ends. As
these have dried cartilage both on their tips and imner 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 (6), and backwards and upwards into a thick irregular 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 the 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 perforation, 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 (d). There are no other indica-

VOL. V1.—PART III. Q

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

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

In Platanista, 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 Jnia. 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
directed 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 Phocena entirely wants these processes ;
otherwise it presents some resemblance to that of Jia in its breadth, flatness, and in
consisting of a single piece.

The pectoral limbs of Jnia are described by d’Orbigny as “larges, longues, et
obtuses ;” and the present skeleton fully corroborates this account.

The scapula (Plate XXYV. fig. 3) does not present that*singularly aberrant character
which is one of the most peculiar features of the skeleton of Platanista, 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 coste (of which the former is
slightly the longer) are much hollowed out, so that the lower half of the bone is
narrower 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 .

“GOH, Oona goose soeosabanodehooeuocs~snocacnuee beadss0eu00") Donen 37
Extreme: breadth (ics sists... cw Ae eaten oemlaeaet aanieeee ieee 48
Breadth of body at root of acromion process ............-...+5 1-2
Length .ofiacrenuion. su... ....ite Beet seen enemas. at 17
Length of coracoid: process: .2:.2<-thcesacesscessbecton> concn 1:3
Length of plenord? fossa. 1252522225, seek ameenboeeitaey s+ aba wees: 1:2
Breadth ‘of glenoid fossa: wap: . Lostic dh, eee aeabh ewes See ksess 0-9

T'lie 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

INIA GEOFFRENSIS AND PONTOPORIA 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 very 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.

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Whidthratenard Glee. cweed cae. s at een eee Renee nat
Wadithtstlowenend). setae cause det oowele std ies Smead ness 16
Mena thyoh TAadius dnwcasik scsalsecthowieiroa thor sions amnidangeceesee 2°5
aWridthaatimiddl en 23th sae antes are ead eatise eee yoann 1:2
Wadithtatwllower em diet set co.ce aucun casceratoeda nee. 1:4
Mensthyofulnaiedsts spackqustageee seston madayseqetesnadagetk 1:9
\Vool ce hemaibts Glew samen searmahatcear Renato cn obmosodberc. 1:0
Wirdtheattl ower end) ) vcccaeanssacaee sac scekcetehe reat eaeed 1:6

The carpal region is large, and composed in the present specimen in great measure
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, RVai2 eV e.
The individual phalanges are thus not numerous; but they are long in proportion to
their breadth.

From the humerus downwards the pectoral limb of Jnia presents considerable re-

semblance to that of Platanista, both agreeing in the great length of the humerus as
Q 2

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

compared with the forearm, and in the absence of the olecranon process. In the
carpus, to judge by Eschricht’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 Jnia 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 Fréminville, an officer m 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 Delphinus Blainvillei; also in ‘l'Institut,’ 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 “ Mammiféres” of d’Orbigny’s ‘ Voyage
en Amérique Méridionale,’ published in 1847, but the introduction to which bears the
date of December 1846, redescribed and figured the skull (plate 23), pointing out
that its peculiarities were sufficient to entitle it to rank as a subgenus, for which the
name of Stenodelphis 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 Fréminvyille.
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 Fréminville*, it
has unfortunately been treated as a certainty in most systematic works}, and thus Pon-
toporia, 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

* «)'aprés un renseignement favori par M. de Fréminville, le Dauphin dont provient ce crane, est long de
quatre pieds, et il est blanc, avec une bande dorsale noire.”

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

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLII. 107

Inia was written, a second skull of Pontoporia, also from the mouth of the Rio 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 skull of Jnia a com-
parison with this nearly allied form.

The skull (Pl. XXVIII.) is that of a perfectly adult animal. ‘he 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 points. The rostrum is curved downwards towards the extremity, much
more so than in the Paris specimen; this is probably the effect of age, as a similar
change takes place in Jnia and some other Dolphins. The mandible partakes also
of this curve. The small, rounded and depressed cranium, and very long, narrow and
compressed beak, give a remarkable appearance to this skull, reminding one, as Gervais
remarks, of the head of a scolopacine bird.

The principal dimensions are :—

pe Rente len POD i isaajote casa naa nepethe aeeuls <a ieadt a dacere 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
IDGINES conusesnsqno eoosas snp dondbeananengsaonsooccgcedonseatcc 15°65
Greatest breadth (across zygomatic process of squamosals) 4:8
Breadth of foramen magnum ...............2ssesseeneeserees iol
Breadth of occipital condyles .................cesesseseeeeeees 2°4
Breadth across antorbital processes of frontals............... 2°6
iBreadthyot rostrum) at Das We op) onan selec nabs eermcpanseie 1:8
Breadth of rostrum at middle ...............sceeeeetereneeeees 0-6
Miamadables Lemp thy esas ctr alele elena seeieleiebtriele lerl-cte is 13:
Mandible, length of symphysis ..........0.++e+eeeeeeneseeee es 8-0
Greatest breadth posteriorly ..............ceseeeeeeeeenee es aso p eto
Height at the coronoid process............-+2:seeeeseeteeee eens 2:3

The supraoccipital is broader and shorter than in Jnia, terminating in front by a much
more open angle, and on each side in alow ridge, coming in close contact with the broad
posterior extremities of the suprafrontal plates of the maxillaries. 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 Jnia 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 MR. 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 Jnia,
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 off.

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, -7" 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 Jnia. 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 hinder ends of
the maxillaries are broader and flatter than in Jnia; but in front of the nostrils they
are much more contracted, and above the orbits have.a small but distinct. longitudinal
crest, °3" 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
isno 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 Jnia. The rostrum is considerably
longer and narrower in proportion to the size of the cranium than in Inia.

The palate-bones resemble those of Znia 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 Jnia rather than that of Platanista.

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLII. 109

The mandible resembles that of both Jnia and Platanista, 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 the
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 :
oe ?, All have broad fangs, much compressed laterally, surmounted by a
crown, 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
direction, 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, which 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 :—

IE@NVETH A (OME TeWBYES | Gsenonansensder corny sernaoceneenscracsepen sc ‘15
Menethy ols Gr Owls i.piczn ah) savenidcvinn ap goinedhiyssaeanaeees “24
Antero-posterior breadth of cingulum .................. 7
Transverse breadth of cingulum............:.::0seseeeeee pie
Antero-posterior breadth of apical part at middle... *05
biransyerse bread by «cereale pesaa saiidoll= Gee onicleree rei cciee 10

This peculiar form of the teeth, which distinguishes Pontoporia from all the ordinary
Dolphins, and affords another evidence of its affinity with Znia, has not been observed
in the Paris specimen. Gervais’s description is as follows :—* Les dents * * * * sont
petites, longues de 5 ou 6 millimétres au plus, toutes plus ou moins aigués, et au
nombre de 53 ou 54 supérieurement, ainsi quinférieurement. Les postérieures sont un
peu moins aigués que les autres, et leur partie terminale est un peu recourbée.”

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.

- III. On the Systematic Position of Inia and Pontoporia in the order Cetacea.

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

110 —; MR. W. H. FLOWER 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 comparisons, between
objects essentially dissimilar, with which anatomical treatises on the Cetacea are too
often encumbered.

Before proceeding further with this part of the subject, I would remark, in passing, that
several resemblances pointed out above between the skeleton of this Cetacean and
that of the Sirenia, according singularly with d’Orbigny’s observations upon its external
form 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 from 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. Mysracoceti or BaLaNnoipEA. Teeth neyer functionally developed, but always disap-
pearing before the close of intra-uterine life. Upper jaw provided with plates of
baleen. Olfactory organ distinctly developed. External respiratory 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 vertebre ; their capitular processes, when
present, not articulating immediately with the bodies of the vertebre.

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

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLII. 11

2. Opontocetr or DeLpHinomEA. 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-sternal
bones. Many of the ribs with capitular processes developed, and articulating
with the bodies of the vertebre.

It is not necessary to pursue further the arrangement of the J/ystacoceti, 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 from which to commence the formation of a
natural division of the Toothed Whales, 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 entirely unossified
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 in 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 Royal
College of Surgeons for the present year (sce ‘ Lancet,’ 1866, vol. i. p. 381).

VoL. VI.—PART III. R

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

these-may also be added Platanista and Inia. Here, then, isa 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 vertebra 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 vertebree and
rib-attachments. Whether Aogia 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, /nia 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 Inia on thee 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 vary
widely from each other, one diverging towards the Physeteridz, the other towards the
Delphinide, 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 fluviatile f.

* Dr. Gray in the “ Zoology of the Voyage of the Erebus and Terror” placed Inia and Pontoporia in one
section at the end of the family Delphinide, following immediately upon Platanista. In his recently published
Catalogue, Platanista constitutes the fourth family (Platanistide) of the Cetacea, following the Catodontide ;
Inia forms a separate (the fifth) family, Iniide ; and Pontoporia commences the sixth family (Delphinide),
comprising all the remaining Dolphins except the Globiocephalide and the Ziphiide.

Gervais (Hist. Nat. des Mammiféres, 1855) unites Platanista, Inia, and Stenodelphis (Pontoporia) to form
one of the five tribes (Platanistins, Delphinins, Orcins, Monodontins, and Phocénins) into which the family

Delphinidés is divided. The primary divisions of the order or families are :—Physeteridés, Ziphiidés, Delphinidés,
and Balénidés.

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

INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLII. 113

I will now endeavour to formularize the distinctive characters of these three primary
groups of the Opontoceti, giving them the rank of families.

1. Puysererip&, Costal cartilages not ossified. The hinder ribs losing their tuber-

cular and retaining their capitular articulation with the vertebrae. The greater
number of the cervical vertebre ankylosed together. Pterygoid bones thick,
produced backwards, meeting in the middle line, and not involuted to form
the outer wall of the postpalatine air-sinus. Symphysis of mandible of moderate
or excessive length. No 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.

Il. Puaranistip2. Costal cartilages not ossified. The tubercular and capitular articu-

IIL.

lations of the ribs blending together posteriorly. Cervical vertebra 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 entire ramus. Orbit very small. Lachrymal bones not distinct from the
jugal. Pectoral limbs large. Dorsal fin rudimentary*.

De.PHinip#. Costal cartilages firmly ossified. Posterior ribs losing their capi-
tular articulation, and only uniting with the transverse processes of the vertebra
by the tubercle. Anterior (2-6) cervical, in most, ankylosed together. Ptery-
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. Symphysis of mandible short or moderate, never
exceeding one-third the length of the ramus. Bones of the skull not raised into
a distinct crest behind the anterior nares. Orbit of moderate size, Lachrymal
bone not distinct from the jugal. Pectoral limbs varying much in form and size.
Dorsal fin usually present.

I. The Physeteride appear to constitute a very natural group}. This may, however,
be divided into two well-marked subfamilies :-—

that Mr. Darwin 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 Pontoporia.

+ Van Beneden insists strongly upon the close affinity of Physeter with the Ziphioids : he says, “ Comme on-le
yoit, les Cachalots sont pour nous des Ziphioides yézitables, portant une rangée de dents fortes et espacées sur
chaque branche de maxillaire ” (Mém, sur une Nouv. Espéce de Ziphius, Mém. de l’Acad. Royale de Belgique,

t. xvi. 1863).

R2

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

1. Physeterine, characterized by the numerous teeth in the lower jaw, and having
no distinct lachrymal bone, including the genera Physeter and Kogia (Gray)*.

2. Ziphiine, 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, Micropteron, Dioplodon, and several extinct
forms.

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

1. Platanistine. 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. Iniine. 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 structure; but as its cranial and dental characters accord most nearly with
those of Inia, it may be placed provisionally in the same subfamily.

Ill. Although the Delphinide 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 groups. 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-
tebre. As these two animals are in almost every part of their skeleton nearly identical,
even to the number of the vertebre 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 Platanistide. 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. Phocena and Neomeris stand by
themselves in the form of their teeth and certain cranial characters. Oyca is distin-
guished from all the others by its excessively broad manus, and Globiocephalus by the
extreme length and narrowness of the same member. Delphinus 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 Huphysetes (Wall. Descr. New Sperm Whale,
&e., 1851); but Gray’s name (Zool. Erebus and Terror, 1846) clearly has the priority.

INIA GEOFFRENSIS AND PONTOPORIA 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, Lagenorhynchus, &c.

In the following tabular view of the arrangement of the Cetacea, many of the genera
lately formed, chiefly by subdivision of the old genus Delphinus, 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.

Suborders. Families. Subfamilies. Genera.

Balenide ............ Balenine@ ......-.+++005 { paeee
I, Mysracoceti* ubalena.
or

Balenorden: Megaptering ...........- Megaptera.

Physalus.
Sibbaldius.

Baleenoptera.

| Balenopteride .........
Balenopterin®.......--+

Physeter.
Kogia.

Berardius.
Ziphius,

| Dioplodon.

| Micropteron.

Physeteride .......--- i Hyperoodon.

or Platanistide ........-+-

II. Oponrocetrt Platanistine.........-+-- Platanista.
Delphinoidea.

Pontoporia ?
Inia.

Monodon.
Beluga.

pan Phoceena.
| Delphinide ........+-.-- Peonsah
Grampus.
Orea.
Delphinine? ......+++- Pseudorca.
Lagenorhynchus.
Delphinus.
Delphinapterus.
\_ Globiocephalus.

* pvarak, xyros ; equivalent to the German “ Barten-Walle.” + 6ltovs, Knros.

116 OSTEOLOGY OF INIA GEOFFRENSIS AND PONTOPORIA BLAINVILLI1.

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

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

a. Anterior process.

6. Lateral process.

ce. Surface for attachment of cartilage of first rib.

d. Surface for attachment of cartilage of second rib.
Fig. 4. Internal surface of sternum.
F

=
isi}

. 6. External surface of sternum.

=
gag

PLATE XXVIII.

Skull of adult Pontoporia biainvillii. All the figures (except fig. 5) half the
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 a end of the
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V. On a Raptorial Bird transmitted by Mr. Axversson from Damara Land.
By J. H. Guryey, £.Z.8.

Read November 14th, 1865.
[Puate XXIX.]

T 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 our knowledge
of the ornithology of that part of South-western Africa.

Mr. Andersson remarks, with reference to the present specimen, which was procured
at Objimbinque, Damara Land, on the 10th of March last, “I have only obtained 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 :—

“Trides bright lemon-yellow; extremities of mandibles black ; basal parts and gape
bluish lead-colour; tarsi and toes bluish white; claws bluish black.

“Entire length 1 ft. 632; im.; length of wings when folded 1 ft. 1}3 in.; length of
tarsus 243;in.; length of middle toe 2;in.; length of tail 735,in.; length of
bill from corner of gape to the tip of the mandible, straight, 15% 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
line 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 bird, including 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 including the upper and under wing-
coverts, as also on the upper and under tail-coverts; the upper surface of the tail bears
five transverse bars of a pale brown, 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-fourths of its length by a brown medial line, starting

118 MR. J. H. GURNEY ON A RAPTORIAL BIRD.

from the angle of the lower mandible, and extending for about 3 inches in a straight line
towards the stemum. ‘The feathers of the breast and sides are of a mingled brown and
white, the latter predominating in 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 observed 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 bird; 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 to 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 their character, and the scales with which they are
covered are (with the exception of those covering the last jomt of each toe) remarkably
small. The middle toe, which is considerably elongated, has a prominent 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 Machwrhamphus alcinus of
Westerman, the type of which is in the Museum at Leyden. The present specimen
has been added to the collection in the Norwich Museum.

* Proc, Zool. Soc. 1866, p. 324, + Westerm. Bijd. t. d. Dierk. i. p. 29.

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VI. On some Fossil Birds from the Zebbug Cave, Malta.
By W. K. Parker, F.RS., F.Z.8., &e.

Read and received for publication Dec. 12th, 1865.
[PuaTe XXX. |

Five years have elapsed since I first examined numerous bony remains from the Zebbug
Cave, the “lamellirostral” nature of which was apparent to Dr. Falconer and myself
from the first. I transmitted a list of them to that lamented paleontologist 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
their 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 the
tip of the beak to the end of the tail, 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-
third larger than my specimen of the tame kind, 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 Bernicle, such as the Bernicla brenta.

On June the 10th, 1861 (the next summer), I received, through Professor Rupert
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 tibia of the largest kind, which is 64 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 paleontologist whose loss we have so lately
suffered, are as follows :-—

VOL. VI.—PART III. 8

120

Skull (posterior fragment)
Ribs (upper part)

Ulna (middle).

Femur (various parts)

Tibia (various parts) .
Tarso-metatarse (various parts)
Phalanges (perfect) .

MR. PARKER ON SOME FOSSIL BIRDS

2 specimens.

Of the smaller kind of Swan (Cygnus musicus?) there are—

Cervical vertebra (2nd or 3rd)
Sternum (anterior part)
Scapula (proximal part)
Humeri (various parts) .
Ulna (various parts) .

Radius (various parts)
Metacarpus (various parts) sas
Phalanx (proximal, perfect) . . . . 4
Phalanx (distal, perfect)

Sacrum (various) .

Femur (shaft-part)

Tibia (various) er
Tarso-metatarse (various ae dc
Phalanges (perfect) .

-Of the small Goose-bones (Bernicla

Coracoid (head) .

Radius (distal and middle partidual
Ulna (middle) : eit ma
Metacarpus (almost neha
Femur (nearly perfect) .
Tibia (lower end)

P) there are—

3 ”
1 specimen.
. 12 specimens.
. 720 53
20,
3 ”
2 ”
1 specimen.
7 specimens.
18 ”
° 7 »
ors ,,
5 ”
1 specimen.
2 specimens.
3 ”
A ass
3 ”
4 ”
5 ”
1 specimen.
2 specimens.
1 specimen.
1 ”
1 RS
Lag ieee

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 corroborated the conclusion I came to as to the skull be-

FROM 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. falconeri, as compared with
those of the large female C. olor :—

Middle thoraci sib— eee ae

a. Acrosstheneck . ... O Dad ee meee rt)

b. Width of outeredge. . . 0O ON hale aa 4
Ulna—

Diameter of shaft. . . . . O Olt ais hoee RO 7
Radius—

Diameter of shaft... ...;.... 0 Sic eseaasoe AO 4
Femur—

Across head and trochanter . 1 1 1 5

Width of middle of shaft . . 0 54 0 8

Width across lower condyles . 1 0 1 oy
Tibia—

Fore-and-aft width of head 1 5 1 9

Thickness of head 0 + 1 2

Width of shaft . 0 + 0 8

Width across lower condyles 0 11 1 4%
Tarso-metatarse—

Extreme length 4 3 5 3

Width across head 1 0 1 24

Width across shaft 0 5 0 6

Width across condyles 1 0 1 4
Phalanx (proximal, middle)—

demathe, | come 78s. apt DST EES 2 Da) Sek i: (Gs

Thickness of proximalend. . 0 Baty Soro eo 74

Thickness of shaft . . .. 0 Bo wel ot, 4h

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 of females; but there are bones still larger in the col-

lection, most likely those of male birds.
s2

122 MR. PARKER ON SOME FOSSIL BIRDS

The coarseness of these bones is well shown when the diploé is displayed (see Pl. 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 (Pl. XXX. figs. 4 & 5).—The diameter of the ulna, as seen in fig 5, is as 7
to 5 compared with fig. 5a; and the strength of the shaft is well shown 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 (Pl. 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
bird: 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 (Pl. 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 strength 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 (Pl. XXX. figs. 16, 17, 18 & 19)—The length of this right shank
is seen to be greater 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), which does not ap-
pear in fig. 19: this mistaken 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 perfection of the figures exonerates me
from detailed description.

Phalanges (Pl. XXX. figs. 20, 21 & 22)—There are only three phalanges which I
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. 20a, 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

FROM 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 bird
had its wings of the full relative size: the immense ulna shows this (see Pl. XXX. figs. 1,
4 & 5).

As the 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 “ Lamellirostres,”
viz. the Palamedea—that in which the lamelle of the beak are arrested in their growth,
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 frag-
ment is well shown in Pl. 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. 3 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. musicus), perhaps C. bewickii, or, it may be, some species nearly allied to these. At any
rate it is interesting to find that C. musicus is still to be found in lands bordering the
Mediterranean, the Rev. 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 in the species of Swans is so great that I feel it to be
unnecessary to describe 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 MR. PARKER ON SOME FOSSIL BIRDS FROM THE ZEBBUG CAVE, MALTA.

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

DESCRIPTION OF PLATE XXX.

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

Fig. 1, Anterior fragment of sternum of Cygnus musicus (2); upper view.
Fig. 2. Anterior fragment of sternum of C. musicus(?); side view.

Fig. 3. Anterior fragment of sternum of C. musicus(?); front view.

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

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

Fig. 5a. 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 view.

9. Femur (left) of C. falconeri; upper view.
Figs. 6a-9a. 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 view of distal end.
Fig. 13a. 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. 20a. Phalanx (proximal, of middle toe) of C. olor; upper view.
Fig. 21. Phalanx of C. falconeri; side view.
Fig. 21a. Phalanx of C. olor; side view.
Figs. 22, 23. Phalanx of C. falconeri; end views.
Figs. 22a, 23a. Phalanx of C. olor; end views.

* The circular hole in this view is of artificial origin.

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1.3 CYGNUS MUSICUS ?

125.74

VII. Synopsis of the species of recent Crocodilians or Emydosaurians, chiefly founded on
the specimens in the British Museum and the Royal College of Surgeons. By Dy.
Joun Epwarp Gray, F.R.S., V.P.ZS., ELS, &e.

Read December 9th, 1862.

[PLates XXXI. to XXXIV.]

THE 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 surrounds 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 Crocodilide or Alligatoride was published in the
‘Annals and Magazine of Natural History’ for 1861 (3rd series, vol. viii.). Since that period
J 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 Royal
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 IY. T

126 DR. J. E. GRAY’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 Alligator, 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 useful to the working naturalist 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 modern art of describing is too long,
often insufferably long, while human life remains as short as ever” (Illust. Zool. 8.
Africa, p. 54).

I know by experience that synoptical papers take far more mental and bodily labour to
prepare than the description of asingle 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

OF RECENT CROCODILIANS. 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 offhand. 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 nomenclature 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 isan 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 my
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. Gavialis gangeticus= Gavialis,
710. Crocodilus cataphractus=Mecistops cataphractus, the type specimen.
711, 712, 714, 716. Crocodilus acutus=Molinia americana, from America.

128 DR. J. E. GRAY’S SYNOPSIS OF THE SPECIES

The chief difficulty in distinguishing the 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 three 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 have 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 paleontologists, as they call themselves, neglecting, or,
at most, only taking the outline of the osteological and zoological characters of the living
species at second hand, and describing the fossil, and often forming genera and species
ona 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. Crocodilus acutus=Oopholis porosus of India.
715. Crocodilus acutus= Crocodilus vulgaris of Africa.
717. Crocodilus vulyaris, much distorted.
718. Crocodilus vulgaris=Bombifrons, perhaps B. siamensis.
719-724, 727, 728. Crocodilus biporcatus= Oopholis porosus.
725. Crocodilus biporcatus= Crocodilus vulgaris.
726. Crocodilus biporcatus=Bombifrons indicus.
750, 751. Crocodilus rhombifer, from Bengal= Bombifrons indicus.
752. Crocodilus palustris ?=Bombifrons indicus.
760-762. Alligator lucius= Alligator mississippiensis.
764, Alligator niger=Jacure nigra.
1 Dr. J. E. Gray ‘On the Change of Form of the Heads of Crocodiles,” Transactions of the Sections in ‘ Report
of the British Association of Science,’ Cambridge, 1862, p. 109.

OF RECENT CROCODILIANS. 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 skullis 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 ‘ Ostéographie,’ devotes five folio plates to the osteology and
dentition of recent Crocodiles, giving details of Crocodilus biporcatus, C. lucius, C.
vulgaris, C. schlegelii, C. longirostris, C. rhombifer, and C. sclerops. These plates
were prepared to accompany an essay that M. de Blainville was preparing for the
* Mémoires de l’Académie des Sciences 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
of Jacare 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 osteological 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, Blainyille, Gray, Ann. Phil.x.195, 1825; Cat. Tortoises & Crocodiles Brit. Mus. 38,1844.
Crocodilia, Huxley, Journ. Proc. Linn. Soe. Zool. iv. p. 1.

The Emydosaurians or Crocodilians may be divided into three families :—

A. The cervical and dorsal plates forming one dorsal shield.

I. Gaviatip#. The large front teeth and the canines in the lower jaw fit, into notches
in the margin of the upper jaw.

130 DR. J. E. GRAYS SYNOPSIS OF THE SPECIES

B. The cervical shield forms a small group, which is separate from the dorsal shield.
II. Crocoprtips. 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. Axuicatorip. 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 AND AUSTRALASIA, AMERICA.
Fam. Gavialide.

Gavialis.
Tomistoma.

Fam. Crocodilide.

Crocodilus. Oopholis.
Bombifrons. Palinia.
Halcrosia. Molinia.
Mecistops.
Fam. Alligatoride.

Alligator.
Caiman.
Jacare.

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 cataphractus. 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 porosus (or
Crocodilus biporcatus of Cuvier), which is found only in the estuaries at the mouths of
the large rivers; 2. The Muggar’ (Bombifrons indicus); and 3. The Garial (or Ghurrial),

‘ See Dr. J. E. Gray “ On the Crocodiles 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 Crocodile is Coombeer. The Rapacious Shark is called the Muggar ;
and by reflection this name is also sometimes given to the Crocodile, because it is a rapacious animal.

OF RECENT 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 Australia. Governor Grey met
with them in the north-west.’—Blyth, Rep. Austral. Vert. in Mus. A.S. C.

If by “ both the Gangetic species of Crocodile” Mr. Blyth means the estuarine Croco-
dile (Oopholis porosus) and the Coombeer or Muggar (Bombifrons indicus), no example
of the latter animals from either Java, Timor, or Australia has occurred to me, and the
animal figured as Crocodilus raninus vy Dr. Salomon Muller is certainly Oopholis porosus ;
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. Duméril and Bibron (Erp. Gén. 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 occurrence of Crocodiles on the north
coast of Australia.

The estuarine Oopholis porosus 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 rhombifer,
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 incorrectness of the assertion of MM. Duméril and Bibron that the
Crocodiles of America are confined to the islands of that continent (Erp. Gén. 25, 47)'.
«' In the ‘ Gentleman’s Magazine’ for August 1866 appears an article, entitled ‘« Notes on a Young 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. William Phillips, who said that it was found lying
dead in a gutter in his farmyard, evidently but lately killed; its bowels protruded from a wound in the belly.

132 DR. J. E. GRAY’S SYNOPSIS OF THE SPECIES

Family I. GAVIALIDZ.

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 front large
teeth fitting into a notch in the front, the canines into a notch on the sides of the
front of the upper jaw. ‘The jaws elongate, slender.

Crocodilide (part.), Gray, Ann. Philos. x. 195, 1825.
Crocodilide § *, Gray, Cat. Tortoises & Crocod. B.M. 36.
Gavialide, Huxley, Journ. Proc. Linn, Soe. Zool. iv. p. 16, 1859.

Synopsis of Genera.

GaviALis. Beak elongate, linear, end swollen. The lateral teeth oblique, not received
into pits.

Tomistoma. Beak conical, thick at the back, the lateral teeth erect, received into pits
between the teeth.

1. GAVIALIS.

27-27 2828

25—25 OT 26-96"
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.

Beak of skull linear, end dilated from the enlarged nostrils. ‘Teeth

Gavial, Oppel. Le gavial, Cuvier.

Gavialis, Merrem, Gray, Aun. Phil. x. 195, 1825; Cat. Tortoises, &c., B. M. 36, 57, 1844. Geoff.
Mém. Mus. xii. Huxley, Proc. Linn. Soe. Zool. iv. p. 20, 1859.

Gavialia, Fleming, Phil. Zool.

Ramphostoma, Wagler, Syst. Amph. 441. Rhamphognathus, Vogt, Zool. Brief. 11. 289.

1. Gaviauis Ganeeticus. (The Garial or Nakoo.)

Narrow-beaked Crocodile, Kdw. Phil. Trans. xlix. 639, t. 19.
Le gavial, Lacép. Q. O. 1235, t. 15. Faugas, Mont. S. P. 235, t. 8. f. 46, 47.
Lacerta gangetica, Gmelin, 8. N.i. 1057. Shaw, Zool. i. 197, t. 60.

The men said it ran out of the stack of wood, they killed it, but they could easily get him another ; he offered
a guinea for another specimen, dead or alive; but the reward was never claimed.

An account of the discovery appeared in the ‘ Field Newspaper’ for 1861 or 1862; and another, with a
figure of the specimen, was published in Hardwicke’s ‘ Science Gossip,’ Jan. 1, 1867, p. 7, figs. 1 & 2. Dr.
Vesalius Pettigrew and Mr. Frank Buckland thonght 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 umbilicus. The figure
shows too long and slender a beak fcr a young specimen of any Crocodile I have seen.

OF RECENT CROCODILIANS. 133

Crocodilus longirostris, Schneid. Amph. 160. Daudin, Rept. 4293. Blainv. Ostéog. Crocod. t. 2. f. 4,
t. 3. f. 6, t. 4. f. C, t. 5. f. 5.

Crocodilus arctirostris, Daud. Rept. ii. 393.

Crocodilus tenuirostris, Cuvier, Ann. Mus. x.t.1. Tiedem. Amph. t.15. Wagler, Syst. t.7.f.111.
Merrem, Tent. 38.

Gavialis gangeticus, Geoff. Mém. Mus. xii. Gray, Syn. Rept. 86; Cat. Tortoises &c. B. M. 57.
Dum. & Bib. Erp. Gén. 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. Geoff. Mém. Mus. H. N.
ie HS

Gavialis longirostris, Merrem, Amph. 37.

Gavialis tenuirostris, Merrem, Amph. 38. Guérin, Icon. R. Anim. t. 2. f. 3.

Ramphostoma tenuirostre, Wagler, Nat. Syst. Amph. 141, t. 8. f. 3.

Le gavial, Lacép. H. N. Q. Ovyip. i. 235, t. 15.

Gavial, Owen, Monogr. Fossil Reptilia of the London Clay, t. 11. 1849 (skeleton).

Hab. Indian rivers. Bengal, Nepal, Malabar.

2. ToMISTOMAs

Beak of the head conical, thick at the base. ‘Teeth = 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 margins not raised.
Gavialis, sp., Miller ; Owen.
Tomistoma, S. Miller, Wiegm. Arch. 1846, i. 122.
Rhynchosuchus, Huxley, Journ. Proc. Linn. Soe. 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. gravesti and Mecistops cataphractus.

Dr. Falconer, when describing the skull of Crocodilus cataphractus, 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 Bornean species, Crocodilus schlegelii, was in fact originally de-
scribed as a new species of Gavial; but the 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 our skull, nor with the figures of the skull in Blainville’s ‘ Ostéographie.’
See also Huxley, Journ. Proc. Linn. Soc. Zool. iv. p. 18.

VOL. VI.—PART IV. U

134 DR. J. E. GRAY’S SYNOPSIS OF THE SPECIES

1. ToMISTOMA SCHLEGELI. (Bornean Gavial.)
Crocodilus gavialis schlegelii, Miller, Naturgesch. Ost. Ind. t. 123. f. 1-5.
Crocodilus schlegelii, Blainv. Ostéog. Crocod. t. 2. f.3; t.5.f.4. Briihl, Skelet. Krok. t. 8. f. 6.
Owen, Fossils of the London Clay, p. 15.

Rhynchosuchus schlegelii, Huxley, Proc. Linn. Soe. iv. (1859) p. 17; Ann. & Mag. Nat. Hist. 1859.
Mecistops journei, Gray, Cat. Tortoises &c. B. M. 38, not synonyma.

Hab. Australasia, Borneo (Miiller, Brit. Mus.).

The two figures of the skull in Miiller 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 there is a young specimen 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. CROCODILID.

The cervical plates forming a distinct shield, separate from the doysal 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.

The 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.

Crocodilide §**, Gray, Cat. Tortoises &c. B. M. 36, 1844.
Crocodilide, Huxley, Proc. Linn. Soc. Zool. iv. 5.
Crocodilus, Cuvier ; Gray, Ann. 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 Mecistops 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 the 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 CROCODILIANS. 155

The head of the Crocodile first increases 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, Tomé-
stoma schlegelii, as is found in Miiller 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 caught.

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
this 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 they are produced beyond it, and form a bony septum between the nostrils.
In the long and 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. Halcrosva.

2. Nasal bone produced, and dividing the edges of the nostril. Oopholis, Crocodilus,
Molinia (americana), Bombifrons, Palinia.

3. Nasal bone not reaching the nostril. Molinia (intermedia), Mecistops.

The intermaxillary bone in Bombifrons and Palinia 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.

Ue

a)

136 DR. J. E. GRAY’S SYNOPSIS OF THE SPECIES

The palatal bone in all the genera is truncated or rounded in front, except in
Mecistops, where it is narrow, short, and acute in front.

The skulls of the genera Bombifrons, Oopholis, and Molinia are easily distinguished in
the young state,—the face of Oopholis being much longer and narrower than that of
Bombifrons, and that of Molinia is longer and narrower than that of Oopholis. The
measurements following are for three skulls which appear to be from animals nearly of the
state of growth, same in inches and lines :—

Bombifrons. Oopholis. Molinia.
in. lines. in. _ lines, in, _ lines.
Length of the skull, entire ...........-.- .4 8 5 8 6
Length of face to front of orbit.............. 2 8 3 6 4 4
Length of forehead to front of orbit.......... 2 0 2 1 2 4
Length Ge palate from condyle to front end of } Th 3 4 3 10
IE IY Age MGtomcdato sobaco otdrred
Length of middle suture of maxilla .......... 1 2 1 13 1 Uf
Length of middle suture of intermaxilla ...... 0 9 1 3 1 6
Widthyat woceipit ie eis: wichy- acess cise’ = cisteleiteiny= 2 6 2 5 2 103
Width at hinder contraction of beak.......... 1 6 1 4 if 43
Wadthvatmoteht ¢/-crtassreis ett cdey-retetartarcrecateys 0 9 0 9 0 9

The dorsal scales present considerable variations in different specimens from the same
locality ; but, allowing for such variations, the genera :uay 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 Mecistops.

3. The dorsal scales quadrilateral, as broad as long; the vertebral series scarcely
keeled, the lateral series irregular and keeled. Halcrosia 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 sia longitudinal
series. Estuarine Crocodiles.
1. Oopuouis. Asia and North Australia.

B. Nuchal plates four, in a transverse series. Dorsal plates as broad as long,
square. ¥Fluviatile Crocodiles.

OF RECENT CROCODILIANS. LS

a. Intermasillary bone truncated behind, with a nearly straight hinder
edge. Face broad, oblong.
2. Bompirrons. Toes webbed. Legs distinctly fringed. Asia.

3. Pauinta. Toes short, free. Legs with only an indistinct fringe-
America.

b. Intermavillary bone elongate, produced, and truncated behind ;
sutures sloping backwards and converging, then transverse or
sinuous. Toes webbed. Legs fringed.

4. Crocopitus. Face oblong, without any ridge from front of orbit,
forehead flat. Africa.

5. Mortnta. Face elongate, forehead convex, smooth, without any ridge
from orbits. America.

IL. Cervical disk strongly keeled on each side, and nearly continuous with the dorsal
shield. Aberrant Crocodiles.

* Face broad, nasal bone produced into the nostrils. Alligatoroid Crocodiles.
6. Haucrosia. Africa.

** Face very long, slender, nasal bones not reaching the nostrils. Gayialoid
Crocodiles.
7. Mecistops. Africa.

1. The nape with a rhombic disk formed of six plates, which 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 bone
produced, truncated, and converging on the sides. Estuarine or brackish-water
Crocodiles. 2

1. OoPuHoLis.

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 SPECTES

a. The dorsal scales in six longitudinal series; the vertebral ones elongated
like. the others.

1. OopHouis rorosus. (The Saltwater Crocodile.)

Crocodilus porosus, Schn, Amph. 159. Gray,Cat. Tort. & Croc, &c. Brit. Mus. 58; ‘P. Z. 8. 1861, 140.

Crocodilus oopholis, Schnu. Amph. ii. 165. ,

Crocodilus biporcatus, Cuvier, Oss. Foss. v. 65, t. 1. f. 4, 18, 19 (young skulls) ; t. 2. f.8. Miiller
and Schlegel, Verh. t. 3. f. 6 (middle-aged skull). Owen, Cat. Osteol. Mus. Col. Surg. 159,
nos. 719, 723, 724, 727, 728. Huxley, Journ. Proc. Linn. Soc. Zool. iv.11. Blamy. Ostéogr.
Crocod. t. 1, t. 3. f.1,t.4.f. , t. 52.

Crocodilus acutus, Owen, Cat. Osteol. Mus. Col. Surg. 157, no. 718.

Champse fissipes, Wagler, Amph. t. 17.

Crocodilus biporcatus raninus, Miller and Schlegel, Verh. t. 3. f. 7 (aged skull) !

Oopholis porosus, Gray, Ann. & Mag. Nat. Hist. 3rd series, x. 267, 1862.

Hab. Asia and Australia; India, Bengal, and Penang (Hardwicke) ; China (Lindsay) ;

Trincomalee ; Borneo (belcher); Tenasserim coast (Packman); Siam, Cambogia (Mouhot).

Var. australis, Giinther.
Crocodile, Landsborough, Explor. of Australia, i. 70.

Hab. North Australia (Elsey & 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 Mr. 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 cf young and half-grown specimens. %. Miiller and Schlegel
figure two skulls, one under the name of C. biporcatus (f. 6), and the other C. bipor-
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 C. diporcatus raninus (f. 8), appears to be from a specimen of
Crocodilus or Bombifrons siamensis. It certainly is not an Oopholis, from the form 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. biporcatus 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 CROCODILIANS. 139

The British Museum received from the Leyden Museum an adult skull of the
Crocodilus (biporcatus) raninus from Borneo; it is 22 inches long, and agrees in every
respect with the Oopholis 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 outer
series elongate-ovate.

2. OOPHOLIS PONDICHERIANUS. (Pondicherry Crocodile.)
Oopholis pondicherianus, Gray, Ann. & Mag. N. H. 38rd series, x. 268.
Crocodilus pondicerianus, Giinther, Rept. 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 plates four, or rarely two or five, in a cross series. The dorsal plates as
broad as long, in four or six series. Fluviatile or River Crocodiles.

a. The intermaaillary bones truncated 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 webbed. Legs distinctly fringed. Asiatic Crocodiles.

2. BoMBIFRONS.

The premaxillary suture straight, or rather convex forwards. The face oblong;
forehead with 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 nearly

horizontally.

140 DR. J. E. GRAY’S SYNOPSIS OF THE SPECIES

1. Bomprrrons inpicus. (The Muggar.) (Plate XXXL, figs. 1, 2, 3.)

The intermaxillary short, nearly semicircular.
Crocodilus vulgaris, var. indicus, Gray, Syn. Rept. 58, 1831!
Crocodilus dubius, Geoff. Ann. du Mus. xii. 122?
Crocodilus suchus, var. D., Dum. Enc. Méth. Rept. 27.
Crocodilus palustris, Lesson, Bélanger, Voy. 305. Gray, Cat. Tort. & Croc. B. M. 62 (young).

Owen, Cat. Osteol. Mus. Coll. Surg. 164 & 752! Giinther, Rept. B. Ind. t. 8. f. a.

Crocodilus bombifrons, Gray, Cat. Tortoises & Crocodiles &e. B. M. 59, 1844 (adult) !
Crocodilus bombifrons (palustris ?), 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. Dumér. & Bibron, Erp. Gén. iv. 108.
Crocodilus rhombifer, Owen, Cat. Osteol. Mus. Coll. Surg. 164, n. 752! (not Cuvier).
Crocodilus ? Owen, Cat. Osteol. Mus. Col. Surg. 159, n. 726!

Hab. India: Ganges (Dr. Sayer); Madras (Jerdon); Ceylon (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; but

sometimes there are larger and broader shields intermixed in and deranging the series,
and at other times the whole vertebral series is formed of wider shields.

This species has generally been confounded with Oopholis biporcatus 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 Oopholis porosus.

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 243 inches long.

In my Catalogue of the Tortoises and Crocodiles in the British Museum, published
in 1844, I 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 Duméril and
Bibron as an adult of Crocodilus biporeatus, 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 RECENT CROCODILIANS. 141

In the same work I separated the Indian specimen from the common African Croco-
dilus, under the name of Crocodilus palustris of Lesson, and pointed out that it seemed
to be the same as the Crocodilus biporcatus raninus of Miiller and Schegel; and I
described two other very young specimens under the name of Crocodilus trigonops, 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 interlocated 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 generally
spread over the Indian peninsula.

Var. Nose narrow, the intermaxillary bones rather longer and narrower.
Hab. Ceylon (skull, Kelaart).

Fig. 1, and 6. Skull of adult C. bombifrons, Gray, 1847. Presented by Capt. Oriel.

There may be two species of Ceylon Muggars, as in one of the heads the intermax-
illaries appear to be longer and narrower than in the others from the same country. I

VOL. VI.—PART. IV. Dg

142 DR. J. E. GRAY’S SYNOPSIS OF THE SPECTES

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. indicus, nearly adult.

Fig. 1. Fig. 2. Fig. 3. Fig. 4.

in. lines. in. _ lines. in. lines. in. lines.
iene GhroL Skule... ci. .steaetererstenete res 20 0 17 3 9 10 4 8
Length from occiput to front of orbit.. 6 9 5 9 3 if 2 8
TLenethotstace ptormrice.« =.) en eet 13 3 11 6 6 3 2 0
Length of lower jaw .............- 27 0 23 0 none. 5 5
Widthiatrocciprtinain cre. «|-tereeria cr 13 5 10 6 ay palit 2 6
Width at hinder notch ............ 9 2 6 9 3 9 1 6
Widthvatmotch Weenies 5 4 ieaedue 2 4 0 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

OF RECENT CROCODILIANS. . 145

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. indicus, half-grown. India, Sir John Boileau.

Ze

Fig. 4. Skull of young, of natural size (C. trigonops, Gray),

Asa good illustration of the difference in the appearance of the skulls of the individuals
of the species, I may give the measurement of two skulls of “ Muggars” from India, of
the same size, in the British Museum Collection :—

Broad variety, Narrow variety.

inches. inches,
Length of the skull along the forehead ............ 94 94
Menotihvol side of fhe skill yep errr ets siete e cele: 102 103
Whidthrot backlotiskullee, y-rtveieeemsctelnallel te stele «chars 5¢ 53
Widthinitrontiof ORDitsi perc wersener) atetereleie re -s)ied-\h« 4} 4
Wadthiioverlarcestioothy. ama geheerekiet cyefest-leh-l-sa 3g 3y
MVIGIAC EK Me Apa oed san co4 0 Ole oc moUe) ce Te 23 2 or 142

The broad-nose varicty (fig. 3) was presented by Sir John Boileau, and the narrow one

by Capt. Boys. °

bo

x

144 DR. J. E. GRAYS 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. Rept. 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. 255, t. 54. Giinther, Rept. B. I. t. 18. f. 3.

Crocodilus galeatus, Cuvier, Oss. Foss. v. 52, t.1.f.9 (from Perrault). Dum. & Bibr. Erp, Gén.iii.113.

Crocodilus palustris (part.), Dum. & Bibr. Erp. Gén. ii. 113.

Crocodilus vulgaris (part.), Gray, Syn. 58. Dum. & Bibr. Erp, Gén. ii, 108? Miller & Schlegel, _
Verh. 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, x. 269.

Hab. Siam, Cambogia (MM. 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, /. 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 head, and has 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 isa 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.

OF RECENT CROCODILIANS. 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,
unequal. Cervical disk rhombic, of six large plates. Dorsal plates large, 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 RHOMBIFERA, (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 rhombifer, Cuvier, Ann. Mus. H. N. x. 51; Oss. Foss. v. 51, t.3.f.1-4. Tiedem., Oppel, &
Lebosch,Nat.Amph.75,t.10. Gray,Syn.Rept.59. Dum. & Bibr.Erp.Gén.iii.97. Sagra,Cuba,
t.4! Huxley, Proc. Linn. Soc. iv.10. Blainy. Ostéog. Croc. t. 5. f. 3 (head?) (not Owen).

Crocodilus (Palinia) rhombifer, Gray, Cat. Tort. Croc. B. M. 63; Ann. & Mag. Nat. Hist. x. 270.

Crocodilus planirostris, Graves, Ann. Gén. des Sci. Phys. de Bordeaux, 11. 348. Gray, Syn. Rept. 59.

Crocodilus gravesii, Bory de St. Vincent, Dict. Class. H. N. iii. 109, t. Dum. & Bibr. Erp. Gén. ii. 101.

Hab. South America, Cuba (W. S. Macleay, Ramon 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. 8. 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 rhombifer from two specimens :—one in the Cabinet of
the Academy of Sciences, in a nearly entire 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 describes.and figures as coming from New Spain, under
the name of Aguez-palin, belongs to this species.

M. Graves, in the ‘ Annales Générales des Sciences Physiques de Bordeaux,’ describes
a Crocodile under the name of C. planirostris, from a specimen which was formerly in
the Colléction of the Academy of Bordeaux, but is now in the Museum of that town. It
was procured from M. Journée, 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 might

146 DR. J. E. GRAYS SYNOPSIS OF THE SPECIES

have come from Africa; and he figured and described it under the name of Crocodilus
gravesii in the Dictionnaire Classique d’Hist. Nat. vol. iii. p. 109, t.

MM. Duméril 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 rhombifer, 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 that island.

2. PALINIA? MORELETI. (Yucatan Palinia.)
Crocodilus moreletii, Dum. Arch. du Mus. vi. 255, t. 20; Cat. Rept. 28, n. 5*.
Palinia? moreletii, Gray, Ann. & Mag. N. H. 8rd series, x. 271.

Dorsal scales keeled, nearly square; scales of the sides and limbs smooth, without
tubercles.

Hab. Yucatan, Lac Flores (IZ. 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 difference 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 uniform, 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 intermaxillary bone elongate, produced and truncated behind; 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. 8rd series, 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 CROCODILIANS. 147

enlarged like a canine; but thisisnotcorrect. 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. (Olive African Crocodile.)
Crocodilus niloticus (part.), Daud. Rept. ii. 267. Wagler, Syst. Amph. t. 7. f. 11. 1, 2.
Crocodilus vulgaris, Cuvier, Oss. Foss. v. 42, t.1. £.5 & 12, t.2.f.7. Blainv. Ostéogr. Crocod. 126.

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, Dict. Class. H. N. vy. 105.
C. lacunosus, Geoff. Croc. d’ Egypte, 167.
C. marginatus, Geoff. Desc. @ Egypte, 365. Gray, Cat. Tortois. 61.
Crocodilus cataphractus, Riippell, MS. Gray, Syn. Rept. 78. Mus. Frankfort.
Crocodilus verd de Sénégal, Adanson, Sénég. Cuvier Oss. Foss. v. 4.
Crocodilus acutus, Owen, Cat. Osteol. Mus. Coll. Surg. p. 157. n. 715, not Cuvier.
Crocodilus binuensis, Balfour Baikie, Proc. Zool. Soc. 1857, xxv. 484. Skull described.
Green crocodile, Gray, Rep. of Brit. Assoc. 1862, Sections, p. 107.

Hab. 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 (Havet, fide Cuvier, Oss. Foss. 44).

Fig. 5.

Figs. 5, 6, 7, 8. Head and nuchal and cervical shields of Crotodilus vulgaris.

148 DR. J. E. GRAYS SYNOPSIS OF THE SPECIES

The largest specimen in the British Museum is nearly 15 feet long. ‘There is a very
fine 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 72 inches, at the notch 42 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
over 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 203 inches; greatest width behind, 12
inches, at largest lateral tooth 63 inches, at the notch 3? inches.

These two skulis rather differ in the direction of the suture behind the maxillary
bones; in the wider specimen it is much more produced behind than 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 though 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 have 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 very 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 various 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 reach the 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. 45.

Geoffroy St. Hilaire, in his ‘ Essay on the Crocodiles of Egypt,’ separated the Egyptian

OF RECENT CROCODILIANS. 149

specimens into two species under the name of Crocodilus lacunosus and C. marginatus.
In the “ Annales du Muséum,” 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, under
the name of Crocodilus suchus, Geoff., in the ‘Monograph on the Fossil Reptilia of
the London Clay,’ published by the Paleontographical Society, 1850, t. 1. f. 2. I do
not see how it differs 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. 61, I separated the adult Cape
crocodiles from the North-African specimens, under the name of C. marginatus, 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 marginatus of Geoffroy.

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 Baguana=the people of the crocodile.”—Zool.
South Africa, Appendix 2, 1845.

MM. Duméril and Bibron in their ‘ Erpétologie Générale,’ 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. Geoffroy, from North Egypt and the Cape of
Good Hope.

Var. d. the Crocodile verd of Adanson, from 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 slightest
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. Duméril and Bibron would have found, if they had examined any of

VOL. VI.—PART IV. ut

150 DR. J. E. GRAYS SYNOPSIS OF THE SPECIES

the twelve specimens which they say they procured. They have named the adult
specimen in the Paris Museum Crocodilus biporcatus.

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. marginatus, 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 Bomébifrons indicus from India.
Can the habitat be a mistake? perhaps the habitat was only intended for the first-de-
scribed species, Cataphractus mecistops, for which it is the true locality.

A skull of Crocodilus 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. 716.

5. Mouinia. «

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 maxille.

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 bones produced to the nostrils. Molinia.

1. Mouiia americana (American Crocodile).

Crocodilus americanus (Plumieri), Schn. Amph. ii. 23. Gray, Cat. Tort. & Croc. &c. B. M. 60.

Crocodilus acutus, Geoff. Ann. Mus. ii. 58, t. 57. f. 1. Cuvier, Oss. Foss. v. t. 1. f.3 & 14, t. 2. £5.
Gray, Syn. 60. Dum. & Bib. Erp. Gén. iii. 120. Owen, Cat. Osteol. Spec. Mus. Col. Surg.
157. n. 711, 712, 714, 716; Reptiles of the London Clay, t. 25.f.10. Brihl, Skelet. Krokod.
t. 8 & 9, t. 10, t. 17.

Crocodilus americanus (acutus, Cuv.), Huxley, Journ. Proc. Linn. Soe. iv. 11, 1859.

Molinia americana, Gray, Ann. & Mag. N. H. 3 ser. x. 272.

?? Crocodilus biscutatus (part.), Cuvier, Oss. Foss. x. t. 2. f. 6. Tiedem. Amph. t. 12.

Crocodilus de St. Domingue, Geoff. Ann. du Mus. ii. 53, t. 27. f. 1.

Hab. Tropical America. Cuba (W. S. Macleay); Jamaica (B.M.); West Ecuador
Nicaragua (Fraser ; Richardson); West coast of America (Belcher); St. Domingo (Cuvier);
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 RECENT CROCODILIANS. 151

Var. with two additional small cervical scutella behind the others. BM.
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. 272.

Hab. West coast of America (Belcher); Mexico (Warwick).

Cuvier in his essay gives the history of this species under the name of Le Crocodile a
museau effilé, ou de Saint Domingue (Crocodilus acutus, nob.), Oss. Foss. vy. 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 XXXII. figs. 4-6.)

Dorsal plates in six rows, all slightly and nearly equally elevated; the keels of the two
yertebral 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 intermedius, Graves, Ann. Sci. Phys. i. 344. Gray, Syn. 59.

Crocodilus journei, Bory, Dict. @H. N.v. iii. Dum. & Bib. Erp. Gén. iii.129. A. Dum. Arch. du
Mus. x. 172, t. 14. f. 3 (head). Huxley, Proc. Linn. Soe. iy. 11.

Crocodile de ? Orénoque, 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.

2? Mecistops bathyrhynchus, Cope, Proc. Acad. N. 8. Philad. 1860, xi. 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 1 Orénoque, 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 schlegelii of Borneo, and therefore called the
Bornean animal Mecistops journei. M. Duméril, in his paper in the Archives du
Muséum, not seeing the mistake, says that I refer the true Crocodilus intermedius to
the genus Mecistops, and suggests that Crocodilus acutus ought also to belong to it.

M. Auguste Duméril, for the purpose of comparing the head of this Crocodile with
that of Crocodilus leptorhynchus of West Africa, gave a figure of the head and front part
of the back of the Crocodile de Journée, 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
y2

152 DR. J. E. GRAYS SYNOPSIS OF THE SPECIES

M. Bory de St. Vincent. If the latter, it is so embellished that one is unable to recognize
its origin.

Figs. 9 & 10. Skull of Molinia intermedia: aduit.

Il. Nape 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 webbed. Abnormal Crocodiles.

* Face broad; nasal bone produced into the nostril, Alligatorian Crocodiles.

6. Hancrosta.

The premaxillary suture transverse, rather convex backwards. Nasal bones produced
beyond the intermaxillary, 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 on each side, the hinder one smaller.
Dorsal plates in four 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.
Osteolemus, Cope, Proe. Acad. N. 8. Philad. 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 are two bones in the eyelid instead of one\as in Caiman palpebrosus.

OF RECENT CROCODILIANS. 155

The skull of the Alligator palpebrosus 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 fossee 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 pit 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.

Hatcrosia ni@Ra (Black African Crocodile). (Plate XXXI. figs. 4, 5, 6.)
Krokodile noir du Niger, Adanson, MS., Mus. Paris. See Cuvier, Oss. Foss. iii. 41.
Crocodilus niger, Latr. H. N. Rept. i. 510, from Adanson.
Crocodilus palpebrosus, var. 2, Cuvier, Oss. Foss. ui. 41, t. 2. f. 6 (part.).
Crocodilus trigonatus (part.), Cuvier, Oss. Foss. ii. 65.
African Black Crocodile, Gray, Rep. Brit. Assoc. 1862, Sections, 107.
Osteolemus tetraspes, Cope, Proc. Acad. N.S. Philad. xu. 550.
Crocodilus frontatus, A. Murray, Proc. Zool. Soc. 1862, pp.139, 213, fig. head, t. 29. by Ford. Strauch,
Syn. Croc. t. 1 (head, young).
Halcrosia frontata, Gray, Ann. & Mag. Nat. Hist. 3rd series, x. 277.

Hab. West Africa: Senegal (Adanson); Gaboon ; Old Calabar ; aca River (Cope).

Figs. 11-14. Head and cervical and nuchal plates of young Halcrosia nigra

154 DR. J. E. GRAYS 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 ina 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
dorsal 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 with 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 34 inches, its width in front of the orbit
25 inches, and at the notch of the canine teeth 14 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 the palm-oil ships, and
frequently in a living 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 ‘ Proceedings’ of
the Society as a new species of Crocodile under the name of C. Frontatus; 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 asa
duplicate, it is not included as. Alligator palpebrosus, or under any name, in M.
Auguste Duméril’s List of the Reptiles of West Africa, printed in the last volume of the
Archives du Muséum of Paris.

This Crocodile has very much the external appearance of the Caiman with bony
eyelids, Crocodilus palpebrosus, Cuvier; and I think it very likely that Cuvier mistook a
specimen of it in the Paris Museum, which Adanson had marked with his own hand
* Krokodile noir du Niger, for a specimen of that species. (See Cuvier, Oss. Foss. iii.
p- 41.) And it is still confounded with that species by the French 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 paupiéres osseuses.

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 more
carnivorous, and said to be fond of human flesh.

OF RECENT CROCODILIANS. 155

Cuvier, in his Essay on the species of existing Crocodiles, first published in the 10th
volume of the ‘Annales du Muséum,’ and reprinted in his ‘ Ossemens Fossiles’ under
the head of Le Caiman & paupiéres osseuses (Crocodilus palpebrosus, nob.), after dividing
this 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 Krokodile 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 therefore 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 Green, 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 writing this phrase? or has he erroneously
ticketed the specimen? How are we to disentangle these errors?” &c., vol. v. p. 41.

Duméril and Bibron, in their ‘ Erpétologie Générale’ (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. palpebrosus, 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 usual inat-
tention to true habitats at that period, said came from Ceylon, was a true Alli-
gator, 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 biscutatus, established on the Gavial
du Sénégal 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 & paupiéres 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 DR. J. E. GRAY’S SYNOPSIS 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. Duméril 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 long, 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 narrowest. Intermaxillary produced behind, and embracing the front end of the
nasal.

Mecistops, Gray, Ann. & Mag. Nat. Hist. 3rd series, x. 273; Cat. Tortoises & Crocodiles B. M. 58.
Huxley, Proc. Linn. Soe. 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.”—Loe. cit. p.158. The
Crocodilus schlegelit is a Gavial.

Dr. Falconer observes, ‘The nasal bones (in Mecistops) are extremely narrow and
attenuated, but, as in the true Crocodiles, they descend between the maxillaries so as to
project into a notch between the intermaxillaries. The same holds good in C. schlegelit,
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 Miiller ranged it’ (p. 363).

Dr. Balfour Baikie states, “In all essentials the skull of the Mecistops shows it is to
be properly classed as a member of the family Crocodilide rather than the Gavialidee.
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 CROCODILIANS. 157

MECIsToPs CaTAPHRACTUS. (African False Garial.) (Plate XXXII. figs. 1, 2, 3.)

Crocodilus biscutatus, Cuvier, Oss. Foss. iii. 52, 65, t. 5 (very young).

Crocodilus bisulcatus, Bory, Dict. Class. N. H. v. 108, misprint.

Crocodilus cataphractus, Cuvier, Oss. Foss. y. t. 5. f. 1, 2 (erocodile 4 nuque cuirassée) ; [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 cataphractus, Gray, Cat. B. M. 58.

Mecistops bennettii, Gray, Cat. B. M. 57.

Gavial of Senegal, Gray, Rep. Brit. Assoc. 1862, Sect. 107.

Mecistops, Balfour Baikie, Proc. Zool. Soc. 1857, p. 58.

Hab. West and Central Africa; ? Fernando Po (Bennett), Gaboon, Lagos. Central
Africa, River Binué (Laikie). .

The species has been described from small young specimens. It grows to a large
size. ‘There is an imperfect specimen which is scarcely adult, in the British Museum,
that was sent from Fernando Po by Capt. R. F. Burton, which must have been 13 or 14
feetlong. Unfortunately it wants the head; the body is 5 feet and the tail 8} feet long.

Figs. 15-18. Head and cervical shield of Mecistops cataphractus.

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 during the rainy season are tumultuous mountain-torrents, with rocky
beds.—Proc. Zool. Soc. 1857, p. 58.

VOL VI.—PART IY. Z

158 DR. J. E. GRAY’S SYNOPSIS OF THE SPECIES

Most probably Mr. Bennett’s specimen came from the coast, and was only received
through agents at Fernando Po.

Cuvier, in his Essay, described, under the name of Crocodilus biscutatus, 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 Sénégal,” 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. Duméril 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 Cuvier’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 Mecistops cataphractus before its elongated
jaws were developed, and that the name of Gavial du Sénégal 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. Duméril 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

1 T recollect this visit with pleasure; for I was deputed by Dr. Leach to show this celebrated naturalist and
wayering politician some of the natural-history 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. He was so interested in these bygone 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, we found it difficult to retreat. Cuvier visited England again in 1830, during the
short revolution which placed Louis Philippe on the throne. While here, the Zoologists invited him to a dinner
at the Albion Tavern: he was greatly pleased with what he called the almost royal magnificence of the

OF RECENT CROCODILIANS. 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 cuirassée” (Crocodilus
cataphractus, nob.).

In 1834 Mr. Edward Turner Bennett (Proc. Zool. Soe. ii. p. 10) gave a notice of a
specimen of Crocodilus 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 24 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 Mecistops
for this animal, and for the first time described a full-grown specimen of it which we
had received from the Gambia as MW. bennetti; for My. Rendal 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. Duméril, expressed in his paper “On the Reptiles of
Western Africa” (Arch. 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 difference 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 Duméril, 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, t.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

entertainment. During the dinner the news arrived that the Orleans party had succeeded ; he and his step-
daughter, Miss Duvaucel (who was in the gallery with some ladies), immediately displayed the national
colours, Cuyier’s political predilections were not strong; for he had held office under Napoleon and under the
Bourbons, 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. When I visited
him in after times, he more than once referred to the events of his visits.

Z2

160 DR. J. E. GRAY’S SYNOPSIS OF 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 1846 (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 Binué (see
Proc. Zool. Soc. 1857, p. 58).

Family III. ALLIGATORID.

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.

Alligatoride, Gray, Cat. Tortoises &c. B. M. 56, 1844. Huxley, Journ. Proc. Linn. Soe. 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 premaxilla and maxille. 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, which have not come under my observation.

Spix divided the Alligators into two genera:—Jacaretinga, with acute nose (1. J.
moschifer, t. 1= Caiman palpebrosus, p. 161; 2. J. punctulatus, t.2=Jacare punctulata,
p. 159); and Caiman, or Jacare, with blunt nose (including 1. C. niger, t. 4=Jacare
nigra, p. 167; 2. C. fissipes=Jacare latirostris, p. 167).

His figures are very good representations of the species—indeed, the best knowh.

MM. Duméril 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, flattened, a ridge
in front of each eye, the upper eyelid jimely striated. Nape with two rows 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. eynocephalus, p. 86, Caiman fissipes, Spix, Bras. t. 3. Head short, broad,
thick, a ridge in front of each eye, the upper eyelid rugose. Nape with two rows of

OF RECENT CROCODILIANS. 161

large square keeled shields. Back scale keeled, the three last cross bands 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 preocular ridges, wpper 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 entirely bony. The five former belong to the genus under con-
sideration.

The preorbital ridge distinct, beak broad with three lateral foveole, eyelid striated,
beak broad and blunt. C. nigra, t. 21.

The nuchal scutella many, in three series. C. fissipes, t. 22.

The nuchal scutella many, in two series. C. sclerops, t. 23.

The preorbital ridge evanescent, beak without lateral foveole, 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 ans, 2 ins 1s inves. mie
(Champsamietay oe clee. « seeie <n. 16 (O Sao 3. «6 4 9 By il
HES) GSaobongebous oss NOs 6 5 2) i 3.5 4 0
BELELOPS! tere tete isis ie hers onel= 6 @ 5 8 2S 8 By 3)
VAUiiEON Badoooesaboboo e Alte) 4 6 220 2 9 558}
ponctulata -.ccte a. es 10. 5 5 4 2} Epo 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 with an internal bony plate. The cervical scutella in pairs,
forming an elongated shield. Nasal bone short. Tropical America.

1. Jacare. The orbits united by a bony cross ridge. Eyelids partly striated or

rugose.
2. Carman. The orbits not united by a cross ridge. Eyelids bony, entirely smooth.

162 DR. J. E. GRAYS SYNOPSIS OF THE SPECIES

Il. The ventral scutella thin, the dorsal scutella bony, not articulated together. The
eyelids fleshy, smooth. The cervical scutella in pairs, separate. Nasal bone
elongate, separating the nostrils. North America.

3. AuLiGAToR. ‘The face broad, depressed.

Section I. The ventral scutella like the dorsal ones, bony and articulated together,
forming a shield. The eyelids with 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.

Jacare, Gray, Cat. Tort.Croc. &c. B.M.64,1844; Ann. & Mag. N.H. 3rd series, x. 327, 1862. Huxley,
Proc. Linn. Soe. 1859, 4.
Jacaretinga, Spix, Lacert.

The pits in the maxilla are the cavities left by the preorbital ridges as they advance.
The intermaxillary bone short, truncated behind, with an elongate-oval or lanceolate
cavity 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 fissipes,
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 other 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, JacARE nigRA. (Black Jacare.)

Crocodilus sclerops, Schn. Amph. 162. Blainy. Ostéogr. Crocod. t. 3. f. 2, t. 4. f. 18.

Crocodilus yakare, Daud.

Alligator sclerops, Cuvier, Oss. Foss. v. 35, t. 1. f.6 & 7, t. 2. f. 3. Brithl, Skelet. Krokod. t. 12.
1h iy ROH Mio iia ILS Heris lle

Alligator sclerops, var., Gray, Syn. Rept.

Caiman niger, Spix, Bras. t. 4 (good).

Champsa nigra, Natterer, Beitr. t. 21 (good).

OF RECENT CROCODILIANS. 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.

Hab. 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 jaw. Dorsal scutella
broad, slightly keeled, equal; the lumbar scutella in four longitudinal series.
Nuchal scutella distinct, in two cross series. Eyelids rugose, with a small internal
bone. Back olive, banded 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' the
jaw. Cervical disk short, broad, formed of four bands of scutella. Sides of jaws
pale, with a series of dark spots.

2. JACARE LATIROSTRIS. (Dog-headed Jacaré.)

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).

Champsa fissipes, Wagner, Icon. t. 17. Natterer, Beitr. t. 22 (good).

Crocodilus sclerops, Wied. Abbild. t. Blainy. Ostéogr. Crocod. t. 3. f. 2, t. 4. £13. Schinz, Rept.
t. 112.

Jacare fissipes, Gray, Cat. Tortoises B. M. 64.

Alligator sclerops, Pr. Max. Abbild. t.

Alligator cynocephalus, Dum. & Bib. Erp. Gén. ii. 86.

Jacare latirostris, Gray, Ann. & Mag. N. H. x. 328.

Hab. Brazils; Pernambuco (J. P. G. Smith); Surinam.

The nose of the young specimen is as long as the width at the eighth tooth. ‘The
nose from 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 irregular pale brown cross bars.

Hab. Pernambuco (J. P. G. Smith).

The smaller specimen is peculiar for the very small size of the ventral shield in front

164 DR. J. E. GRAY’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. JACARE MULTIScUTATA. (Brazilian Jacaré.)

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
Jive series of scutella.

* Face depressed, broad ; sides of the jaws with a series of large coloured spots.

4. JAcARE LoNaiscuTaTa. (Long-shielded Jacaré.) (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, Gray, Ann. & Mag. N. H. x. 328, 1862.

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 Jacaré.) (Plate XX XIII.)

Dorsal scutella broad, uniformly keeled, in eight longitudinal series in the middle of
the body; ventral scutella in twelve longitudinal series, the hinder ones smaller, 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.
Hab. Lake of Santa Cruz de la Sierra. British Museum.

OF RECENT CROCODILIANS. 165

** Face attenuated, rather high on the sides; sides of the jaws one-coloured.

6. JACARE PUNCTULATA. (Dotted-jawed Jacaré.)
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.

Crocodilus sclerops, Schn. Amph. 162. Cuvier, Ann. Mus., & Oss. Foss. v. t. 1. f. 6 & 7, t. 2. f. 3.
Tiedem. Amph. 60, t. 5. Guérin, Icon. t. 2. f. 2 & 10. Gray, Syn. Rept. 62. Dum. & Bib.
Erp. Gén. iii. 79.

Crocodilus americanus, Laur. from Seba, t. 104. f. 10.

Crocodilus caiman, Daud. Rept. ii. 394.

Caiman (Jacaretinga) punctulatus, Spix, Bras. t. 2 (good).

Champsa sclerops, Wagner, Syst. t. 7. f. 1, 2, and f. 42. Natterer, Beitr. t. 22 (heads good).

Alligator punctulatus, Dum. & Bibr. Erp. Gén. i. 91.

Jacare punctulata, Gray, Ann. & Mag. Nat. Hist. x. 329, 1862.

Hab. Brazil (Spix); Surinam; Argentine Republic (/. Christy).

Natterer figures two other species, under the name of Champsa vallifrons (t. 24),
(Jacare vallifrons, Gray, Cat. B. M. 65), and Ch. punctulata (t. 25) (Jacare punctulata,
Gray, Cat. B. M. 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 very
nearly the same.

A stuffed specimen from the Argentine Republic measures 6 feet 9 inches long, the
head from the occiput is 104, and the nose from the ridge 634 inches. In another, from
the Zoological Society’s Gardens, 5 feet 10 inches long, the head from the occiput is
10 inches, the nose from the ridge 64 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. JACARE HIRTICOLLIS. (Rough-necked Jacaré.)

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.

Hab. 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. VI.—PART IV. 2A

166 DR. J. E. GRAYS 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 pairs, with sometimes one or a pair interposed between the second
and third pairs. i

Skull with the superior temporal fosse 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. Mus. 66,1844; Ann. & Mag. Nat. Hist. 3rd series, x. 330.
Huxley, Proc. Linn. Soc. iv. 3.

This genus has been divided into two species—one having the cervical shields two, and
the other four in a cross series; in all the latter there are 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 either 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.

T 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 are regarded as distinct, and the second one is called C. trigo-
natus. Yet MM. Duméril & Bibron, in their work, persist in following Cuvier’s
first idea of their being only varieties, and in regarding Adanson’s specimens as
belonging to the second variety, and also in doubting if the “ two varieties,” are both
from America.

The specimen in the British Museum proves most distinctly that there are two very
distinct ‘Alligators with bony eyelids found in ‘Tropical America; which agrees well
with the character that M. Cuvier and MM. Duméril & 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 author also figured a third species, which he calls A. gibbiceps, which, if it is
separable from A. trigonatus, must be distinguishable from it by very slight characters.

The Black Crocodile (Ialcrosia palpebrosa) of West Africa has so much resemblance

OF RECENT CROCODILIANS. 167

to this animal that Cuvier considered Adanson’s West-African specimen a variety of
this species.

Duméril & 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 Alligator palpe-
brosus, var.

A. Head shelving on the sides. Nuchal scutella in a single cross series, cervical scutella
five pairs; dorsal scutella highly keeled, irregular, in six series; the lumbar
scutella in two 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 TRIGONATUS. (Rough-backed Alligator.)

Crocodilus trigonatus, Schn. Amph. 161. 6. Tiedemann, Amph. 66, t. 67.
Crocodilus palpebrosus, var. 2, Cuvier, Oss. Foss. v. 40, t. 2. f. 1.
Caiman trigonatus, Gray, Cat. Tortoises &c. B. M. 66; Ann. & Mag, N. H. x. 330, 1862.
Alligator palpebrosus, Brihl, Skel. Krok. t. 19. f. 3.
Jhampsa trigonata, Natterer, Beitr. t. 26 (good).

Hab. Tropical America.

The largest specimen in the British 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 pairs; 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, Gray, Ann. & Mag. N. H. x. 330.

2. CAIMAN PALPEBROSUS. (Banded Alligator.)

Brown ; tail black-banded.
Crocodilus palpebrosus, var., Cuvier, Oss. Foss. v. t. 1. f. 6-17 and t. 2. f. 2.
Champsa palpebrosa, Natterer, Beitr. t. 27 (good).
Caiman (Jacaretinga) moschifer, Spix, Bras. t. 1 (skull).
Caiman palpebrosus, 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.

Hab. Tropical America.

Natterer figures the head of a species under the name of C. gibbiceps; but I do not
see how it differs from 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 DR. J. E. GRAYS SYNOPSIS OF THE SPECTES

Section II. The ventral scutelia thin, the dorsal scutella bony, not articulated together.
The eyelids fleshy, smooth. The cervical scutella in pairs, separate. Nasal bone
elongate, separating the nostrils. North America.

3. ALLIGATOR.

Head depressed, broad, without 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 dorsal scutella not articulated together, in six longitudinal series ;
the ventral 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. 66; Ann. Mag. N. H. x. 330, 1862. Huxley, Proc. Linn. Soc. iv. 3.
Champsa, Wagler, Syst. d. Amph. 140.

ALLIGATOR MISSISSIPPENSIS. (Alligator.)

Alligator, Catesby, Carol. t. 63.

Crocodilus mississippensis, Daud. Rept. 1. 412.

Crocodilus lucius, Cuvier, Ann. Mus. x., and Oss. Foss. v. t.1.f.8; t.2.f. 4. Tiedem. ier 58, t. 4.
Merrem, Zool. 34. Owen, Cat. Osteol. Spec. in Coll. Surg. p. 165. n. 760, 761. Blainy. Ostéog.
Crocod. t.2. f.1,t.5.f.1. Brihl, Skelet. Krokod. t. 8. f.5,6, t. 9. f.3,t. 10. f. 3, 4, t. 11. f. 2, 3,

£. 20.

Alligator mississippensis, Gray, Cat. Tortoises B. M. 66; Ann. & Mag. Nat. Hist. x. 331, 1862.

Crocodilus cuvieri, Leach, Zool. Misc. ii. 117, t. 102.

Alligator lucius, Merrem, Tent. 34. Dum. & Bibr. Erp. Gén. iii. 75, t. 25, 26.

Alligator cuvieri, Bory de St. Vincent, D. C. H. N. v. 104.

Hab. North America, New Orleans, Texas.

Var. 1. The nose very broad and short. ‘The largest specimen of this variety in the
British Museum is nearly 4 feet long.

Var. 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
ridge.

OF RECENT CROCODILIANS. 169

EXPLANATION OF THE PLATES.

PLATE XXXT.
, 2, 3. Skull of Bombifrons indicus. Adult.
6. Skull of Halerosia nigra. Half-grown*.

PLATE XXXII.

, 2, 3. Skull of Mecistops cataphractus. Adult. Length 21 inches.
5, 6. Skull of Molinia intermedia. Adult. Length 30 inchesf.

PLATE XXXII.
Jacare ocellata. Young: stuffed. Natural size.

PLATE XXXIV.
Jacare longiscutata. Young: stuffed. Natural size.

* Called on the Plate “ Halerosia frontata.” + Called on the Plate “ Crocodilus intermedius.”

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VII. Note to Memoir on the Indian Cetacea collected by Sim WaAurer Enwror.
By Professor Owrn, P.2.S., F.ZS., ke.

tead May 9th, 1867.

ly relation to my paper on Indian Cetacea, read before this Society on the 26th of
June 1865, and published in the Society's ‘ Transactions’', I have received the following
letter from Sir Walter Elliot, K.S.1., F.Z.S., to whom I was indebted for the specimens
upon which my obseryations were based.

Travellers’ Club, 15th April, 1867.

“Dear Prov. OweNn,—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, im the Zoological
Society's ‘Transactions.’ In replying to some inquiries of Mr. Flower, at the College
of Surgeons, regarding the skull of Physeter 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 first brought the crania to you in 1865, 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 that 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

1 Vol; vi. p. 17.

172 PROFESSOR 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 drawing 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 correct, this must be a very different genus. The mouth is small, 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 (¢.¢. in front of the dorsal) depressed; behind the
D. much compressed; the part nearest the tail rising into a sharp ridge.’ On the face
of the drawing I wrote in ink ‘ new sp. Tel. name, wongu: adult ? : Waltair, March 1,
1853;’ and under the mouth I noted the dentition ‘}.4.’ 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.
Total lengthy eek: Chek iad. S725 ARBOR, Fe ee TIE Oe (eos.
ft. in. ft. in
From muzzle topspiraclé ©... 3... shes 2 cee ew Oe af
rom spiracle iho dorsalene. it o.,zshevestteusy cia er ee)
From commencement of dorsal to end of caudal .. 3 10

eee pe

? I then thought this circumstance was accidental.

PROFESSOR OWEN ON INDIAN CETACEA. ie

“I
wo

Length of dorsal from insertion in front to tip .............0.. 000. aan
Bread thy ods itone rye tss-tsisro'ctsis tte Sere, oes Dene MM Cana eee Ou
Lateral measurements.

Length from snout to insertion of pectoral.................. 1 5
Length from insertion of pectoral to vent ...............:.. oF ai
Length from vent to centre of caudal...................... 2 3
(OG
Inferior measurements.
en othefromisnout to went 252 eacimer st ee tg Giana eee ae
Length from vent to centre of caudal..................-... PGS
—- 7 °3
EFOMSNOUG LOLCYC ES, dvcc, craic, soba tre eis chen erie eae enante qeriasteeege eee Oe:
Vength'ofigape’ =... ns... Sauce ee lorhie te. bee olen othe tickdis siti te ican
Girth where largest (in front of dorsal) ............... Ee Ae pas 4
eng thi: of pectoral aopociyecp aang aetmyd see. obs e hs eee ee Lnjy2
Breadth; of, caudal) ......-...22. an. ey CRSA a ele 1 10
ene ihvotavente toes ya- tg citurcats a bra citgs «ein eae waa gs eS Ox 10
Length of small apertures at either side of ditto..................... Ome

«<'This very remarkable animal does not agree with any known genus or species.
The fishermen call it wongu. The snout is rounded and blunt, the mouth, small and
placed far below it, the teeth $.$=20. The eyes considerably above. the mouth, and
nearly 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 curved 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 others
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 possession for some time.

“Tn the course of these migrations some liberties appear to have been taken with
VOL. VI.—PART IV. 2B

174 ; PROFESSOR OWEN ON INDIAN CETACEA.

the drawings themselves. On the cancelled figure of the Physeter, 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.

‘T 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.R.S.”

ee cere

IX. Contributions towards a more complete knowledge of the Skeleton of the Primates.
By St. Grorce Mivarr, F.L.S., Lecturer on Comparative Anatomy at St. Mary’s
Hospital. Part 1. The Appendicular Skeleton of’ Simia.

Read December 13th, 1866.
[PLates XXXV. to XLII]

THE 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 wil! share the
fate of the Dodo and Dinornis, while we may hope that tropical geology will one day
cause a careful description and complete delineation of the bones of Stmia to be much
prized by some future paleontologist.

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,
1 Trans. Zool. Soc. vol. i. 1835. ? Ostéographie, “ Primates: Pithecus,” p. 27, 1839.

3 Recherches d’Anat. Comp. sur le Chimpansé, 1841. * Trans. Zool. Soe. vols. iii. & iy.
> Paper read before the Zoological Society on the 8th November, 1864. ° Trans. Zool. Soe. vol. v.

176 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

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 Zroglodytes 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 T’roglodytes,
where the triangular surface is very indistinctly marked or absent (Pl. XXXV. fig. 1 s).

The spine, apart from the acromion, forms a more elongated triangular plate of bone
than in Zroglodytes, 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
roughness continuing backwards almost to the triangular space before mentioned, and
thus differing from the same part in Zroglodytes (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’.

1 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, 1864, vol. i. p. 565). W. Vrolik also says
it is ‘broader and more analogous to the scapula of Man” than in the Chimpanzee (Todd’s Cyclopedia,
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” which in ordinary mammals would
be “ posterior.” % Duvernoy, Archives du Muséum, 1858, t. viii. p. 24. * Duvernoy, loc. cit.

> This is especially the case in the variety described as Pithecus morio by Prof. Owen, No. 1179 4 in the
Osteological Collection of the British Museum. ° Not so in the type of the variety Morio.

7 Owen, Trans. Zool. Soe. vol. i. p. 364.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. UCT

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. ‘he 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 Zroglodytes ; 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 (Pl. XX XV. 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 very slightly exceeds the rest of the
supraspinous fossa in depth.

The infraspinous fossa is concayo-conyex, 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. Simia, 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 which 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? (Pl. XXXYV. fig. 2). There can scarcely
be said to be a trace of the suprascapular notch*®. The vertebral margin is no longer,

1 Owen. Trans. Zool, Soc. vol. i. p. 364.
2 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. 3 This notch is almost indistinguishable in the same Andaman Islander,

VOL. VI.—PART IY. 2¢

178 MR. ST. G. MIVART 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 very 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 7roglodytes.

The axillary margin is unlike that of the higher forms in that it is decidedly
convex (Pl. XXXYV. fig. 1) in almost all cases, though in the variety Morio it is
more nearly straight (Pl. XX XV. fig. 5). The ridge near the glenoid surface, for the
attachment 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
(Pl. XXXV. fig. 2).

The inferior vertebral angle is rather more obtuse’ than in the higher forms, 7. e.
than in Man and Troglodytes.

The glenoid surface is pyriform (Pl. XXXYV. figs. 4 & 54), 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. XXXYV. fig. 2).

This process is subject to considerable individual variation as to its length® (PI. 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.g. in the specimen No. 55. 12. 26 in the British Museum.

“ 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 3C, however, it is much longer.

Present in both scapule of No. 3C in the British Museum ; also in the Gorilla No. 1011 I.

Such is the case in the specimen No, 3 H in the British Museum.

sw a

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 179

clavicle decidedly exceeds one-fourth of the length of the spine (as measured from the
atlas to the coccygeal end of the sacrum), while in Man and 7voglodytes 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 Chimpanzee, and less
also than that which often exists in the Gorilla. ‘The sternal curve, convex forwards,
is very much more extensive than is the backwardly convex curvature of the outer
or acromial part’. The acromial end bends decidedly somewhat downwards. ‘The
curvature does not appear to be greater in young than in old individuals’, nor in
small adult specimens as compared with larger ones (Pl. 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 Man 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 third 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
(Pl. XXXVIL. figs. 1 & 2d) for the deltoid*; but towards the sternal end of the
bone a slight one (for the attachment of the pectoralis 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-

1 De Blainville has found it to exceed the scapula in length (Ostéographie, “ Primates: Pithecus,” p. 30).

? Archives du Mus. t. viii. p. 25.

3 As shown in the immature specimen No. 3H, in the osteological collection of the British Museum.

4 Judging from the representation given by Professor G. Sandifort in his treatise on the anatomy of the
Orang Outang, in ‘ Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche overzeesche bezittingen,
Leyden, 1840, p. 48, and tab. 3. fig. 2C; also from that of Cuvier in the ‘ Recueil de Planches de Myologie,’
pl. 15. fig. 2x.

5 Mr. W. S. Church describes part of the pectoralis major as arising from the clavicle in the Orang (Nat.
Hist. Review, vol. i. p.513). Cuvier also so represents it (Recueil de Planches de Myologie, pl. 15. fig. 27).

2¢c2

180 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

rior in position, and presents a more or less marked and depressed triangular surface
(Pl. XXXVII. fig. 3, between p and 7), 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 (Pl. XX XVII. fig. 37),
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, XXXVIL fig. 1p).

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 separated 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 concave 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. XX XVII. figs. 2 and 3¢), 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.
The line for the trapezoid portion of the same ligament is generally very prominent
(Pl. XXXVII. fig. 34); and behind and external to it there is sometimes’ a small, yet
marked fossa; but the concavity beneath the acromial end of the bone is never so
marked as it often is in Zroglodytes*.

The acromial end of the bone is very little expanded in Simia; and in this the Orang
presents a marked contrast to the Gorilla; and it also differs from Zroglodytes 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 acrumial end restricted, but the tubercle for the
conoid ligament considerably enlarged.

Humerus. (Plate XXXVL._)

This bone is of great size in the Orang, but nevertheless is not, as in Man and T’roglo-
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

1 E£.g.in Nos. 3 A and 3C in the British Museum.
? See skeleton of a Gorilla, No. 1011 I, in the British Museum.

MR. ST. G. MIVART ON 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 (Pl. XXXVI. fig. 2).

As in Man and T'roglodytes, 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 T'roglodytes.

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 (Pl. 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 rough-
ness sometimes extends so far downwards as to join the ridge extending upwards
from the internal condyle (Pl. XXXVI. fig. 4c).

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 & 2 8).

The head of the humerus is very large’ and rounded, its greatest diameter decidedly

1 As in both the humeri of No. 3B in the British Museum.

2 Professor Owen says, “it is larger in Simia satyrus than in man, its extent equalling a complete hemi-
sphere ” (Trans. Zool. Soc. vol. i. p. 364). De Blainville remarks, “sa téte articulaire est surtout singuliére
par son énormité, son diamétre étant bien supérieur 4 celui de la téte du fémur” (Ostéographie, “ Primates :
Pithecus,” p. 30),

182 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

exceeding the breadth of the two tuberosities, in which respect Simia differs from both
Man and Troglodytes (Pl. 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 (Pl. 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 connecting the two condyles, is in general much more acute than in Man
and Troglodytes. There is nevertheless considerable individual variation, the angle
bemg sometimes as near a right angle as in the Gorilla (Pl. XXXVI. fig. 5), while in
other instances (Pl. 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 (Pl. XXXVI. fig. 3). The surface for the infra-
spinatus looks more outwards and less upwards than in Man (Pl. XXXVL. 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 approached by the margin of
the articular surface of the head than in the higher forms; so that there is a small deep
pit (Pl. XXXVI. 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 generally 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. Soc. vol. y. p. 4, pl. 3. figs. 1, 5, 8.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 183

Whether, however, it descends more or less than in the Gorilla, it always differs from
Man and agrees with Zroglodytes 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 //omo'.

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 (Pl. 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*, with 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 (Pl. XX XVII. fig. 5).

The lower end of the anterior surface is more concave transversely than in Man,

1 See Trans. Zool. Soc. vol. v. pl. 3. figs. 1, 5, 8.

2 Tt is imperforate in both humeri of No. 31 and in those of No. 3H 50. 8. 15. 1 in the British Museum.

3 W. Vrolik remarks “Tl me parait que cette courbure est un produit de lage”? (Recherches d’Anat. Comp.
sur le Chimpansé, p. 13).

184 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

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 Simia passes insensibly into the posterior one, presents a
rough tract and a slight excavation for the supinator 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 (Pl. XX XVIL fig. 6).

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. fig. 5 a) for the insertion of the supinator longus. The styloid process
itself is not so pointed as in Man and Troglodytes (Pl. XX XVII. fig. 5 /f).

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 pollicis and that for the extensor ossis

metacarpt pollicis (Pl. XX XVII. 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 (Pl. XX XVII. fig. 69).

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 the scaphoides,
than is the case in Man (Pl. XX XVII. fig. 8).

Ulna, (Plate XXXVIIL.)

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 (Pl. XX XVIII fig. 1),

? Owen, Trans. Zool. Soc. yol. v. p. 7.

MR. ST. G. MIVART 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. XXXVIIL 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 Tyoglodytes.

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 (Pl. XX XVIII. fig. 1) surface for the 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 (Pl. XX XVIII. fig. 14). 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 concavity is more extensive than in them, reaching as it
does somewhat more nearly to the superior limit of the olecranon (Pl. XX XVIIL. 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 pollex and index) is generally as flat as in Man, and more
so than in Zroglodytes; 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
from the olecranon to the styloid process, and gives attachment to an aponeurosis com-
mon to the flexor profundus 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, which
at that part is considerably roughened for a greater or less extent close to the radial
border (Pl. XX XVIII. fig. 1a). Very rarely, however, the radial margin is enormously
produced, In the Orang, unlike the higher forms, the upper part of this margin does

1 As in the specimen in the Collection of the British Museum, which bears the No. 32, from the MS. catalogue
of the Zoological Society’s Collection.

VOL. VI.—PART IY. 2D

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 (Pl. XX XVIII. fig. 4) between it and the ridge
which does so limit 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. Itis not in any way so
prominent as in that genus, being even less so than in Man (Pl. XXXVIIL. figs. 2 & 6).

The greater sigmoid cavity (Pl. XX XVIII. fig. 1 7) 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 flewor sublimis digitorum is, as in the Gorilla’, well developed.

The fossa for the brachialis anticus is very marked and deeper than in Man or the
Chimpanzee, or than is sometimes the case in the Gorilla (Pl. XX XVIII. fig. 1c).

The fossa for the anconeus 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 (Pl. XX XVIII. fig. 4f).

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 anconeus, the surface for the extensor ossis metacarpt
pollicis 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 pollex) extends up almost to the lesser sigmoid cavity (Pl. XX XVIII. 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 (Pl. XX XVIII. fig. 4¢) and bounding anteriorly the surface for the
anconeus.

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’ (Pl. XXXVIII. figs. 1 & 22). 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. Soe. vol. y. p. 8.

° Noticed by Professor Owen, Trans. Zool. Soc. vol. i. p. 364. He refers to Webster and Treadwell’s « Boston
Journal of Philosophy,’ vol. ii. p. 570, and the ‘ Philosophical Magazine,’ vol. lxviii. p. 186, 1826.

MR. ST. G. MIVART 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 (Pl. XX XVIII. figs. 5 & 6). This last-
mentioned process appears to vary much as to size, from individual to individual
(Pl. XX XVIII. figs. 2 & 6s); 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 ulnaris 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 TJroglodytes. 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 radius are less
in the Orang than I have found them to be in the Chimpanzee, though greater 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 hand
than does the vertical carpal arch of the last-named genus (Pl. XLII. fig. 1).

As in the higher forms, the carpus in the Orang articulates directly with the radius
only.

Scaphoides. (Plate XLII. figs. 2, 3, 4.)

This bone is very much narrower antero-posteriorly (from dorsum to palm), and
relatively much more transyersely 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

1 Owen, Trans. Zool. Soc. vol. v. p. 7.

2 In a mounted specimen of the Gorilla in the British Museum, this process is very short indeed.

3 Pointed out by W. Vrolik, Recherches d’Ant. Comp. sur le Chimpansé, p. 13; and Todd’s Cyclopedia,
vol. iv. p. 203.

4 See De Blainville, Ostéographie, “ Primates: Pithecus,” p. 16 ; Professor G. M. Humphry, Limbs of Verte-
brates, 1860, p. 4; Professor Huxley, Hunterian Lectures, Medical Times, 1864, vol. i. p. 565; and Dr.
Lucae, Abhandlungen yon der Senckenbergischen naturforschenden Gesellschaft, 1865, vol. v. p. 311. The

2D2

188 MR. ST. G. MIVART ON THE SKELETON OF 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 surface, 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 (Pl. XLII. figs. 2-4, 6) appears much produced, because of the
narrowness of the bone. It is not, however, really very 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 articular surfaces. The smaller and more proximal of
these (Pl. XLIT. fig. 3c) joins the semilunare, and, as in Zroglodytes, is larger than the
corresponding surface of Man. ‘The one nearer the palmar (or more distal) surface is
more concave, and articulates with 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 third or proximal face of the intermedium. External again to
this (7. é. on its radial side), and on the inferior aspect of the base of the tuberosity, is
a small surface (Pl. XLII. fig. 4) 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 vascular forarnina.

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 having 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 surface 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

skeleton 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. (1867) plate xiv. fig. 1.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 189

unites with the distal concavity 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 (Pl. XLII. fig. 5).

The radial end of the intermedium is obtusely pointed (Pl. XLII. figs. 5 & 6, d), but
its ulnar end is truncated (Pl. XLII. figs. 5 & 6,c) and presents a small articular surface,
more or less concave, which joins the semilunare. The angle at the junction 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 Man or Troglodytes. 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, ¢. ¢. from dorsum to palm.

The surface for the cuneiforme (Pl. XLII. fig. 8 ¢) 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 groove (Pl. XLII. fig. 11 ¢), 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 (Pl. XLII. fig. 10¢) 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

1 Trans, Zool. Soc, yol. v. p. 10.

190 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

small and less marked than in Man. The surface which joins the unciforme (Pl. XLII.
fig. 11 a) 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 (Pl. 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 Gorilla, 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 (Pl. XLII. fig. 17).

The saddle-shaped surface for the metacarpal of the pollex is always (as also in
Troglodytes) 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 (Pl. XLII. figs. 15 & 17, a) there is a very
distinct, though smali, saddle (the surface being strongly concavo-convex), while in
other instances (Pl. XLII. figs. 16 & 18, a) 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 (Pl. XLII. fig. 174) 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;
while 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. LO.

2 The saddle is unusually little marked in the skeleton of a Gorilla, No. 5779 A, in the Museum of the Royal
College of Surgeons; and in the detached and articulated manus of a Chimpanzee, No. 744, in the same collec-
tion, it is absolutely wanting. Professor Huxley has noticed the absence of a saddle-shaped surface in this
species. See ‘ Medical Times,’ 1864, vol. i. p. 428.

3 Figured by Prof. Vrolik in Todd’s Cyclopxdia of Anat. and Phys. vol. iv. p. 204, fig. 1247. Mr. W. H.
Flower also informs me he observed its existence in the wrist of an adult male Orang in the Leyden Museum.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 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 Sima 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 in excess (Pl. XLII. fig. 22a). 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 (7. ¢. from dorsum to palm), as has the corresponding surface for the scaphoides
in Man (PL XLII. fig. 23d). The surface for the trapezium is concave. The proximal
radial angle is a little produced, but not so much as is the case in Man.

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 Zroglodytes; also there isa 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 (Pl. XLII. fig. 28a). On the ulnar side of the same surface
is a similar notch, which is much more marked than either in Man or Troglodytes
(Pl. XLII. fig. 284). 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 margin than is the casein Man. ‘The head of
the bone, as also in Troglodytes, has a more radiad aspect than in Man; it articulates
above with the mtermedium and semilunare. 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 (Pl. XLII. fig. 26c). The surface for the
unciforme is, as in Zroglodytes, 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 7roglodytes, 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 surface 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
from above downwards and the less relative transverse extent of the articular surface
for the cuneiforme (Pl. XLII. fig. 32 ¢).

1 Trans. Zool. Soc. vol. v. p. 10. > Loe. cit.

192 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

Metacarpus.
The length of this segment of the limb (estimated by the third metacarpal), compared
with that of the spine, is greater than in 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 7’roglodytes, 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 are 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 7’roglodytes, espe-
cially in the Gorilla’. The processes on each side of the proximal ends of the palmar
surfaces of the heads are much less marked than in any higher forms, especially than
in Troglodytes. 'The fossee 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. 57-40.)

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, does 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 Zroglodytes, the tubercle for the flexor carpi
radialis (P|. 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 continued between the lower
facet for the third metacarpal and the articular surface for the trapezoides. The ulnar

* Owen, Trans. Zool. Soc. vol. v. p. 11. 2 Owen, loc. cit.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 193

lateral facets are, as in Troglodytes, divided by a deep groove; and, as in all the higher
forms, the ulnar angle of the proxinal end of the palmar part is strongly inclined ulnad.
Sometimes (Pl. XLII, fig. 34 11.) the proximal articular surface has a crescentic form,
with the concayity of the crescent towards the dorstum. ”

Third Metacarpal. (Plate XLII. figs. 41-44:)

As in Troglodytes, so also in Simia, the proximal radial angle of the dorsum is less
produced than in Man (Pl. 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 (Pl. XLII. figs. 43 & 44).
‘The proximal articular surface (Pl. XLIL fig. 34 m1.) somewhat appwadkes 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 Zroglodytes. The palmar part of the same surface is much
more convex, and is longer, from dorsum to palm, than in Tvoglodytes, 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 than in any above (Pl. XLIL figs. 47 & 48).

Fifth Metacarpal. (Plate XLII. figs. 49-01.)

This metacarpal differs from the corresponding one in Man and Troglodytes in that
its proximal surface is longer (from dorsum to palm), and is moré convex, and at the
same time less concave than in them (PI. XLII. figs. 54 v. & 51).

Digits. 7.

The proximal phalanges of the four outer digits of the manus are much curved, with
the coneayity palmad (more so than in Man or ast yee though not so much so as
are the homotypal segments of the pes.

They are also very broad, and have projecting lateral ridges (Pl. XLII. fig. 1),
which are more developed than in Man, though not so much so as in T’roglodytes’.

The proximal phalanx of the pollex is more slender than in higher forms, and thus
differs notably from 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

1 Owen, Trans. Zool. Soc. vol. i. p. 365.
VOL. VI.—PART IY. 25

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 with that of the Gorilla.

The index, with its metacarpal, 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 (Pl. 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 by 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 XX XIX.)

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
the Gorilla, and very much less so than in Man, being in fact much as in the
Chimpanzee, though perhaps on the whole somewhat broader’.

The external surface (P]. XX XIX. 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 than in Zroglodytes; and the bone
is somewhat less developed posteriorly than in that genus.

‘ Lueae, loc. cit. p. 308,

* As often before remarked or represented, Owen, Trans. Zool. Soc. vol. i. p. 365; De Blainville, « Ostéo-
graphie,’ Primates, Pithecus, p. 30; Huxley, ‘Medical Times,’ 1864, vol. i. p. 565; and Huxley & Hawkins,

‘Atlas of Comparative Osteology,’ plate x. fig. 3. Also Duyernoy, ‘Archives du Mus.’ vol. viii. p. 27; and
Lueae, loc. cit, p. 305.

* As mentioned by Professor Owen, Trans. Zool. Soc. vol. i. p. 363, and by Professor Huxley, ‘ Medical Times,’
1864, yol. i. p. 565.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 195

The internal surface (Pl. XXXIX. fig. 2) looks forwards, 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 (Pl. XX XIX. 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 (P\. XX XIX. fig. 2m). That part cf 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 bemg 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 (Pl. 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 7roglodytes) no such widening takes place; but
on the other hand, there is sometimes at the anterior end of the crest (Pl. XX XIX.
fig. 4 & 5a) avery 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 third
(Pl. XX XIX. 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 differ much from the form of the anterior margin of the
ilium of Man (Pl. XX XIX. fig. 2).

As in Troglodytes, the anterior superior spinous process in Simia is not so marked
and distinct a process as in Man; but the anterior inferior spinous process (Pl. XX XIX.
fig. 2 5) is sometimes almost, if not quite, as prominent as in him. Sometimes, however,
it is not more marked than in 7’roglodytes.

Within this process, and above the acetabulum, the ilium presents a smooth surface
for the psoas 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

1 M. Duvernoy says “un peu creux,” ‘Archives du Muséum,’ vol. vill. p. 28.
2E 2

196 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

upper end of the auricular surface. This upper part of the posterior margin is, as also
in Zroglodytes, much longer than in Man.

The lower part of the posterior margin (below the auricular surface) is very decidedly
concave, but, as in 7voglodytes, the concavity is nothing like so strongly marked as in
the human ilium (Pl. XXXIX. figs. 1&2 9). Indeed it is rather less marked than
appears to be generally the case in 7'roglodytes.

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 7'roglodytes, though it is never-
theless considerably narrower than in Man. This surface presents a narrow groove
running from without inwards, and concave from behind forwards, the concavity being
much increased by a very large process. This process (Pl. XX XIX. figs. 2,3, & 4p),
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 surface of the horizontal ramus (Pl. 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 (Pl. XX XIX. 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 Chim-
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 7'roglodytes, thinner from within 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 the one already mentioned, and no process
whatever near the symphysis, which, as also in 7reglodytes, is much longer than in Man
(Pl, XX XIX. fig. 2s).

The descending ramus of the pubis resembles that of the Gorilla and Chimpanzee in
being much wider than that of Man. ren

‘ Trans. Zool. Soe. vol. i. p, 363. - W. Vrolik calls it «« Epine pubienne ou éminence ilio-pectinée,” ‘ Recherches
d’ Anat. Comp. sur le Chimpansé,’ p. 10.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 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 acetabulum, including (as
before said) the whole of its depressed tract. Below the socket for the femur the
other part of the antero-external face (Pl. XX XIX. fig. 4) presents (as in 7’roglodytes)
a wide surface of bone—concave from above downwards, ‘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-external surface of this part of
the ischium (Pl. XX XIX. fig. 1) is similarly elongated as compared 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 (Pl. XXXIX. figs. 1-4) 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 7roglodytes. 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 (Pl. XX XIX.
fig. 6), and less in that of the spine of the ischium than it isin him. It is, however,
decidedly more prolonged up backwards towards the last-named process, than in 7’rog/o-
dytes (Pl. XX XIX. tig. 3 7).

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 surface 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
(Pl. XXXIX. fig. 4).

The acetabulum is deepest superiorly, and more predominantly deep there than in
Man, and somewhat more so than even in 7roglodytes.

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 Jroglodytes 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. Soe. vol. v. p. 14.

198 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

when the outer surface of the ilium (Pl. XX XIX. 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 Zroglodytes 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 7’roglodytes—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 7’roglodytes, 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 7’roglodytes’.

The body or shaft of the bone differs much from that of the femur of Man, and
greatly resembles that of 7’roglodytes. 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 (Pl. XL.
figs. 8 & 4). It also differs from Man’s, and agrees with that of Zroglodytes, in the
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 Sima and Troglodytes, though least so in the Gorilla.

The anterior intertrochanteric line (PI. XL. fig. 1 ¢) is continued into the spiral line
(Pl. XL. figs. 2 & 3f), 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. 3c in the British
Museum, the inclination is considerable, approaching that of Man.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. iS)s

(Pl. XL. fig. 2h). 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 [roglodytes, so also in Simia, 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 maximus (Pl. XL. fig. 49). The lower end of this depression does not reach to
the middle of the bone’s vertical extent.

As in 7roglodytes, the popliteal space is flatter than is the case in Man.

The entocondyloid prominence (Pl. XL. figs. 1 & 21) is somewhat more developed
than in 7roglodytes, 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 (Pl. XL. figs. 1 and 2 4).

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 greater 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 7’roglodytes. In shape it is very like
the corresponding part in the higher forms, 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 (Pl. XL. figs. 1, 2, 4, 6) beyond the line of the external margin
of the shaft’.

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.
When looked at from above (Pl. XL. figs. 1, 2, 5, 4, 6¢) 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. Church notes its origin as
‘extending from 23 inches below the great trochanter to within the same distance of the external condyle”
(Nat. Hist. Review, 1862, vol. ii. p. 86).

2 See Owen, Trans. Zool. Soc, vol. vy. p. 15, plate 7.

3 Owen, Trans. Zool. Soc. vol. v. p. 15. * Owen, loc. cit. plate 7.

200 MR. ST. G. MIVART ON 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 lme behind is rather more
sharply and strongly prominent than in any of the higher forms (Pl. 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,
ee, sometimes for a short space near the intertrochanteric fossa (Pl. XL. figs: 1

, 4, 6, 7a).

“The sharp projection of its anterior margin is more like what exists in Zroglodytes
than in the general structure of Man.

It is commonly asserted that the ligamentum teres is absent in. the Orang g, 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. 77) 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 therefore be the case that this ligament is pueuaaie: absent
in the Gorilla, and occasionally present in the Orang. T

The rotular surface (Pl. XL. fig. 10) does not,-as in ee 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 7Z’roglodytes. It is even less
concave transversely than in the Gorilla and Chimpanzee, and therefore still more
widely differs in this respect from the rotular-surface 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 Z'’roglodytes* (Pl. XL. fig. 5m & 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 Y’roglodytes, the whole 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 two as in Man*
(Pl. XL. fig. 5m & n).

‘ Owen, Trans. Zool. Soc. vol. i. p, 365, and De Blainville, ‘ Ostéographie,’- Primates, Pithecus, p. 31,

* No, 37 in the osteological collection of the British Museum.

* #.g. in all four femora of the specimens Nos, 5179 a and 51798 in the Museum of the Royal College of
Surgeons. * Owen, Trans, Zool, Soc, vol. v, p. 16, plate 7. fig. 3,

> Owen, loc. eit. p. 16, plate 7. fig. 3. ° Owen, loc. cit. p. 18.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 201

Patella. (Plate XL. figs. 8 & 9.)

The patella of the Orang has its surface marked with vertical grooyes, as in Man,
but it agrees with that of 7’roglodytes 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 subequality 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.

Tibia. (Plate XLI. figs. 1-5, 8, 9.)

The length of the tibia, as compared with that of the spine, is much as in 7’roglodytes,
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 with the radius, the tibia
of the Orang is much shorter than that of Zroglodytes; 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 (Pl. 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 7roglodytes*, 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 greater projection, tibiad, of the internal tuberosity and in the stronger vertical
concavity of the inner surface of the bone leading down from it to the shaft, the Crang
resembles the Chimpanzee, and differs from the Gorilla, and still more from Man.

The tubercle, as also in 7roglodytes, 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
(Pl. XLI. fig. 2), as also in 7roglodytes, relatively, and often absolutely, greater than in
Man. The latter articular surface, again, as in 7'roglodytes, is also larger relatively
than in Man (Pl. XLI. fig. 5%), 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

1 W. Vrolik, ‘ Recherches d’Anat. Comp. sur le Chimpansé,’ p. 15.
? Owen, Trans. Zool. Soc, vol. v. p. 19.

bo
sy

VOL. VI.— PART IV.

202 MR, ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

(Pl. XLI. fig. 47), and, as in Zroglodytes, 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 (Pl. 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 Man, 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 which 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 (Pl. 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 7'roglodytes, 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 7roglodytes—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 the tibialis
posticus and flexor longus hallucis than I have observed in any of the higher forms
(Pl. XLI. fig. 2p).

As in Troglodytes, so also in Simia, the tibial malleolus is more truncated at its apex
than is the case in Man.

The distal articular surface of the shaft of the tibia (Pl. XLI. fig. 9) 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 7'roglodytes, 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 forms a more
open angle with the distal surface of the shaft than even in 7'roglodytes, and its vertical
extent is also much less (PI. XLI. fig. 1).

* Trans. Zool. Soe. vol. v. p. 19. * Owen, loc. cit. p. 20.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 205

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 of the
astragalus (Pl. XLI. figs. 5 & 9s).

The shaft of the tibia is, as in the higher forms, triangular above its middle; but
below that it is, as in Z’roglodytes, much more rounded than in Man.

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 (Pl. 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 which corresponds to it in Simia, if the
faint ridge (Pl. XLI. fig. 57) 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 7’roglodytes, is much less strongly marked than in Man.

The medullary foramen (Pl. XLI. fig. 2) 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 Gorilla.

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 Zroglodytes,
except that in Sima the crest is rather more sharp towards its upper end.

The inner border, as also in Troglodytes, is so little marked as to be hardly 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 (Pl. XLI. fig. 5/7) appears to correspond with
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 T’roglodytes.

Fibula. (Plate XLI. figs. 1, 3, 6, 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 straighter 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 (Pl. XLI. figs. 7 & 10) is more rounded than in

! Owen, Trans. Zool. Soc. vol. y. p. 19. 2 Owen, loc. cit. vol. i. p. 366,
9p?
ARS

204 _ MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

Man, and, though slightly concave, has not the marked depression existing in
Troglodytes.

As in the Gorilla and Chimpanzee, the outer side of the head has a very marked
prominence (Pl. 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 Siméa, one single anterior ridge appears to
answer to both the external and internal anterior lines of Man! (Pl. XLI. fig. 1 w). -
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 (Pl. XLI. fig. 3v) 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 posterior 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 (Pl. 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 (Pl. XLI. figs. 3 & 7 y).

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
offered by the interosseous ligament.

As in Troglodytes, the peroneal: malleolus is shorter and blunter than in Mises 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 Mr. A. T. Norton, in his convenient and carefully prepared
‘ Osteology for Students,’ recently published, ? Owen, Trans. Zool. Soc. vol. y. p. 20.

MR. ST. G. MIVART ON 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 very 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 Zroglodytes 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 margin 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 (Pl. 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 (Pl. 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 (Pl. XLIII. figs. 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 caleaneum 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,’ 1864,
vol. i. p. 565. * Dr, Lucae remarks this, Joc. cit. p. 304.

206 — MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

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 (Pl. XLIII.
fig. 2e), 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. XLIIIL. 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 (Pl. XLII.
fig. 4¢) 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 (Pl. XLII. 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. XLIIL. fig. 12d).

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 in the Gorilla. It encroaches more on the neck of the bone than in the
higher 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 in 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. XLII. 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 T’roglodytes, and while more
sharply limited on its peroneal side than in them, is less so on its tibial side (P]. XLIII.
figs. 9 & 15 9).

The posterior articular surface for the calcaneum has, as in the Gorilla, its anterior
and inner margin more convex (Pl. XLIII. fig. 97), and its posterior and outer one more
concave than in Man, its crescentic form being 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. vol. 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. 565.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 207

anterior articular surface by a groove for the astragalo-calcaneal ligament. ‘This groove,
though somewhat deeper than in Man (Pl. XLIII. fig. 9 4), is not nearly so much so as
in the Gorilla.

The anterior articular surface for the os calcis (P1. XLII. fig. 9h) is much elongated,
and is concayo-conyex, but it is not very distinctly marked off from the rest of the
articular surface of the head of the astragalus.

Naviculare. (Plate XLIII. figs. 14-16.)

This bone in the Orang has a rather more marked proximal concavity than 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 bone less antero-posteriorly expanded; and
the tuberosity (Pl. XLIII. fig. 15 e) is less produced than in Man, and much less so than
in Troglodytes.

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 still more so than in Man. It agrees
with that of Troglodytes, and differs from that of Man in the marked concavity of the
surface for the ectocuneiforme (Pl. XLIII figs. 14-164), 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 Gorilla, and still more
so than in Man. In the Orang there is generally a larger facet (Pl. XLIII. fig. 16 c)
for the cuboid than in Troglodytes. 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 than in Man or

Troglodytes (P|. XLII. fig. 16 d).

Entocuneiforme. (Plate XLIIL. 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. 25 & 24). Sometimes, however (Pl. 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 Z7o-
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? (Pl. XLIII. figs. 25-26 ¢).

1M. Vrolik remarks that it is so placed “ que le gros orteil doit s’écarter des autres.” ‘ Recherches d’Anat.

Comp.’ p. 15. * Noticed by Professor Huxley, see ‘ Medical Times,’ 1864, vol. i. p. 565.

208 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

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 (Pl. XLIII.
figs, 22-26 6, 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 (Pl. 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 (Pl. 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 (Pl, XLII. fig. 50), 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 rather convex articular facet
(Pl. XLII. 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,

Ectocuneiforme. (Plate XLIII. figs. 32-36.)

In the Orang the ectocuneiforme has its proximal articular surface (Pl. XIII.
figs. 82 & 366) much more oblique and much more conyex 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 (Pl. XLIII. fig. 34¢), The distal articular surface is
somewhat T-shaped (Pl 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 (Pl. XLIII. fig. 33 ¢) 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 (Pl. XLIII. fig. 34 d).

Cuboides. (Plate XLIII. figs. 17-21.)

This bone is shorter antero-posteriorly, as compared with its transverse extent, than is

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 209

the case in Man; in this the Orang resembles the Troglodytes, as also in the greater
distinctness of the distal surfaces for the fourth and fifth metatarsals respectively
(Pl. XLII. fig. 19a, 4). 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 in the Gorilla and in Man. The posterior surface of the
cuboid (Pl. XLII. 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 (Pl. XLII. figs. 17 & 18c). As in Troglodytes, the under
surface of the cuboides, behind the prominent ridge, is less extensive than in Man; and
(as also in Troglodytes) 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 naviculare
(Pl. XLII. fig. 20¢, 4). 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 Simia 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 3271 to 100, while in Man, the
Gorilla, and Chimpanzee I have found the proportion to be as 29°7, 28-5, and 30°6 to

100, respectively.

The four outer Metatarsals. (Plate XLII. figs. 37 & 40-64.)

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 those 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 in Sima are rather more curved, with the concavity downwards, than
are the metacarpals; they also diverge distad somewhat less. ‘Their shafts are scarcely

more laterally compressed than are those of the metacarpals; yet they are slightly more

VOL. VI.—PART IV. 26

210 MR. 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 from the condition presented by Man; and alto-
gether the metatarsus and metacarpus are less differentiated in Siméa 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 pollex, 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 Zroglodytes,
the process for the attachment of the tendon of the peroneus longus is ESS! developed
(Pl. XLIII. fig. 39 a).

Second Metatarsal. (Plate XLII. figs. 40-43).

‘The proximal surface of this metatarsal in the Orang is much shorter vertically and
tapers less downwards than that of Man (PI. XLIII. fig. 871.). It is also more
concave transversely than in him, and is somewhat convex vertically 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.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 211

This proximal surface resembles that of the homotypal metacarpal one more in the
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 (Pl. XLIII. fig. 37 m. a).

On the peroneal side of the bone there are two articular facets for the third meta-
tarsal (P]. XLIII. fig. 434, 4'), which are sometimes connected at their proximal ends
by a very small vertical articular surface for the ectocuneiforme.

As in Troglodytes, the tibial articular surface (for the entocuneiforme) in the Orang
is relatively larger than it is in Man (Pl. XLIIL fig. 42).

‘There is a more or less marked tubercle at the proximal end of the plantar surface of
the bone, between the articular surface 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 (Pl. XLIII. fig. 37 111.) 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 (PJ. 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 (Pl. 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 (Pl. 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. 3

Fifth Metatarsal. (Plate XLII. 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 that
2@2 ;

212 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

of Man. The external proximal process, or tuberosity (Pl. XLIII. figs. 52 & 53 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 than
in Man.

The proximal surface of this metatarsal is very much more concave and less convex
than is the corresponding surface of the fifth metacarpal.

Digits.

The proximal phalanges of the four outer digits are very much curved’, with the con-
cavity downwards (PI. 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 of 7roglodytes from their serial homo-
logues, and of course very much 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 narrower than are their homotypes; but again the difference is
less than in Troglodytes, and greatly less than in Man.

The second phalanx of the hallux is often absent®, but when present is much shorter
than is its homotype of the pollex*, 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 the most they are but a trifle shorter, thus agreeing with Zroglodytes
and differing much from Man.

The hallux with its metatarsal, when compared in length with the spine, I have found
to be only as about 15-6 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 42 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 pollex is strikingly different

" Professor Huxley, ‘ Medical Times,’ 1864, vol. i. p. 565.

? See Lucae, loc. cit. plate iii. figs. 5 & 9. * See Lucae. /oc. cit. plate ii. tigs. 5 & 9,

“ Camper found this to be the case in seven out of eight Orangs (Quyres, tom. i. p. 54). Two phalanges,
however, are recorded in two cases by Professor Owen, Trans. Zool. Soe. vol. i. p. 367. W. Vrolik, loc. 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 Blainville, loc. cit. p. 32, records five cases, in each of which the hallux had two
phalanges,

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 213

from that existing in the superior forms, the former only being about four-fifths the
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 (Pl. XLII. fig. 1). The order of
projection is similar to that of length.

The proportion borne by the longest digit (without its metatarsal) to the longest meta-
tarsal, is greater than in Troglodytes, and of course very much greater than in Man.

The hallux only reaches as far forwards as from three-fourths to five-sixths of the
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 does 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 Hylobates.

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 Hylobates.

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.

1 See Huxley and Hawkins, ‘Atlas of Comparative Osteology,’ plate x. figs. 1, 2 & 3, and plate xii. figs.
la, 2a, 3a, & 4a; also Dr. Iucae’s paper before referred to, plate iii. figs. 2,5 & 9, plate ii. fig. 5, plate 1.
fig. 2, and plate iv. figs. 1, 5 & 6.

2 See a paper “On the Skeleton of the Primates,” communicated to the Royal Society on November 22,
1866, read on January 10, 1867, and to be published in the ‘ Philosophical Transactions,’ yol. elvii. (1867)
p. 299, plates xi.—xiy.

214 MR. ST. G. MIVART ON THE SKELETON OF 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 Vycticebine 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 pollex; the
great difference between the length of the pollex 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
very 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 Boa top | Greatest Greatest
of glenoid sae Length Length Length length of breadth of

to inferior yer- | f axillary | of vertebral | of superior glenoid glenoid

tebral angle. Bp ae ee eA surface. surface.
No. SATS. mise: 7:45 6°75 5°75 3°65 1-65 1:07
OLD terns. lee 7-00 6:45 5:20 3-70 1:55 1:00
SSO ean 7:50 6:90 6:00 3:40 1:75 1:17
Average of, the 731 6-70 5°65 3:58 1-65 1-08

three ......

Variety Morio .. 5-49 4:82 3°98 2.98 1:22 0-81

1 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, 3B, and 3C.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

Posterior | Angled formed Angle formed by| Angle formed by| Angle formed by
Pertapral by glenoid glenoid surface | spine of scapula | spine of scapula
angle. surface with with spine of with axillary | with vertebral
axillary margin. scapula. margin, margin.
No. 3A 35 110 65 42 105
jy (G8) powabAce 35 113 70 41 103
pO Ou erences 35 115 70 40 108
Average of the
ae le \ 35 112 68 41 105
Variety Morio 30 120 70 41 106
Average axillary margin: 100 :; average vertebral : 84-3.
Average axillary margin: 100 ;: ayerage superior margin : 53:3.
Clavicle.

Length Length Breadth Greatest Greatest
measured ina} measured about diameter of | diameter of
straight line. | along curyes,| middle. |acromial end.| sternal end.

INOS EAt Eat stots 7-50 7:80 61 1:10 1-27
— Od Go ouadro 6:55 6°75 60 1:02 1:30
» BO essen 7-20 7-40 “60 1-05 1:42

eee eee Lt 7:08 731 60 | 1-05 133

three) seas :

Variety Morio 5°28 5:40 44 CLE “78

Average length of scapula from summit of glenoid surface to posterior yertebral angle; 100

length of clavicle measured in a straight line : 96-7,

Extreme
breadth of
head.

Humerus.
Length measured
in astraight line} Transyerse | Extreme
from summit of | diameter of | breadth of
head to lower middle of tubero-
end of inner mar- shaft. sities.
gin of trochlea.

IOs. G2 ining one 14-00 93 159
— 88 Shores 14:10 92 1:68
NG REE ME 14-40 -90 1:67

Average of the 14-16 91 64.

three) ......

Variety Morio 11:35 67 1:26

Extreme
breadth
between
condyles.

2-50
2:59
2°69

2:59

2:05

Average length of scapula ; 100 ;:
Average length of clavicle : 100 :;

average length of humerus : 193:5.
average length of humerus : 200-1,

:: the average

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

Radius.

Extreme length : |
to end of otyleid, Diameter of Diameter of pier ot
PS ee head. distal end. shaft.

Bs) Docc goose 14:15 91 1:44 :

Bs gad bocce 13-90 93 1:55

Bil) aku j owen 14-70 92 1:60

Average of the on 4

three...... } ea a oe
Variety Morio .. 11-75 72 1:09

Average length of scapula : 100 :: that of radius :; 194-6.
Average length of humerus : 100 ;: that of radius : 100-5.

Ulna.
Extreme length to G |
d of styloid pro-| , Greatest lTensth of sty-|
Sean, aha a heed of To ares
straight line. beat A
SHAW Aware tates 14-70 91 34
Gy sy totveoiae ont 14:20 1:00 29
Bi(Ol Awe one - 15°20 1:10 28
Average of the i P 3
three...... } Heh ch me
Variety Morio .. 11:98 66 24

Average length of humerus : 100 :: that of ulna, measured in a straight line : 103:9.

Os innominatum.

From superior|From superior’ Distance from}; farce or
anterior to | anterior to | Distance be- | Distance be- | posterior in- |- ’27°° f Sorel
Extreme | superior pos-| superior pos- |tween anterior] tween pos- | ferior spinous apa ie te
length. _| terior spinous| terior spinous| spinous pro- | terior spinous} process to hi Eat ee
process along| process in a cesses. processes. spinerof |O UAHE LUOETOS
; crest of ilium.| straight line. ischium. ity:
Oye Whar Ses) 9-90 6°25 5:10 3:50 2:65 2°62 1:90
BBE coine oae a 9:69 6:45 4:82 3:40 2°75 2:58 1:83
Ohm cloriadioo 6 10°25 6-42 4-60 3:25 2:40 2:50 2-10
Sees mot 9-94 637 4-84 3:38 2-60 256 1-94
TEC... see
| Variety Morio .. 7:84 4:55 3°51 2°75 2:50 1:90 1:75

MR.

ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

VOL. VI.—PART IV.

Distance from oa Angle of ante-
anterior infe-| Length of | Greatest ver- Grestest htkty 4 < eee rior part off
rior spinous | symphysis | tical diameter arate peal Ilio-pubic Tlio-iachial ilium with

process to pubis. of acetabulum. Mer 9 acca angle. angle. upper margin
symphysis. ulum, of pubis.
3A. 410 1:97 1:70 | 1:60 130° 175° 145°
SSM apeeie oy oh cease 3°90 1:45 1:68 1:52 132° 163° 137°
Bh Cas Sehr 4-50 1-80 1:75 1:68 140° 167° 150°
r f th = 7
ies ae a 4-16 174 1-71 1-60 134° 168° 144°
Variety Morio .. 3°25 uel 1:38 1:30 128° 172° 150°
Femur.
Breadth be- | Breadth be-
Transverse | Antero-poste-
Extreme Etter diameter of | rior danse i sieaee, Paley mene aaa
Sg head. middle of | of middle of | “Joid emi- | gins of the
2 nences. condyles.
Oj. ogo Oo 10-40 1-40 1:05 76 2:33 2°23
(3) 8) eRe 10:70 1-40 95 “72 2:27 2-11
SIC ie access 11-20 1:45 95 ‘78 2°33 2:29 |
Average of the ) Fi : é a : :
ice f 10-76 1:41 98 75 2°31 2-21
Variety Morio .. 8:90 1:12 “71 aye 1:79 1:70 |
Average length of femur ; 100 ;: that of 0s innominatum : 92:3.
Average length of humerus ; 100 :; that of femur ; 76-0.
Tibia. Fibula.
Antero-poste-| Greatest
Extreme ea, rior diame iene ieee
length. f middle of | diameter o length.
tA surface. [°° Shaft. | distal end. p
SHEN on hha aonior 9-20 2:30 83 1:60
eB mesa sea\uekoions 9:05 2:22 lf 1:70 8:75
Ci Uamate ce c oie ei 9°80 2°39 ‘78 1-72 9-12
Mrorsciet thal tikea5 2-30 79 167
three. ...-
Variety Morio .. 7:24 1-77 Dd 1:25 7°23
Average length of femur : 100: : that of tibia : 86°8.
Average length of radius ; 100 :: that of tibia : 65°6.
2H

218 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.
Manus.
oh Length in (Length follow-
| Length of Ist} Length of Length of Length of Length of |straight line of ing curve of
metacarpal. second. third. fourth. filth. longest proxi- longest proxi-
| mal phalanx. | mal phalanx.
Boob 1:90 4-15 396 | 384 354 2:85 315
2 Ocee eine 2-00 4:22 418 3°97 3:60 3:00 3°55
| Average ..| 1-95 4-18 407 | 3:90 35 2:92 3:35
Pes.
| Length in |Length follow-
Length of lst} Length of Length of Length of Length of |straight line of} ing curve of
metatarsal. second. third. fourth. fifth. longest proxi- | longest proxi-
mal phalanx. | mal phalanx.
| OB ize 1:85 390 | 3°80 3°63 | 3°45 2°75 3-14
| BGrsmdoet 1:92 3:83 3-78 3:63 332 | 275 3-15
i
| Average 1-88 386. | 3-7 3-63 3:38 275 314, |
DESCRIPTION OF THE PLATES.
PLATE XXXV._ Scapula of Simia.
Fig. 1. Outer surface.
Fig. 2. Internal surface.
Fig. 5. Outer surface of the variety Morio.
Fig. 4. Glenoid surface 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 avromian process ;
c, the coracoid process; g, the glenoid surface; s, the triangular surface at the origin
of the spine; ¢, the tubercle for the trapezoid ligament ;
angle; v 2, the inferior vertebral angle.

v1, the superior vertebral

PLATE XXXVI. Humerus.

Fig. 1. Anterior surface of the right humerus of the specimen No. 3 C in the British
Museum.

Fig. 2. Posterior surface of the same.

Fig. 3. Outer surface of the humerus of the specimen No. 3 B.

Fig. 4. Inner surface of the humerus of No. 3 A.

Fig. 5. Superior surface of the humerus of No. 3 B.

Fig. 6. The same of that of No. 3 C.

Fig. 7. Distal surface of the humerus of No. 3 B.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 219

In these last six figures, a represents the articular head; 4, the bicipital groove ;
¢, rough surface, probably for the coraco-brachialis; d, the lesser, or ulnar, tuberosity ;
é, the greater, or radial, tuberosity; /, the external condyle; m, the perforation in the
coronoid fossa ; 0, the internal condyle; p, the capitellum; g, the trochlea; s, musculo-
spiral groove.

PLATE XXXVII. Clavicle and Radius.

Fig. 1. Anterior surface of the right clavicle of the specimen No. 3C 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; e¢, the tubercle for the
conoid ligament; d, the prominence for the deltoid; p, ridge probably for pectoralis
major; 7, rough surface for costo-clavicular 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 surface 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; 6, tuberosity; f, the styloid process; 4, surface
for ulna; fh, the surface for the scaphoides; %, groove for extensor ossis metacarpi
pollicis ; 1, that for ex. carpi radialis longior ; m, that for the ex. carpi radialis brevior ;
_ 0, that for the extensor secundi internodii pollicis; p, that for the eatensor communis
digitorum; u, the surface for the semilunare.

PLATE XXXVIII. U/lna.

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.

Fig. 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, yadial or outer margin; 6, nutrient foramen; c, fossa for brachialis anticus; d,
lesser sigmoid cavity; e, ridge running downwards and backwards from that cavity;
f, surface for anconeus; g, greater sigmoid cavity; 4, the distal articular surface ;
s, styloid process; «, ridge for attachment of pronator quadratus.

2H 2

220 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

PLATE XXXIX. Os innominatum.

Outer aspect of right os innominatum of No. 3 C.
Inner aspect of the same.
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; 4, anterior inferior spinous process; ¢, posterior
superior spinous process; d, posterior inferior spinous process; ¢@, ilio-pectineal line ;
f cotyloid notch of acetabulum; g, greater sciatic notch ; h, spine of ischium; 7, lesser

ez
meats) Ae)
a oe al

sciatic notch; /, tuberosity of ischium; m, auricular surface; 0, obturator foramen ;
p; spine of pubis; 7, 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 ligamentum teres.
Fig. 8. Anterior surface of patella of No. 3 C.
Fig. 9. Posterior surface of the same.

a, articular head; 4, greater (or peroneal) trochanter; c, lesser (or tibial) trochanter ;
d, posterior intertrochanteric line; e, anterior intertrochanteric line; f, its continuation
as the “spiral line;” g, depression of the gluteus maximus; h, line leading down
towards the entocondyloid prominence; 7, pit for ligamentum teres; k, ectocondyloid
prominence; /, entocondyloid prominence; m, external condyle; x, internal condyle;
0, rotular surface; p, popliteal space; s, pit for popliteus.

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, 8.

Fig. 3. Posterior aspect of right fibula of No. 3 I.

Fig. 4. Internal (or tibial) aspect of right tibia of No. 5 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.

MR. ST. G. MIVART 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; 5, tubercle; c, external tuberosity; d, internal tuberosity; ¢, arti-
cular surface for external condyle of femur; f, articular surface for internal condyle
of femur; g, spine; h, groove behind spine; 7, pit for semimembranosus; k, upper
surface of tibia for fibula; /, ridge leading downwards from the front of that surface ;
m, ridge leading downwards from behind that surface; n, medullary foramen ; 9, internal
malleolus; p, groove for tendons of tibialis posticus and flexor longus digitorum; q, axti-
cular surface for superior face of astragalus; r, articular surface of malleolus; s, lower
articular surface of tibia for fibula; ¢, process of head of fibula for biceps; wu, anterior
margin of fibula; v, posterior external margin of fibula; w, posterior internal margin
of fibula; 2, 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 which are united by their ligaments only.
. Left scaphoides of skeleton No. 3B 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; 0, radial tuberosity; c, surface for arti-
culation with the semilunare; d, surface for the intermedium; ¢, that for the radial
end of the intermedium ; f, that for the trapezium.

Fig. 5. Left intermedium of No. 3B: first surface.
Fig. 6. Second surface of the same.

a, surface for articulation with the trapezoides; J, that for the magnum; ¢, 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. 3 B: its radial side.
Fig. 8. Ulnar side of the same.
Fig. 9. Palmar side of the same.
a, surface for articulation with the radius; 4, that for the scaphoides; c, that for
the cuneiforme; d, that for the magnum.
Fig. 10. Left cuneiforme of No. 3 B: its ulnar surface.
Fig. 11. Radial surface of the same.
a. Surface for articulation with the unciforme; 4, that for the pisiforme; ¢, groove

bo

Fig.

222, MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

separating these two surfaces; d, surface for the semilunare; e, that for the fibro-
cartilage of the wrist-joint.
Fig. 12. Left pisiforme of No. 3 B: 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. 5 B: its outer (or extensor) surface.
Fig. 16. The same aspect of the right trapezium of the skeleton No. 3 A in the British
Museum,
Fig. 17. Left trapezium of No. 3 B: its palmar surface.
Fig. 18. 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.
a, saddle-shaped surface for articulation with first metacarpal; 6, surface for second
metacarpal; ¢, 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.
a, the larger (or ulnar) surface for articulation with the second metacarpal ; 0, smaller
(or radial) surface for articulation with the same metacarpal; c, facet for the magnum ;
d, 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; 6, notch on ulnar side of the same surface;
ce, palmar articular surface for second metacarpal; d, dorsal articular surface for the
same metacarpal; e, surface for the trapezoides; ff’, surface for the unciforme.

Fig. 29. Left unciforme of No. 3 B: its dorsal surface.
Fig. 50. Palmar surface of the same.

Fig. 51. Radial aspect of the same.

Fig. 32. Ulnar aspect of the same.

Fig. 33. Distal surface of the same.

a, its palmar process; 6 0’, 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.
g. 55. Dorsum of first metacarpal of the same manus.
Fig. 36. Radial side of the same (first) metacarpal.
37. Dorsal aspect of second metacarpal of the same manus.

MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 223

Fig. 38. Palmar aspect of the same metacarpal.
Fig. 39. Radial side of proximal end of same.
Fig. 40. Ulnar side of proximal end of the same.
a, tubercle for the fleaor carpi radialis; b, groove between the two surfaces for the
third metacarpal.

Fig.
Fig.

Fig.

Fig.
Fig.

a,

41. Dorsum of third metacarpal of the same manus.

2. Palmar aspect of the same metacarpal.
3. Radial aspect of proximal end of the same.
. 44. Ulnar aspect of proximal end of the same.

radio-proximal angle of dorsum.

bo

Im OF Re OO

. 45. Dorsum of fourth metacarpal of same manus.

. 46. Palmar aspect of the same metacarpal.

. 47. Radial aspect of proximal end of the same.

ig. 48. Ulnar aspect of proximal end of the same.

. 49. Dorsum of fifth metacarpal of the same manus.
. 50. Palmar aspect of the same metacarpal.

. 91. Radial aspect of proximal end of the same.

PLATE XLIII. Pes.

Plantar aspect of the pes No. 5079 in the Museum of the Royal College of
Surgeons, the bones of which are united by the ligaments only.

Right os calcis of skeleton No. 3 B in the British Museum: its peroneal
aspect.

Tibial aspect of the same.

Anterior (distal) aspect of the same.

Posterior-aspect of the same.

Dorsum of the same.

Plantar surface of the same.

anterior articular surface for astragalus; 0, posterior articular surface for astra-
galus; ¢, 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.

Fig.

im
Fig. 12
13

. Peroneal aspect of the same.
. Anterior (distal) aspect of the same.
. Posterior aspect of the same.

a, neck; 6, articular surface for naviculare; ¢, surface for tibial malleolus; @, groove
behind the last; e, rounded surface posterior to the groove; f, surface for peroneal

224 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

malleolus; g, groove for flexor tendon; h, anterior articular surface for os calcis; 7, pos-
terior articular surface for os calcis; 4, groove for astragalo-calcaneal ligament.
Fig. 14. Right naviculare of No. 3B: its anterior (distal) aspect.
Fig. 15. Tibial aspect of the same.
Fig. 16. Peroneal aspect of the same.
a, surface for entocuneiforme and mesocuneiforme; 6, 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; 4, surface for fifth metatarsal; c, pivot-like process
of tibio-plantar angle; d, ridge of plantar surface ; é, articular surface for ectocuneiforme ;
f, articular surface for naviculare; g, groove for tendon of peroneus 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. 3C.
a, articular surface for first metatarsal; 6, articular surface for the second metatarsal ;
c, surface between the articular surfaces for the first two metatarsals; d, surface for
mesocuneiforme; é, surface for naviculare.
Fig. 27. Right mesocuneiforme of No. 3 B: its dorsum.
Fig. 28. Tibial aspect of the same bone.
Fig. 29. Peroneal aspect of the same.
Fig. 50. Anterior (distal) surface of the same.
Fig. 31. Posterior surface of the same.
a, articular surface for metatarsal; 6, articular surface for naviculare; ce, articular
surface for entocuneiforme; d, articular surface for ectocuneiforme.
Fig. 32. Right ectocuneiforme of No. 3B: 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. 56. Posterior aspect of the same.
a, articular surface for metatarsal; 6, 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. 3B.
a, facet for entocuneiforme.

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MR, ST. G. MIVART ON THE SKELETON OF THE PRIMATES.

Fig. 38. Dorsum of first metatarsal of No. 3 B.
Fig. 39. Lateral view of the same metatarsal.

a, process for attachment of tendon of peronwus longus.

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, 45. Peroneal side of proximal end of the same.

a, articular facet for entocuneiforme; 4, 6', articular facets for third metatarsal.
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 tie same.

Fig. 47. Peroneal aspect of proximal end of the same.

Fig. 48. Dorsal aspect of fourth left metatarsal of the same skeleton.
Vig. 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. 55. Plantar aspect of the same.

Fig. 54. Tibial aspect of proximal end of the same.

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VOL. VI.—PART IV.

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X. Description of the Remains of three extinct Species of Elephant, collected by Capt.
Spratt, C.B., R.N., in, the Ossiferous Cavern of Zebbug, in the Island of Malta.
By Grorce Busx, RS. ; partly from the Notes of the late H. Fauconsr, M.D.,
F.RS.

Read June 27th, 1865.
[Prates XLIV. to LIIT.]

§ I. Introductory.

IN 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
suffice 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 43 to 5}, and
in width from 14 to 23 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 (Cygnus falconeri, 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
Zebbug Cavern,” which will be found in the 23rd volume of the Quarterly Journal of the Geological Society (1867).
Two Reports, also, on other ossiferous Cayerns in Malta, in which numerous remains of Elephants and other
animals have been collected by Dr. Leith Adams, haye appeared in the Reports of the British Association for 1865
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 fissures 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 Z. melitensis.
—June, 1867.

VOL. VI.—PART VY. 2k

228 MR. BUSK ON THE REMAINS OF

was of gigantic size (Myoxus melitensis, 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
asimilar character have been discovered, containing chiefly bones of one or more species
of Hippopotamus unmixed with those of the Elephant, and regarded by Captain Spratt,
for certain geological reasons, as having been introduced into the 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 paleontological 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 the 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 Cygnus falconeri, has been undertaken by Mr. W. K.
Parker, F.R.S., whilst there is reason to hope, from a letter from Dr. Leith 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 in 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 recently 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 1862. In 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 he proposed the name of Elephas melitensis.

”

* 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 Cygnus falconeri is already pub-
lished, anted, p. 119.

THREE EXTINCT SPECIES OF ELEPHANT. 229

Naturally much struck with such an extraordinary deviation from the otherwise univer-
sally received conception of the 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 paleontologist.
This I have endeavoured to perform, though haud 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, therefore, 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 iv 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 maay 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 iny extreme
surprise, that it apparently comprised the remains 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

2k 2

230 MR. BUSK ON THE REMAINS OF

of more than one dwarf species. Further attentive examination and comparison of the
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 lived to resume his investigation of the Maltese fossils,
which for the last two years or more had been completely interrupted by the attention
he had devoted to the fossil remains from Gibraltar, 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 circumstances, 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 LE. 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 palontology, especially as regards fossil proboscidia, is so deeply indebted,
and to term it E. falconeri.

As regards the large form associated with E. melitensis and E. falconeri, 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 EH. antiquus; 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.

THREE EXTINCT SPECIES OF ELEPHANT. 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.

§ Il. Exepuas, sp.? (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 nearly, 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 (?)
humerus, and (3) a nearly complete spine of probably the 17th or 18th dorsal
yertebra. 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
(Pl. 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 off, the fracture extending chiefly to the left side. The fractured sur-
face looks asif 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 primigenius 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 gntter,

252 MR. BUSK ON THE REMAINS OF

is 2'-15. In two lower jaws of a mature African Elephant, at the College of Surgeons,
the 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
the same point is about 1-9, varying from 18 to 2"-0. As regards the width of this
part alone, therefore, 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 compared with a very perfect mandible of £. primigenius
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 character will allow us, that the Maltese large Elephant had, as regards
the mandible, more of the characters of H. primigenius than of either of the existing
species. The mental foramen, which, as has been said, must have been about 07:5
in diameter, is placed about 1-0 behind the edge of the symphysial 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 670;
whilst in the Mammoth already referred to it is 68. 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 6".

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 (0™185), 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 :—

* Ossem. fossiles, 4™° éd. 1834, 8vo, tom. i. p. 504.

THREE EXTINCT SPECIES OF ELEPHANT. 233
Bs africanus) (VOUng) 3.2 20-2 cn-ies eae caer aos 1 to 4:00
PP abOsews (LEMMAS) Mi caerah clade hace ehoebtesnee (ol 1 ,, 4:53
By qndieus (Sumatra) .32.04...cked siteaweelae 2a: ej 042
EL Primagenius oc. ssenccveccerensenewntensncees etc) (

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 5-6 long ; and the neural spine itself measures along the anterior border
5"2, and along the posterior 4-7. Its smallest antero-posterior diameter is 1-3, and at
the base 2":3. Its least tr. d. is 0:5; and the ap. d. of the expanded outer end 1-65. 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'-9 in breadth, and it is of an oblong form. ‘The corresponding spinous process in
the African Elephant (No. 708, 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
1-2, and at the base 3-7, the least tr. d. 1-0, and ap. d. 3'"8, at distal end 15.

Of the teeth of this species, I have been able to detect only innumerable fragments
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 layers have been removed ; we may conclude, therefore, that the fragment
does not represent the real diameter of the tusk, which may consequently be regarded as
having 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 shaft of the left femur.

The exoccipital bone (figs. 3 and 4, Pl. 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 REMAINS 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 :—

Taste I.—Measurements* of Exoccipital Bone, in Maltese and African Elephants.

| | Width be-

| Height or | tween bor. | Length wrath of basi. Thickness Foe, ene
greatest [Breadth of| ders of eon- of basiocci- occipital syn- | TeB1ON | 7g ati- | con dyloid
length of exoccipital.| dyloid fossa | Pital syn- | Ghondrosis, | of mastoid | rele
exoccipital at foramen |chondrosis. [\cellss4: +] CUBE San: aren
‘| | face. surface.
| magnum. | |
Young African Ele- ) | z 2 é
eae 4: 35 16 19 95 13 22 1:35
phant, No. 708, B.M. { z
Young Multese fossil. 47 39 16 18 ot el reetiag 2°35 | 155

]
|
|

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 al 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
striking 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. 7087, B.M.), in an
exoccipital having a greatest diameter of 57:1, 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 EF. africanus. I have not
had an opportunity of comparing the exoccipital of the Indian Elephant of the same
age. In the young African Elephant the opening into the paramastoid cells is tri-
angular, and a transverse septum may be observed 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 cireumference 5-9. In
a young L. 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 show 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, pelvis, 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.

§ ILI. ELeraas MELITENSIS.

Bones or fragments of bones belonging to the larger of the two dwarf species of
Elephant to which I have assigned the name of E. melitensis constitute a very consi-
derable part of the collection. They are exceedingly numerous; but amongst them
are very 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 EL. melitensis.

Separating the mature from the immature bones, and excluding the teeth, there 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 ver-
tebre ; (7) the neural spine of one of the anterior dorsal vertebra ; (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 Y. 21

236 MR. BUSK ON THE REMAINS OF

head of the humerus (side uncertain) ; (12) the upper end, minus the olecranon of the
right ulna; (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 very 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. fig. 15) is about
4’”-6 long, and about 2:2 in ap. d. at the 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 inch 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 13. The ap. d. of the ramus ona 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 1""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 peculiar
pock-like pitting which is exhibited on that surface within the base of the coronoid
process in all Elephants, but which is much 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 EXTINCT 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 tne 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 which
the comparison was made, the tr. d. of the head in the Indian Elephant was 4",
and that of the neck 1-9; whilst in the African the head was 3-7 in tr. d., and the
neck 1/-5. In general form also, a considerable difference may be remarked. Viewed
laterally, the ascending ramus in the African 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
yery 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. EB. indicus. B. E. africanus.
2L2

238 MR. BUSK ON THE REMAINS OF

portionately also of much larger size, isso 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. melitensis. ‘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 in the
presence of the sulcus on the posterior border, in the much thickened anterior margin
of the 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. melitensis
1-8; 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 (Pl. XLVII. fig. 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
wide 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 35. 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. 2678, C. 8.) 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, melitensis. In a vather younger specimen of the Indian
Elephant, termed £. swmatrensis, in the British Museum, the same measurements
are 3!-3 and 2'-4 respectively ; and in an African Elephant of mature age, 4"-7 and 3'-2.
In the latter instance a considerable difference is apparent in the form of the facet. In
E. indicus (vax. swmatrensis) the surface is kidney-shaped, having a deep sinus on the
inner border, whilst in E. africanus that border is entire; but I am not aware that this
difference exists in all cases between the India and African species.

2. The seventh cervical vertebra (P]. XLVI. fig. 9) isa beautifully perfect specimen of
that important and highly characteristic element 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 E. swmatrensis, are as follows :—

* Other fragments of the atlas of this species are shown in PI. LI. fig. 35, and, as I believe, but am not sure,
in fig. 33. :

bo
(S)
ile)

THREE EXTINCT SPECIES OF ELEPHANT.

TABLE I1.—Dimensions of 7th Cervical Vertebra.

| l ] ]
| Dransverse | Transverse | Transverse | Transyerse Verti 1 |Lransverse Vertical
| Heich Length of diameter at|diameter at) diameter | diameter | ¥°"4C@l diameter ofjdiameter o
eight.| Spine. | articular | transverse | at costal | of body Snes | vertebral | vertebral
| | Processes. | processes. facets. anteriorly, |°* 2°¢Y*| canal. canal,
E. melitensis...) 68 25 3°55 57 3-5 29 | 2:85 20 ils
E. africanus....| 820) 4:9 715 9°38 61 635 | 54 3°85 21
E. sumatrensis ..| 75 25 56 54 4:65 | 3:5 2°38 19
| |
E.indicus ....| 14:0 70 65 10:25 | 68 55 4:25 3°75 30

The body, which is about 1°53 in its greatest thickness, is almost circular, slightly
convex in front, and well hollowed behind, especially towards the lower part. The
neural canal is triangular and, viewed anteriorly, very nearly equilateral; the laminz
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
lamin, 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 subtriangular, 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 4.
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 they do in £. africanus, the height,
instead of 1:"7, would have been only 1/09. 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 compare the 7th cervical vertebra of E. melitensis with that of the Sumatran
Elephant (younger than the African example), the differences are still more striking,
especially in the 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 more than half its transverse width. The neurapophyses or
lamine, instead of being thin and flat, or riband-shaped, are very thick and square, and
the neural spine in proportion yery 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 MR. BUSK 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 Sumatran 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 EH. melitensis has more of the African than Asiatic character.
In order to render more distinct some of the diversities presented by this vertebra in
the different instances cited, I have subjoined the accompanying woodcuts, which are
drawn to a scale of one-third the uatural size—

E. indicus. E. melitensis.

all of which represent the posterior view of the neural arch.
3. A dorsal vertebra (PI. XLVI. fig. 10), either the 6th or 7th, but in all probability

THREE EXTINCT SPECIES OF ELEPHANT. 24]

the former, if the ribs are articulated as in the Indian Elephant, seeing that the remains
of an articular surface are visible on the anterior aspect of the left transverse process.
The bone is remarkably perfect, wanting only the extremity of the neural spine and
a small portion of the right transverse process.

Its principal dimensions are 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 1-85. Thickness 1-5. Width of anterior surface of body
2'15, of posterior 2”-2; anterior costal facets 1/-1 09; of posterior 1:05 x 0"-65 ;
ap. d. of neurapophyses 1” 2; extreme distance between the outer borders of the
two posterior articular facets 1"6. 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
wregular 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 sufficiently so to afford a very good idea of its characters. Its dimen-
sions are:—Height of body 1-7; thickness or ap. d. 15; 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 1-65. The body is suboval, very concave
behind, and nearly flat in front. Both transverse processes are broken short off, as is
also the greater part of the apparently small neural spine.

In the second lumbar vertebra of E. (var.) swmatrensis, 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 1"-95, and the transverse diameter of the canal 2"-2, and its height 1'-68.

5. The only other fragment belonging to the spine, and appearing from its dimensions
to correspond very closely with the three vertebra just described, is one of the anterior
dorsal spines (Pl. XLV. fig. 7). It is broken off 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 (Pl. XLV. fig. 8) measuring about 4-5

242 MR. BUSK ON THE KEMAINS 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
longest, 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. M., denominated
E. sumatrensis, has the greatest diameter of the head 1-5, 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 EF. swmatrensis. 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
(Pl. 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 the acetabulum it is evident that the curve of
the articular surface must have had a radius of about 1-6. The cup was consequently
fitted to receive a femoral head of about 5+ inches in diameter. The anterior surface
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, which 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. The outer angle is round and smooth;
and the inner, or that which forms the ischial border of the obturator foramen, though
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 EL. falconeri, as it shows, perhaps as strikingly as any other
single part, a considerable distinction between the two forms.

Tam not aware that the circumstance has been previously 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. indicus. In E. africanus, again, the margin is tolerably even all round, whilst in

THREE 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
difference in the outline of the foramen in the two species is shown in the accompanying
figures, taken from specimens of £. africanus and (so termed) E. swmatrensis in the
British Museum.
7

E. indicus. E. africanus.

I have had no opportunity as yet of examining this part in E. primigenius, or in any
other fossil species; but, from the figures of the Elephant’s pelvis given in pls. xii.
& xvi. of the ‘Ossemens fossiles, it would seem that the ischial eminence above
noticed and the peculiar constriction of the upper part cf the obturator foramen are
as well marked in fossil bones which in all probability belonged to the Mammoth
as they are in the figure given, pl. 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. faleoneri the configuration of this part more nearly resembles that of
E. indicus and E. primigenius.

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.—PART VY. 2M

244 MR. BUSK ON THE REMAINS OF

1. The portion of scapula (Pl. XLVIII. fig. 23) consists of the greater part of the neck
and the adjacent bone, about three inches in its greatest 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 1-2. The articular surface is perfectly smooth,
and its curve in the vertical or longest direction has a radius of about 2", and in the
transverse of about 1-75. The entire fossa may be estimated at about 23 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 :—

ETON GUGUSA Si OUNS) cree: seer sess nates eee eee ec eT: 3°5 xX 2°6
FE VROUCUS (HNATOTE) 022 itn s epeenocves neem abana. <5

E. indicus (var. sumatrensis, young) ............06 47 x30
EE pnemig angus) (OUvien) ieesnase sacs rest osacmscces 8:5 x 4:4
HE. OP TUCHMUS (HEIMULE [crn vescinecadeeenseescenaneseos 67x43
Hf. MELLENStS: (MAtULE) —.. or aconee sconce snes ve sere. 24x 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 (Pl. XLVIIL. fig. 22) isin 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 slightly 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,

THREE 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 pl. 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 £. primigenius, indicus, 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. indicus; whilst in E. africanus, and still more so in
E. melitensis, 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
Pl. XLVIII. figs. 24 and 24a. 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 with 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
1-65, and its ap. d. at right angles to the same line, 1-65; so that it is circular; whilst
the radius of the curve of the concavity is about 1’"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 (q, 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 0-95,

2M 2

246 ; MR. BUSK ON THE REMAINS OF

1-25; eaternal 115; 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 (7) in the
accompanying figure, which is intended to show the outline of the transverse section _
the shaft at a distance of about 36 below the middle of the

radial sulcus, or at a distance equal to about twice the trans- 8 ast i
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 ue za
nearly flat below, passing above into a rather deep sulcus be-

tween the inner articular head and the base of the olecranon. is

The posterior angle is very acute, but above it is broken ob-

liquely off. The internal angle is thick and rounded, and pa

the external rounded and slightly carinate.

4. Asecond well-marked fragment of the ulna is shown in fig. 25. It is the o/ecranon-
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) A very 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 4-1, and its ap. d. 3-1; whilst the length of the outer facet is
3-4, and its width 2'"1.

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 ..........e0c0e008 639 634
Antero-posterior diameter of inner facet ......... “400 634
Rensthrofouterifadcineniey eds. xiviaieae tetowse. “447 °4.23
Wadith! Gk 055 Recah teen Acad Ath toh 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 H. melitensis, to the African form. These differences are—the
radial sulcus is more rounded and shallower in £. africanus, 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 EZ. 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 drachialis anticus, which is scarcely indicated in E. melitensis.

(3) Another character in which H. melitensis approaches EF. 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 Pl. 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 1"°5, and the circumference 55 ; 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 pelvis &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 MR. BUSK ON THE REMAINS OF

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 [ plus gréle] 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 +, 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 upright 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 (évasée) 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 EH. 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,

* Op, cit.1, p. 571, + Ostéographie des Mammiferes, iii. p. 42.

THREE EXTINCT SPECIES OF ELEPHANT. 249

in 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 603 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 E. africanus, and fig. 10 that of H. indicus of about the same age.

10

2. On the posterior aspect the difference 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
circumstance 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 which, 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

250 MR. 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 hollowed 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
meagurements given in the Table.

. 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. melitensis.

1. It differs from the African, and agrees with the Indian, in the convexity in the
yertical 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.

6. 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. 25]

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 reserve what I have to say until
I come to that bone.

§ IV. Enepuas FALCONERI.

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.

. Portions of several vertebre.

. A portion of the second rib.

. A portion of the scapula.

. The greater part of the right humerus.

. 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.

a OF OF LD

1. Aélas.

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 . falconeri. But
upon due examination it will be found that one only, represented in fig. 32, really ap-
pertains to that species, and that the other is a portion of a very young atlas, belonging,
as I believe, to #. 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 and
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 EZ. 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 Y. 2N

252 MR. BUSK ON THE REMAINS OF

about 40 inches. This would be a height rather greater than that which I have assigned
to E. 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 Z£.
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 Z. falconeri, and of one of younger age and somewhat lower stature, or,
it may be, of individuals of different sexes,—a supposition for which some support may
be found in the circumstance 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 larger 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 margins of the two
anterior articular facets, or what may be termed the transverse diameter of the con-
dyloid cup, may be estimated at about 33, 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 ich 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, R. C. 8.), the latter being to the former as 75 to 100; whilst
in the atlas of EL. 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
Museum,—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 the African type in E. melitensis, and a corresponding
resemblance to the Indian in E. falconeri. ;

2, Other Vertebre.

Several fragments of the spines of dorsal vertebrz are contained in the collection, of
which two are represented in Pl. LI. figs. 54 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 surface the 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 E. melitensis before described, and which is figured in Pl. XLVI.
The corresponding facets in that vertebra measure 0-7 x0”-5; whilst the transverse
width across from the outer edges of the facets, at the widest part, is, im the one case,
1-1, and in the other 17-55, 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. Ribs.

The only portion of the ribs distinctly recognizable from its dimensions is a fragment
(nearly 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 of E. melitensis, described in page 241, and figured in Pl. XLV. fig. 8.
It nevertheless affords several distinctive characters beyond its mere size, which is at
once obvious.

1. In the rib of E. melitensis 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. melitensis the anterior surface of the bone in the expanded portion
is very concave, and in E. falconeri nearly level*. 3. In E. melitensis a very acute and
prominent ridge or angle descends for a considerable distance from the anterior part of

* Tn the Indian Elephant, both in the very young animal and in one nearly full-grown (Chuny), the anterior
surface of the rib at the part indicated is, if any thing, rather convex, and quite unlike the condition presented
in EZ. melitensis. As in that species, however, the anterior and inner border of the bone in the upper or curved
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 Z. melitensis.

2N 2

254 MR. BUSK ON THE REMAINS OF

the 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 £. 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, isa small portion of the right (Pl. 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 fosse are
very small. The glenoid fossa is narrow, elongated, and pyriform in shape. It measures
about 1:71”, and the border slightly overhangs at the upper end. The radius of
the longitudinal curve is 1”:1, and of the transverse 1”,—curves that would seem to
correspond pretty closely with the computed size of the head of the humerus of £.
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 HZ. 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 surface 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 the 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 this 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 are 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
yaries perhaps a good deal in the Elephant; 3, the entire absence of the least trace of a coracoid tube~

THREE 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 P]. XLIX. One of these(fig. 26) is amongst the most perfect and instructive of all
the bones collected in the Zebbug Cavern. It isa 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 uninjured; 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 itis, 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 EZ. 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 figure, in which the spine is repre-
sented in a vertical position.

256 MR. BUSK ON THE REMAINS OF

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. r

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—with the following results :—

As regards differences in the proportions, expressed numerically, it would seem (a) that
the antero-posterior diameter of the head is about the same, (6) that the transverse di-
ameter of the head in the Indian is as about 102 to 92 in the African, (¢) 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 (¢) that in the African the antero-posterior diameter at the same part is considerably
less, or as 60 to 49, whilst (f) the circumference 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 56 in the African, and of the outer as 56 to 48, and (7) 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

THREE 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

the 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 africanus. 13 & 15.—Elephas indicus.

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 figures.

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 Z. falconeri, as we have no means of actually
measuring the head, we are unable to compare 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 bone 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 inner 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

THREE EXTINCT SPECIES OF 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

18

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 of 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 EF.
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 jydged 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 EF. 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 Z. 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 £. melitensis,
the extreme tr. d. of the surface in the specimen standing in the ratio of not
more than 63 to 100, whilst in EZ. 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 EF. 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 £. melitensis
and E. primigenius.

The second portion of the hiner (represented in fig. 27, Pl. 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. VI.—PART V. 20

a

260 MR, BUSK ON THE REMAINS 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. falconeri 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 fig. 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 find
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 1-2. The olecranon is about 1:1 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 12 or 1’°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 this hollow is very circumscribed,
and immediately below it the surface is flat from side to side, and a little lower down
convex. ‘Lhe outer surface is concave and quite smooth; the internal, except 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
project at all in front. The adjoining figures represent the outlines of a transverse
section of the shaft, in fig. 19 at about 2”-1, and in fig. 20 at about 2-7 below the

level of the articular surface at the bottom of the 49 re ‘ 20

radial notch; and the comparison of these with 2 Apne? -¢
the outline of a transverse section of the shaft i
of E. melitensis at a rather lower point (p. 246) prides

will serve to show how closely in form they all 2 iP

resemble each other. In other respects also: the
ulva of E. falconeri, save in size, appears to agree pe pe

THREE EXTINCT SPECIES OF ELEPHANT. 261

very closely with that of E. melitensis, 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 remarkabie
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 find 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 the 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 H. melitensis the disparity is still greater in the same direction.
In EL. 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 HL. 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 H. 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*. Butit is nevertheless interesting to institute some comparison 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 of the upper articular surface in

PAE

a a

262 MR. BUSK ON THE REMAINS OF

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. africanus, E. indicus, E. melitensis, and E. faleoneri. The two latter are of the natural size, and the others
enlarged to the same width from the figures in the ‘ Ossemens Fossiles,’

21

E. indicus. E. africanus,

23

E. melitensis. E. falconeri,

THREE EXTINCT SPECIES OF ELEPHANT. 263

parative flatness or rather convexity of the surface below the radial fossa, and in the
consequent want of elevation of the outer and inner angles which constitute the borders of
a deep concavity in £. 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 brachialis 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 Royal 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, 97-8; and according
to the African Elephant in the British Museum, 97-2. We may conclude therefore
that the probable length was about 10 inches. 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
entire 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 (Pl. 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 1” 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/:55.
In form and proportions it differs in no respect from the corresponding bone in EL. indi-
cus (Chuny), which has a length of 30. 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. falcoueri a length of between
11” and 12”, or about the same as that which [have deduced from the other data which
have been already discussed.

10. Pelvis.

The pelvis of Z. falconeri is represented by a considerable portion of the left os innoni-
natum, which is shown in Pl. L. fig. 51. The fragment includes the entire acetabuluny,
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

264 MR. BUSK ON THE REMAINS OF

character between the pelvis of the Indian and African Elephants has been already fully
referred to (p. 242). Above the acetabulum isa 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”°1, length 27°53; 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 1”-8. The body of the ischium is unfortunately broken off obliquely on the outer
side; but about 1:25 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
what 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 inthe Mammoth. The
length 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. melitensis, which in that respect, as before pointed out, more resembles the
African than the Indian species. Another particular in which 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 shallow.

11,12. Femur.

The collection contains at least three well-recognizable portions of the femur of £.
falconeri, belonging to individuals of widely differing ages. ‘Two of these specimens are
shown in Pl. L. figs. 29,29 a, & 30. The latter of these is the upper portion of the shaft
of aleft 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. 26). 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. 265

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
tr. d. at upper end

femur. The various dimensions afforded by the specimen are:
2’-4., ap. d. about 0-95; tr. d. at lower end 1°25, ap. d. 0-85: 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 E. melitensis, taken,
as nearly as can be judged, at the same part of the shaft. The anterior surface of the

BEE NPR A By

2o)
E, falconeri. 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 ¢rochanter 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 Z. 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 MR. BUSK ON THE REMAINS OF

infratrochanteric fossa, the existence of which, so far as I am aware, is peculiar to /.
falconert. 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. Tn 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 Z. falconeri 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 1”°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 15, and, from those of
a Ceylon Elephant also in the British Museum, 1-7, and, from those of the African,
1-6. 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 molarsin wear. In all 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 #. falconeri is that represented in figs. 29 & 29 a,
It consists of the entire 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

THREE 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 Z. 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 Z. 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,

27 as

1. Transverse section of femur of Z. melitensis
(Pl. XLV. fig. 6).
. Transyerse section of femur of EZ. falconeri
(PL. L. fig. 29).
3. Transverse section of femur of £. falconeri
(Pl. L. fig. 30).
as. Anterior surface.
ps. Posterior surface.
2s. Internal surface.
es. External surface.
ea. External angle.
ap at f. Pretrochanteric fossa.

to

ea

* 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 Y. 9p

268 MR. BUSK ON THE REMAINS OF

of the femur of Z, melitensis, the section 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
EF. falconeri, especially in the presence of the pretrochanteric fossa (indicated by the
letters @¢f), and of the posterior, in the great elevation of the postero-external
angle € a.

‘The accompanying cut represents the outline of the lower epiphysial surface.

—— ae

23 —~

~

Ps

The various dimensions of the bone are given in Table V.; and, taking these data in
comparison with the 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 perhaps affords some ground for believing that
the general proportions of the length of the limbs of Z. falconeri to its height, at any
rate when young, were not widely different from those of the Indian Elephant.

13, 14. Bones of the Hind Foot.

(1) Astragalus.

The only specimen of the Astragalus, contained in the Zebbug collection, is that
whose upper surface is shown in Pl. XLVII. fig. 14.

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 Elephant’s astragalus,

THREE EXTINCT SPECIES OF ELEPHANT. 269

it would seem, is developed 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 7. 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 the 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 m 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 upper edge of the scaphoid facet projecting im 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 inits 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 of 2. falconeri 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 &. africanus
as 602 to 1000, in #. 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 Z. africanus as 900, in
the mature /. 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, which, in the Indian and
2P 2

pa

270 MR. BUSK ON THE REMAINS OF

African Elephants, as compared with the total width of the astragalus, stands in the
ratio of about 758 to 1000; whilst in #. 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 #. 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 #. indicus, the width of the astragalus is about 4”, and the length of the
humerus 28”, and of the femur 337-5, in accordance with which the humerus of /.
falconeri should be about 14”, and the femur between 16” and 17”. Again, in #.

* 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 above account of the astragalus drawn up, represent what I regard as the
astragali of H. melitensis and E. falconeri. They have belonged to fully mature animals, and are drawn of
the natural size :—

E. falconeri.

THREE EXTINCT SPECIES OF ELEPHANT. 271

africanus the breadth of the astragalus is 5:2, and the length of the humerus and femur
respectively 36” and 42", according to which, in 2. falconeri these bones should measure
145 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 #. 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 #. falconeri is the fourth left metatarsal
(Pl. LI. figs. 40, 40 a, 40 4). The bone is quite entire, and only slightly chipped
on 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 proportions than
most of the other bones referred to #. falconeri), and also with the portion of the
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 #. 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 #. 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 /. falconeri and Ff. 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 /.
falconeri may have been proportionally longer than in Z. indicus, and consequently
from their sizes those of the humerus and femur computed somewhat above the reality.

_§ V. Vury youne or Immature Bones.

Having adduced the evidence afforded by the mature or nearly mature bones in

272 MR. 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 Elephant, 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 which are represented in PI. LH. figs. 42',
49! a, 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'} 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

* The fact is, that both are broken across the weakest part of the bone.

+ To save words, as I am unwilling to employ a specific name, I use the Nos.*of the figures to designate the
two bones.

THREE EXTINCT SPECIES OF ELEPHANT, 275

is broken off 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 1-95, and at the constricted part above the condyle 0”:9; whilst fig.
44 in the corresponding directions measures 1”-85, and 1-1. It is true that the latter
isa 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 fig. 42/ the
condyloid articular facet measures 1” x 0-5, and in No. 441-105. And in No. 42’
the surface of the exo-basioccipital synchondrosis is 0'"6 x 0'-3, and in No. 44 0'-6 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 @ 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 ¢), in
fig. 42’ a, is separated, as it were, from the cerebellar fossa, or rather 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 (jf) (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 different
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 affording 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 differ 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 MR. BUSK ON THE REMAINS 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 Pl. LIT.
might possibly have belonged to extremely young foetuses of a larger 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-

A

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 (¢f) and the exo-basiocci-
pital synchondrosis (40 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 mc) 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 corresponding 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 fragile ;
whilst in the Maltese specimen the bone is firm and solid, and bendently much further
advanced in ossification.

For the purpose of comparison, as to size, with other bones of the pigmy Elephants,
I have thought it might also be useful to give the subjoined outlines of various other
bones of the same African uterine fcetus.

THREE EXTINCT SPECIES OF ELEPHANT.

Premaxilla.

Deciduous
incisor.

x
Humerus.

Various bones of uterine fuctus of 2. africanus.

WOT WV lee Vs

24

276 MR. BUSK ON THE REMAINS OF

2. Upper 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 Pl. LII. fig. 46, with what
appears to be the point of a very young permanent tusk zn 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 04 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 semitransparent 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 entire 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 the 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.

Femur.

Various bones of uterine foetus of Z. africanus.

Radius.

bo
r=)
i)

77

278 MR. 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 PI. LII. fig. 43. 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 borderis 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
neryous 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 LIIIL., the penultimate milk-molar, as I should presume it to be, of ZH. meli-
tensis; but I am by no means certain of this. The thickness 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,
sufficient 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 1’-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

THREE EXTINCT SPECIES OF ELEPHANT. 279

have been unable to perceive any greater 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 Pl. LIL. figs. 42 and 42a; 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, 0-35, 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 1 0"1.

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 the
greater incurvation also, as it were, of the diastemic border, the left fragment, when
viewed from aboye, 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 entire to
afford some diagnostic characters; two of these are figured in Pl. LI. figs. 49& 50. The
former is the almost entire shaft of the left humerus; it is apparently slightly rolled at
each end, and an angular fragment has been recently broken off 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 Pl. XLIX. fig. 26, that
little doubt can be entertained with respect to their belonging to the same species. One

280 MR. 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 the 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 feetal bone. Ihave little hesitation, therefore, in refer-
ring this humerus, fig. 49, to a very young EZ. 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 bicipital groove, bounds internally a perfectly
flat surface in front of the deltoid crest. The same angularity, it should be stated, is
visible, though of course less marked, in the young humerus of L. 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 45 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 £. 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 bone 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. XLVIL 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 differ 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

THREE 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.

nis (29 ) (1s) 2
eu GaG
at 2Z

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 shown by their
dense texture and aspect of greater age, as compared with the far larger radius of the
uterine foetus of E. africanus, of which an outline woodcut (No. 37) is given in p. 277.

(2) Hinder Extremity.

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. XLVIL. figs. 15, 16,17, 20, and 21. Of these I regard 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 :—

5 Least
| Length. Upper end. | Lower end. Lae
ipl Geer 2-5 Of ‘9x7 1:6 |
Fig. 21.... 2-4 “9x6 ‘Tx 6 1-4

282 MR. BUSK ON THE REMAINS OF

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. 16 0.
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

43

20,up ,

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 shown in
figs. 16 and 17, from its greater robustness, belongs to H. melitensis, 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, tibiz, of a totally different type.

Two of these tibie are shown in Pl. LI. figs. 38, 38a, and 39, 39a; 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. 39a@*; 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 ELEPHANT. 283

comparison. Both bones are so much worn at either end, apparently by water-rolling,
that no portion of either epiphysial surface remains; but the portions removed at either
extremity cannot be very great, so that the length of the epiphysial shaft in fig. 38 may be
estimated perhaps at about 3/2; on comparing which 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 ali other tidi@ 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 the
means existed, which unfortunately they do not in this country, it would have been
interesting to compare these immature tibize with those of the Hippopotamus, to which,

at a guess, one might be inclined to assign them*.

§ VI. Dentition. (Plate LITT.)

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 which com-
prises the teeth.

It is well known that that distinguished paleontologist lad 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 paleontologists 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. LILI. have all been lithographed from Mr. Dinkel’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. Warned by the great blunders committed by Nesti, Fischer de Wald-
heim, and other paleontologists, who have been misled by the characters of milk-teeth
to identify them as the remains of pigmy species of Elephant, I have been chary in

* With reference to this, it should be remembered that a diminutive species of Hippopotamus still exists.
VOL. VI.—PART Y. 2k

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 the molars. ‘The specimen is repre-
sented of the natural size by figs. 1 & 1 a,16. 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 fang, 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 length 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 :—

Binitine dens the..ce scree eaeseene ase i toa
Wengthioherown 45sec eee 0-6
Width of crown ...............06. O"-4
Thickness of crown ............... 0"-3
Diameter at the collar ............ 0'3
Diameter at broken end ......... 0-25

“These minutiz 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 foetal African Elephant referred to by Dr. Falconer, there are nume-
rous bones of another, very much smaller and obviously 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 cap, which appears to be formed of two layers, an
external (probably of osteine), 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 1-8, and the greatest
diameter of the crown about 0!-45, 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 layeris smooth, and
throughout a line of thickness shows 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 aconical point. ‘The dimensions are :—

eniathinesseasre anaes 2 5-0
But -end tes. sare ener te Toll:

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.

* Itis a rather curious circumstance that the specimen above described by Dr. Falconer, and which, from some
words which 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 precisely corresponding dimensions, and with a ticket upon it in
Dr. Falconer’s handwriting, “ Hlephas melitensis, tusk, Zebbug Cave,” is in the collection; it is the one figured
in Pl. LI. fig. 48. The specimen, as will be seen, is far more perfect than that described by Dr, 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 the collection may have belonged to the same individual. It is true that the specimen figured in PI. LIT,
has been broken across ; but the fracture, instead of five, is more than seven inches from the truncated extremity,
and probably fully 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 subsequently broken off. In the presence,
therefore, of this more complete specimen of an obyiously 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 very strongly suleate ; 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 1-1 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

2R2

286 MR. BUSK ON THE REMAINS OF

“3. Lower Mitk-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
cemposed of three collines, with a posterior talon. It is clear that the tooth did not
belong to a feetal 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 posterior talon, ‘The dimensions are :—

in.
Extreme length ............ \ ts asteigan okt eelar aoe 0-40
Width of crown in front ......00cc:cccseeeeeees 0-25
Greatest: width Jss5 5 jess 4 :i8'5- aed asaeebese ey 0°32
Greatest height of crown.................. ae ere 0:40 *.

inch, making the entire length of the tusk between 10-5 and 12", whilst the depth of the pulp-cavity is not more
than 2!'-7 in the present state of the specimen; andas this is remarkably perfect 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 ivory is brought to the market (4), it probably indicates that
the tooth was of considerable age and consequently belonged to a mature animal. Dr. Falconer 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 haye just noticed, I find two other fragments of what L conceive to be very young permanent incisors.
One of these, or rather a portion of one of these, is seen attached to the premaxillary bone represented in fig. 46,
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 brown colour. The remaining depth of the pulp-cayity is 0'-9 ; 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 1!"3, and greatest diameter 0'-4. At first sight it does not appear
altogether impossible that the two fragments of the small tusk may be parts of one and the same; but, in the
first place, the fractured 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 premaxillary 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 haye therefore substituted it in
the specimen for the other.

* The dimensions above given are not quite half those of the corresponding tooth in Z. africanus, and as nearly
as possible half those of the second milk-molar in Z. indicus, 2. primigenius, and E. antiquus ; so that, admitting
it to be really the second and not the true first milk-molar, it is obvious that the Maltese specimen must have

THREE EXTINCT SPECIES OF ELEPHANT. 287

“De Blainville (‘ Ostéographie,’ Eléphans, pl. ix. fig. 1) has given a figure of a lower
jaw of a very young African Elephant, in which a pre-antepenultimate 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 both
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-antepenultimate 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 crown 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; imdeed 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 im front (of anterior ridge) .......2.0.. 02.0000. see 0°35
Gxreatestiwidtihy hehindeceeneeceeeee see te hens). cee aaaeene ae 0-5
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 crown of
this tooth; but judging from a germ specimen, to be described in the sequel, it con-

belonged to a diminutive species. It is a curious circumstance, however, and one well worthy of note 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 subequal in size and strongly divergent. Dr. Falconer states that there is
some indication of the existence of a distinct small anterior fang—though I am myself by no means satisfied of
this, but on the contrary 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 haye 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.
Tn 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 MR. BUSK ON THE REMAINS OF

sisted of five ridges with front and back talons. From the very narrow width in front,
and the rapid increase backwards, it is inferred 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 five ridges with a small posterior talon. The
disks are wide and open in the antero-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 supporting the last three ridges and talon is pre-
sent, nearly entire ; but the front 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
fang. The grinding-plane is slightly concave in the antero-posterior direction, proving
it to have been an inferior molar; and I infer that it is the equivalent tooth of the
specimen last described, 7. ¢. 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 ................4. 1:30
Width of crown at front ridge ............... 0:57
Greatest width of front ridge ............... 0-70

“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 grooves 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 OF 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 narrow 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 greater 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 :—

HEME CHR OL, CHOWAN ceo ce soto ete: scale snteonzoetenice ornate cae 2:2
Nill laminas trom tirsnse coneriss cere hace te sere sue semen aif
Whiditihvmmsthretmard dilewfeepcra.ses-cesa-c-seceeeseesee 0-7
Wadthvatvetshthinid serew.ares: cease seeere -eacs eee: 0-7
Height of crown at the last ndge .................. 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, conjoined 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. 6 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 distinct talon in 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 inferior penulti-
mate shown in 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 :—

leathers CLONval eee eee: «onc ceehntet «e+ 02s «sacs 1-4
Wiel tiomtattee tar ices cee ee teattbe o sieeve se 0°6
Greatest widthpbehind 5... scesss<cxaneencsiece et 0°8
Height (greatest) of ridge plates............... 0-95

The tooth is of the left side. There are in the collection a number of detached plates
of an unconsolidated germ tooth of the same age.

290 MR. BUSK ON THE REMAINS OF

“Of the last upper milk-molar (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 8a, which consists of the intermediate portion, comprising six
ridges, but mutilated both in front and behind. It is evident that of the hind portion
only one or two collines are wanting, these being 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 affected 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 in age with
fig. 5 of the lower jaw. ‘The dimensions are :—

Length of fragment of crown ...............68. 1-4
Wadthvim fronts. wad. ddaa steeds ee kee eee = 0-95
Width behind ............. onicterad eke meee: 0-85
Extreme height of crown-ridges ....... SiS ARR 2°3

There are no means of determining with certainty what was the precise number of
ridges that entered into the composition of this tooth*; but assuming from the data

* Besides the teeth noticed 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 machwrides, 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 all 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 milk- or as a
true molar, opinions may be divided.

THREE EXTINCT SPECIES OF ELEPHANT. 291

given above that only a single ridge and talon have been broken off at either end, the
perfect tooth would have presented eight ridges, besides talons.

“Figs. 7 and 7a 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. 8; but the form and size are irreconcilable with fig. 6; it is therefore
inferred to be a part of m.-m. 4; it presents no special characters for a description.

“5. True Molars.—The evidence above adduced from various 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 those
of the lower jaw :—

“ Lower True Molars.—¥ig. 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 crown 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 1-85. The length of the grinding-surface is about 2!'; and on it are exposed the macherides 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 pressure by a succeeding tooth, either 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 surface of the enamel is finely and irregularly fluted, and in some places, though
very rarely, an extremely faint indication of transverse wrinkling 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 any satisfactory 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, differing 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 ifs form and the thick-
ness of the plates, as shown more clearly in the entire tooth than in the fragment figured 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 equally (if
not more) likely.

We have to consider therefore whether the tooth described in this note, is m-m. 4, or m1 A

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 Z. indicus, they will be found on the average,
as regards length, to measure about 5-1 and 6-6 respectively, or to stand in relation to each other as -772 to
1:000 ; in Z, primigenius, 3''-6 and 5!'-2, or as °692 to 1-000; in Z. antiquus 5!'-3 and 6'-7, or as *791 to 1-000 ;
and in £. meridionalis 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 (7), 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
considerable difference will be found to exist. For instance in &. i ndicus, the last milk-molar plates are alhout

VOL. VI.—PART V. 25s

292 MR. BUSK ON THE REMAINS OF

supported upon the 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. his is the typical form which the last true molar (m. 3) com-
monly assumes in 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 Z. antiquus 0!-52 and 0!'-6, in E. primigenius
0-33 and 0'-43, and in a single instance of E. meridionalis 0''-56 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. africanus, but should estimate the difference as much
greater even than in any of the species above mentioned, or as about 0!-65 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
m-m. 4 of the same species as No. 9, would have proportions widely different from those which that tooth
possesses in all 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 will, 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 width. The former, again, is 2!-9 long and the latter 2-2, showing an equally
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 yery much strengthened. Fa

The 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 £. indicus the average length of these teeth may be stated as about 6"-8 and 8!"- 8, in Z. africanus as 6!"-4
and 7!'3, in E. antiquus as 6'"7 and 8''-3, in E. primigenius as about 5!-3 and 8'-2, and in E. meridionalis as
6"'-5 and 8''-8,—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 series—that is, are 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 very 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 degree
even than is indicated by the figures. The residuary worn surface of the crown
exhibits seven abraded ridges, of which the 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 crown gradually diminishes back to the talon, as normally
occurs with the last true molar of the Elephants. The dimensions are :—

in.
streme) leneth) of trae mien tip ycc baa. ean --pares nonce « em aenaeess qian asia 4:2
Wadthyoly crown abesecondinid ep pee. eueh suiersionseee erie. --merieee ae ee er oer 1:3
Widt huotcnowil atiseveit Weld Oe meeere ret caactesseiis- cesta cber see cideinae\alae 1:2
IWidthtotrerown be laine. .tpprpiee aasep ee ects cie eb ema) sels Seine 0°85
Space occupied by second, third, and fourth disks...............:+.:+66+ 11
Length of residuary grinding-surface, including seven ridges ......... 2°3
Extreme height of crown near middle, where unworn ...............++- 2:1

The fangs are broken off along the base near the roots; the contour of this part of the
tooth when reversed is very much 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 very
distorted, descending nearly vertically in a spoon-shaped concavity 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 dis-
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,—

The height of the crown at the outer side of the distorted portion being ...... 2:9
iAmdgof thewininerenr-eoenee teem eee ee ree etietar elses tects seloidoe «0Ail- wap sel--Iatd-leel-\sceerise 195

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
has 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 ............ceeseeeeeee 3:3
Width at anterior plate ............ecsesseeeseeeeeees 1:35
Width sbelin dle s-ccccecude aoc csismenceen woeaensameneses 1:2
Extreme height of Crown.........cscseneeeneeeeeeeeees 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 a,
which is inferred to be another example of a last lower molar of the left side, in a
different 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 direction, with 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 wide intervals of cement: this is well shown on the side aspect of fig. 11 a.
The posterior talon forms a prominent splent, consisting of about a couple of digita-
tions. - All the fangs have been removed by fracture close to the base. The crown
surface in front has been ground down to the level of the fangs; and there is no trace
remaining 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 ELEPHANT. 295

twelve or thirteen 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 that presented by any other species, except certain varieties of Elephas
antiquus. The dimensions are :—

Wenptiie of Crown aeece see ce = ee0 Baad eae sopsin’ sana 4-4
Width in front at fourth rae Seana Sy atirt eas 1-4
Greatest width of fourth ridge ............08. aie asiete 1-45
Width at eighth ridge ....... pieces dic cvaw kes tule 1:2
Width at last ridge ..............0. Spousocsddecdsdonoee 11
Height of crown at seventh ridge .................5 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. 12. There is not the slightest indication
of a disk of pressure anywhere upon the hind talon, the surface of which is perfectly
preserved with its coating of cement from the fang 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.

“ Upper 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 slightly
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,
presenting 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}, 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 macherides.
+ They are not quite so thick as they are represented in the figure.

296 MR. BUSK ON THE REMAINS 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

Eixineme, lesa tof ery ince. busin acca iedats veka ees oe er
Widthwin front ss Neng RSs. xceieer quatre ean rote geste iene ese ON(atleepiate))
Wadthem she imiddlenaysnismet lp ieee a cement lec
Width.behind icc; “Valse 2S eee a eee
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 paleontologist 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 Library 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 found 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 fang that supports
them. ‘The corresponding 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 proved by
the circumstance that the grinding-plane of the crown intersects the most anterior of the
extant fangs. The rest of the fangs from this point backwards to the posterior talon are

* There can, I think, be no doubt that the deep hollow below the hind talon is due to the pressure of the
succeeding capsule,

+ There can be little doubt as to the true position of this tooth as m.1. In form and general character it is
the exact counterpart, except in size, of an m. 1 of #. antiquus—identified as such by Dr. Falconer.

THREE EXTINCT SPECIES OF ELEPHANT. 297

all present, but more or less fractured or abraded. ‘The molar is vertically fractured
across through the middle, involving the loss of the greater part of one colline; but as
the fragments fit at the base, this circumstance does not interfere with the precise appre-
ciation of the 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 by 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 cornua at the sides, such as are commonly seen in Elephas antiquus.
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 surface 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 which I am acquainted.
The enamel of the plates is rather thick, quite as thick in proportion as in the existing
Indian Elephant or #. 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.

“T have reckoned that what remains of the crown is composed of ten ridges; and,
taking into account that the most anterior portion, supported upon the large front fang,
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 collines, 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 MR. BUSK ON THE REMAINS 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 :— in
Extreme length of crown, measured from back talon to anterior edge, exactly 4:0

Width of ditto at second-ridge \' £2 2eogPog (6, yea) HOGG Es Sota ceil
Width of dittolat third ndgé” ss" 2) 07202 GE PONT ok RGRAY BIOS Sor Se
Width-orditto at Sixth ridge. 2/07! f: (CAN ARE SostOTaare eat cakes (06, yeaa
Greatest’ width of crown -2 ene ugnbnyind eqemesigoh oogerenngt a5
Width at Othinidge 2127 ORE OTe RO OG Ze, 1s ao), Tale
Width at last ridge . . . See AM AE PERSO
Greatest height of crown, taien at afieadon of 10th riage = RIETIQ? “WIS HADQIG
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 alimentary 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. “ Ridge-formula.—It now remains to consider how the data furnished above 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 eitige

“(1) Milk-molars.—The antepenultimate milk-molar m.-m.2 (fig. 2) is seen to
have been composed of three collines, like the corresponding peas of the African
Elephant ¢, 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 Euelephas 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.

+ In the only specimen to which I haye 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 OF ELEPHANT. 299

in E. antiquus the number is ten, and in H. primigenius and E. indicus it amounts to
twelve.

(2) True molars.—Of the antepenultimate true molar (m. 1) there 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 (m. 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 twelve, while in E. primigenius and EF. 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 lower from nine to twelve. In £. anti-
quus the number is sixteen ; and in E. primigenius the number of plates reaches twenty-
four f.

“From the above data the ridge-formula of the molar series is deduced to have been

Milk-molars. True molars.

B4+5+4+8 | 8+9412

Sees sb 7 SESE
This formula at once brings the small Zebbug species within the subgeneric group of
the Elephants which I have called Lowodon, 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 alayer of enamel. But, though allied, the
two forms are specifically very distinct. Besides the signal differencé 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 disks 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 ridge

* Unless, as I believe, we may regard fig. 9 as such.
+ The numbers of plates were not filled in in Dr. Falconer’s MS.

VOL. VI.—PART V.

bo
I:

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 African species than does the corresponding
molar of E. antiguus. ‘Che 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 with the corresponding molar of the African species.”

Having thus given Dr. Falconer’s descriptions of the teeth, 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 point 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 evidence was 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 macherides.
What the difference may be, if any, in the numerical formula, Iam 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 Pl. LIT.

If we compare, for instance, the m. 1 represented in fig. 9 with either of those shown
in figs. 11-13, but more especially with the first, of which a side view is also given, such
a difference will at once be perceived in the form of the macheerides 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 gives an average
thickness of each plate of about 0:27, whilst in fig. 11 seven plates occupy a length of
3'-0, equivalent to an average thickness of about 0-43. Again, if we look at the form
of the macherides 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.

THREE 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. primigenius.

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, Iam 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-
clusion is byno 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 showitself. For instance,
the mean thickness of the true molar plates in E. africanus varies from 0-85 to 1" or
more, and in £. indicus is about 0:55, whilst the thickness of the plates in the m.-m. 3
of the two is pretty nearly the same, or about 032-35; in E. primigenius, although
the thickness of the plates in m. 1 is not more in most cases than about 0!-43—-45, thase
of the 5rd m.-m. average about 0°34, or rather more than in E. africanus, though con-
siderably less than in E. antiquus, 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 HZ. 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 macherides, the decision would probably have been
easy enough ; but it will be observed (leaving out of the question the m.-m. Z, fig. 2) that
only three of the specimens were worn sufficiently for this purpose; and as, from the
form of the machzrides 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. Iam not sure how far such a character may be depended upon;
but, in the case more especially of milk-teeth uncovered with cementum, I think what

272

302 MR. BUSK ON THE REMAINS OF

may be termed the sculpturing of the surface may be regarded as of considerable value.
Taking 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 the exposed plates im fig. 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, 5, 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 present
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 EL. meli-
tensis, | 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 when 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. 5 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. antiquus, and proportionally a good deal wider than the same tooth in Z.
africanus and E. meridionalis. 1t 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 transyerse wrinkling appears to me, from subsequent observation, to be a very uncertain character ; but
the large vertical markings afford, undoubtedly, characters of some importance.

oo
i=)
wo

THREE EXTINCT SPECIES OF ELEPHANT.

DESCRIPTION OF THE PLATES.

PLATE XLIV.

Fig. 1. Symphysis of mandible of the largest Maltese Fossil Elephant (Elephas
(p. 231).

Fig. 2. Neural spine of the seventeenth or eighteenth dorsal <ertebra 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).

. 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 H. melitensis (p. 238).
Figs. 10, 10a. Sixth or seventh dorsal vertebra (p. 240).
Figs. 11, lla. Second or third lumbar vertebra (p. 241).

PLATE XLVII.

Fig. 12. Portion of atlas of E. melitensis (p. 238).

Fig. 13. Portion of ascending ramus of mandible of ditto (p. 236).

Fig. 14. Upper surface of astragalus of ZL. falconeri (p. 268).

Fig. 14(d7s). Dorsal aspect of fragment of scapula veferred doubtfully to E. falconeri
(p. 254).

Fig. 14 (dis)*. View of the glenoid fossa.

Fig. 15. Posterior view of very young tibia, probably of EZ. melitensis.

Fig. 15a. Proximal articular surface.

Figs. 16, 16a. 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 tibie (pp. 281, 282).

304 MR. BUSK ON 'THE REMAINS OF

PLATE XLVIII.

Fig. 22. Proximal end of humerus of EL. melitensis (p. 244).

Fig. 23. Fragment of scapula of ditto (p. 243).
Fig. 23a. The glenoid fossa.

Fig. 24. Proximal end of ulna of E. melitensis (p. 245).
Fig. 24a. 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 (dis). 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 (p. 260).

Fig. 28a. Articular surface. |

PLATE L.
Figs. 29, 29a. Anterior and posterior aspects of femur of FE. falconeri (p. 266).
Fig. 30. Fragment of upper end of femur of ditto (p. 264).
Fig. 31. Portion of pelvis of ditto (p. 263).

PLATE LI.

Ia

Vig. 32a. Anterior aspect.

Figs. 33, 33a. Similar aspects of another atlas of much younger age and doubtfully
referred to HL. melitensis (p. 251).

Figs. 34, 34a. Anterior and posterior views of a fragment.of an eighth to tenth dorsal
veitebra of E. falconeri.

Fig. 35. Fragment of atlas of £. melitensis (p. 238).

Figs. 36, 36a. Lateral and posterior aspects of neural spine of a dorsal vertebra of

E. fulconeri.

7. Proximal end of second rib of ditto.

Figs. 38, 38a. Anterior and posterior views of a very young or fectal tibia (? Hippo-
potamus).

Figs. 39, 39a. Lateral and posterior views of another tibia of appaiently similar
character.

Figs. 40, 40a, 404. Fourth metatarsal of /. falconert (p. 271).

Fig. 41. Proximal phalanx of third manual digit of ditto (p. 265).

THREE EXTINCT SPECIES OF ELEPHANT. 305

PLATE LIL.

Figs. 42, 42a. Fragment of right ramus of mandible of very young animal.

Figs. 42', 42/a. External and internal aspects of a very young or feetal oxoccipital
(p. 272).

Fig. 45. Fragment of right side of mandible, rather older than that shown in Fig. 42
(p. 278).

Figs. 44, 44a. External and internal aspects of a very young or fetal exoccipital
apparently different from that shown in Fig. 42' (p. 273).

Fig. 45. The entire symphysis of a very young or feetal mandible, displaying the socket
of the m.-m. | and, on the right side, part of that of m.-m. 2 (p. 278).

Fig. 46. Portion of the premaxillary with germ of permanent incisor (p. 276).

Figs. 47, 47a. Distal extremity of fibula? of E. falconeri?

Fig. 48. Tusk of E. falconeri (p. 285).

Fig. 49. Very young humerus of E. falconeri (p. 279).

Fig. 50. Another very young humerus (p. 279).

PLATE LIII.

Figs. 1, la, 16. Deciduous incisor of E. melitensis? (p. 284).

Figs. 2, 2a. Antepenultimate lower milk-molar, m.-m. 2 (p. 286).

Figs. 3, 3a. 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, 5a. Last lower milk-molar of the left side, m-m. 4 (p. 288).

Figs. 6, 6a. Penultimate upper milk-molar in germ, m.-m. 3 (p. 289).

Figs. 7, 7a. Fragment of germ of last upper milk-molar, m.-m. 4?

Figs. 8, 8a. Portion of last upper milk-molar, m.-m. 4? (m. 1, mih7) (p. 290)?

Fig. 9. First upper true molar (m. 1)? of E. falconeri? ( p. 298).

Fig. 10. Crown surface of tooth trom Maccagnone (p. 301).

Fig. 11. Last (2) lower molar of the left side, m. 3 (p. 294). (E. melitensis?)

Fig. 12. Greater part of the last lower molar of the right side, m. 3 (p. 291).

Fig. 13. Posterior half of last lower molar of the right side, m. 3 (p. 293).

506 ON THREE 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.

Tas_E IIJ.—Atlas.

Various measurements.

a er ath
fo} | Ae!
7 2 | |
a | | os

Gs} ‘5 ioe.

= ay (Eee

z eo |828

5 Sq | oes

| f | Be |d8s,

g I 4

&. indicus (Chuny) ...... 16-0 :
A (Sumatran) ..| 10:5 | 64 | 7-3
(2678, R.G.8.) ..| 12:5 | 7-0 | 7-8
3 (Cuvier) ...... 13:5 Staal Mae
» (Blainville) ..../ 14:4 | 7-7 78
E, africanus, BM. ........ 13:5 | 85 | 95
AG (Blainville) ..} 11°8 66 | 76
IEE BLE ENUSTR Face aya atcn 5 gence etic fetes) ne
| BE. falconert 0.6... .25 2s. 5:0} 2:9 3:3

tse lle | ct Sod se le z
ee) a | 2 le 12 ce
Se Fol ea peal 38
soa Pease PMS ae
=e | as one S
ore S| ce cual Sepals ae
Ss > pe Cees Spee nO
ee |e Bee eee See riesal me mene
SSB) ag] a | aie) Ss] ga
a 4a}/<4 i4 /d a
ade ah le
5:2 | 2-4 |/1-9 |3-4 165 |3:3x2-4
6°55 | 2:7 |2-1 13-8 | 7-6 | 4:0x3-0]
ae oh ae lh oe beeeests | S226
88 |36 | 25 |41 | 8:3 |45x30
beta ech beaten] (cee boar es
Ey = oeiligsxsall he Cea hmres' | OOS lh
| NZI ha sh Uefa eth lata} Se atsal

Height and width
of ring.

32% 2-4
35x30

43 x 3-2

Radius of longer

curve of condy-

loid facet.

TasLe 1V.—Dimensions and Proportions of Astragalus.

eed ieee, | Pon
9 al
| ig se
| 3 ; | 2 |
feoks E hes ia
& & aaa
ae Be i € = ‘3 3 lee 2
} =A | : a | o
Ec ah lb ee al Bi ga i aa
al\a\s| 4 3 3 8 8 |e5/%
=e pest es ec = E F Bl epees
|g] S| cs 2 = |
(a{ale & & or 6 eles | é
| m. indicus (Chuny)../51/6-0|..|...... Pian ares ne ORR ‘850.
|, (Sumatran). .| 3-4 /3-9 24 |3-1x26/ 29x 1-8|2-5% 1-5 | 2-916 | 22% 1-8 -871 “615
(2707, R.C.8,))43 | 49 28 | 37x33 | 34x21) 26x13 | 35x22 | 27x 1-4 +871 -573
| 2 (young) B.M.|30 3-5 21 |28x21/27%1-8/ 20x11 25% 1-6) 1-6 x-75 +856 600
“3 africanus, B.M. ..|46 |5°2 3-1 | 40x 3:7 | 42x 2-5| 3:5 x1-8| 3-0 2-5 | 3-0 x 2-7 -884 +596
E. melitensis... +0 27 |B1 20 | 24x22 3-0x 1-9) 1-9 x 1-1) 18x 1-4 | 21x 1-0 +870 645
BE. falconeri oo. 1-9 |22 1-4 /1-7x16 21x10/14x -6/1-4x 8/15 x -65 -863 -636
By iy dns 2. es [tes pale eg Sete Gin s951( Lis. 0d lee 1-4 x70 -857 571

1-2 |1-7x1-4/1-9x-95| 22...

facet to its tr.d.

facet to its tr.d,
Proportion of y.d. of scaphoid

| Proportion of a.p.d. of tibial

‘733 |-620
‘891 “610 +
‘750 “648 *
-900 -602
912 -633 |
“941-500 -

“874-500 -

Proportion of tr.d. of scaphoid

facet to total tr.d. of bone.

phoid

Radii of curves of sca’
facet.

won

Nias 2
wank San

Tauue V.—Dhe Principal Measurements of the Humerus and Femur, as affording data for computing the Height and Proportion of Elephants.

[To face paye 306.)
oe Femur. ; Height of Animal.
i]
| Tee a 4 3 z According to
: 3 aa g|s § |) ; Accor! ni ale, z
? i ie | ale HAE ee ee nen” | Anqanee | coum” | 3
€ = ra = s g 3 ihe, g z 2 Sl | Ee ee sealed data, ROS. ane A
2 a | 2/2 |: S12] BH gs i A z i 3 7 zlgs)e |ce| & ia a
2 5 & > is ‘ e s = 7 2 = = == - —-
A a See Wa Sl) eel | PPP) 2) ele la al el blghle lanl i i Palle
¥ | 3 A ea eile 2 | 2 | 3) 3] s ig é Soil salle: |litog = OW EM eal relelpsts [praise a ||| eal ene (ered lt areal sree limes
A eS Sa Wee ey) ea eh el |e PASM alcatel i)< Mesa cir rice Weal deel Celtis
a SS Sy Be eee Se as = J sz | = Z| Z| + 5 So Sis || © 3a 12 ll SEV bezpa| Viva ater Aisle esi ce 2
& = ad ral a ee | se ; a d a eh lhe aes % z a Ws ey a | a | ee) 35 az| | 8 @ € e
EOECIEE RS Ue See Ee ee EOI De Met EU ERI sca es | E Eee ete ies ET ee Ia
| |
Se le | « «|e
E.indicus (Chuny) ....| 350 | 80 |101 | 86 |120 | 30 | Go }150 |in0 | 52 | 63 | 55 | 40 | 120 42 | o1 | 47 | 32 |127 }130 |] 90 || sa | 86 ]78 |... |... | .. | 119 | 115 | 103 | 103 | 92 | 96 | 103
CIC WG cel on Wie .
» S744B, ROS! 350 | 79 | 104} 98 | 110 | 39 | 60 |170 |e} 85 | 63 | 57 | .. |120 | 41 | 43 | 43 | 60 | 5a | 38 jis |i30]/91 | 79 | se )/73] | .. |). | | 20 115 | 105 | 103 | 95 | 96 | 105
2 fo | rer liracn a itras
5 (GSES) conf) BEE AE YP sa | oo GIP EEO | so IP co Pca IGE ce Woe Woe a Weel ce U] Bea Wee We |] eM valli ce ce Heco I] oe Woo Hone |) bo )) re) ee | Os |) eS | oi |] see
Wee Ol a |
» » Ceylon, BAL..| 3L0 | 65 | 70] .. | 82] 26 | 44 Jane} 9p) 73 | .. | .. | . jare] .. | ..- |) 385 | 50 | 875) 27 (106 | 110 | 79 | 70 | 72} o2 | .. | .. | .. | .. \iao7 | 104} 94 | o1 | 85 || a5 || 85
CN Old Wc 6 le .
» Sumatran, BM} 278 | 52] 69] 68 | .. | 23 | 36 |100| 72/54] .. | .. | .. |100| 26 | 24°] S37) 41 ]32 | 23 | 69 | 90] 50 | 50 | 57 )5a | .. | | |). | oe | ov | a | oh | | | og
» » (young) 272308) 161 | 36] 40/32] .. | 14 | 92) 50] sr] 47 | 25 | 24] .. | .. | 19 | 10 3 2 , : |
CO) SS oceliec
» oy (young) BAL. 45] 50)... | .. | 17) 28 | 73] e2| 56 |s1 | 32]. | 75] .. || .. we feosfe2es ee |e) | vo owe ee | | eres) @ on ler, || asp! (Boe os) ba | au ee
| @-| 6 . . .
E. africanus, BM... 79 | 92/93 }110 |) 33 | 40/131) .. | 78) 56 ) 48 | . | 120] 40 | 44] 425 | bo | 42 | 34 |it9 | isa | 63] 80 | a4 joo} . | .. 1 .. | . | 119) 116 | 103 | 103! os
* my s. | | |
B.MaltaQaygo) .-<---| 27 | .. |. |. Be lest || aot] (es alt ec Plies . |. | 28 |27 ney | ert || ee ; : [Pera |
* | Jv esy)|\rer lieu) ecilire
E. mulitensis, fig. 4 | a7 | 45 1s }15 || 20 | 56.) 63 | 49 | 45.) 44 |
0 | | | he «|
i. pi igi GS) e-s-| | 18-20 11) |] hey |) 3 |) ies & | 56 44 |
| | * * | | | he . . . {
E. falconeri, figs. 26, 27. my jae | } al ea Peal eat ie |. 30) 36 32 | 20
| . | | | |
yn 185.30. J. } .. | a0 | a5 | 44 25 ]18 | 14 [12 ) se} .. | .. | 13 13 | 09 23 / 95 | 18 | 22 | 092) _ : F
* | iy i *
E, primigenius, BM... 320 | 73) .. | 96 }103} 48 | ....}136 | 95] 8a | 59 | 52 | 30 |iz0) . | ,. 38 ces {ro If Fs |) can co || eat eee [Pee | ee ine cap | ogee | ce 52 5 |, [ems

N.B. The numbers marked with an are computed, the others taken from direct measurement. ‘They indicate inches.

‘Panne VI.—Dental Measurements in various Elephants.

oT t m.-m. 8. m1. m. 2. m. 3,
} i
= = = ] =
5 = . > ins 5 & 7 io = ! a 5
= pe ]
Sie. | P. | T. Si, || |Z] Siz. | PB, Z| sao |r| nl} Sy sie [P| |e] sme [P| NIE | sie |e yn|e| sie |e |r |S su |p |n}s] sie |e} fio, [P| |e) see | P| n|S
a 4 4 a a 4 4 4 4 4
ip isedican ‘8x55 27x13) 8 ‘87 ) 25x 8 | 2 | 31) 52x15 | 12) 2 }-40 QOOece. 13) 1 G3x21 | 12) 2 |-48] 68x20 | 12) 2 52!) sox mI lie 95x30 | 16) 2 | 135x3-2 | 24). 155 | 15-0x 27) .. |-55
y indicus oe. |
ee esck| aetterss 27x15 | 8 87 | 28 x13) 0 | 1 |-31 |) 45x15 | 12) 1 12} 1 }:35 | 68x24 |/72)| 2 |-52 | 78x 12] 2 |-60 | 75x30 | 16) 2 |-44| 90x39 | 16) 2 | 12-030] 22] .. |-48 | 107x2:5 | 17 61
24x13 | 8 . | 255x135] 8 | 1 |-32) 47x10 | 11) 2 }40) 53x1-7 | 12 70x25 |} 12) 2 |:53 | 75x24 88x30 10-17) 1 |56 |) 95x 16| 2 55 |) 5 o IVT x27 | 138 | 2 |-7
i) |
24x14 | 7 F30} | eeceaee PSY Is 63x10 | 12] 1 |41} 68x25 | 12] 2 |-52 Be) {| 110x381 | 16 “66
i ae .. | 24x12] 8] .. |-30]) . 52xL6 | 13 40 || G:4%2:5 | 12] 2 |-49 14} 1 }-45 || a 84x29 | 13] 1 |-60
” 23x12] 8 | 2 |-97]) . 51x17 | 13} 1 |-40} 60x20 | 14) 1 |-42 14 | -. |:48 ||
2 20% Eh || arrillife ||" S Cuil es +. |) b4x295) 13] .. |-42) 63x19 | 14) 2 | 47
\ ea f S| a
‘an i f
» “9x -6 (d) 22x11 | 6-7 “BL | 20x-9 6} 1 |-32 s+ |e. |] G7x20 | 9 | 1 }-63) 6Ox20 |} 9 | 1 }-71) 63x20) 6 | 1 1 j10 || 85x29 9} .. 94] 93x31 | 9 . | Lo
ie (=
B.ofricanus) -- ies Bo AIT ot bee | eel ees ss! cll oa'l| 20 (ioe 9) 1 |74) Gsx22 | 8 |/a) 1 | rexes | a | 1 1 |-83/ 98x81 | 8] 1 |10 Ps
, “D55075 4 sal! f en] hse 66x22] 9 | 1 |-74) 68x20 | 9 | 2 |176 | S0x20 | 8 | 2 1 |-83 | 11-03 10/} 1 |10
: “33x63 e+ {32 \| : + |=. |] 63x20] 9 | 2-75) o4xo0 | 9 | 1 || 78x27 | 8 | 2 [78 Ba |. rot
” alls ale G8x28 | 7 | 2/80) oc 1 |e we |e] Bb OT 11
R |
2 /-35 | 50x20) 9 | 2 |:50. 11} 1/6 | 83x24 | 12) 2 63) 52x24 72 | 2 | -63 108%3°0 | 16 | 1 |-67 }135x29 | 16 | 2 |-63
2/33) 47x18 | 10) 2 [42 . 5 | 80x24 | 12] 2/61) 93x26 | 13 |1-2)-66
= |. || 58x21 1/10} 2 |-49 | 5
Ls Ll
imigenh 2 |-30 | 34x18 | 11) 2 |-30 13)} .. |:39 | 80x30 {16 |°2 |-47)) 22... 120x40 | 26) .. |-34 |
B prinignis 2 |-30 | 7x20 | 11| 2 |-30 12] 2 |40| soxs2 | 16) 2 |-47
i 2 |-31 |) 83x18 | 11] 1 |-30 Ball cs | 67x33 } 16) 2 |-39
2 (31 ] 40x20 | 11) 2 }-33, 12 +39 |
2, -. |. |) 43x2: 11} 2 |-35 I
8 2 ..|.. |] 87x15 | 11) 2 }-30
pion, 2 46x18 | 8 | 2/51] 46x10 |8 | 2 |-51 8 | 2 /%2| oox40 | 9 | 2 |o0 | 78x53 1 | 80 | 11-0%43 103 x3
rs 2 47x10 | 8 | 2 |52| 47x18 | 8 | 2 |-50 8 | 2 71) 88x85 | 10] 1 {88 ]| . - |. ] 96x36 180x433
2 oe > [7 | 2 | -: | 89x20) |! 8 +50 IS] 2 (Pu ee ceo hea [ficm ice rs | |) 66x35 W5x37
y 2 Be cil Sil eel], coe OAs ae ca 5 | 1 |70 E Fal toe boa cece 13x 3S
eel
E. Zing, 1 | -4x-32 | 3) 1 | 11 : ‘ 9 | 21 3 22-75 T4x1 | ' af pall 0x13 | 8
att Ale sal|lhos A alte 5 taeeed 29x11 ; 3 = sll 61035
a BS 4 : 4 ie ahi : 30x13 : a fi us 16«12
2 a) bale ey ea z Pare eee | 5 ; Si liagilb = <aWe 2x13
\ M u i iL |

P, Plates,

'T. Talons.

A.pal. Mean thickness of plates,

= Upper jaw.

Exrraxatios op Lerrnns, &e,

TZ Lower jaw.

d. Right side,

© Lefl side.

‘The numbers connected hy bracket-lines refor to teeth belonging to the same individual.

J Erzleben. del et lith M&W Hanhart. imp

Grant Looted Ul 6 Figs,

M&N Hanharl ump

J Erxleben.. del ct lite

A Nt

af ah, ce aaa :

M&N Hanhart mp

J.Erxleben del et lith.

&N Hanhart imp

‘

N

J Exxleben del. el: ith

M&N Hanhart imp

J Byxdeben del eb tith

hart imp

M&N Rant

From nat on Stone by J Evxleben

nip

M&N Hant

TFrxlebert del.et liu .

-

zt ump

WN Hanha

M

From life on Stone by J Erxieoen

SILI LOO bp Hie OLD C2) z

MAN Hanhzect: imp

J Exxleben. del ct lith

MAN Harihart imp

1Erxleben de} et tith

rc
co
=
~I
co

XI. On a Species of Dormouse (Myoxus) occurring in the Fossil state in Malta.
By A. Leita Apams, M.B., F.GS.

Read May 9th, 1867.

[Puate LIV.]

IN the ‘Joumal 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 machwrides pre-
senting well-defined, undulating ridges), and the absence of the small grinder in the
upper jaw separate it from the Sciwrina, and assimilate it to the subfamily Myovina,
whilst its large proportions represent a species distinct from any other known Myozus,
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 differ 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 Myoaus 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 Myoaus melitensis.

This Rodent 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 the lower portion

VOL. VI.—PARY V. 2.U

308 DR. A. LEITH ADAMS ON A FOSSIL MYOXUS.

of the fossiliferous deposit of the Mnaidra-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 circumstance 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
indicaté that they had been introduced into it by carnivorous mammals or birds.

EXPLANATION OF PLATE LIV.

Fig. 1. Side view of the skull of Myoaus 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. 4

Figs. 8, 9. Humerus.
Fig. 10. Scapula. ;
Fig. 11. Articulating surface.
Figs. 12, 13. Femur.
Fig. 14. Tibia.
N.B. Except in fig. 5 all the parts are represented of the natural size.

M&NHsnhart lth

4.4- Dian

L.A,ad nal.del

MYOXUS MELITENSIS.

f 309 ]

XII. On the Osteology of the Cachalot or Sperm-Whale (Physeter macrocephalus).
By Wiuu1am Henry Frower, 7. 2.S., PRS, PLS. GS. E.ZS., Conservator
of the Museum of the Royal College of Surgeons of England.

Read November 14th, 1867.
[Puates 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 Fossiles,’ 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 Plantes. 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 vertebre. 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 vertebra, 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 Anatomiques
sur la Structure intérieure et le Squelette de plusieurs espéces de Cétacés,’ 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 auditis, 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 Balena, while the consolidated
mass of cervical vertebre of a whale of this genus is described as that of a Cachalot.

3. Lacépéde (‘ Histoire Naturelle des Cétacés, 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
vertebre.

4, Beale (‘ Natural History of the Sperm-Whale,’ London, 1839) 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. 2x

ERRATUM.

Vol. VI. p- 369, line 18, for “vostro” read “ cervice.”

310 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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 8S. 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 in 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 pelvic 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 very instructive skull of a feetal
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,’ 1864.

8. The petrotympanic bones of a Cachalot from the same Museum are figured in
Owen’s ‘ British Fossil Mammals.’

9. Dr. Gray (Proc. Zool. Soc. 1864, p. 590, and 1865, p. 440) has given figures, taken
from photographs, of the cervical vertebra 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 Hobart 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

* Tt is stated by Dr. G. Bennett (Gatherings of a Naturalist, 1860, p. 162) that the real author of this work
was the late William Sharpe Macleay. But as there is no indication of this in the work itself, as Wall’s name
alone appears on the titlepage, 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.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 311

complete skull of a very young animal, four lower jaws of adults, four perfect pectoral
limbs, some caudal vertebra, and several detached pelvic and petro-tympanic bones.

b. 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 1844 of a dealer. Unfortunately the locality whence it was obtained is not
recorded.

jf. A disarticulated skull, about the same size as the last, in the Museum of St.
Bartholomew’s Hospital.

III. 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.

IV. 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 will
be useful before entering into details.

1. The Tasmanian specimen. As before stated, this was captured off the coast of
Tasmania in the year 1864. 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 phalanges. 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
eaudal vertebra; 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 vertebre were placed close together, and in a straight line, was 30’ 4”.
The cranium measures 16’ 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
34 102”, and the entire skeleton 51’ 73”.

2x2

312 MR. 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 vertebre 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 vertebra 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 vertebre, the
hyoid bones, the pelvic bones, and many of the phalanges are missing. ‘The vertebral
formula isC. 7. D.11. L. 8. C. 214% The length of the column, the bones being
placed close together, is 28’ 7”; the cranium is about 17’ 9” (the ends of the premaxille
being broken, it cannot be measured exactly); so that, allowing for the terminal caudal
vertebra, 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, 133" and 123” 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 vertebre are united, only slight traces of the original
separation still remaining in the anterior lumbar region; and there is only a slight
indication of the 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 pairs of ribs are present; but the vertebra which I have reckoned as the

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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 difficulties 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 Anatomical 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 vertebre, 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 vertebre 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, vertebra, yibs, and other bones closely resemble those of the three
former skeletons—so much so that one cannot avoid the suspicion that the specimen has
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 vertebre 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 vertebre. 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, Fileet-street, described in the Catalogue
(1794) as “The Astonishing and Complete Sketeron of a full-grown Sperma-cett WHALE, being the real bones
joined together; near 70 feet in length. The Head, or Skull alone, measures 16 feet.” I am indebted to
Mr. Gore, of Bath, for this reference.

314 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

column, and insufficient spaces left between the bones; so that even these measurements
afford but an approximative comparison.

Cranium.

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 vertebre being placed in contact) in the Sydney skeleton (according to
Wall) is as 46 to 100, in the Tasmanian as 57, in the Caithness as 60, and in the York-
shire as 67 to 100. ‘The first is scarcely more than half-grown, the second adolescent,
the last two adult.

As seen in the section, Pl. 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 breadth 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 broadly expanded maxille, which rise from the median line towards the margin of
the skull, instead of falling 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 Balena mysticetus is rather longer in proportion to the vertebral column, but it is less massive.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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 (Pl. LVII. fig. 1).

As the individual bones of the foetal Cachalot’s skull have been described by Professor
Owen *, I will confine my present account chiefly to the structure of the different
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 exaimined.

Commencing with the cranial cavity, as the central point around which the whole
head is developed, and of which a view was afforded by a median section of the skull
(see Pl. LVI. fig. 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 13" 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 cavity 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 40° 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 above 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. Roy. Coll. Surg. yol, ii. p. 442, 1853,

316 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

In 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 skull 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, 2" by 1" 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, 2” 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 narial 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 east taken from the interior of the cranial cavity, figured at p. 372, this small foramen has yielded
no impression, The relative size and position of the four others are well seen.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. d17

outwards and forwards. In the young skull the division of this canal into three
branches takes place close to the cranial cavity. The first 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 much
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 Skull——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 in 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 77” by 112”.
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 (Pl. 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 (Pl. 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 irregularly 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 differ 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. 2Y

318 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

ends abruptly in a narrow bifid extremity on a level with the 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 the 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 (n). 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 Pl. 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 fcetal 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 the 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 (aon).
Their flat upper surface is formed of a very thin plate of bone of remarkably dense and
brittle 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 (7 f') is represented by a fissure 10” in length, and 2” in
breadth, placed between the blow-hole and the 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 (P\. LV.).—The temporal fossa is very 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 squamosal
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

MR. 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 (p) 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, 64” high and 11” long, with very prominent and distinct
boundaries, complete, except for a space of 14” 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 (7). 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
divisions 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 narrower 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 united into a single sharp edge, much
upturned 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.—tThis region is chiefly remarkable for the extent and massiveness of

the pterygoids, although falling short of that in the Hyperoodon in this respect. They
Diva

320 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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 fissure, of which the sides are
almost in contact, except at the bottom, where it widens into a triangular space.
The external surface of the pterygoid is slightly hollowed, as in Hyperoodon 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 diame 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.9)
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 affirmed and denied, are situated in or near this
groove.

Ten such teeth were sent with the present skeleton. Some of
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 3%5"
to 34g" of an inch in diameter, separated < a slight constriction
font the expanded root, which constitutes the largest part of the
tooth. The surface of the crown 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 Fei amiga oom eres:
having been broken off. The pulp-cavity is completely closed at tooth, nat. size,
the base of the tooth, which, in most of the specimens, is more or less surrounded by
rough, irregular, spiculated outgrowths.

Fig. 1.

* A 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 pterygo-palatine 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
Owen. 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 petrosalf. 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 corresponds to the strong tenon-like
process of corresponding situation and function in the Whalebone-Whales. The con-
tiguous edge of the squamosal has a laminated character, the ridges and grooves on its
surface exactly fitting 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 29. As might be expected, there is scarcely
any appreciable difference 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 Heats Tae of Mh processes oat ffotitals) 87
Extreme height (top of occipital crest to bottom of pterygoids). . . 65
Length of rostrum itr tip to line drawn across bottom of antorbital
notches). . . amore: aie St, ee eee LG
Width of commencement of rostrum lane ART Tideetea) sist dG
Width of rostrum at quarter distance from base, in straight line . . 58
Right maxillary . CO Wig Recs aie Siete cbse oa ee
NGC kG WARGO E ee ee te cy Cn tes ci yo. eh
Riolesprema seat sites a teed bs Aas ae Ga 5
Ihe tonne maa lei ny oe cali ee ee ee tt eA A ae 6
Space between premaxillaries . . . . .-.-- +--+: . 4

Op. cit. plates xxiv., xxv., and xxvi.
Brit. Foss. Mammals, p. 526. Natural size, though stated by mistake to be half the natural size.
In the specimens figured by Camper and Owen this process has been broken off.

++

322 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

inches.
Width of rostuméiat mmiddlen.. oh’ tivyntindrek aaa atoms 24h
Richt: maxillaryigy Noo esa] shu eld obi eeiiae ee 13
Leftanaxillaity ns: iassahs Tithe Sa ival ih necioee
Rightpremaxillaryu edie lo andi dich dame as 9
efi premasallary! cejisms: tripe. ceend-ay a as elisa aks 9
Space between premaxillaries . . . ss hs biggest & 2
Width of rostrum at three-quarters distance fern base cn sialic milin Seek
Right maxillary Fron eee eat pasa cie eere 2
Lefiiasxallany., duns ptiilles 0 fires ean) shitisnct Pema ees: 2
Right premaxillary . pacneeubabecn rep Saree pleas 8
heft; promoxillary fond erontidmudit ast yetta ee seey Gk tsetse te 83
Space between premaxillaries Bc 4
Premaxillaries extending beyond maxillaries. . . . . . . . . 22
5 3 se yomer . . rite Ke eA
Antero-posterior length of orbital process of fr satel bride? nibh cenage
Length of jugal . . . . . Sh rate ead oe
Height of occipital crest shane the barton of fhe srekt rene
basin behind ethmoid . .. . ; jaegn ee eb Ae,
Height of occipital crest above upper es of cotagiied magnum . . 30
Width, of, occipital condyles us] :-\n ani testes 0) gn onlll aidaioe Zag
Height of right condyle (vertical) . . . . Som tebeiah a nee
Width of foramen magnum at upper end, peewee emiagles paTiog §

The crania of the different skeletons presented no very marked distinguishing features,
beyond such as might depend upon age or individual 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.

OS
bo
eo

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

relied hlee leer ene in athena RS
ce ga | = Be = 2
S| eS Tiespecs | Seal ced ares ete
gs | 22) 85 | 2 | #8 | 24/28
Ae Seer Sa a SNS
Length of skull without rostrum ..) 14”) 24" 34”| 57"| 55"| 56"| 56”
Proportion. = 01.28. | 100 | 100 | 100 | 100 | 100 | 100 100
Length of rostrum .............. | 20 38 | 80 | 189 | 146 | 156 | 171
JERE ORM ia.g Yigalemaolma serene ote | 143 | 158 | 235 | 244 | 265 | 279 | 305
Extreme breadth .............. 18 36 60 94 87 92 | 102
Proportion P24 x cate 24 </ovex one eaysushe seve 128 150 = 176 165 | 158 | 164 182
Bxiremerherz hirer eerie: 15 26 44 66 65 70 67
Priiciien eae ere ue ee 107 108 129 | 116 | 118 | 125 | 120

Many of the differences of the skull, dependent upon age, are well illustrated in
Pl. 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 Kogia) 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 skull, the dimensions given cannot be relied on as quite accurate.

524 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

successful whaling-campaign. It will suffice here 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 Pl. LVI.
fig. 1), characters which separate them from the Whalebone-Whales. It differs from
the true Delphinide 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 Platanistide, 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.

lengthy. | Symphiysi) tein
Mandible of the young skull from Tasmania ................ 49" 21" 31”
Eroporiontnn es ems ere reer n ronan oe oe 100 43 | 63
| Mandible of Tasmanian skeleton ..........0..000seceeeeee- 174 102 7. ii
Proportions eyes, S raepe Meso eee REE ONT nee eee ee eR tt erartinks | 100 59 | Riera
Largest mandible from Tasmania, presented by Mr. Crowther ..| 194 124 75 |
| Prop ORbIO Nee ope. si trecas ce hos ohare sch cae. gl, «oi sooo) ee ee 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. Soc. vol. vi. p. 89 et seqq.
+ See Murie, ‘ Proc. Zool. Soe.’ 1865, p. 390.
¢ The length is taken from the apex to the middle of a line drawn across the posterior ends of the rami.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-\WHALE. 325

skeleton the teeth are all retained in their place by the tough fibrous gum, in which
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 13” 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 are 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 sufficient number
were preserved to show their general characters. These are simple, cylindrical, nearly
straight, obtusely pointed, 13” long and rather less than }" in their 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 26-26 and 24-25 respectively ; and a very
old jaw, with massive and much worn teeth, locality unknown, has but 21-20; anda
small, but adult specimen (female ?), 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.

Hyoid 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 (4h) and thy-
rohyals (th) (Plate LX. fig. 1).

The stylohyals (sh) are large, subcylindrical, and slightly curved, truncated at both
ends, 25" long, and 4" to 43” 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. 24

326 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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 203” 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 the Yorkshire Cachalot, is a very peculiar
feature among Cetaceans.

Vertebral Column.

The vertebrae of the Cachalot, especially as contrasted with the large Whalebone-
Whales, 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 chevron
bones on the lower surfaces of the bodies of the caudal vertebre. ‘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 Pl. 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
that 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. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

roughened by numerous concentrically placed ridges.

327

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 vertebre 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 vertebree 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 5’ 6”.

The following Table gives the weights of all the vertebra, as well as the length of
the body, the greatest width (measured between the extremities of the transverse pro-
cesses), and the greatest vertical height (from the summit of the spinous process to the

most depending part of the body).
vertebr of the Caithness and the Yorkshire specimen, as far as circumstances would

permit :—
TAsMANIAN SKELETON.
|| Length |

Weight. | “of [peraue Maen

|| body.

Ibs. oz. | in, in. in.
fet cervical sts. ono. a Bi Os |Get See | ass
padi cervical (united) || gg g {| 54 32 we:
Bt, COrsall oc. cha eteccae ise i 23 263 | 212
Bait) es eee lo aris odes pd 29 0 53 254 22
Picdaatseg! be asats: aw reas 29 0 || 7 | 238 | 24
ET AS abe an a ol 29 0 | 62 | 212 | 25
S119 elect aa a ABA baie 30 0 6 213 253
Giliers Fancracouts Bane cheat! 31 0 || 68 | 203 | 263
FS nae eee 31 8 | 7i | 192 | 263
Sites ltt saan sae) 32 8 || 78 | 191 | 263
Gilt. iccnenknein ete Ys 31 0). Me 8h Ol BO lh BEF
a ic eae SAE ete athe 34 0 || 88 | 233 | 28
Titan? Spa sees oinaoreoinorca 36 0 93 31 29
datgamnber 2.4o- cama 34 8 || 92 | 31 | 303
S| eee Hea ee 37 8 | 9F | BI} | 305
Belste oe cn ease 38 0 || 102 | 33 | 323

Corresponding measurements are given of the

CarTHness SKELETON,

| YorKSHIRE SKELETON. |

Length Extreme|Extreme
bodys breadth. | height.
in. in, in.
62 36 213
9 363 24
5 28 223
6 24} 233
64 234 26
64 22 28
f= || eB ose
72 21 283
1% 203 29
ie 20 29
8 222 30
82 274 313
9 31} 32
93 32 33
10 32 343
104 32 353

of Extreme| Extreme
| body. breadth. | height.

in. in. in.

63 40 igs

9 514k

43 | 97

df 25

6 25

63 24

63 23

62

6}

ih

a that,

8 263

83 | 31

4) 292 ae
10 2oi es 36
103 295 | 354

* The arguments in favour of including this vertebra among the caudal series are given in a former paper,
Description of the skeleton of Inia geoffrensis, p. 100, of the present yolume,

DA?

|

328 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

TASMANIAN SKELETON. Carraness SKELETON. || YORKSHIRE SKELETON.

|
| |
| s Length /fstreme Extreme|| 28th |pxtreme Extreme | Le"8*2 |Extreme|Extreme
Weight. || of |preadth.| height. ||, °f |breadth.| height. ||, ° breadth. | height.
body. body. 2 body.
lbs. oz. rey in eset in. || in. in. in. in. in. in.

4th lumbar teaceentrrec: | 39 8 103 | 33} 302 || 11 | 33 352 =|) 102 | 292 | 355 |

Stl. bsyeneecee ak: chit 40 0 103 | 34 32 114 | 33 36 || 102 | 30 34

Gin. 9 eo Myce Sytem ens 38 8 11 334 314 | 143 S| 33 344 11 30 32
heevthy 25h ee pe 3t43" 0 113 | 33 32} || 12 32 = 11? | 31 314
|< Stila, i ine ah ren ee ie | 43 0 | 112 | 32 31 || 12 + + eR AE 2

listicatdalle,crcasis eo ai 46 0 12 31 314 |) 12 Tt tf 12: | 314 | 303
| S2inds ep eRe eRe | 49:0 12 293 81k || 12 t + 314 | 30
3)As nee ee OT. Ome oe | 58 O 12s | 2s 30 || 12 tT fe 12 27 30
(Pe Atheecns esr Siac aie rere 65 0 125 | 262 29 12 t 28 12 255 | 29
lsc Bichicgugisele sd bd vliecenditt 66* 0 || 12t | 23 | 282/12. | 24 | 26 | 12. | 203 | 27
Gillis soy dobdodeabas od 58 0 123 a 253 || 113 | 222 | 243 11Z | 183 | 23
pull et eerie ine Sete 55 0 12 173 24 | 112 | 20 22 11+ |-16 22
eSth iain anamateentis ays | 45 0 112 | 16 22 || 103 174 | 21 103 | 133 | 21
PAC aa eee eee cee | 40 0 11z | 142 21 || 102 | 153 | 19 10 13 193
[ALOU PsP 8 27. feeeres si | 34 4 10% | 123 20 ¥ -/ 138%" | 185 2 | 113 | 173
iT hate Seer eer aera ran eso ape 28 0 | 173 | 8 124 | 16 Tah loll 143
11 ci ae ok | 20 0 7 «| 11 143 63 | 12 133 a |) LOe | 122
(EL Siheeeee. ¢ ont! Ste nee. }10 5 6 103 123 | 43 | 103 4 47 3 92
PA Gh ais) cao itakemteee tanceal| eae te 43 s zi 4 92 8 33 + 8
TEM ose. eon ase enncees | 3 02 4 73 2 33 95 64 35 33 ue
hGths 8 a Stee eRe };) 215 4} 72 7 z 83 6 3g 4 63
Ai eee reer ees Bete tne Qa, 4 7z + || 33 73 53 33 7% =
Sth pees art sr.snt ten avers Higa iil 32 65 z | 3 63 53 3 + 43
lL O that acre beer. 4 1 0 3a 53 43 23 63 | 44 28 + 47
[i 2OSN ee mice Stes nti 0 5 3 5 4 22 54 4 23 + 3
Date ee ee re seco ee eee 0 3i 25 33 4 24 5 3h 25 33 3
DON dea pag ye cto ah hasiehis)< He 0; 92 2 3y 23 3h ; 2 23 28
BTC suse eters: city. tesrneue S 0 12 On 24 2 13 4 12
24th ,, 0 02 2 2 1} =

The Cervical Vertebre 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
vertebre 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 vertebre ; 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 vertebre (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, Hyperoodon, and Micropteron, they are
all united into one solid mass, as in Balena among the true Whales. In Platanista,

* The vertebre in this region, as is usually the case in Cetacean skeletons, contain much more oil than at
other parts, which partially accounts for their great weight. Elsewhere the vertebrae were free from oil, and
dry at the time they were weighed.

+ 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 Balenopteride. In
the other Delphinoids the atlas, axis, and generally one or more of the other vertebre
are confluent; if any are free, it is at the hinder end of the series. 7

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 vertebre. 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 43"
for the reception of the condyles of the occiput. The posterior surface is remarkable
for its flatness.

Fig. 2.

Anterior surface of atlas+.

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 divided 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 vertebre ; the latter fits over the
rough surface of the axis to which the odontoid ligaments are attached, and affords a
passage for them.

* Jn two skeletons of male Narwhals in the Mus. Roy. Coll. Surg. the bodies of the 2nd and 3rd cervical
vertebre are firmly united.

+ The woodcuts of the vertebre (as the figures in Plates LVIII. and LIX.) are all reduced to ;4, the size of
nature. The anterior surfaces are represented in every case.

330 MR. W. H. FLOWER 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 22”.

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 fits 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 Hyperoodon the foramen is
complete; in Orca the same; in Globiocephalus very nearly so; also in some of the
true Delphini, as D. tursio, In Beluga, Platanista, and Phocena 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 Beluga, Monodon, or

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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.
in. in.
Bixtremel breadth s i iiiecosecaettteerepsia ocien sleraucie eerecionersteeea acer 374 36
Width between outer edges of anterior articular surfaces ...... 23 26
Extreme width of each of these surfaces .........-.-ee0e eee: 2 10
Extreme height of each of these surfaces..............00000 144 163
Width: of nenral canal rere eerie eect ore eerten « 92 11
Height) oftmeuralicanally qervteuieierecriet tiem stacey yous iene cere oe 10 103
Height of contracted lower portion of the canal .............. 4 2s
Width of contracted lower portion of the canal at the upper end. 3i 1
Contracting*below toms iiss stat sec tecincis ocelot oe eee ene 1 il
Height of portion of bone below the neural canal ............ 64 UG:
H=xiremeheightamemrddlevline strc lee crite eisai see 184 213
Vertical height of transverse processes at outer end .......... 14 a0
Greatest antero-posterior thickness of the same .............. 5 ;
Heightiob neuralvarcheinpmiddley erect atti aise eine: 1 3h
Antero-posterior thickness of neural arch in middle .......... 3z oc
Length of inferior surface of bone in middle line............ 6 62
Width between the outer edges of posterior articular facets .... 25 mises

* 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 erypt 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 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

The remaining six cervical vertebra 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 vertebre. The union of the seventh to the antecedent vertebre 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 vertebre 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.

Anterior surface 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 $” 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

Be

(ss)
et)

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE,

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 the
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 off, 8” broad, and 62” high. ‘The inferior margin is very indistinctly marked
in the middle,—the anterior surface of the axis above the rudiment of the odontoid
process gradually passing into the flattened floor of the neural canal, which continues 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 greater 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 vertebre (and in the axis of the
Whalebone-Whales), coalesced and with the intermediate space filled 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 transverse process of the cervical vertebrae ( 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 vertebre, 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 vertebre are free, as Beluga and, especially, Platanista. 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), 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 ‘'asmanian Sperm-Whale there
is no trace of an inferior transverse process on the smooth sides of the bodies of any of
the cervical vertebre as far as that which appears to be the sixth, inclusive. The lower

VOL. VI.—PART V1. 3A

334 MR. W. H. FLOWER ON THE OSTEOLOGY OF 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 vertebre 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 (diapophysis, 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 lamelle of bone which constitute the lateral parts of the neural arches of the
third, fourth, fifth, and sixth vertebre are 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 lamelle, 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 vertebra examined have the whole of them escaped destruc-
tion in the cleaning-process; sufficient remains, however, in every case to show that
their general arrangement is the same. The intervals between them are of nearly equal
width (3), but diminish in height from before backwards, the first being 24”, 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 anterior surface, and with a
small pointed spine in the middle line surmounting 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 Toothed
Whales at least, quite peculiar to Physeter. Even Hyperoodon shows no trace of them,
as the conjomed neural arches of the first six cervical vertebre rise smoothly and
gradually into a greatly elevated spinous process. The axis of the Balenoptere presents
rugged lateral processes somewhat similar to those of the Sperm-Whale.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 335

As the posterior surface of the body of the cervical vertebre is ankylosed with
the first dorsal, little can be said of its characters. The centrum, however, is very
deeply concave, receiving the convex anterior surface of that of the first dorsal; and
there is a smooth articular facet (posterior zygapophysis) at each side of the neural
arch. This facet is more vertical (inclining less backwards at the upper end) than
in other Cetacea.

Measurements of the conjoined six posterior Cervical Vertebre.

(Tasmanian,| Caithness.
in, in.
lBixctreme breadth ey. tem + overs he tee aa sie RE, Swen e-ve 32 361 |
Pxtremerhelclitevastastpiete ils aie sail eeee = teeta ioctl - ..| 222 24
Breadth between outer edges of articular facets for atlas ...... 23 25
Breadth of anterior opening of neural canal................-- = 10
Height of anterior opening of neural canal ............-.-+-- 5 7
Height of body of bone below this ..........+.seeeeeeeeees + 104
| Antero-posterior length of superior surface (floor of neural canal). 4 63
Antero-posterior length of inferior surface .......-....++++++ 53 85
Antero-posterior length of lateral surface..............+++++5 9 113
Breadth of posterior surface of body of seventh cervical vertebra .| 17% 15
Height of posterior surface of body of seventh cervical vertebra . . 102 125

The Dorsal Vertebre (Pls. 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 vertebra. The lateral parts of the centrum are free, as also is the whole of the
neural arch.

The body of the first is by very far the shortest in the antero-posterior direction; the
second is nearly twice as long ; and they continue to increase gradually 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 degree. In the
succeeding vertebre the anterior and posterior surfaces are nearly flat and parallel.
The bodies, at first very broad in proportion to their height, rapidly become narrower ;
in the fourth the breadth is already less than the height ; the sixth, seventh, and eighth
are the narrowest; after these there is a slight increase of width.

The neural canal in the first two vertebre is triangular, but rapidly assumes a trans- _
versely oval form, and gradually 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 MR. W. H. FLOWER ON THE OSTEOLOGY OF 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 vertebre. 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

Fig. 4.

Anterior surface of the fourth dorsal vertebra and head of right mb.

scarcely 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 vertebre.
As far as the sixth their general direction 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 first to the ninth vertebra inclusive.
In 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 flattened surface 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 inner 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-

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 337

spicuous and important process on the neural 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 Delphinus delphis (Cat.
Osteol. Prep. Mus. Roy. Coll. Surg. vol. ii. p. 450), that these processes belong to the
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 teuth vertebra inclusive, arising in all from the side of
the neurapophysis, at nearly the same height throughout. They gradually decrease in

Tenth dorsal vertebra.

length from the first to the tenth (see Pl. LVIII.). The first is compressed from before
backwards ; the next two subcylindrical; the following four are very short, thick, and

338 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

almost confluent with the bulky, rounded metapophyses. After this the transverse process
gradually 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 off. On the eleventh dorsal vertebra there

Fig. 8.

Eleventh dorsal vertebra and right 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 oval, with the long diameter antero-posterior.

On the upper part of the sides of the bodies are conspicuous articular facets 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

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE, 339

edge, for the seventh rib, is raised upon a small tubercle ; while the posterior surface,
for the eighth rib, is still further reduced and flat. In the eighth (fig. 5) the anterior
facet is still more raised, and the posterior is nearly obsolete. In the ninth (fig. 6) the
articular surface 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 the pos-
terior edge for the tenth rib. In the tenth vertebra (fig. 7) the above-described tubercle.
developed from the anterior articular facet, takes the shape of a long massive process,
6" in length and nearly as much in antero-posterior thickness, springing 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, concave 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 (fig. 8) has no articular
facets on the side of the centrum; but the process arising from this part on a level with
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 vertebra, 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 (diapophysis) of the cervical vertebra, 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 vertebra, which, from their situation, would be taken to represent
the inferior processes (parapophyses) of the cervical region. We find, 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 hand, a new process, springing from the side of the
centrum of the eighth or ninth vertebra, and being at first only a development of the

340 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEKM-WHALE.

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 Hunterian Collection in the
College of Surgeons, upper transverse processes are regularly developed from the arches
of the dorsal vertebre 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 mb 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 vertebra united so as to form a complete ring.

It will be very important to ascertain whether Kogia 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 transverse process springing from the 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 the 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. ‘The principle, however,
is the samef.

Iumbar Vertebre.—The eight lumbar vertebre present a remarkable similarity to
one another, both in form and dimensions. Their bodies increase slightly but pro-
gressively 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.

+ For the condition of the rib-attachments in Platanista and Inia, in some respects intermediate between
those of the Physeteride and Delphinide, see “ Description of the Skeleton of Inia geoffrensis,” &c., Trans.
Zool. Soc. vol. vi. pp. 98-103.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 341

keel running 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 vertebra is pentagonal, or like a lozenge with the upper angle cut. off.
The first of the series is rather different in form, having no distinct keel. In the older
individuals, as the Caithness specimen, the keels of the lumbar vertebre are developed
to a still greater extent than in the Tasmanian skeleton.

Fifth lumbar vertebra.

The 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; the
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 the
first lumbar vertebra they arise from the lamine of the neural arch by the sides of the
canal, and their upper edge is 63” above the top of the body; in the last they spring

9

VOL. VI.—PART VI. 3B

542 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

from the fore part of the spine, altogether above the junction of the lamina at the root
of the neural arch, and their upper margin is 11” above the top of the body. There
are not any corresponding processes on the hinder edge of the spine, as in Platanista.

The Caudal Vertebre are twenty-four in number, of which twelve or more appear, by
the form of their inferior surfaces, to have 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 vertebre vary
greatly. The first differs but slightly from the last lumbar vertebra. The centrum 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 with the
thirteenth, is so extraordinary that, unless there were abundant evidence to the con-
trary, one might be tempted to suppose that several vertebra 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.

Fig. 10.

Fourth caudal vertebra, without 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

i 29
given at p. 328.

Mk. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 343

The neural arch forms a distinct canal as far as the fifteenth caudal vertebra; in the
sixteenth and following vertebree 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 vertebre (with elongated
bodies) the neural lamin arise from little more than the anterior half of the body, but
posteriorly, as the bodies decrease and the lamin 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 thickened, 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 vertebre 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 surface, 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 58," high by 34,”
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 vertebre 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 vertebre 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 their place is taken by a
slight groove. In the Caithness Cachalot the transverse process is rudimentary in the

9

3 BA

544 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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 vertebre 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 oval foramen 385" by 335" 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 vertebree, being found
eyen 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 24” 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 narrow bony septum (less than 1” in width). In the next
three vertebre the septum is still narrower ; in the sixteenth it suddenly widens to 2"; and
thence onward the inferior openings are gradually placed rather further from each other,
until in the last few vertebre 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 toeach other as the vertebree diminish in size.

The articular surfaces for the chevron bones are remarkably large and prominent,
with elevated edges, and rough cup-like depressions in their 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 (anunited) hemapophyses
fit closely. In the Caithness Cachalot, these bones are ankylosed to the body of the
vertebra, though not meeting each other at their free ends. In the second vertebra,
the hinder facets are of larger size; but none are 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 vertebree, the two pairs of facets are well
developed, the anterior ones at first comparatively small, but towards the end of the

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 345

series almost equalling the hinder ones in importance. On the thirteenth vertebra arti-
cular facets are faintly indicated on the anterior edge only.

Having now examined the modifications of the different parts of the caudal vertebre,
their bodies, neural arches, transverse processes, arterial foramina, and articular facets
for heemapophyses, a few words must be said on the peculiarities of the different vertebre
of the hinder part of the series taken as a whole.

The first twelve, or chevron-bone-bearing vertebre, 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 hemapo-
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 intervertebral substance being 2” less in diameter than
the anterior.

The succeeding vertebra is of very simple character, being reduced to a centrum with
a slight tubercle covering the neural canal to represent the spinous 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 vertebre 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. The quadrangular space between these perforations bears at each
corner four tuberosities, an anterior and a posterior pair, remnants of the neurapophysis,

546 Mk. W. H. FLOWER ON THE OSTEOLOGY 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 hemapophyses in the anterior caudal vertebre.
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 vertebre. 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 vertebre is a strongly marked longi-
tudinal groove, bounded above and below by large rough tuberosities, of which the
upper 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 vertebree, and begins
suddenly in the first of this group.

It now only remains to speak of the special peculiarities of the last four caudal ver-
tebre. 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 vertebre of the last 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 vertebrae where 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 SPERM-WHALE. 347

middle by these grooves on each side below, and by the one connecting the two fora-
mina (here much reduced in diameter) on the upper surface, that 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 the irregularities of their prominent edges correspond
exactly, and they have evidently been in close apposition.

The last twelve vertebra 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 with 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 58”, while the twelve cor-
responding vertebre 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,
51” in greatest depth, and 4” in length. The second had another, consisting of two
distinct oval plates 21” deep by 23” long. There were no further traces of hema-
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 vertebree
bearing well-developed ribs. They agree, moreover, in possessing eight true lumbar
vertebre. 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 chevron bone. Of remain-
ing or caudal vertebra, 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 vertebre being 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 vertebre, any more

* Wall, op, cit.

348 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-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? 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, that 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 vertebre) 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 vertebra, 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 vertebra 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
vertebre 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.

PHYSETERIDS.
Physeter macrocephalus australis (Mus. Roy. Coll. Surg.), C. 7, D. 11, L. 8, C. 24=0.
P. macrocephalus australis (Wall), C. 7, D. 10, L. 8, C. 24=49.
P. macrocephalus borealis (Yorkshire), C. 7, D. 11, L. 8, C. 283=49.
Kogia grayit (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. rostratum (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. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

PLATANISTIDA.
Platanista gangetica (Eschricht), C. 7, D. 11, L. 8, C. 25=51.
Inia geoffrensis (Brit. Mus.), C. 7, D. 18, L. 3, C. 18=41.
Pontoporia blainvillii (Burmeister), C. 7, D. 10, L. 7, C. 18=42.

DELPHINID.

Beluga leucas (Mus. Louvain), C. 7, D. 10, L. C. 33=50.
Monodon monoceros (Mus. Roy. Coll. Surg.), C. 7 11, L. 6, C. 26=50.

uD:
Phocena 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 3 (Mus. Roy. Coll. Surg.), C. 7, D. 13, L. 14, C. 32=66
P. communis ¢ (Mus. Oxford), C. 7, D. 12, L. C. 46=65
Neomeris phocenoides (Mus. Leyden), C. 7, D. 13, L. 18, C. 830=63
Orca gladiator (Mus. Leyden), C. 7, D. 11, L. 10, C. 23=51.
O. gladiator (Mus. Cambridge), C. 7, D. 12, L. C. 33=52.
Pseudorca crassidens (Reinhardt), C. 7, D. 10, L. 9, C. 24=50.
Globiocephalus melas (Mus. Roy. Coll. Surg.), C. 7, D. 11, L. 12, C. 28=88.
G. melas (Middlesex Hosp.), C. 7, D. 11, L. 13, C. 28=59.

v

G. ? (Tasmania, Mus. Roy. Coll. Surg.), C.
Delphinus sinensis (Mus. Roy. Coll. Surg.), C. 7, D. 1
D. guianensis (Van Beneden), C. 7, D. 12, L. 14, C.
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=66.

D. 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=70.
Lagenorhynchus leucopleurus (Mus. Tee C. 7, D. 15, L. C. 59=81.
L. albirostris (Mus. Cambridge), C. 7, D. 14, L. C. 67=88.

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22

349

It will be seen from this Table that the total number of vertebre varies extremely
in different genera, and even in species of some natural genera, the variation being
chiefly in the lumbar and caudal regions. The well-marked group Physeteride, 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 formule 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 vertebre. 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. ane

350 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

the transverse processes, which points to a close affinity with ZHyperoodon 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 vertebrae
present but little of the peculiar elongation, so characteristic of those groups.

Chevron Bones.

Ten chevron 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 vertebree with those on the different chevron bones, the
missing ones were ascertained to be the fifth, sixth, and the thirteenth and fourteenth.

The first (See Pl. 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
inches. inches.

Meng thy ass kata eos. RA 8 Oe Oe See 6” 52
Diameter iat DAse carte cavities wnt ci en is me eke |) eae 2:8
Diameter at middle, antero-posterior .......... | 19 2-3,
=F sy PLANS VETSO ss osha hie WSs a yeucts 13 TEES)

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 lamin are united
below, and form a prominent spine. The second is comparatively 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 somewhat shorter, but more mas-
sive and rounded at the lower end. ‘The entire length of the second bone is 102", 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 vertebrae
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
bone 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 ON THE OSTEOLOGY OF THE SPERM-WHALE. 351

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
examination, owing to the imperfection of the materials and the difficulty, in incomplete
sets, of ascertaining with precision which are the corresponding bones.

Ribs.

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 Balena of correspond-
ing magnitude. This character is best illustrated by stating that in the skeleton of the
nearly adult Balena mysticetus, in the Mus. of Roy. Coll. of Surgeons, 46’ in length,
the twelve pairs of ribs weighed 7 cwt. 60 Ibs.; in the Cachalot, several feet longer
and of corresponding age, the eleven pairs weigh less than 3cwt. 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 in 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. 93 oz., and those of the left side 164 lbs. 53 oz.

Weight and Measurements of the Ribs (excluding the rudimentary eleventh pair).

|
Extreme H Circumference)
Weight. length in ean coy" at inferior | Curve*,
straight line. extremity.
lbs. oz in. in. in. in.
; (et pie eh ae pl 27 6 45 14 29 103
ise Marcapersgetetsieyee ene oe 27° «4 47 14 28 11
2 { Raa ogee Be «Gar 25 13 634 113 203 22
le core rtateacs aan 25 1 63 113 20 21
3 RRS Satna sea cve ohare ens 25 «(0 70 3 173 28
Tia acy cpg tastes opto 25 13 713 92 18 27
A 1M mote Sooo ae 21 8 i 94 16 29
dO pea lode ahi ceo 23 2 74 oF 163 29
5 Lin POO pe OSE 19 8 68 83 14 30
iat oysters crn eietereasgaeves PAL ya iii 72 83 15 29
6 RAL SRA. RG A 19 5 674 9 123 29
lprebceo coos Shacdon i) 7 683 82 12 28
a i eaten omecictn, oi 4 WG 655 8? 10 26
fy | Te ihectes, oe esl: 16 15 663 8i 103 25
8 IW deosabobgoReD es 15 3 63 83 8 22
Seat. Mecectateetote cheer et 13 15 65 8 83 225
9 IC BO. 3 OND RD TES 10 6 61 7 ip 19
| DI a es Mlanwoee to 10 5 63 64 83 18
10 | R. 975 56 ff 87 11
Uae eae TOMS OU OD 8 4b 5573 63 8 114

* 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.
302

352 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

The first rib (Pl. LX. fig. 2) is very different from the others in form. Though
by far the shortest, it weighs absolutely more than any of the others, this being
occasioned 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 off 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 much
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 different 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
have 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 the middle of the bone, the truncated end pre-
senting a regularly oval surface.

The third 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

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 353

following changes may be observed 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 with 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 vertebre,
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-att
direction.

The eighth repeats much the same condition at the upper end; it may be distin-
guished, however, by its greater slenderness, 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 are
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 surface, 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 13}" in length,

354 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

and 23" in greatest diameter at the proximal end; the other is 11” in length, and 23”
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 corre-
sponding vertebre clearly indicate their 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
halfgrown.

The skeleton at Paris has fourteen pairs of large ribs, with indications on the trans-
verse processes of a fifteenth. Their articular surfaces, as well as the corresponding
processes of the vertebra, are much decayed and broken; but, as far as can be ascer-
tained, they appear to follow each other in a regular and natural sequence, and afford
no certain evidence that the skeleton has been artificially compounded. ‘The first pair
have rudimentary capitular processes, larger on the left than the right side. These
are followed by ten pairs with well-developed capitular processes reaching to the bodies
of the vertebre in the usual way. The last three are connected only with the lower
transverse processes, which are strongly developed and hollowed out at the end for
their reception.

In all the true Dolphins (Delphinid) the anterior ribs (about half of the series) have
long heads, by which they are connected with the body, or root of the arch, of the ver-
tebra in front of that to which the tubercle is attached. Near the middle of the series
this head suddenly ceases to be developed, and the ribs articulate only to one vertebra, by
the tubercle. It will be seen from the preceding description that in the Cachalot the
condition of the upper end of the ribs, and their mode of connexion with the vertebra,
is quite different. Of the Cetaceans whose osteology is thoroughly known, Hyperoodon
comes nearest to Physeter in this respect, as already mentioned when speaking of the
transverse processes of the thoracic vertebra. ‘The form of the first rib of the Cachalot,
however, is very peculiar—the absence of a distinct head reaching the body of the
vertebra having no counterpart among Toothed Whales, not even in Hyperoodon.

Sternum.

The sternum (Pl. LX. figs. 3 and 4), is a large, massive bone, though of rather spongy
texture. Its general form is roughly triangular, the apex being turned backwards.
The broad anterior edge, nearly equal to the sides in length, is tolerably straight. It
is composed of three distinct portions—two large anterior lateral pieces, and a small
posterior median piece. The former have probably each consisted of an anterior and
posterior portion, though the traces of this distinction are well-nigh obliterated. The

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 355

latter, in like manner, appears to be the result of fusion across the middle line of two
lateral portions; so that the sternum was originally ossified from three pairs of distinct
centres, as shown by Wall in the figure of the young skeleton at Sydney.

The contiguous borders of the large anterior pieces do not come in contact for the
whole of their length, but leave an oval median aperture, 11” long, and 8" in greatest
width, the fore end of which is 7" from the anterior margin of the bone. The greatest
breadth across the entire sternum, close to its anterior end, is 424” in a straight line.
Its extreme length is 47". The posterior piece, of an irregularly quadilateral form, is
10%" wide near the front, and contracts to 63”, expanding again slightly at the hinder
extremity. Its extreme length is 142", being more produced backwards on the left
side than on the right. As seen in the side view (Pl. LX. fig. 4), the whole bone is con-
siderably curved in the longitudinal direction, and is very unequal in thickness. The
superior lateral angles form very large and massive prominences, bearing rugged oval
surfaces for articulation with the expanded lower end of the first rib. Below this the
bone becomes comparatively slender, and then thickens again at the junction of the first
and second piece, where is situated a small cup-shaped articular surface for the cartilage
of the second rib. The second piece retains throughout a considerable vertical thickness.
At its hinder extremity is the surface for the attachment of the cartilages of the third
rib.

The sternum of the Caithness Cachalot presents the same general form, but is broader
in proportion to its length. The two anterior pieces are united together for a space of
8" in front of the median foramen, but are still disjoined behind it. The foramen is
smaller and more circular, being 6" in length by 5” in breadth. The most notable dif-
ference, however, is in the hinder piece, which is represented only by a median spheroidal
nodule of bone, 4” in diameter, which fortunately still remains in situ, imbedded in a
mass of dried cartilage. The greatest breadth of this sternum is 46", its length 32".

The sternum of the Yorkshire skeleton comes nearer again to the Tasmanian, both
in general proportions and in the development of the hinder portion. Its extreme
breadth is 44", and its length, including the hinder piece, which is 12”, is 45". As in
the last-described adult specimen, the anterior portions are united across the median
line in front of the foramen, but not behind. The foramen itself is of a different form
from that in either of the others, being very long and narrow, 14” by 43”.

All the specimens appear to agree with that described by Wall in having surfaces for
the direct attachment of the cartilages of four pairs of ribs—the first to the anterior
corners (where the bone is broadest), the second to the junction of the ankylosed first
and second pieces of the sternum, the third to the hinder end of the second piece, and
the fourth to the third piece, or, in the Caithness skeleton, to the cartilaginous mass in
which this is imbedded.

The sternum of the Cachalot resembles that of its nearest congeners, Hyperoodon and
Ziphius, in general principle of construction, being formed from three pairs of ossific

356 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

centres, but differs in its rugged massiveness, in its broad triangular shape, and espe-
cially in the tardy union of its lateral parts across the middle line.

Neither of the three skeletons of Cachalots in this country has any vestige of
ossified sternal ribs. Wall says, “ the sternal parts of the ribs are all cartilaginous” *.
We may therefore conclude that Physeter resembles its congeners Kogia, Micropteron,
and Hyperoodon in this important character.

Pectoral Limb.

Besides the limb-bones belonging to the skeleton, from which unfortunately some of
the phalanges were lost, Mr. Crowther has kindly placed at my disposal two pairs of
complete fins of adult male Cachalots, with all the bones in their natural connexion.
Nothing is therefore wanting in the materials for completing our knowledge of this
portion of the skeleton.

In relation to the entire size of the animal, the pectoral limb of the Cachalot is com-
paratively small, the length of its osseous parts from the head of the humerus to the
terminal phalanges being 4’ 10", or about 3}; of the entire skeleton. The terminal
portion is broad, and rounded or almost truncated at the extremity, the digits being
spread apart, and all five well developed, especially the second, third, and fourth, which
do not differ greatly in length. This form of hand is also found in the other Physeteride,
the Platanistide, and Beluga and Monodon among the Delphinide. It has its greatest
contrast in the narrow lanceolate fin, with adpressed digits, of which the second and
third far exceed the others, characteristic of Delphinus, and which reaches its extreme
development in Glodiocephalus.

An observation in Wall’s memoir} has given rise to the idea, since repeated in other
works, that there is a want of symmetry in the two pectoral limbs of the Cachalot, the
bones of the right being described as “considerably larger than those of the left.”
However this may be with the specimen in the Sydney Museum, there is certainly no
appreciable difference either in size or form in the bones of the two pectoral limbs in
any of the skeletons that I have examined.

The scapula is higher in proportion to its breadth than in any other Cetacean;
indeed it is the only one known in which the height actually exceeds the greatest
transverse breadth. The whole of the outer surface (corresponding to the infra-
spinous fossa of the ordinary mammalian scapula) is remarkably concave, and the
internal surface is in a corresponding degree convex{. The spine, as is usual in the
Cetacea, has a very narrow base of origin, placed near the neck of the bone, and so
close to the anterior edge as to reduce the supraspinous fossa to almost imperceptible

* Op. cit. p. 28. t+ Op. cit. p. 5.

= In the Sydney skeleton, according to Wall’s figure, the internal surface of the scapula is placed outwards.

The same is the case with the skeleton at Burton Constable, the articulator having doubtless been misled in
both cases by the above-mentioned peculiarity.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 307

dimensions; on the other hand it is very long, projects slightly outwards, and then
directly forwards, and is expanded vertically at its acromial termination, The coracoid,
which arises immediately below the spine, but rather to the inner side, or just above
the anterior edge of the glenoid fossa, is a stout, compressed, truncated process, some-
what dilated at its extremity, and of about half the length of the acromion, The
glenoid fossa is a very broad and irregular oval, with prominent margins. On the pos-
terior internal part of the margin the surface of the bone is rough, as if the process of
ossification were not complete ; and in the cartilaginous mass which covered it a detached,
rough, irregular epiphysial nodule was imbedded.

The scapule of the different specimens examined present slight variations in form
and proportions, as will be seen by the following table of dimensions. ‘There is nothing,
however, in this bone or in the other osseous structures of the limb to distinguish the
Tasmanian from the northern Cachalots.

Dimensions of Right Scapula.

Tasmanian.| Yorkshire. | Caithness.| Paris,

% ; hf ; A a sn OF inches, inches, inches. inches,
xtreme length, from highest part of superior border to anterior
margin of “glenoid fossa Nas z Sreraeiemrs i pe eherene\ Rist Netetle gresel ess oes 394 a
Length from ‘centre of superior border to middle of glenoid fossa... 32h 36 34 35
Length of anterior margin, from anterior superior angle to anterior 361 41 36
margin onplenoid! fossa y-eioctars creado une sasha atone tee eatec z
Length of posterior margin, from posterior superior angle to pos- 262 26 971 26
ferormmarcinvor re lenoid fossa) eae errs eerste iieteie a ue
From anterior superior angle to origin of acromion process..,..... 21 224 21
Breadth between anterior-and posterior superior angles .,........ 29 343 293 27
Breadth immediately above the root of the acromion process ...... 12 133 124 13
From posterior inferior angle (hinder margin of glenoid fossa) to 931 261 99
GuilOr Xow 55 Saag aacmasAOosongobronuenevemoonon z 2
ene throtacromionte te syaten eisiereeiien- thie sticker et ciere eeraleeae stare ei 133 163 13 eh
Vertical height of acromion at narrowest part (near its root)...... 54 6 54 43
Vertical height at broadest part (near its extremity) ............ 93 11 11 8 |
From posterior inferior angle, to end of coracoid process.......... 164 193 162 21
ene tibyot coracoid: processpra\astt-recre. eke eaearetaia fete er nenar: 7 12 82 11
Vertical height of coracoid process at its narrowest part, near middle. 23 3F +
Vertical height at its thickest part, near its extremity .......... 3 5 5 34
Length of glenoid fossa, including its thickened margin.......... 94 9 8 10
Breadth of glenoid fossa, including its thickened margin ........ 8 8 8

In the superior height of the scapula compared to its breadth, Physeter presents much
the same deviation from the other Dolphins as Balena does from the other Whalebone-
Whales. The scapula of Hyperoodon is intermediate between that of Physeter and
the low broad form so characteristic of the typical Delphinidw. In Kogia the scapula
appears to follow more closely the ordinary type.

Humerus.—The shaft of the humerus is much compressed from side to side, The
head is not marked off by a very distinct neck, and the broad. single tuberosity is not
particularly prominent. The most distinguishing peculiarity of this bone, found in all

VOL. VI.—PART VI. 3D

358 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

the specimens examined, but not seen in the Dolphins generally, is a rough rounded
tubercle, projecting downwards from near the middle of the radial border, and connected
by a broad ridge with the lower part of the tuberosity. Immediately beyond this
tubercle, the border of the shaft is deeply hollowed out. The distal end is divided
nearly equally into two rugged and irregular hollowed facets, meeting at an angle of
about 150°. The radial facet is considerably broader than that for the ulna.

In both the aged animals from the British coasts, the extremities of the radius and
ulna are ankylosed to the contiguous surfaces of the humerus; and this is partially
the case on the left side in the Tasmanian skeleton, but not on the right; nor has
it occurred in either of the other specimens sent by Mr. Crowther. It must be observed,
however, that in neither of these has the epiphysis forming the head of the humerus,
and which includes nearly the whole of the tuberosity, coalesced with the shaft. In
none of the specimens is there any trace of a distal epiphysis to the humerus. A con-
siderable difference will be observed in the form of the two humeri figured in
Plate LXI.

The radius and ulna, in nearly all the specimens examined, are united together at
their proximal extremity, and in one case (the Caithness skeleton) also at the distal end.
In the greater part of their length there is a considerable space between them, chiefly
occasioned by the narrowing of the middle part of the ulna; for the corresponding border
of the radius is nearly straight.

Each bone has an epiphysis at either extremity, that at the proximal end being a thin
plate, comprising only the articular surface, and early ankylosed, the other being larger
and more tardy both in ossification and in union with the shaft. In all the specimens
in which the head of the humerus is still free, this epiphysis is also ununited ; and its
rugged, irregular surfaces, and small size compared with the extremity of the bone to
which it is applied, show that the portion preserved is only the ossified nucleus of a larger
cartilaginous mass. In the absence of the epiphyses the surfaces of the lower ends of
the bones are deeply concave.

The ulna is lighter and more compressed, as well as shorter, than the radius. It is
very broad at the distal extremity, and, being contracted at the middle, has rather an
hourglass-form. From its free border, near the proximal end, rises a well-developed,
compressed olecranon process, expanding as it rises, and terminating in a semicircular
margin.

The radius has far less character about its outline, the borders being approximatively
parallel.

The free or distal end of the lower epiphysis of both bones is divided into two facets,
conforming with the outline of the contiguous carpal bones.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEKM-WHALE. 859

Dimensions of the Bones of the Arm.

Tasmanian |'Tasmanian,|Tasmanian,| y, . ith- -
| aralatont fee i No. 3, | Yorkshire. Cate Paris.
Humerus. inches, inches, inches. inches. | inches. | inches.
Teentep ys 37 eerste tates regio suabier dense uta <uaene | 192 223 18} 203 20 172
Breadth (vertical, or from the radial to the Q3 9 8 g1
ulnar border) ofibeade- <a eyee + aioe coe } ce: 3 pe eS oer
Breadth at narrowest part...............+-- 6 Tx 6 63 74 6
Breadth) atilowerend,...25..: 2.000000 oers | eel OS 11 93 114 103 9
Thickness: at ead, /i.5 sve )s:ste0e sis 9 26 fe ere yates nae 83 8s tz 8
Hifi cknessvaG middle eererse res state secre aetaeice ee ae 4 4
Radius.
Length (between middle of each extremity
canitdae epiphysis mminacr eter oe fete : } ame Ee te = Eon) EN
Breadth at upper end.................. isle 72 6 73 ae 53
iBreadinbatymid dle src tremors aetna peels 6 7 53 64 63 54
Breadth avdowerends wens re sess 2 ee clare 72 97 83 2 i
Ulna. 9
GETS Ble crept er tegegee ae ciel tes alescuss-l aclaternreive 123 14 11} 123 12 103
Breadth at upper end, including olecranon ....| 10 12 103 12 11 10
IGE nnn co aes opacenoonbaanocaee 54 5 53 43 5 5
Brea dthea tal ower nd eyetngegd eters cere err gi 9 83 9 82 8
Greatest width of interosseous space............ 2 23 2 Hs Z| eres

The carpus (Pl. LXI. fig. 1) is very remarkable on account of its width and short-
ness, the bones of which it is composed being so extended laterally as to appear almost
as if ina single row. As in most of the Toothed Whales*, ossification of the carpal
elements advances more rapidly than in the Whalebone-Whales; and in the older speci-
mens the contiguous surfaces of the bones are brought into close apposition. In the
younger individuals each bone is surrounded, except on its smooth free (dorsal and
palmar) surfaces, by a layer of cartilage; and consequently its exterior presents the
peculiar and characteristic appearance described at p. 326. Moreover, in many cases, a
kind of epiphysial ossification has taken place in this cartilage, so that the bones are
surrounded by a more or less complete case of thin osseous matter, which appears
ultimately to unite with them (see Pl. LXI. fig. 4). This mode of ossification of the
carpal bones, by a peripheral as well as a central nucleus, is, I believe, peculiar to the
Cachalot. It has occurred to a much greater extent in the carpus of the entire
skeleton than in the detached limb figured at Pl. LXI. fig. 1; indeed, in the latter, the
peripheral ossifications were so small and so slightly adapted to the contiguous surfaces,
that, their attachments having been lost during maceration, they could not be replaced
with certainty, and so do not appear in the figure.

All the specimens agree in the number and relative position of the carpal elements.
Besides a bone projecting freely from the ulnar side of the carpus, probably correspond-

* Orca is an exception.
3D2

360 MR. W. H. FLOWER-ON THE OSTEOLOGY. OF THE SPERM-WHALE.

ing to the pisiform*, they are five in number, differing little from each other, either in
size or form. Although, as before said, they are so extended as to appear almost to
form a single row, it may be readily seen that the central bone and that placed at
either extremity really belong to what is ordinarily considered the proximal row of the
carpus, while the two others constitute its distal row, thus conforming to what may be
called the typical form of the Cetacean carpus.

The determination of the homologies of the carpal bones of the Cetacea with those
of other Mammalia is beset with difficulties, ‘and has consequently led to some differ-
ences of opinion among those anatomists who have attempted it. The most recent essay
on this subject is that of Dr. Van Bambeke}, who, however, laboured under the great
disadvantage of having very insufficient materials at his command. I have been able to
examine a considerably larger number of specimens, comprising nearly all the known
genera, but must still admit that the determination of homologies of parts from the
comparison of adult, or nearly adult, individuals is but provisional, and awaits, for its
verification, the opportunity of tracing their development through the earliest stages
of existence.

The results of these observations on the carpal bones in the Odontoceti (to which
group it is alone necessary to refer to illustrate the anatomy of the Cachalot) may be
stated in the following propositions. It may be premised that every species appears
liable to certain individual variations, and that sometimes the different sides of the same
animal are not precisely alike either in the arrangement or even the number of the
carpal ossifications. Such cases have often afforded a valuable clue to the identifica-
tion of particular bones.

1. The pisiform bone is represented in Delphinus tursio by a small ossification on the
ulnar border of the carpus, and attached to the lower end of the ulna itself. In Inia a
similar ossification projects from the same border of the carpus, but on a level with the
bones of the second row}. In Physeter it occupies a more normal position. These are
the only instances that I have met with of the occurrence of this element in an ossified
state in any of the Toothed Whales, though it must be admitted that, as it is a part so
liable to be removed in cleaning the skeleton, it may be more frequently developed than
our prepared specimens would indicate.

2. Excluding the above, the carpus of the Odontoceti appears never to consist of
more than six bones, three belonging to the proximal and three to the distal row.

3, The three bones of the proximal row are constant, and may easily be identified as
corresponding to the scaphoid, semilunar, and cuneiform of anthropotomy, or the

* This was wanting in the limb figured, having probably been removed during the partial process of cleaning
it had undergone before I received it, The bone represented in the figure is taken from the hand of another
specimen, in which it was still in situ.

+ “Sur le Squelette de l'extrémité antérieure des Cétacés.” Mém. de l’Acad. Roy. de Belgique, t. xviii. 1865.
+ As in the Chelonians,

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 361

radiale, intermedium, and ulnare of Gegenbauer. ‘The middle one is usually the
largest and most thoroughly ossified. In the genus Orca alone there is no ossification,
even in the adult animal, corresponding to the cuneiform.

4. The three bones of the distal row are generally represented by distinct ossifica-
tions in Hyperoodon, Beluga, and Monodon. These appear to correspond with the
trapezoid, magnum, and unciform of human anatomy.

5. In most cases the bones of the distal row of the carpus are reduced to two, which
appear to correspond best with the trapezoid and unciform, the magnum being either
absent or amalgamated with the trapezoid. I here differ from Dr. Van Bambeke, who
considers that the two bones of the distal row represent the magnum and the trape-
zoid, the unciform being absent. My reasons are :—

a. The magnum in mammals generally is a smaller and less important bone than the
unciform.

é. In those animals in which the carpus approaches most nearly to that of the Cetacea
in configuration and functions, but retains all its elements distinct, as the Manatee, the
magnum and the trapezoid are particularly reduced, while the unciform is large, and
occupies the position of one of the well-developed bones of the cetacean carpus.

c. In those Cetaceans in which all the bones of the second row are developed, and
can be distinctly recognized, as in Hyperoodon, the magnum is small.

d. In the skeleton of a Beluga I have found the magnum present, while in the
carpus of another animal of the same species, otherwise completely ossified, its place
was occupied by cartilage.

e. Whenever the magnum is present as a distinct bone, it is placed exactly above
the middle of the third metacarpal bone, and has a carpal bone on each side of it,
articulating with the second and fourth metacarpals respectively. In the Cetacea having
but two bones in the second row of the carpus, the middle of the proximal end of the
third metacarpal corresponds with the interval between these bones, which articulate
respectively chiefly with the second and fourth metacarpals, each taking a portion of the
third. Sometimes this interval is so large as to suggest the absence of a bone.

f. It is, however, not improbable that the bone which in general position corresponds
with the trapezoid, as it forms the chief support to the second metacarpal, may also
contain the representative of the magnum. ‘This seems clearly to be the case in the
carpus of Micropteron sowerbyense, according to a sketch kindly sent me by Professor
Yan Beneden, in which a single, transversely elongated bone corresponds to the bases
of both the second and third metacarpals. This view is also confirmed by the extremely
reduced condition of the trapezoid in the Manatee.

6. The trapezium appears never to be present as a distinct bone, although the first
metacarpal so often assumes the characters and position of a carpal bone that it may
easily be taken for it. ‘The rudimentary and simple character of the bones of the manus
of the Cetacea is well illustrated by the difficulty of interpreting the nature of the

362 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

bone which generally appears at the radial end of the distal row of the carpus.
Although in the manus of some forms, e. g. Globiocephalus, and some Delphini, one
could scarcely hesitate, at first sight, to identify it with the trapezium, the following
considerations induce me to agree with Dr. Van Bambeke in naming it the first meta-
carpal.

a. Its characters are in many species intermediate between those of a carpal and a
metacarpal bone. This is the case in Phocwena, Pseudorca, Physeter, and some Delphini ;
while in Monodon, and especially in Delphinus tursio, it has perfectly acquired the cha-
racteristic elongation of a metacarpal.

b. There is a great tendency for the metacarpal at the other extremity of the
series (the fifth) to assume many of the characters of a carpal bone, especially at its
proximal end.

c. The most crucial test appears to be afforded by its early ossification, corresponding
in this respect with the true metacarpals. Thus in the genus Orca, while there is but
a single ossific nodule in the middle of the cartilaginous carpus, the bone in question
is as well ossified as are the undoubted bones of the metacarpal segment.

It is quite possible that the trapezium may be contained in the bone which I have
called above “ scaphoid,” and that this should, therefore, be named “ trapezio-scaphoid ; ”
but of this I am not able to furnish any proof.

7. The cuneiform always directly supports the fifth metacarpal, and frequently some
part of the fourth. Moreover, in those hands in which the ulnar side of the carpus is
greatly reduced, e.g. Globiocephalus, the fifth metacarpal has even a connexion with
the ulna itself. This condition of the Cetacean carpus is illustrated in other mammals
in which the manus approaches or attains the form of a paddle,—as in the Seals, and
more distinctly in the Sirenia, the fifth metacarpal is in more or less direct relation.
with the cuneiform of the proximal row. The same occurs also in some terrestrial
mammals, as the Armadillos.

The carpus of the Cachalot follows the simplest type of the Cetacean manus, the
chief peculiarity being its shortness and lateral expansion, by which the bones of the
second row are, as it were, forced up between those of the first, so as to bring them all
nearly on the same level. There is, however, no difficulty in identifying them. Accord-
ing to the foregoing propositions, they will represent:—the scaphoid or trapezio-
scaphoid (Plate LXI. fig. 1, s), the lunar (/), the cuneiform (c), the unciform (w), the
trapezoid or trapezo-magnum (f), the pisiform(p). The symmetry of their arrangement
is well seen in the figure*.

The digits are five in number. The bones of which they are composed are tipped
at each end by a thin layer of cartilage, which in no instance among the specimens
examined contained an osseous epiphysis; nor is there any sign of a terminal epiphysis
having been united to any of these bones. This is somewhat remarkable, as in some

* In Wall’s description the distal epiphyses of the radius and ulna are taken for bones of the carpus.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 365

of the Toothed Whales, e.g. Globiocephalus, the metacarpals and phalanges are com-
pleted by very large epiphyses. The cartilaginous plates terminating two contiguous
bones are not blended together as in the Whalebone-Whales (where the phalanges
appear only as separate ossifications in a continuous rod of cartilage), but are quite
distinct, and, when a longitudinal section is made, show a free space between them, not
unlike a synovial cavity; but the dry condition of the specimens prevented a very satis-
factory investigation of this point.

The digits spread considerably from each other, giving breadth to the hand; the fifth
especially stands far apart from the others; but the first, or thumb, is, as usual,
adpressed towards the second digit. The first is by far the shortest; the second the
longest; the third almost equal to it; the fourth slightly, and the fifth considerably
shorter. The phalanges generally are elongated, compressed, and narrower at the
middle than at the ends, the last peculiarity being more characteristic of the meta-
carpals and proximal phalanges than of those situated more distally.

The exact enumeration of the phalanges of the digits, for any given species of Ceta-
cean, is never very easy, as the terminal bones are often slow and irregular in their
ossification, sometimes being represented only by cartilage, or by minute nodules of
bone readily lost in the process of preparation.

As none of the hands belonging to the skeletons examined were quite perfect, I
have given a figure of this part from another specimen, in which all the bones are
retained in their exact relative position and distance apart (Plate LXI. fig. 1). The
pisiform bone and the terminal phalanx of the second finger were wanting in this other-
wise complete specimen; they have been added to the figure from another of the sepa-
rate pectoral limbs sent to the Museum by Mr. Crowther.

The first digit has a short metacarpal, the broad upper extremity of which articulates
above with the scapho-trapezium, and by its side with the second metacarpal. ‘This is
succeeded (in the specimen figured) by a single slender tapering phalanx, the apex of
which reaches rather below the articulation between the metacarpal and first phalanx
of the second digit. Although in the present specimen this is a single bone, it appears
usually to consist of two, the terminal one being much the smaller, and generally more
or less ankylosed to the other. In some cases I have found them completely distinct.
In one of the hands belonging to the Tasmanian skeleton, these phalanges are not only
united together, but also to the distal half of the contiguous second metacarpal (see
Plate LV.).

The second digit has a large metacarpal, which may be distinguished from all the
others, by its superior size, and by a small lateral articular surface on the radial side of
its proximal extremity, where it is in relation with the first metacarpal. ‘This bone is
followed by five phalanges gradually diminishing in size.

The third digit has likewise five phalanges, besides the metacarpal; and the fourth
has four. The metacarpal of the fifth digit differs from the others in being narrower in

364 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

proportion to its length, and having less of the hourglass-shape, its outer border being
nearly straight. It is followed by three phalanges, which rapidly decrease in size.

Length of Bones of Manus.

|
Tasmanian Tasmanian} Tasmanian] y, :
skeleton. | No, 1. No. 2, Vomabie,
inches. inches. inches, inches.
1st digit, metacarpal... . 23 3. 23 43
phalanges ....| 53 ri 33 & 3 | 33 & 23
2nd digit, metacarpal....} 54 iG 5 53
Ist phalanx ..| 5 6 4 43
2nd _, ol 5 3} 3
3rd, Sie 3 22 134
4th “_,, 2 13 Biers
Sth, sob agcot 2goe il Aoiets
3rd digit, metacarpal....} 53 64 5 53
Ist phalanx ..| 42 6 37 43
2nd_ sa, ..| 33 4} 34 4
Srdiek 45 .-| 22 3 2} 23
4 4th 5, Gall onan 13 13 nici
5th =, 22 Rene: Fi 1 5H 6b
4th digit, metacarpal....] 4? 6 43 43
Ist phalanx ..| 4 5} 32 4
Oni Gas, dhekalie Bd 34 24 33
3rd, oe 2 23 2 13
4th ,, Mele mers 13 i re
5th digit, metacarpal....| 43 53 4 43
Ist phalanx .. Ft 3g 23 3
2nd, as 23 24 2
3rd-_,, Pa 1 Te50
Pelvis.

In all known Toothed Whales the sole rudiment of a pelvis is formed by a pair of
elongated subcylindrical bones (ossa ischia) placed horizontally and nearly parallel to
the vertebral column*, opposite to the junction of the lumbar and caudal regions, and
giving support to the crura of the penis or clitoris, as the case may be. In no species
of this group have any such accessory bones or cartilages representing the hinder
extremities, as those discovered by Reinhardt in Balena mysticetus, and subsequently
by other observers in several of the Whalebone-Whales}, been hitherto observed. It
must be remarked, however, that they have never been looked for with much care in
any of the larger members of the group, as the Cachalot or Hyperoodon.

The description and figure given by Wall{, which represent each lateral half of the
pelvis of a young female Cachalot as composed of two bones, placed end to end, does not
accord with the general observation upon the more common Dolphins, that each of the
ischial bones ossifies froma single centre. Unfortunately I have no materials to con-

* In the Porpoise, their anterior extremities diverge from each other,

+ Eschricht, Nordisch. Wallthiere, p. 186; W.H. Flower, Proc. Zool. Soc. 1865, p. 704.
+ Op. cit. p. 32, and pl. i. fig. 4.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 365

firm this description. In addition to the single bone which accompanied the skeleton,
Mr. Crowther has presented to the Museum two other pelvic bones, taken also from
adult male animals; but neither of them bears traces of any original segmentation.

They all present the general characters common to the corresponding bones in other
Toothed Whales, but have, as is so frequently the case, strongly marked individual
peculiarities. They are all more or less compressed, slightly expanded at one end,
which was tipped with cartilage, and present some modification of a sigmoid curve.
The bone which belonged to the skeleton (see Pl. LX. figs. 5 and 6) is 14" long; at the
middle its diameters are 1-3 and 095; at the expanded end 2‘ and 1"-1. Its surface
generally is simple and smooth; but at 4" from the smaller end one of the margins rises
into a triangular elevation surmounted by two short rough processes having different
directions.

Of the two other bones (which, though apparently belonging to opposite sides of the
body, are stated to have been taken from different individuals), one (Pl. LX. fig. 7),
though closely corresponding in actual length and thickness to the last, is so strongly
curved that in a straight line its two extremities are but 12" apart. Its surface is more
angular, presenting several strong longitudinal ridges and grooves. Near the middle of
its greatest convexity isa small but prominent spine 0'"4 in length; but otherwise there
is no appearance of the lateral process so well marked in the last. Both ends were
tipped with cartilage.

The third bone (fig. 8) is the largest of the three. It is distinguished by a very
regular sigmoid curve, by its smoothness, and the absence of all spine or process, and
especially by. its great compression, and corresponding width in the opposite direction.
Its length is 14""1, its diameters at the middle 2"-4 and 0-9, at the broader end 3'-7
and 0'-9. As in the first-described specimen, the more pointed extremity is smoothly
rounded and evidently complete; the condition of the other end shows that it had a
cartilaginous continuation.

The principal part played by these bones in the economy of the animal would
prepare us to find that they presented considerable differences according to the sex of
the individual to which they belonged; but in those genera in which the entire magni-
tude of the male and female differs but little, we do not generally find a very marked
difference in the pelvic bones. Thus, in two perfectly adult Porpoises of nearly equal
dimensions, the length of the pelvic bones of the male was 54", of the female 41”"*. In
the Hunterian collection is a dried preparation of the penis of a Hyperoodon with the
ischial bones still attached to the crura. ‘The animal, from the size of these and other
parts preserved in the Museum, as well as from a statement of Hunter, must have been
much larger than the ordinary Hyperoodon rostratum, and probably belonged to the
supposed species called H. latifrons, Gray, regarded by Eschricht as the adult male of

* It appears, according to Eschricht’s observations, that in the genus Orca the pelvic bones show consi-
derable sexual differences. (Recent Mem. on Cetacea, Ray Soc. 1866, p. 176.)

VOL. VI.—PART VI. 3 E

366 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

the former. The right bone is 10"-2 in length in a straight line, the other half an inch
shorter. They differ in many particulars from the same bones in the Cachalot. They
have a single, strong, and tolerably uniform curve, are but slightly compressed, much
thickest in the posterior half, and gradually tapering forwards, and, though presenting
some well-marked longitudinal ridges, have no angular processes projecting from. the
surface. The disparity in size between these bones and those of a perfectly adult female
of HZ. rostratum in the Museum of the University of Oxford is extraordinary, the latter
being but 5” in length, and very much more slender in proportion. As the male
Hyperoodon does not exceed the female in bulk so much as the male Cachalot is said
to do the opposite sex, we may expect to find fully as great a difference in the pelvic
bones of the two sexes of thisanimal. In the Museum of the Royal College of Surgeons
is a bone (No. 2460, Osteol. Series) presented by the late Dr. Buckland, and. described
in the catalogue as “the left pelvic bone (ischium) of a Cachalot (Physeter macroce-
phalus).” It may possibly have belonged to a female of this species, though it presents
but little resemblance to those of the males above described, and, indeed, is more like
that of the Hyperoodon. Its length is 83”; its general form that of a club, being
dilated towards one extremity, and much attenuated at the other. It is but slightly
compressed, and has a single strongly marked curve.

Conclusion.

In the foregoing description of the skeleton of the Cachalot I have made but little
comparison with animals of the genus Kogia, having in fact had no opportunity of doing
so. The only two skeletons at present existing are both at Sydney, and-no adequate
description of them has yet been published. From such indications as we have (further
elucidated by some photographs kindly sent me by Dr. Bennett and Mr. Krefft), there
can be no doubt that they belong to a genus which, both in external and osteological
characters, is perfectly distinct from, though nearly allied to, Physeter. In the sketch
of the classification of the Cetacea, appended to the description of the skeleton of
Inia*, the genera Physeter and Kogia are united to form the subfamily Physeterine,
which, with the Ziphiine (including Hyperoodon, Ziphius, &c.), form the very natural
family Puysererip%. A detailed examination of every part of the skeleton of Physeter
has perfectly corroborated the position then assigned to this genus. Materials are at
present greatly wanted to complete our knowledge of the osteology of the Ziphiine ;
hence it is impossible to say to which of the genera of that section Physeter approxi-
mates most closely.

After a concise and masterly analysis of the almost inextricably perplexed literature
of the zoology of the Cachalots}, Cuvier came to the conclusion that, up to the time at
which he wrote, but a single species could be considered to be truly known. Since then
the claims of at least one distinct species to a place in the zoological system have been

* Trans. Zool. Soc. vol. vi. p. 110, 1867. + <Ossemens Fossiles,’ edit. 1836, vol. viii. p. 189.

be |

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 36

urged, and on far safer and more scientific grounds than the vague descriptions and
partly imaginary drawings on which most of the earlier Physeteres and Catodontes were
founded. Of these the most important is that set forth in the oft quoted memoir of
Wall, where the detail with which the skeleton of the southern Cachalot is described
and compared with what was known of the Cachalot inhabiting the northern seas, has
succeeded in establishing, to the satisfaction of most zoologists, the species P. (Catodon)
australis as distinct from P. (Catodon) macrocephalus. The diagnostic characters
relied upon are as follows:—1l. The entire head as compared with the body is rela-
tively smaller. 2. The skull is shorter in proportion to its width and height. 3. The
lower jaw is proportionately shorter. 4. The form of the sternum is different. It
has been already shown that the first. three of these characters depend simply upon
the immature condition of the specimen described. In the fourth the author has
been misled by Beale’s description of the incorrectly articulated skeleton at Burton
Constable, in which the body of the hyoid is appended to the hinder end of the
sternum. Putting aside these supposed distinctive characters as valueless, there is not
one other, presenting any approach to a specific distinction, pointed out throughout the
memoir. Catodon australis, therefore, as founded and characterized in Wall’s work, can
have no existence as a zoological species.

It will be gathered from the foregoing memoir that, although numerous discrepancies
have been met with among various bones of the different skeletons examined, in some
cases so marked that, if two individuals alone were known, they might easily have been
considered specific, the comparison with a third example has nearly always proved a
corrective to such a supposition,—and that, taken as a whole, the Yorkshire skeleton
differs from the Caithness skeleton as much as the Tasmanian does from either. I am
therefore quite unable, from the materials at present available, to point out any constant
difference of specific. value between the Cachalot inhabiting the Australian seas and
that occasionally visiting our northern coasts.

It must not be inferred from this statement that I deny the possibility of their being
specifically distinct. Similarity of osteological characters does not prove unity of
species. Who would have suspected the specific distinction of the Lion and the Tiger, or
the Quagga and the Zebra, if these animals were only known by two or three skeletons
of each? But, at the same time, no new species should be admitted into the. system,
unless its distinction is established either by well-marked and constant (1) anatomical
characters, (2) external characters, (3) geographical distribution, or (4) peculiarity of.
habit and mode of life. On no one of these points has it yet been shown satisfactorily
that the southern and northern Cachalots differ.

With reference to the geographical distribution, it may be remarked that, unlike the
Right Whales, the Cachalots are essentially inhabitants of the tropical and warmer parts
of the temperate seas, and that they pass freely from one hemisphere into another.
Between the North Atlantic and the Australian seas there is. no barrier interposed to

3E2

368 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

animals of such great powers of locomotion: they are known to round Cape Horn; and
in fact there is scarcely a spot between the two seas where they have not actually been
encountered. Whatever may have been the case in former times, the Cachalot can
hardly now be considered a regular inhabitant of the seas bordering Europe. Those
that occasionally appear are either solitary stragglers, or more often dead and partially
decomposed carcasses, floated northwards, probably by the Gulf-stream. The occurrence
of a “school” of Cachalots, like that at Citta Nuova, in the Adriatic, in 1853, is quite
exceptional *.

Many museums contain lower jaws of very small Cachalots, which, judging from the
condition of the bones and teeth, are perfectly adult. One, in the Museum of the
Royal College of Surgeons, is 6’ 10” in length. The symphysis is 40"; and it has
twenty-two teeth on each side. Another, in the Oxford University Museum, is 7! 02”
in length. These are usually considered to be the jaws of the female of the common
species; if this is not the case, they must indicate a second species of the genus. An
entire skeleton, or even a cranium of a female Cachalot, is still a desideratum, and one
which ought soon to be supplied, as, owing to its comparatively small size, it would
not be beyond the means, as to cost or space, at the disposal of many museums.

It is a singular circumstance that the deformed and twisted jaws before mentioned
appear all to be of a size corresponding with those just referred to.

There is good reason to believe that, as with all the other large Cetaceans, the mag-
nitude of the Cachalot has been greatly exaggerated. Leaving out of the question all
earlier and even less trustworthy descriptions, Beale states that he was present at the
capture of a Cachalot which measured the length of eighty-four feet +; while F. D.
Bennett. says, “the largest size authentically recorded of the Sperm-whale is seventy-six
feet in length, by thirty-eight in girth; but whalers are well contented to consider sixty
feet the average of the largest examples they commonly obtain” {. It is probable that
the natural and often unconscious proneness to exaggerate the size of such an object,
especially where measurement with anything like scientific accuracy is almost im-
possible from the very circumstances of the case, must be allowed for, in the former
of these statements. The only indications on which we can absolutely rely are the
osseous remains, which perfectly corroborate the latter part of the information given by
Bennett. No single specimen of all the different skeletons, or fragments of skeletons,
some of which are quite aged, examined by me give any evidence of a greater length
of skull and vertebral column than fifty-five feet, if quite so much. The soft parts
of the head, and the portion of the flukes projecting beyond the median notch of
the tail, which corresponds to the termination of the bony column,might bring the
animal in the flesh up to sixty feet; and from the tolerable uniformity in the size
of all the adult skeletons, skulls, and lower jaws (now in considerable quantity)

* Heckel, in Wiener Sitzungsber. d. Math.-Naturw. Cl. Bd. ii. (1853) p. 765.
+ Op. cit. p. 15. t ‘ Narrative of a Whaling Voyage,’ vol. ii. p. 154.

MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 369

known, I venture to question whether the Cachalot frequently, if ever, exceeds that
length, when measured in a straight line. Mr. Crowther assures me that the specimen
described in this memoir was considered a full-sized animal. But the most important
evidence upon this head is derived from a magnificent lower jaw, also presented to
the Museum by that gentleman, which was considered in the colony ‘‘ unique on
account of its great size.” This jaw measures in a straight line, from the tip to a line
drawn across the hinder edges of the rami, 16' 2", or 1 inch longer than that of the
Yorkshire skeleton, and but 20" longer than that of the Tasmanian skeleton. The fact
that the animal to which this jaw belonged was considered by experienced men a giant
among Cachalots gives a good indication of what size the ordinary individuals attain.

Lastly, a few words on the zoological designation of the Cachalot, presuming that
there is at present but one well-established species. Following the rules laid down by
the Nomenclature Committee of the British Association, we find in Linnzus’s ‘ Systema
Nature’ (12th edit. 1766), the genus Physeter including every animal, then known or
imagined, with which the Cachalot can possibly be identified. Of the four species
assigned to this genus, the one called macrocephalus is without doubt that especially
founded upon the common Cachalot, notwithstanding the error in the diagnostic ex-
pression, “fistula in rostro.” The P. catodon was a small species, probably the Beluga;
the P. microps and tursio had high dorsal fins; while the references under the head of
P. macrocephalus to Clusius’s description, the statement as to the size and the number
of the teeth, and especially to the “ spermaceti e ventriculis cerebri,’ all point indis-
putably to the great Sperm-Whale.

Artedi’s name of Catodon has been revived as the generic designation of the Cachalot
by several zoologists, whose faith in Sibbald is so great as to retain in the system, upon
the strength of his description and figure alone, an animal of which, as Dr. Gray says,
‘there is not a bone, nor even a fragment of a bone, nor any part that can be proved
to have belonged to a specimen of this gigantic animal, to be seen in any Museum in
Europe” *. If the Linnean genus Physeter is to be kept in abeyance until the redis-
covery of Sibbald’s “ Balena macrocephala tripinna” +, it is to be feared that it may
ultimately disappear altogether from zoological literature.

* «Cat. Seals and Whales in Brit. Mus.’ (1866), p. 215.
t ‘Phalainologia Nova,’ 1692,

370 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

DESCRIPTION OF THE PLATES,

PLATE LY.

Skeleton of the Tasmanian Cachalot, as mounted in the Museum of the Royal
College of Surgeons. Scale 3;. The ribs and pectoral limb of the distant side are
omitted for the sake of greater distinctness. The small eleventh rib is almost entirely
concealed by the one in front of it.

It is but due to the long-continued and valuable services rendered to the study
of comparative osteology by my namesake, Mr. James Flower, the articulator to the
College, whose mechanical skill, combined with extensive and accurate anatomical
knowledge, has enabled him to bring the art of mounting skeletons to the greatest
degree of perfection yet attained, to mention his name in connexion with this, his
greatest work.

pm. premaxillary. so. supraoccipital.
m. maxillary. eo, exoccipital.
d. g. dental groove. sh. stylohyoid.
pl. palatal. bh. basihyoid.
f. frontal. th. thyrohyoid, :
s. squamosal. p. parietal ?
j. jaugal.
PLATE LVI.

Fig. 1. Vertical longitudinal section of the skull of the same Cachalot.

Fig. 2. A similar section of the skull of a very young Cachalot, from Tasmania.

Fig. 3. A similar section of the skull of a nearly adult Hyperoodon, in the Museum of
the Royal College of Surgeons.

All +5.
pm. premaxillary. bo. basioccipital.
m. maxillary. as. alisphenoid.
v. vomer. bs. basisphenoid.
e. ethmoid. ps. presphenoid.
if. foramen corresponding to the pt. pterygoid.

* infraorbital.” pl. palatal.

f. frontal. n. nasal,

so. supraoccipital.

Mk. W, H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

PLATE LVII.

veo
=I
—

Cranium of a young male Cachalot from Tasmania, presented (as were all the other
specimens from which these figures were taken) to the Museum of the Royal College
of Surgeons by W. L. Crowther, Esq. The animal was supposed by the whalers to be

about ten months old.
Scale 4.
Fig. 1. Upper surface.
Fig. 2. Lower surface.
ty. tympanic bone. aon. antorbital notch.

The other letters as in the former Plates.

PLATE LVIII.

Vertebral column of the skeleton of the Tasmanian Cachalot, seen from above.

1
Scale Ta:

PLATE LIX.

Vertebral column and chevron bones of the same skeleton, seen from the side.

fourth and fifth chevron bones were missing.
Scale =};.

PLATE LX.
Fig. 1, Hyoid bones of the same skeleton.

sh. stylohyal. bh. basihyal. th. thyrohyal.
Scale <4.
Fig. 2. First rib of the right side of the same skeleton. Anterior view.
Scale 345.
Fig. 3. Sternum of the same skeleton. External surface.
Scale +5.
Fig. 4. The same. Side view.
Scale 35.
Figs. 5, 6. Two views of the pelvic bone of the same skeleton.
Scale %.

Figs. 7, 8. Two other pelvic bones of Tasmanian Cachalots, referred to at p. 369.

Scale ~.

PLATE LXI.

Fig. 1. Bones of the anterior extremity of another Cachalot, from Tasmania.
Scale ¢.
A. humerus. 7. radius. -
u. ulna. 0. olecranon.

The

372 MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE.

e. lower epiphysis of the radius. 7. lunar.

e’. lower epiphysis of the ulna. uw. unciform.
s. scaphoid or trapezo-scaphoid. ¢. cuneiform.
‘t. trapezoid or trapezo-magnum. p. pisiform.

Fig. 2. Scapula of the skeleton of the Tasmanian Cachalot, from below.
Seale +4.
Fig. 3. Outer surface of the scapula and bones of the arm of the same skeleton.
Seale 51.
Fig. 4. One of the carpal bones of the same skeleton, showing the peripheral epiphysial
ossification.
Scale 3.

Side view of a cast of the interior of the cranial cavity, } natural size. (See p. 316.)

Fig. 13.

Cast of the interior of the cranial cavity, seen from above, .

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at?

XIII. On a Picture supposed to represent the Didine Bird of the Island of Bourbon
(Réunion). By Aurrep Newton, WA., FLS., F.ZS., &e.

Read February 14th, 1867.

[Piate LXII.]

PICTORIAL evidence contributes so largely to what we know of the Dodo and its
allies, that in calling the attention of the Society to the old water-colour drawing now
exhibited I should not deem any apology requisite, were it not that this drawing has
been already displayed at a meeting only a few months ago. But as on that occasion
the exhibitor, Mr. Tegetmeier, did not place on record the remarks he made (P. Z. 8S.
1866, p. 201), and as I have reason to believe these remarks did not touch the points to
which I am about to advert, I trust I may be excused for again submitting the drawing
to the inspection of the Society. I must first of all express my thanks to Mr. Tegetmeier
for the opportunity he has given me of examining the drawing, and also to the owner of
it, Mr. C. Dare, of Clatterford, in the Isle of Wight, for his kindness in permitting it to
be copied for our ‘ Transactions.’

It will be remembered that in the late Mr. H. E. Strickland’s work, ‘'The Dodo and
its kindred,’ the former existence of at least three distinct species of Didine birds was
very clearly demonstrated. Of these the true Dodo (Didus ineptus) was presumed to
have been peculiar to the Island of Mauritius, the Solitaire (Pezophaps solitarius) to
that of Rodriguez, and the third (which Mr. Strickland left unnamed) to that of
Bourbon, or, as it is now called, Réunion. Of the first two there were then no incon-
siderable remains known; but of the third it was believed that nothing existed, save a
few scanty notices, which were industriously compiled by that lamented naturalist from
the narratives of various voyagers. The earliest of these, Tatton, who visited Bourbon
in 1613, speaks of “a great fowl of the bigness of a Turkie, very fat, and so short-
winged that they cannot flie, beeing white.” In 1618 Bontekoe passed three weeks in
the island; and his account confirms the former statement. He calls the birds
“ Dodeersen,” the name often applied to the true Didus ineptus, whence we may suppose
they generally resembled that species; but he does not mention their colour. Carré, in
1668, speaks of the Bourbon brevipennate, “Il ne ressembleroit pas mal a un Coq
WInde, s'il n’avoit point les jambes plus hautes. La beauté de son plumage fait
plaisir 4 voir. C'est une couleur changeante qui tire sur le jaune.” In the following

VOL. VI.—PART VI. 3F

374 MR. A. NEWTON ON THE DIDINE BIRD OF BOURBON.

year a French colonist from Madagascar, the Sieur Du Bois*, gives a more detailed, but,
I suspect, a not very accurate, account of the species, under the name of “ Solitaires.”
Here it is said of them, again, that they “ont le plumage blanc.” Thus two out of
the four eye-witnesses speak to the plumage of the Bourbon Didine bird being white; a
third calls it ‘‘a changeable colour, which verges upor. yellow,” which, as Mr. Strickland
justly observes, “is rather vague, but seems to imply a pale yellowish or cream-coloured
tint, which another author might easily have described as white” (‘The Dodo,’ &c.,
p. 60). The fourth witness does not mention the colour at all.

This fourth witness, Bontekoe, however, furnishes some other evidence of value. He
calls the birds by the name of the true Mauritian Dodo; and, from his description, they
undoubtedly much resembled that species in form. But further, one edition of Bontekoe’s
work, published at Amsterdam by Gillis Joosten Zaagman in 1646, contains a figure
professing to be that of the Bourbon “ Dodeers.” ‘This is reproduced in fac-simile by
Mr. Strickland; and though that gentleman says (op. cit. p. 63) * there can be no doubt”
it “refers to the true Dodo of Mauritius,” I see no reason whatever for arriving at that
conclusion. ‘This figure is unlike all the original representations of the true Dodo in
several minor points, but especially in one respect. The first four primaries are directed
downwards, and at the extremity forwards. Now, in every picture and figure of the true
Dodo that I know of, all the primaries are directed backwards.

I think, therefore, we may not unreasonably infer :—

1. That the Didine bird of Bourbon in general shape resembled the true Dodo (Didus
ineptus) of Mauritius.

2. That the plumage of the Didine bird of Bourbon was white, with some admixture
of yellow.

3. That in the Bourbon Didine bird the first four primaries of the wing were not
directed backwards, but downwards and forwards.

A glance at the picture now exhibited (P1. LXII.) will show how far it fulfils these
conditions.

But, on the other hand, I must not pass over what seems to be a formidable objection
to the supposition I have laid down. Du Bois describes his “* Solitatres” as having “le
becq fait comme celuy des Bécasses, mais plus gros.” Nothing more unlike a Wood-
cock’s bill can be imagined than that of the bird represented in Bontekoe’s figure and
the drawing here! But not one of the other eye-witnesses refers to such a peculiarity.
Two of them liken the Bourbon bird to a'Turkey, the third to a true Dodo; surely, then,

* When Mr. Strickland wrote, in 1848, he was only able to cite this witness from a MS. copy of a journal
presented by Mr. Telfair to this Society, and still in our library, in which the name of the author was not
given, but merely his initials. From a note of M. Milme-Edwards in a recent number of the ‘Annales des
Sciences Naturelles’ (vol. vi. pp. 42-44, July 1866), we learn the name of the author of this journal, which,

we are informed, was published at Paris in 1674.

MR. A. NEWTON ON THE DIDINE BIRD OF BOURBON. 3

if it had possessed a Scolopacine bill, the fact would have been mentioned. I venture
to suggest that Du Bois must have written some other word, and that “ Bécasses” is a
false reading, or else that a treacherous memory supplied the statement.

It now remains for me to remark on the picture exhibited. It represents apparently
a flooded meadow, in the pools of which various aquatic birds are distributed, while the
Dodo is standing, with an expression of alarm on his countenance, on a scanty bit of dry
ground. By his side is seated a fine Bernicla ruficollis, somewhat too brilliantly coloured
perhaps; and the other birds portrayed are Cepphus grylle (engaged with a small eel or
snake), Mergus castor (a female or immature male), Ciconia alba, Clangula glaucion,
Fulix fuliqula, and Mareca penelope (a female). ‘These are all drawn with much atten-
tion to detail, and generally very fairly coloured. ‘The Dodo and the Goose form the
principal figures in the composition. The beak of the Dodo, as represented here, also
demands a word of comment; instead of terminating in the formidable dertrum to
which we are accustomed in the pictures * of the Saverys and that of Goeimare (Trans.
Zool. Soc. iv. p. 197), its tip is rounded off, as if it had undergone the operation known
among falconers as “coping.” Now I cannot help thinking that in this point we have
some grounds for believing that the subject of the figure must have been a bird kept in
captivity. ‘The Dodo was no doubt able with its powerfully-hooked beak to inflict very
serious injury; and it is not at all improbable (so it seems to me) that the keeper of
such a bird would consult his own safety, and, by trimming an offensive weapon so
likely to be used against him, deprive it of the means of doing harm. On this account,
therefore, I think there seems to be a strong probability of this drawing having been
taken from a living subject which had been brought to Europe and kept in some
aviary. It is further to be remarked that the inside of the Dodo’s mouth in this
drawing is coloured of a bright red, and a red ring is seen surrounding the eye, though
whether this be intended for the iris or an orbit it is not so easy to say. I believe that
all the pictures of Didus ineptus concur in representing the iris of that species to be
yellow.

And now as to the artist by whom this drawing was executed. In its left hand

corner are to be plainly seen the letters pif and on consulting Brulliot’s

‘ Dictionnaire des Monogrammes’ I find (p. 821, Nouv. Ed. Sec. Partie. Munich: 1853)
that this is the signature of Pinrre Wir7rHoos, “ qui peignait a la gouache des fleurs,
des insectes, et des plantes avec beaucoup d’art et de vérité,” and who died at Amster-
dam in 1693. It is therefore quite possible that the figure I have before mentioned in —
Zaagman’s edition of Bontekoe, published, according to Strickland, shortly after the

* Professor Schlegel has already suggested that the picture of the Dodo in the British Museum was drawn
from a bird the beak of which had become unnaturally elongated in captivity (Verslag. en Mededeel. der Kon,
Akad. van Wetensch, 1854, p. 237, note).

376 MR. A. NEWTON ON THE DIDINE BIRD OF BOURBON.

[Ss]

year 1646, at Amsterdam, and the present drawing were both taken from the same
source, probably a bird brought from the Island of Bourbon, and kept alive in that
town.

From a recent paper by Mr. W. J. Sterland in a popular periodical* I learn that a
portion of the picture now exhibited + was copied and engraved some years ago in the
‘Tllustrated London News’ (no. 821, vol. xxix. p. 303, Sept. 20, 1856). On referring
to the place indicated I find that the woodcut there given is a most wretched misre-
presentation, while the accompanying notice by Mr. W. W. Coker and Mr. Gould does
not at all bear upon the subjects to which I have here adverted.

* Hardwicke’s ‘ Science-Gossip,’ No, 25, Jan. 1, 1867, pp. 5, 6.

t+ P.S. April 1868.—The original picture, I am informed, has been recently deposited in a “ museum ” at
Carisbrook Castle, in the Isle of Wight.

duit quPyUeH NP Ww

UUT TAS Lr

XIV. An account of the Fishes of the States of Central America, based on collections
made by Capt. J. M. Dow, F. Gopman, Esq., and O. Satvin, Esq. By ALBERT
Ginruer, A., M_D., Ph.D., PRS. F.ZS.

Read March 22nd, 1864, and December 13th, 1866.

[Puates LXTII. to LXXXVII.]

§1. Introductory Historical Remarks on the Collections forming the basis of this
Memoir.

BEFORE proceeding to the enumeration and description of the fishes known to exist
in the States of Central America, I may be permitted briefly to notice the circumstances
which enable me to submit to the Society the results contained in the present memoir.

Mr. Salvin started in the year 1859 on his second excursion to Guatemala, chiefly
with the intention of working out the ornithological fauna of that country. But having
had his attention directed by me to the fact that its cold-blooded vertebrates were almost
entirely unknown, he made and brought home a small collection of reptiles and fresh-
water fishes, which proved to be of sufficient interest to encourage him to pay still more
attention to this subject on a third excursion, which he undertook in company with Mr.
Godman in the year 1861. By far the greater part of the materials which form the
basis of this memoir were obtained on this occasion. Not only did the two travellers
extend their excursions to various parts of Guatemala, but Mr. Salvin also visited
Panama, where he met and collected in company with Capt. Dow, of the Panama
Railway Company’s Steamer ‘ Guatemala.’

Capt. Dow, indeed, had commenced to collect fishes previously to this, having sent
several collections to the Smithsonian Institution in Washington, and to the Zoological
Society of London, whence they were transferred to the British Museum; and for the
last three years he has continued his researches with such zeal and liberality that I
cannot abstain from acknowledging here the great services. he has rendered to the cause
of science.

The collections made by these gentlemen contained not less than about 1500 examples,
in a perfect state of preservation, many of considerable size. In addition to these, I
have examined a few which had been purchased of a dealer for the British Museum
having been collected at Puerto Cabello in the Bay of Honduras, and, finally, those
collected by Dr. Seemann, originally deposited in the Collection of Haslar Hospital,
and now in the British Museum. The latter have lost much of their scientific value, as,
unfortunately, no record was kept of the localities where they were obtained; and only
in a few cases have I been able to avail myself of specimens of this collection, viz. where
the original label, with the name of the collector, has been accidentally preserved.

VOL. VI.—PART VII. 3G

ws
=I
lo 2)

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

§ 2. Topographical Features of the Localities explored.

As regards the topographical features of the localities explored by Messrs. Dow,
Godman, and Salvin, I have been favoured by the latter gentleman, by whom also the
accompanying map has been prepared, with the following notes :—

Lakes.

AmatitLan.—The Lake of Amatitlan is situated in lat. 14° 29' N., long. 90° 35! W., in
the Republic of Guatemala. Its elevation above the sea-level is about 4500 feet. Being
only a short distance on the southern side of the main ridge, it collects the waters of a
few small streams, which it discharges at its southern extremity, into the river Michatoya,
a mountain-torrent for half its course, then expanding, like all the rivers of Guatemala
which flow into the Pacific, into a broad shallow stream with a shifting sandy bed.
The lake is very deep, and its water clear. The volcanoes of Pacaya and Agua rise
amongst the mountains of its southern border, the whole forming a landscape of great
beauty. Fish are caught during the rainy season near the outlet into the river
Michatoya, and are sent to the market of the City of Guatemala.

AtitLAN.—The Lake of Atitlan is elevated 5000 feet above the sea. Like the last-
mentioned it lies in Guatemala on the southern side of the main ridge, in lat. 14°45’ N.,
long. 91°14’ W. It has no visible outlet. The water is clear and fresh, and the lake of
great depth. The hills on three sides attain to a height of 2000 feet above the lake.
On its southern border the two large volcanoes of Toliman and S. Pedro rise, their bases
being washed on one side by the lake, giving one the idea that one of them (that called
Toliman) has in rising acted as a dam and stopped the outflow of the waters of a
mountain-valley. A few small streams enter the lake, the water of which rises during
the rainy season, to fall again in the dry. On the mountain-slope below, several streams
take their rise, supplied probably by the filtration of water from the lake; but it would
appear, from the alteration of the water-level in accordance with the season of the year,
that it is chiefly influenced by evaporation, A number of Indian villages surround the
lake; at one of them, Panajachel, a small collection of fish was made. Fish never seem
to grow to any size in this lake, the Mojara (Heros) being quite diminutive, The
Indians fish with round nets amongst the reeds that grow at the mouths of small
streams. ‘The lake itself is about twenty-two miles long, and twelve miles wide,

Duewas.—This lake is little more than a depression in one of the elevated (5000 ft.)
plains forming the tablelands of Guatemala. Its depth is nowhere more than 6 feet,
and its banks are everywhere clothed with reeds. A small stream connects the lake with
the river Guacalate. Here, too, fish are caught by the Indians in round nets, which are
held by both hands, pushed in amongst the reeds, and suddenly brought to the surface,

HvamucuaL.—This name applies properly to a series of small lakes situated in about
lat. 14° 32’ N., long. 92° 13! W., close to one another, about six miles from the mouth of
the river Tilapa on the Pacific coast. The place isnot shown on any map ; but it is near

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 379

the large Lake of Tamachian, with which, in the rainy season, all these smaller lakes are
connected. During this period of the year the river Tilapa overflows its banks and
inundates the whole country round. In the dry season water remains in depressions of
the land, forming the lagoons of Huamuchal; but in years of great drought even these
dry up, the fish being destroyed; but a fresh supply finds its way from Lake Tamachian
during the next inundation. The water is slightly brackish. The fish are taken in
drag-nets, salted, and sold to Indians coming from the Altos of Guatemala.

ManaGua.—According to Mr. J. Bailey this lake is about fifty or sixty miles long, by
thirty-five miles wide. Its depth varies from 2 to 10 and 15 fathoms, but in its deepest
part reaches to as much as 40 fathoms. Its elevation above the sea is 156 feet. On its
south-western border the lake is separated from the Pacific by a series of comparatively
low hills, the lowest section of which, through the Plain of Leon, is only 230 feet above
the ocean-level. The high mountains of the Republic of Honduras approach the north-
eastern border of the lake. On its south-eastern side an opening communicates with
the Lake of Nicaragua. Commencing with the Fall of Tipitapa, of 22 feet height, the
river widens into the Estero of Panaloya, and thence into the larger lake.

Nicaracua.—The same authority gives a length of one hundred and five miles to this
lake, and a width of about forty-five, its depth being about 15 fathoms. ‘The surface of
the lake is studded with numerous islands, some of them, as Omotepec, being volcanic
cones. The elevation of the lake above the mean ocean-level is given as 128 feet. The
same line of low hills which divides Lake Managua from the Pacific separates Lake
Nicaragua from the same ocean ; but at no point is the elevation so low as at that above
indicated. The river San Juan, a deep-stream with several rapids, flows out of the
south-eastern end of the lake, and falls into the Atlantic Ocean, at the port of Grey-
town, or San Juan del Norte.

Peren.—The Lake of Peten is situated in lat. 17° 10! N., long. 90° W., and is one of
several lakes formed at the base of the Promontory of Yucatan. Its length is about thirty
miles, its width eight miles, and elevation above the sea 500 feet. The water is quite
fresh, clear, and of considerable depth. Neither the Lake of Peten nor the adjoining Lake
of Yasha has any outlet ; and in both the water is rapidly increasing in expanse—so much
so that several streets of the town of Flores, which stands on an island in Lake Peten,
have been absorbed within a few years, and the posts of huts, which formerly were on
dry ground, may now be seen standing in deep water. This increase of water can only
be accounted for by supposing that a common subterranean outlet has been stopped up,
or that the land of this district is experiencing a gradual subsidence. All the fish
obtained here were caught with a hook and line, or speared. All the natives, even quite
small children, are very expert in using a light spear formed of bamboo cane with an
iron barb at the end.

YzaBau.—This lake, which is also called the Golfo Dulce, is about thirty or forty
miles long, and ten to fifteen miles wide, and has a tolerably uniform depth of about 35

3a 2

380 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

to 40 feet. It is situated in lat. 15° 30’ N., long. 89° 15' W., at the bottom of the Bay of
Honduras. One large river, the Polochic, enters this lake ; and it has a narrow but deep
outlet to the sea, called the Rio Dulce, which is navigated by small schooners plying
between Belize and the town of Yzabal. It was near this last-mentioned place that a
few species of fish were obtained.

Rivers.

BayaNno.—This is a river which rises in the narrow part of Central America, and flows
into the Pacific a little to the southward of the Bay of Panama.

CaHaBon.—The town of Cahabon, where a few fishes were obtained, is situated on an
affluent of the river which bears this name. The main stream rises in the same marsh
as the Polochic, but takes another valley, in Vera Paz, and again joins the Polochic,
when they both flow into the Lake of Yzabal, and thence into the Atlantic.

CuaGres.—This is the principal river of the Isthmus of Panama. It flows into the
Atlantic. The fish were obtained near the railway bridge at Barbacoas, about halfway
across the isthmus.

Cuisoy.—Of the numerous names this river bears, I have chosen this for the principal
stream which forms the large river that flows out into the Laguna de los Terminos, in
the Bay of Campeachy. This branch is also known as the Rio Negro; and after
receiving the water of the Rio de la Pasion, or Rio de Santa Isabel, as it is also called,
the two are usually called the Usumacinta. Fishes were collected from this river near the
Indian village of Cubulco; and a number were also procured by poisoning with herbs a
small stream near Saouchil, an Indian village below the town of Coban, in Vera Paz.

GuAcALATE.—Is one of the numerous rivers which drain the southern watershed of the
main ridge into the Pacific. It flows past Antigua, the old capital of Guatemala. Fishes
were obtained about 3500 feet above the sea, where the river is still quite a torrent.

Moracua.—tThis river, the second largest in Guatemala, rises in the main ridge, and
flows, with high mountains on either side, nearly due eastward into the Atlantic. Fishes
abound in this river; and nearly every year a considerable length is poisoned, and a
large quantity obtained. On one of these occasions a collection was made a little below
the bridge over which the highroad from Guatemala to Vera Paz passes. Another
collection came from lower down the stream, below the village of Tocoy.

San Geronimo.—Is a tributary of the Chisoy before mentioned. A small collection
was made near the village of San Geronimo, in a plain at the foot of the mountains
whence it takes its rise.

Santa IsaBEL.—A small stream flowing into this river, one of the principal branches of
the Usumacinta, was poisoned, and a number of small fishes obtained.

San Satvapor.—A few small fishes were caught by Capt. Dow in a warm stream near
the capital town of this republic.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 381

Marine localities.

BeEtizE.—All fishes from Belize were from the market, and were caught amongst the
coral reefs which line this coast.

Carpon! Isnanp.—Is situated at the mouth of the fine harbour of Realejo, in Nica-
ragua. Fishes were found at low tide in the pools amongst the rocks, and caught with
a landing-net.

CurtapaM.—The whole coast of Guatemala, bordering the Pacific Ocean, is studded
with a number of lagoons formed at the mouths of the numerous rivers which flow
down from the neighbouring mountains. All these rivers are charged with volcanic
sand, which is thrown back by the heavy surf that rolls in on this coast. The body of
water brought down during the dry season is often insufficient to reduce this sandbar ;
and it frequently happens that all outlet to the sea is stopped. The accumulation of
water during the rainy season breaks this barrier; but it again forms when the water
subsides. About the period of the cessation of the rains the natives cut an artificial
channel, which, at first widening of itself, often remains open some months, each tide
bringing a great quantity of fishes into the lagoon, which are there netted by drag-nets.
The water is almost salt, but varies in this respect according to the size of the river
which enters it. A few fishes were also obtained by a hook and line from a canoe in
the open sea.

LisertaD.—This is an open roadstead, the port of the City of San Salvador. Whilst
we were lying at anchor here a few fishes were caught with a hook and line.

PanamMa.—Most of the fishes taken in the Bay of Panama were found in the pools
amongst the rocks at low tide. A reef running out from the town was an excellent
locality ; one spring tide Capt. Dow and I secured twenty-four species in the course of
half an hour.

San Josi.—Is the port of Guatemala on the Pacific side; a few fishes were caught
here in the open sea in a canoe.

§ 3. Definition of the Boundaries of the Fauna treated of in this Memoir.

Although we may presume that our account contains a tolerably. complete list of the
species inhabiting the localities visited, particularly as on several occasions poison (the
best means for securing a complete series of the fishes of a certain locality) was resorted
to, yet there is still a wide field for future explorers in a country where several forms
(such as Heros, Pimelodus, and the Cyprinodontes) are so much developed and specialized.
Of the fishes of Yucatan we still know absolutely nothing. The list of the marine fishes
of the Atlantic coast will, without doubt, be considerably swelled, as the gentlemen
mentioned paid much less attention to the Atlantic marine fauna (which would have
yielded comparatively few novelties) than to the freshwater fauna. And knowing how
little advantage is derived from, and how much confusion is caused by, receiving into a

1 This name is misspelt ‘ Cardoya” in several places in the 3rd volume of the ‘ Catalogue of Fishes.’—A. G.

382 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

fauna species which may be eapected to belong to it, although they are not yet disco-
vered within its limits, I have excluded all species not actually known from Guatemala,
although they have been obtained north and south of it. A collection made by Mr.
Godman at Belize was of great value in determining’ this part of the fauna.

Numerous species of fishes have been described trom Mexico’; and if we were better
acquainted with their geographical distribution, it would have been useful to treat at
least of the southern portion of them, in conjunction with the Guatemalan species.
Unfortunately but a small proportion of the exact localities are known, so that at
present no line can be drawn to indicate where the preponderance of nearctic types
over tropical ones terminates. Thus, confining myself to the fishes occurring between
the political boundary of Guatemala in the north and the Isthmus of Darien in the
south, I would repeat that, previously to the receipt of the collections forming the basis
to this Memoir, only a small number had been described, as will be seen from the
following remarks :—

§ 4. Historical account of Publications previous to this Memoir.

It would be of but little advantage to enumerate the few isolated species incidentally
described in general works or memoirs as occurring in Guatemala or Panama. How-
ever, I must mention that the first traveller who collected fishes in these states appears
to have been Baron von Friepricustuan. I am not aware that any account of his
travels has been published; but in a paper published by the late Jacop HeckeL in
‘Annalen des Wiener Museums,’ vol. ii. 1840, a single species is described, which is
stated to be from Friedrichsthal’s Central-American Collection, and which I have
recognized as belonging to the Lake-Peten fauna (Heros friedrichsthalii). The greater
part of the collection made by this gentleman evidently remained unpublished until
1864, when Dr. F. Srerspacuner determined from it four other species (Denkschr.
Akad. Wiss. Wien, xxiii.), viz.:—Heros wrophthalmus (Gthr.), Heros triagramma=H.
salvini (Gthr.), Heros melanopogon, and Petenia splendida (Gthr.). As we have received
four of these species from Lake Peten, it is very probable that Baron Friedrichsthal
visited and collected in that locality.

In the second place I have to mention Dr. SzEMANN, who, as naturalist attached to
the expedition of the ‘Herald, brought to England a collection of Central-American
fishes. ‘These, as I have mentioned above, were originally deposited in the collection of
Haslar Hospital, but no record as regards the origin of the specimens was kept, so
that most of them are lost for the purposes of this Memoir.

In the year 1861 I received the first collections from Mr. Satvin and Capt. Dow.
The species belonging to the families treated of in the 3rd volume of the ‘ Catalogue of
Fishes’ were described therein; and a separate account of those sent by the latter

' Prof. Troschel enumerates some 130 freshwater and marine species in Miiller’s ‘ Reisen in den Vereinigten
Staaten,’ &c,

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 385

gentleman from the Pacific Coast of Central America was published in the Society's
‘Proceedings’ for 1861 (Nov. 26); it contained fourteen species, ten of which were new.

In the following year the 4th volume of the ‘Catalogue of Fishes’ was published,
containing the descriptions of those species of Pharyngognaths and Anacanthines which
had arrived from our travellers, who were then engaged in collecting.

In the year 1863 Mr. Gin published a descriptive enumeration of a collection of
“Fishes from the western coast of Central America, presented to the Smithsonian
Institution by Capt. J. M. Dow.” He distinguished in it the following twenty-five
species, of which I consider eighteen to have been new to science (Proc. Ac. Nat. Se.
Philad. 1863, p. 162) :—

Diapterus dowii, sp. n.= Gerres dovit.

Pomacanthodes zonipectus, Gill.

Centropomus armatus, sp. 0D.

Epinephelus analogus, sp. n.=Serranus analogus.
Promicropterus decoratus, sp. n.=Rhypticus decoratus.
Bairdiella armata, sp. n.=Corvina armata.

—

Ophioscion typicus, sp.n.=Corvina ophioscion.
Amblyscion argenteus, sp. X.

Caranx panamensis, Gill,=Caranx speciosus (Forsk.).
10. Carangoides dorsalis, sp. n.

11. Carangus marginatus, Gill,= Caranz hippos, L., var.

SHAAN wD

12. Oligoplites inornatus, sp. 1.=Chorinemus inornatus.

13. Exocetus dowiti, sp. n.

albidactylus, sp. n. t=. bahiensis (Ranz.).

15. Upeneus grandisquamis, sp. 0.

16. Trichidion opercularis, sp. u.=Polynemus opercularis.
approximans = Polynemus approximans (Lay & Benn.).
18. Mugil quentherii, Gill,= WM. brasiliensis (Agass.).

19. Batrachoides pacifici= Batrachus pacifici (Gthr.).
Dormitator microphthalmus, Gill,=Eleotris maculata (B1.).
Leptarius dowii, sp.n.=Arius dovit.

. Sciades troschelii, sp. n.= Arius troschelit.

Allurichthys panamensis, sp. n.

4. Atractosteus tropicus, sp. n.=Lepidosteus tropicus.

. Urotrygon mundus, sp. 2.

Dw pow wo bw ww
a

or

At later periods Mr. Gill has described some other species incidentally, which will be
referred to in the general list.

A small collection made by Prof. M. WacNer on the Isthmus of Panama, between
7° and 9° lat. N., and 77° and 83° long. W., was examined by Messrs, Kner & STEIN-

384 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

DACHNER, who gaye a preliminary account of it in ‘Sitzgsber. bayer. Akad. Wiss.’ 1863,
pp- 220-230, and more detailed descriptions in ‘Abhandl. bayer. Akad. Wiss.’ 1864 (1865),
pp. 1-61. Prof. M. Wagner added, besides, a detailed account of the hydrographical
peculiarities of this part of Central America (pp. 65-92). The species treated of in
these Memoirs are the following :—

. Pristipoma humile, sp.n.

. Dajaus elongatus (K. & St.)=Agonostoma nasutum (Gthr.).
. Dajaus monticola (C. & V.).

. Acara ceruleopunctata, sp. n.

. Heros altifrons, sp.n.

. Heros sieboldii, sp. n.

. Eleotris pictus, sp. n.

. Engraulis macrolepidotus, sp. n.

poeyi, sp. 0.

. Xiphophorus gillii, K. & St.,= Pacilia, sp. ?

. Macrodon brasiliensis, K. & St.,=M. microlepis (Gthr.).
. Saccodon wagneri, sp. n.

. Pseudochalceus lineatus, sp. n.

. Chalcinopsis striatulus, sp. n.

chagrensis, sp. 0.

. Chalceus atrocaudatus, sp. n.
. Letragonopterus eneus (Gthr.).

gronovii (C. & V.%).

. Bagrus ()arioides, sp. n.= Arius multiradiatus (Gthr.).
- Pimelodus modestus (Gthr.)

cinerascens (K. & St.)=P. wagneri (Gthr.).

. Loricaria uracantha, sp. n.

lima (Kner).

. Hypostomus plecostomus (K. & St.)=Plecostomus, sp.
. Ancistrus cirrhosus (C. & V.).
. Acanthias vulgaris (Risso *).

Finally, having received in 1864 the last collections made by Messrs. Godman & Salvin,
1 gave preliminary notices of the new species in the ‘ Proceedings’ of this Society, em-
bodying the-numerous contributions to our knowledge of the Siluroids and Characinoids
in the fifth volume of the ‘Catalogue of Fishes,’ to which were added the Cyprinodontes
and Scombresocides in the sixth (1865-66).

§ 5. General List of Central-American Fishes.

After these introductory remarks on the contributions to the ichthyology of Central

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 385

America preceding this Memoir, I at once proceed to give a list of all the species known
to exist in these countries. There are comparatively few which I do not know from
autopsy; their names are printed in italics. An asterisk (*) marks those which are
described or remarked upon. The second column contains chiefly the names of the
localities where they have been found within the limits of Central America. The
localities of species occurring on both sides of the Isthmus are printed in italics; of
these I shall treat again subsequently. Finally, the letter M signifies that a species is
marine, B that it is known from brackish, and F that it is from fresh. water.

ACANTHOPTERYGII.

Fam. PERCIDA.

Crentroromus, Cuv.

1, *appendiculatus, Poey . . . Chagres R. (Cuba, Mex., Surin.). . . . . . . FL&M.
OF AmediussGiira 1s ©. 4 /o) SeChiapamen rane) os) ee) Oe” ere Bs.
Se tnigrescens) Gin 1-28.» Chiapams eee ee. Ee ee. nee B.
4, *parallelus, Poey. . . . . Chagres R.(W.Indies,Bahia) . . . . + . . F&M.
Be Sarmatusy Gillie + os: 4 ee Ohiapamten git lake a oe ee Teel eB.
6. *ensiferus, Poey . . . . . Belize (Cuba, Jamaica,Guyanas) . ...... B.
Crentropristis, Bris. de Barnev.
7. *macropoma,Gthr. . . . . Panama. M
Serranvs, Cu.
8. *creolus,C.&V.. . . . . Albl. & Pac... M
9: striatus, BL... Ce Atlant: PA ae eR RUGS Vi
TOMcoronatisOng iene, ete eee Atlantis isn ts) me Ss eebhat eer cane civ,
11. undulosus,C.&V. . . . . Atlant. . M
10h) *pellicauday Gils 2 a4 ey) heen Racific Coast s)pee lye s+ 555+ eR PML
Ip opuclbpit, GHG a) a tac, hone Cane pe 6) peep on pe Oe nie oo pelo lUEGE
Prectroroma, Cuv.
TA, tote EYE, cp a sake 8 ty to ZAI CO IATIRE IG ARR apres EOnrOy) “Cel Ich ue yma ibe! -ostcy alll
Ruyericvs, Cuv,
liner SdecoratusnGilee tt 1 shen RacihiciCoastees)-. % 4) en as oe PSs eee
Mesorrion, Cu. a
16, chrysurus, Bl. . . . . . Atlant. . M
Nf Gea, OL ae viel Sot Bysdie! AUR Rey detrios ah hope mite a Oe ties (eecee omen eao wewe )
Ne wuninotatussCn 6) Vaaen meena S Ph OCr item ry 8h tenes oss OME
lO WaratistG//i7 ee ee meee CO iapamupbanamamen ., 0 o) autres cell. dork
20. vivanus,C.&V. . . . . . Atlant. . M
Avocon, Lacép.
DiesdovitnGi i eal eee ae acielOOash | Get. is fe. ees ce ek ee NE

VOL. VI.—PART VII. 3H

386 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

Fam. PRISTIPOMATIDZ.

Pristrpoma, Cuv.

22. *melanopterum,C.§V. . . All. & Pac..
23. *virginicum, Ll. . . . . . Aél.& Pac..
24, *dovil,Gthr. . . . . . . Panama:
25. *chaleeum, Gthr.. . . . . Panama.
26. *humile, Kner & Steind. . . Rio Bayano
27. *macracanthum, Gthr.. . . Chiapam ...-:.-... .
28. crocro,C.§ V. . . . . . Rio Motagua (Trop. Amer., Atlant.)
29. *leuciscus,Gthr.. . . . . San José, Chiapam, Panama .
Conopon, C.& V.
30. *pacifici, Gitar. . . . . . Chiapam
Hamuton, Cuv.
31. chromis, Brouwss.. . . . . Atlant. .
$2) canna, €.&V.. -. = . . Atlant. .
33. xanthopterum,C.§ V. . . . Atlant. . ro Sea
34. *brevirostrum, Gthr. . . . Panama, Puerto Cabello
35. *margaritiferum, Gihr.. . . Panama.
Losotss, Cuv.
36. auctorum, Gthr. . . . . . Atlant. (India)

Fam. SQUAMIPINNES.

Cuztopon, Cuv.

S7-astmatous,.2.. 5 © - «|. - wAtlant. eo is
38) capistratus, 2. 5. 2) » Atlant. ~ ee 2 Uf o9
39. *humeralis,Gthr. . . . . Panama (Sandwich Isl.)
Pomacantuvs, Lacép.
40. paru,Gthr.. . . . . . . Atlant. (Colon)
41. *zonipectus,Gill . . . . . Pace. .
Erurerus, Cuv.
42. faber, Bl. . . . . . . . Atlant. (Belize)

i Fam. MULLID.
Urrnevs, C.& V.

_ 48. *tetraspilus,Gthr. . . . . Panama.
44. *grandisquamis, Gill . . . Panama.

Fam. SPARIDZ.

Sarcus, Cuv.

45. unimaculatus, Bl. . . . . Atlant. (Belize)

46. avies,C.&V. . . . . . . Atlant. (Belize)
Curysopurys, Cuv.

47. *calamus,C.§V.. . . . . Atl. & Pac. (Panama)

PIMELEPTERUS, Cuv.
48. boscii, Lacép. . . . . . . Atl. & Pac. (Chiapam & Panama) .

as

Ss

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

Fam. CIRRHITIDZ.

Crrruiricutuys, Blkr.
49, *rivulatus, Val. . . . . . Galapagos Islands, Panama

Fam. SCORPAANID/.

Scorpana, Art.
50. plumieri, Bl., Schn. . Atl. & Pac. (Panama)

Fam. POLYNEMID.

Potynemus, L.

51. *melanopoma, Gthr.. . . . San José aah
52. *approximans, Lay & Benn. . Pacif., Chiapam, Panama
53. *opercularis, Gill. . . . . Paeif.

Fam. SCLENID.
Larimus, C. & V.
54. *breviceps, C. & V,
Microrocon, Cw. & Val.
55. undulatus, Z... :. . x . - Atlant. .
56. *altipinnis, Gthr. .
Umsrina, Cuv.

Atl. & Pac. (Panama)

57. *elongata,Gthr. . . . . . Chiapam

58. *nasus,Gthr. . . . . . . Panama.

59. *analis,Gthr.. . . . . - Panama.
Corvina, Cw.

60. ronchus,C.&V. . . . . . Atlant. .

61. *chrysoleuca, Gthr. . . . . Panama.

62. *vermicularis, Gihr.. . . . Panama.

63. *armata, Gill. . . . . « Paeif.

64. *ophioscion,Gihr. . . . . Panama.
OrouitHvs, Cuv.

65. *squamipinnis,Gthr. . . . Panama. .

66. *albus,Gthr. . . . . . . Chiapam ... .

67. *reticulatus,Gthr. . . . . San José, Chiapam .

Fam. ACRONURID.

Acantuurvs, Schn.
68. chirurgus, Bl.. . . . . . Atlant. .

Fam. CARANGIDZ.

Caranx, Gthr.

69. crumenophthalmus, B/.. . . Atl. & Pac. .

70. amblyrhynchus,C.&V.. . . Atlant. .

71. *leucurus,Gthr. . . . . . Panama. ..... .
72. *speciosus, Forsk. . . . . From Panama to East Africa .

Chiapam, San José, Panama .

oD”

bo -

B.
&

387

S55

5 SS

wes Ss BS

&

S555

388 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

73. carangus, Bl. .

74. *hippos, L. .

75. *caballus, Gthr.

76. *caninus, Gthr.

77. *dorsalis, Gill .
Arcyriosus, Lac.

78. vomer, L. :

79. setipinnis, Mitch.
Cuorinemvs, C. & V.

80. occidentalis, L.

81. saliens, BZ. .

82. *altus, Gthr.

83. *inornatus, Gill
Tracuynotvs, C. & V.

84. ovatus, L. .

85. *fasciatus, Gill

Petamys, C.& V.

86. *sarda, Bi. .
Cysium, Cuv.

87. maculatum, Mitch.
Ecuenets, Art.

88. remora, L. .

89. naucrates, L.

Barracuus, Gthr.
90. *pacifici, Gthr.

91. surinamensis, Bl. Schn. .

*THALASSOPHRYNE, Gthr.
92. *maculosa, Gthr. .
93. *reticulata, Gthr. .

Poricutuys, Girard.

94, porosissimus, C. & V.

ANTENNARIUS, Commers.
95. *leopardinus, Gthr. .
96. *tenuifilis, Gthr. .

Gostus, Art.
97. soporator, C. & V..
98. paradoxus, Gthr. .

Atlant. & Ind. Occ, (Chiapam & Belize) .
Tropics generally . nares
Panama . :

Panama . ee

San Diego (Cal.), Panama .

ee
&

Atl. & Pac. (Belize, Chiapam, Panama)
Atl. & Pac. (Panama)

Atlant. . OE ee Le
Atlant. & Pac. (Chiapam, Isabel) .
Panama . De Sat OE
Panama .

Atl., Pac., & Ind. Oc. (Panama) .
Panama, San José

ee: bs
ee gael

Fam. SCOMBRID&.

AEG POE PRD. ne 5 ye, ee ES AO
Atlant: (Belize) 2s ty. 1 OS eS Me

Atl., Pac., & Ind. Oc.
Atl., Pac., & Ind. Oc.

Fam. BATRACHID.

Panama, West Coast of Africa
Atl. & Pacif. (Panama) .

Puerto Cabello
Panama.

See eee

All. & Pac. .

Fam. PEDICULATI.

Panama .

ss

Panama.

Fam. GOBIID/.

Ags bac. (panama) o. . s+ «ta aegis Bs
Panama meee or OY Pos ba cinachee MS

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 389

99. mexicanus, Gthr.

100. seminudus, Gthr.
Eucrenocosivs, Gill.

101. sagittula, Gthr. .
Srtcyp1um, C. & V.

102. plumieri, B/. .
Exeorris, Cuv.

103. *maculata, Bl.

104. somnolenta, Girard

105. dormitatrix, Bl. .

106. *longiceps, Gthr.

107. *picta, Kner .

108. *seminuda, Gthr.
Ambtyopvs, C. & V.

109. *brevis, Gthr.

Buirnnivs, Artedi.

110. brevipinnis, Gthr.
Sararias, Cuv.

1li. atlanticus, C.& V. ;
Cuinus, Gthr.

112. nuchipinnis, Q. & G.

113. delalandu, C. & V.

114. *macrocephalus, Gthr. .

Cremnosartes, Gthr.

115. *monophthalmus, Gthr.

Spnyrana, Artedi.
116. picuda, Bl.
117. forsteri, C.§ V. .

ATHERINICHTHYS, Gthr.
118. *pachylepis, Gthr. .

119. *guatemalensis, Gthr. .

Muett, Artedi.
120. *brasiliensis, Agass.
121. *incilis, Hancock
122. proboscideus, Gthr.
Aconostoma, Benn.
123. *microps, Gthr. .

Mexico, Rio Motagua
Panama .

Panama .
Atl. & Pac. (Panama)
Ail. & Pac. (Huamuchal)
All. & Pac. (Cardon)
Atl. (Rio Motagua, Yzabal)
Lake of Nicaragua
Rio Bayano
Panama .
Panama .

Fam. BLENNIID.
Pacif.
All. & Pac. .
Atl. & Pac. .
All. & Pac. .

Panama .

Panama .

Fam. SPHYRANIDZ.

Ate Rave ramrur eee oe.
Ind. Oc. & Pac. (Chiapam)

Fam. ATHERINIDZ.

Panama .
Huamuchal

Fam. MUGILID.
Atl. & Pac. .
Atl. (Chagres) .
Atl. & Pac. (Cardon) .

Rio Guacalate .

.M.&B.

ws

.M.&F.
.M.&F.

F.

390 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

124, *nasutum, Gites s. 2)

125. *monticola, Bancroft .
Myxvs, Gthr.
126. harengus, Gthr. .

Rivers of both sides of C. America (R. San Geronimo,

R. Motagua, Panama)

W. Indies and rivers of both sides of C. America .

Panama .

Fam, FISTULARIID.

Fistunaria, Lacép.
127. tabaccaria, L.

All. & Pac. .

Fam. GOBIESOCID.

Sicyases, Miill. & Trosch.
128. fasciatus, Ptrs. .
Gostesox, Lacép.
129. *rhodospilus, Gthr. .
130. nigripinnis, Ptrs.
131. nudus, BI.

Puerto Cabello

Panama .
Puerto Cabello
Atl. & Pac. (Cardon)

PHARYNGOGNATHI ACANTHOPTERYGII.

Fam
Pomacrntrvs, C. & V.

132. *rectifrenum, Gill .

133. leucostictus, Miill. & Trosch. .
GuypHipopon, Cuv.

134. saxatilis, Z.

135. concolor, Gill

136. declivifrons, Gill
Hetiastes, C. & V.

137. *marginatus, Casteln. .

Lacunotamvs, C. & V.
138. falcatus, LZ.
Cossyruus, Giinth.
139. rufus, L. :
140. diplotznia, Gill .
141. *pectoralis, Gall .
Puatyciossus, Gthr.
142. bivittatus, Bl.
143. *dispilus, Gthr. .
Psruposuttis, Blkr.
144. *notospilus, Gthr.

. POMACENTRID.

Pacif. & All.
Atl.

JA i eh tt eRe

All. & Pac. (Cardon)

Pac. (Cardon) .

All. & Pac. .

Fam. LABRID.

Atl.

Atl. State 5k le eg

Panama, Lower Calif. . Ae ae ee
Panama, Lower Calif., St. Helena (? Cuba)

Atl.
Panama .

Panama .

M.

shall tl

SSS FS

= 5S SES

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

Juyis, Gthr.

145. lucasana, Gill
Scarus, Blkr.

146. abildgaardii, Bi. .
Psreuposcarvs, Blkr.

147. sancte crucis, Bi.

148. guacamaia, C.& V. .

Gourres, Cuv.
149. plumieri, C. § V.
150. *axillaris, Gthr. .
151. *brevimanus, Gthr..
152. rhombeus, C.§ V. .
153. squamipinnis, Gthr.
154. aprion, C. & V.
155. *dovii, Gill

Acar, Gthr.

Panama, Lower Calif. .

Atl.

Atl.
Atl.

Fam. GERRID.

Atl

Chiapam

Chiapam Dig PEE

Atl. & Pac. (Chiapam) . Af ss ED
(Jamaica) Atl. & Pac. (Chiap. & Panama) .
Atl. & Pac. (Panama)

Pac. -:

Fam. CHROMIDES.

156. *ceruleopunctata, Kner & Steind. Chagres River

Heros, Gthr.
157. *parma, Gthr. .
158. *margaritifer, Gthr.

159. *melanopogon, Steindachner
59 Ip

160. *melanurus, Gthr.
161. *macracanthus, Gthr.
162. *spilurus, Gthr.

163. *nigrofasciatus, Gthr.
164. *multispinosus, Gthr.
165. *longimanus, Gthr.
166. *urophthalmus, Gthr.
167. *aureus, Gthr. .

168. *affinis, Gthr.

169. *labiatus, Gthr.

170. *erythreus, Gthr. .
171. *lobochilus, Gthr. .
172. *citrinellus, Gthr. .

173. *altifrons, Kner & Steind. .

174. *friedrichsthalii, Heck. .
175. *salvini, Gthr. .

176. *trimaculatus, Gthr. .
177. *dovu, Gthr.

178. *motaguensis, Gthr. .

Mexico, R. Chagres & R. Motagua

Lake Peten .

? Lake Peten

Lake Peten . :
Chiapam & aarti :

Rio Motagua, Yzabal . :
Lakes of Amatitlan & Atitlan :
Lake of Managua .

Lake of Nicaragua .

Lake Peten .

Yzabal, Rio Miotena

Lake Peten .

Lakes of Managua & Nicaea
Lake of Managua .

Lake of Managua .

Lake of Nicaragua .

Western Veragua .

Lake Peten .

Santa Izabel, Lake Bate
Chiapam, Huamuchal .

Lake of Nicaragua .

Rio Motagua

F., B., & M.

392 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

179. *managuensis,Gthr.. . . . . Lake of Managua . F.
180. *microphthalmus, Gthr.. . . . Rio Motagua F.
181. *oblongus,Gthr. . . . . . . Rio Motagua F.
182. *nicaraguensis, Gthr.. . . . . Lake of Nicaragua. F.
183. *godmanni,Gidr.. . . . . . River ofCahabon . F.
184. *sieboldii, Kner & Stemdachner . New Granada F.
185. *guttulatus, Gthr.. . . . . . Lake of Amatitlan. : F.
186. *irregularis, Gthr. . . . . . Rio Usumacinta, 8. Geronimo . F.
187. *intermedius,Gthr. . . . . . Lake Peten . F.
188. *angulifer,Gthr. . . . . . . Yaabal, F,
*Perenia, Gthr.
189. *splendida, Gthr. . . . . . . lake Peten . F.
*Neetropivs, Gthr.
190)-enematopus, Gikr-. .. < «. ey, Wakeiof Managua... 1. 288.) Slow fe (SEE:
ANACANTHINI.
Fam. LYCODIDZ.
*Micropesmvus, Gthr.
UGieetdipusGihn 2° = |4 < ), SeeBanamals (bain 2d. eine Soa
Fam. OPHIDIID.
Brortuta, Cuv.
192:\*Pmultibarbatasschleg. ~ <9’ MPac.’coast: s = >see? ieee _ opaMe
Drvematicutuys, Blkr.
LO3eamareinatus, Ayres’. sees * Panamal>. 2 3: os . .oRRmemeqer sees
Oruipium, Cuv.
L94tebrevibarbesCuv; 0" 0 PMA seUP ac: 4s 2 2... We eee eee
Fam. PLEURONECTID.
Crrnaricutuys, Blkr.
195; *spilopterus, G7ir- " : © sa eA éPac.(Chiapam)- ; = = 72 eee MGB:
196. *guatemalensis, Blkr. . . . . Guatemala
Hemiruomeus, Blkr.
197. *ovaliss Gthrs go Bown. 2 ot BRR Rach Oeek en) Cae 5 eee AEM:
Psruporuomevs, Blkr,
198... *brasiliensis: Ranzant +: SRP RARISS PS Se ae ie. -2
Soxea, Gthr.
1995 seutamiGihr. 4 2 = = ls) Sebanama:

Apnoristia, Kaup.
200! “omatayiactp? to 2» APM mel RON ECAC ont: WU; neh. SRR gts reese emnton ene VIR

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. ~

PHYSOSTOMI.
Fam. SILURID.

Amiurus, Rafin.

201. *meridionalis, Gthr. .
Pruetopvs, Gthr.

202. modestus, Gthr.

203. guatemalensis, Gthr. .

204. godmanni, Gthr.

205. *wagneri, Gthr.

206. *managuensis, Gthr. .

207. micropterus, Gthr.

208. nicaraguensis, Gthr. .

209. petenensis, Gthr. .

210. motaguensis, Gthr.

211. salvini, Gthr.

212. polycaulus, Gtir. .
Artus, Gthr.

213. guatemalensis, Gthr. .

214. *assimilis, Gthr.

215. platypogon, Gthr. .

216. seemanni, Gthr.

217. cerulescens, Gthr.

218. troschelii, Gill .

219. *dovii, Gill .

220. melanopus, Gthr. .

221. multiradiatus, Gthr. .
iiuricutuys, Baird & Gir.

222. *nuchalis, Gthr.

223. *panamensis, Gill .
Piecostomus, Gthr.

224. *?sp., Kner & Steindachner
Cuztostomus, Heck,

225. *aspidolepis, Gthr.

226. *?cirrhosus, Val.
Loricarta, Lacép.

227. *uracantha, Kner & Steindachner .

228. lima, Kner .

Rio Usumacinta

Rio Chagres, Esmeraldas
Huamuchal

Lower Vera Paz, Rio Motagua, Mexico .

Pacific & Atlantic rivers of Panama .
Lake of Managua .

Rio San Geronimo .

Lake of Nicaragua .

Lake Peten .

Rio Motagua

Rio San Geronimo .

Rio San Geronimo.

Chiapam, Panama .
Lake of Yzabal .

San José .
9

Huamuchal .
Pac.

Pac. «,e }:
Rio Motagua
Rio Bayano .

Panama
Panama

Rio Chagres .

Veragua
Rio Chagres .

Atlantic & Pacific rivers of Panama .
Atlantic & Pacific rivers of Panama .

Fam. CHARACINID.

Macrovon, Mill. & Trosch.
229. *microlepis, Gthr..
TETRAGONOPTERUS, Cuv.

230. fasciatus, Cuv. .

VOL. VJ.—PART VII.

W. Ecuador, Rio Chagres

From Brazil to Mexico (Huamuchal, Rio Guacalate,

Rio Motagua, Rio Chisoy)

393.

ta

394 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

231. microphthalmus, Gthr. . . . . Lakeof Amatitlan, Pacif.Coast of Guatemala, Peru.
232. panamensis,Gihr.. . . . . . Panama, Yzabal
233. brevimanus, Gthr.. . . . .-.- Rio S. Geronimo, Yzabal.
234. petenensis, Gthr.. . . . . . Lake Peten,W. Ecuador .
235. humilis, Gitar. . . . . . . . Lake of Amatitlan . Us che ep epee
236. *eneus,Gthr. . . . . . . . Mexico, Pacific & Atlantic rivers of Panama
Cuatcinopsis, Kner. ;
237. *dentex,Gthr.. . . . . . . Rio Motagua and Usumacinta, Yzabal; Ecuador .
238. striatulus, Kner . . . . . . Pacific & Atlantic rivers of Panama .
239. chagrensis, Kner . . . . . . Rio Chagres .
Anacyrtvs, Gthr.
240. *guatemalensis, Gihr. . . . . RioChagres, Huamuchal .

Fam. SCOPELID.
Saurus, C. & V.

PAN foetenssalas se fee ino aopreerAll. 6 Puc:
242. myops, Bl. ... . . . . . . Atl. & Pac.

Fam. SCOMBRESOCID.

HemrruamrPuves, Cuv.

243. unifasciatus, Ranzani . . . . Atlantic, Pacific, & Indian Oceans
Exocetus, Artedi.
244. *callopterus,Gthr. . . . . . Pace.
245. albidactylus, Gill Daas agate:
Ranz.). : Smiths &
odbc dovir Gillis a my. - st eon. 0. Pace

Fam. CYPRINODONTID.

*CHARACODON, Gthr.

247, *lateralis,Gthr. . . . . . . ——?
Haptocuitus, M‘Clell.
248. *dovii,Gthr. . . . . . . . Punta Arenas (Costa Rica) .
Funpuuvs, C. & V.
249. *labialis, Gthr.. . . . . . . Rio S. Geronimo, Yzabal .
250. *punctatus,Gthr.. . . . . . Chiapam .

251. *guatemalensis, Gthr. =i Ecuailor

252. *pachycephalus, Gthr. . . . . Lake of Atitlan.
Betonesox, Kner.

253. belizanus, Kner . . . . . . Lake Peten, Honduras, Mexico
Gampusia, Poey.

254. *nicaraguensis, Gthr. . . . . Lake of Nicaragua.

ANABLEPS, Artedi.
255. doviil, Gill . . . . . . . . Chiapam .

Jf Lakes of Duefias & Amatitlan, Rio Guacalate, W.

badd dat bat

F.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

Paeciiia, Gthr.

- 256, mexicana, Steindachner . ane .. F&B.
257. thermalis, Steindachner . San Salvador, Mexico F.
. 258. chisoyensis, Gthr. . Rio Chisoy F,
259. *elongata, Gthr. Panama .
260. *petenensis, Gthr. . Lake Peten . MLL ithe
261. dovii, Gthr. . Lakes of Nicaragua & iensihan Mexioa A ltrs al ac
262. *gillii, Kner Rio Chagres . F,
263. spilurus, Gthr. . =o
Mo.uiengsia, Lesueur.
264. *petenensis, Gthr. Lake Peten . F.
Xipnoruorvs, Gthr.
265. *hellerii, Heck. . Rio Chisoy, Mexico F.
Girarpinus, Poey.
266. *pleurospilus, Gthr. . Lake of Duefias F,
Fam. CYPRINID.
Scierocnaruus, Gthr.
267. meridionalis, Gthr. Rio Usumacinta F.
Fam. CLUPEID.
Cuanos, Lacép.
268. salmoneus, Forst. . Indian & Pacific Oceans (Chiapam) M.&B.
Atsuta, Gronov.
269. conorhynchus, Bi. Tropical & Subtropical seas (Panama) M.
Mueators, Lacép. .
270. thrissoides, Schn. . Atlantic . M.
PrisTiGasterR, Cuv.
271. *macrops, Gthr. Panama M.
272. *dovii, Gthr. Panama M.
Cuiurea, Artedi.
273. *libertatis, Gthr. . Libertad . M.
Cuatozssus, C. & V.
274. *petenensis, Gihr. Lake Peten . F.
Enerautis, C. & V.
275. brownii, C. & V. é Atlantic & Pacific (Libertad) M. &B.
276. *poeyi, Kner & Biciudeonmes Rio Bayano . aes F.
277. *macrolepidota, Kner & Steind. . Rio Bayano . F.
CrrencravLis, Gthr.
278. *mysticetus, Gir. Pacific coast of Panama . M.
Fam. GYMNOTID®.
Cararus, Mill. & Trosch.
279. *fasciatus, Pall. Rio Motagua F.

ps Duefias, Rio Chisoy, Huamuchal, Lake of

395

12

oo

396 Dk. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

Ornivurvs, Lac.
280. triserialis, Kaup.
281. boro, Ham. Buch. .

282. breviceps, Richards. .

Morana, Cuv.

283. lineopinnis, Richards.

SympBrancuvs, Bl.
284. marmoratus, Bl.

285. immaculatus, Bi. .

Diovon, Kaup.

286. sex-maculatus, Cuv.
Trtropon, L.

287. *politus, Girard

288. *geometricus, Gthr. .

Osrracton, L.

289. cornutus, L.

290. bicaudalis, L.
Bautstes, Hollard.

291. vetula, L. ;

292. *frenatus, Lacép. .

293. niger, Osbeck
ALEUTERES, Cuv.

294. monoceros, Osbeck

LEPIDOSTEUS
295. *tropicus, Gill .

Muste.uvs, Bonap.

296. *dorsalis, Gill .
ACANTHIAS

297. vulgaris, Risso .
CaRrCHARIAS

298. *maculipinnis, Poey .

ZycH#na, Cuv.
299. tiburo, L

Rutnosatus, Miill. & Hede
300. *leucorhynchus, Gthr.

Fam. MURAINIDZ.

Atlantic & Pacific . ;
Indian Ocean, West Indies .
Pacific coast. .

Atlantic & Pacific (Panama)

Fam. SYMBRANCHID.

M., B., & F.

Atlantic (Rio Chisoy, Huamuchal, Lake Peten),

Pacific Coast of Guatemala .

PLECTOGNATHI.

Indian & Pacific Oceans (Panama)

San José .
Panama & Galapasos fale

Tropics
Atlantic .

Tropics

Indian & Pacific teats iGangaies Isl. Ws

Ind., Pac., & Atlant. Oceans

Ind., Pac., & Atlant. Oceans

GANOIDEI.

Huamuchal .

ELASMOBRANCHII.

Panama

Atl., Ind., & Pac. Oceans (Panama)
Cuba, Chiapam .

Atl.

Panama

M., B., & F.
M.

S S85

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 397

Pristis, Lath.

301. antiquorum, Lath. . . . . . Atl. & Pac. Oceans (Chiapam). . . ... =. Mz
Urotoraus, Miill. § Henle.

BO cavunaus Grillas epee heer sui 25) Pac HW ee ee a ee ee eM
Aftozatis, Mill. & Henle.

303. *latirostris, A. Dum... . . . Gaboon,Panama ........... Mz

§ 6. Partial Identity of the Fish-faunas of the Atlantic and Pacific Coasts of
Central America.

It will be seen that, as far as our present knowledge reaches, of these 303 species,
173 are truly marine forms, 57 being found on both sides of the Isthmus.
25 have been found in brackish water, of which 3 are found on both sides of the
Isthmus.
101 are freshwater fishes, 17 being found in rivers of the Atlantic and Pacific sides.

There will be but very few species which are entirely limited to brackish water, and
which may not be with equal propriety added either to the marine or freshwater fauna.
Thus, five of the 25 species hitherto known from lagoons with brackish water belong
to freshwater genera ; and, admitting two groups only, we have

193 marine fish, 59 of which are found on both sides of Central America=304

per cent.

106 freshwater fish, 19 being found in rivers of the Atlantic and Pacific sides=18

per cent.

From the circumstance that our collectors paid more attention to the freshwater
than to the marine fauna (at least of the Atlantic coast), we may assume that the pro-
portion between the two groups will be increased by future researches in favour of the
marine fauna, but that the proportion between species peculiar to one side and those
common to both will be lessened, inasmuch as every collector will discover other
Atlantic forms on the Pacific side, and vice versd.

The very curious fact of the partial identity of the species of both coasts of Central
America was first distinctly stated by myself in the Society’s ‘ Proceedings’ for 1861
(p. 370), when, out of fourteen species collected by Capt. Dow on the Pacific side, five
were found to be Atlantic forms. To these various others were added by me in the
‘Catalogue of Fishes;’ and Mr. Gill confirmed this observation in Proc. Ac. Nat. Sc.
Philad. 1862, pp. 140, 249. Professor Wagner, in his memoir quoted above (p. 384)},
has made the same observation; but the species enumerated by him, fourteen in number,
are, with one exception, freshwater forms, the geographical distribution of which must
have been brought about at periods and in ways different from those of the diffusion
of marine species.

Knowing now that at least 30 per cent. of the marine fish are found on both sides of

‘ See also ‘ Record, Zool. Literat.’ ii. p. 177.

398 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

Central America', we cannot account for this fact by resorting to such occasional means
of dispersal as the accidental transmission of spawn from one shore to the other by birds
or water-spouts, or even the close proximity of the sources of rivers flowing in opposite
directions. If we do not adopt the view that species were created at the spot where we
find them now, similar creations being produced under similar physical conditions, we
have but one way of explaining the partial similarity of these marine fish-faunas,
namely, by assuming that the Isthmus did not form a continuous barrier between the
two oceans at a former period, but that one or more open channels existed. I am not
aware that geology has, up to this time, furnished us with proof positive that this is
really the fact; but considering the volcanic nature of Central America, and the absence
of all fossiliferous strata, it does not appear too bold an hypothesis to assume that North
and South America were formerly connected by a chain of islands similar to that of the
Antilles, and that subsequently an elevation (as in other parts of the globe) took place,
resulting in the final continuity of dry land: the long-continued activity of the
numerous volcanoes may have been another, though secondary cause in filling up the
channels on the Pacific side. If such a bodily elevation of Central America has taken
place, it is easy to show where some of the broadest channels existed, namely, where
we find the greatest depressions running from one ocean to the other. The northern-
most of these depressions exists between Tehuantepec and the river Coatzalco ; the
second is indicated between Puerto Cabello and the Gulf of Fonseca; the third by the
Lake of Nicaragua (the remnant and deepest part of a very broad channel) ; a fourth
between Chagres and Panama. (See map, Pl. LXIIL, where these supposed former
depressions are coloured green.) As far as I have been able to ascertain, the greatest
elevation of the first of these lines of depression would be 1500, of the fourth 287 feet
only. If we presume that only one of the channels was open at a period when the
present marine fauna was already in existence, it will fully explain the existence of
identical species on both sides of the isthmus, especially if the difference of the tides
was as great as it is now’, causing strong currents from one ocean to the other.

Such an instance of a disconnexion of a marine fauna by elevation of land as I am
inclined to assume in the case of Central America does not stand quite alone. We owe
to the researches of Prof. 8. Lovén and Dr. Malmgren‘ the knowledge of the fact that
marine animals (Crustacea, Annelids, and Fishes) inhabiting the glacial ocean are found
in the great freshwater lakes of Sweden and in the Bothnian Gulf, and that this is to
be explained only by the former continuity of the Baltic with the Glacial Ocean.
During the second half of the glacial period the greater part of Finland and of the

’ Mr. Darwin (‘ Origin of Species,’ 3rd edit. p- 378) was not acquainted with this fact, which by no means mili-
tates against his argument, but merely modifies it. 2 M. Wagner, J. c. p. 87.

3 At Chagres the mean elevation is 1:16 foot, while at Panama the highest flow is 22 feet. (Seemann, Voy.
of H.M.S. ‘ Herald,’ i. p. 236.)

« Lovén, Skand. Naturforsk.-Siillskap. férst. offentl. mote d. 9 Juli 1863: Stockholm, 1864, Malmgren,
‘ Kritisk Ofversigt af Finlands Fiskfauna,’ see ‘ Zool. Record,’ i. pp. 136-188.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA, 399

middle of Sweden was submerged, and the Baltic was a great gulf of the Glacial Ocean,
and not connected with the German Ocean. By the gradual elevation of the Scandi-
navian continent, the Baltic became disconnected from the Glacial Ocean, and the great
lakes separated from the Baltic.

The Isthmus of Suez appears to have been a much more permanent barrier between
the faunas of the Mediterranean and the Red Sea. R. A. Philippi has drawn up a list
of species of shells common to both faunas; but it was founded on a collection made by
Ehrenberg, in which the shells from both seas had been mixed’; and P. Fischer’ has
lately shown that the two faunas.are quite distinct. As regards the fishes, I have men-
tioned (on former occasions) a few occurring in both seas (Sargus noct, Sargus rondeletit) ;
but the number is so small that one might be tempted to account for it by the tempo-
rary existence of an artificial communication between the two seas.

Looking at the results of the separation of the Baltic from the Glacial Ocean on the
one hand, and of that of the Pacific from the Atlantic on the other, we find them very
different. As soon as the continuity of the Baltic with the Glacial Ocean was inter-
rupted, the amount of fresh water carried into the former by rivers exceeded the quantity
lost by evaporation of its surface, and the salt water gradually changed into brackish,
and in the northern parts into fresh water. By far the greater part of the animals
became extinct; but a few survived’, however, in spite of the greatly altered physical con-
ditions, without altering their specific characters, still agreeing with the typical forms in
every point, except in size, remaining smaller, leaner, almost starved. The same thing
might happen if by a rising of the chain of the West-Indian islands the Gulf of Mexico
or the Caribbean Sea were at a future time converted into inland seas with narrow out-
lets into the open ocean.

The separation of the Atlantic and Pacific Oceans was, of course, not accompanied by
a change of the water; and any difference that existed in the physical conditions of both
seas, as, for instance, the formation of corals on the Atlantic side, and their total absence
on the Pacific, existed already before the communication between the oceans was closed ;
so that the life of species was not in any way affected by the discontinuance of this
communication. Let us for argument’s sake assume that the part of the isthmus
between the Lake of Nicaragua and Panama was once an island, @ pew prés of the form
of Cuba, inhabited, like Cuba, on its northern and southern coasts by a certain species of
fish. The only effect of a gradual rise of the land on the life of this species would be to force
it to retreat further and further from the original coast, and to accommodate itself to the
new one—an effect to which, if felt at all, the individuals on the northern and southern
coasts would be equally exposed. Thus there is in this case no apparent external cause
for an alteration of the species; and, indeed, the specimens examined by me from opposite
coasts of the isthmus are absolutely identical, and there is not the slightest indication that
one of them has been modified or degenerated into a climatic or local variety. I trust that

1 Martens, in ‘ Zoolog. Record,’ ii. p. 237. > Journ. Conehyl. xii, 1865, pp. 241-248,

3 Seven or eight species of the northern part of the Baltic are believed to be of Arctic origin.

400 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

geology will furnish us with the proof of the former partial submergence of a part of
Central America, as it has done with respect to the northern part of Scandinavia. We
should then be able to speak with more confidence of the permanence, or rather endu-
rance, of the characters of a specific type, and arrive at. a somewhat more definite idea
of the age of species which must have existed before those geological changes were
completed’.

Sir CHAr.Es LYE. has directed my attention to collateral evidence from other classes
of the animal kingdom, by which the partial identity of the faunas of the two coasts is
shown, although not in an equally conclusive manner. The majority of malacologists
appear to have presumed @ priori their distinctness, and consequently described Pacific
shells generally as distinct from Atlantic species. However, Dr. Mércu, in a paper in
which he describes or enumerates about 36() Panama species, makes the following
remarks (Pfeiff. Malakozool. Blatt. 1859, p. 107) :—

“The tropical [molluscan] faunee may be classed in two principal divisions, the Indian
and the Atlantic. ‘To the latter belong, 1, the Guinean (Senegalian); 2, the Antillian ;
and 38, the Panaman, which, although belonging to the Pacific, appears to be most
analogous to the Guinean. A great number of species, especially of Bivalves, have
been regarded as identical with those from the eastern (Brazilian) shore. I believe I
can prove that they are different. Certain irregular mollusks cannot be separated
diagnostically; but I can recognize them by their general habit. It is at all events a
fact that no species stamped with definite characters (wohlausgepriigt) is identical on
both sides of the isthmus. The Panama species may be divided into:—1l, those
analogous to West-Indian ; 2, those analogous to species from Guinea and Senegal ;
3, those very remotely analogous to East-Indian species.”

* I may on this occasion recur to a remark made by me in Proc. Zool. Soc. 1858, p. 381, with regard to the
sea-snakes observed in the Bay of Panama by M. Sallé, Capt. Dow, and Mr. Salvin. There is now not the least
doubt that the snakes seen were Pelamys bicolor, and that they are, moreover, very common there. I find that
Dr. Seemann (Voy. ‘ Herald,’ i. p. 265) already mentions them. But I am much inelined to think that this
most common Indian species has migrated eastwards, and that its arrival on the West-American coast is of
very recent date. Dampier and the other bucaniers who have left us records of their adventures, and who
passed weeks and months in the Bay of Panama, could not have failed to observe them, and to mention them
in their notes, just as they did on other occasions. It is also probable that these snakes would have spread
into the Atlantic Ocean, had they been so numerous on the Pacific side at the time when a communication
existed between the two oceans.

Whilst this paper was passing through the press, I found two notices of the existence of water-snakes
on the western coasts of South America, in seas considerably more southwards than the Bay of Panama. The
notes are in Capt. Sharp’s Voyage in “‘ The History of the Bucaniers of America.” London, 1699, 8yvo, vol. ii.
p. 50: “As we sailed” [near Cape St. Francisco, which is nearly under the equator] “we saw multitudes of
Grampusses every day; as also Water-snakes of divers colours.” And p. 72, when sailing in lat. 19° S., the
author mentions “ A huge shoal of fish, two or three Water-snakes, and several Seals.’ I find in another part
of the same work a note which I believe to be the first description of Zapirus bairdi. The part has a separate
title-page, ‘A Journal of a Voyage made into the South Sea by the Bucaniers or Freebooters of America from

the year 1684 to 1689. Written by the Sieur Raveneau de Lussan.” Lond. 1698, 8yo. The Indian name of
the Tapir is given as Manipourye, page 16.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 401

These remarks appear to me to convey very strong testimony in accordance with my
own observation on the ichthyological fauna, inasmuch as the author refers the Panama
Mollusks generally to the Atlantic fauna. He, indeed, denies the perfect identity of
the species, admitting merely an ‘“‘ analogy” between them; but then it is a question
whether malacologists do not go too far in making specific distinctions, when they are
not even able to express those distinctions ‘“ diagnostically,” recognizing the forms
merely “by their general habit.” Shells are, after all, that portion of a mollusk the
formation and development of which is most influenced by the peculiarities (physical
and chemical) of the surrounding medium and locality; and only too many specific
forms have been distinguished on account of slight differences in the sculpture and
shape of the shells, the importance of which disappears on comparing a large series of
examples. However, as I am not prepared to form an opinion with regard to the shells
of Central America from my own examination, I am bound to receive the testimony of
so celebrated a malacologist as Dr. Morch; and should his observations prove to be
fully correct, they will give an additional interest to this fauna, as proving that the
shells of Mollusks suffer change under circumstances in which the specific characters
of fishes remain unaltered.

With regard to fossil shells, Mr. J. C. Moorn, who has examined several. collections
from tertiary beds in San Domingo, has made the observation that “‘ many bear a strong
resemblance to shells now living in the Indian Seas and the Pacific, and that one or two
appear to be identical” (Quart. Journ. Geol. Soc. 1853, p. 131), and “ that a channel
or sound may have existed in the equatorial parts during some portion of the tertiary
period, by which some few of the tropical shells may have migrated from the one ocean
to the other” (ibid. 1850, p. 43).

Of the other marine animals, the Corals have been made the object of elaborate
researches, the various authors arriving at somewhat different conclusions. First,
Mr. Duncan, in a paper “On the Fossil Corals of the West-Indian Islands” (Quart.
Journ. Geol. Soc. xix. 1865, p. 455), has shown that “in all the calcareous formations
which are coralliferous, and are considerably elevated above the level of the Caribbean
Sea [being probably of miocene age], there is a very limited series of Corals with
generic relation to those now existing and characteristic of the West-Indian Coral
Fauna, but a predominance of forms resembling those of the present Coral-seas of the
Pacific, South Sea, and the Indian Ocean.” This identity of the Corals proves an
identical condition of the physical circumstances, and evidently a wide continuity of
the West-Indian and Western seas.

On the other hand, Prof. Verritt, when speaking of the living Polyp-faune of the
Atlantic and Pacific sides of Central America (Proc. Bost. Soc. Nat. Hist. x. 1866,
p- 323 et seq.), states that their differences of character are very remarkable; that at
Panama none of the reef-building corals of Aspinwall, Florida, or the West Indies
occur, nor even any of the genera of the families to which they belong, with the

VOL. VI.—PART VII. 3K

402 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

exception of a small Porites and Stephanocora ; that these and other differences do not
favour the theory entertained by some geologists, viz. that there has been a communi-
cation between the two oceans at this point, and that the Gulf-stream flowed across the
isthmus into the Pacific, within comparatively recent geological times.

It is not within the scope of this paper further to discuss the point on which Messrs.
Duncan and Verrill are at variance, as we cannot assume that the present fish-fauna
existed at so early a period. From the observations made on the fishes and shells we
are obliged to conclude that down to a very recent period a connexion between the two
seas has been kept open by channels and straits wide enough to allow of the passage
of these animals. Why corals, or at least a part of them, should not have been dis-
persed by their floating germs in a similar manner, is a circumstance which we cannot
explain.

The occurrence of identical species of freshwater fishes in rivers running to the two
opposite oceans is a matter of much less difficulty, and, besides, has been very generally
observed in various parts of the globe. ‘The same agencies which in other countries have
effected a wider dispersion of one species than of another must have been at work here
also. Prof. M. Wagner has, in his Memoir quoted above, so fully treated of this part of
our subject, with particular reference to the hydrographical peculiarities of the isthmus,
that we need not dwell further on it.

§ 7. Definition of the Characteristics of the Fish-fauna of Central America.

In defining the zoological characters of Central America, expressed in its fish-fauna, I
confine myself to the freshwater fishes proper. Here the nearctic types become extinct,
and are represented by five generic types, four of which, although with numerous species
in the north, have but a single one here—Lepidosteus, Amiurus, Sclerognathus, and
Haplochilus. Fundulus, extending a little further southwards (with one species in
Western Ecuador), is represented by four species in Guatemala. Not one of these
species is identical with a North-American.

Much greater is the affinity with neotropical types; and their representatives are much
more numerous: there is one species of Acara, one of Macrodon, seven of Tetragono-
pterus, one of Anacyrtus, twelve of Pimelodus, one of Plecostomus, two of Chetostomus,
two of Loricaria, one of Anableps, one of Carapus, the latter being identical with a
species from Guiana. Types in common with the West-Indian Islands are—Agono-
stoma with three species (one of which is said to be identical with a Jamaican species),
Girardinus and Gambusia with one, the two latter genera being also represented in the
Southern States of North America. The Siluroid genus Arius, which extends over the
tropics generally, is represented by nine species.

Finally, the following genera are peculiar to Central America, or at least have attained
there to the greatest development :—Zeros and the allied Meetroplus and Petenia with
thirty-four species, Hlurichthys with two, Chalcinopsis with three, Characodon with one,

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 403

Xiphophorus with one, Mollienesia with one, Pecilia with eight, and Belonesox with
one species.

The affinity of this freshwater fini with that of Mexico, will be found to be greater
than with that of any other country (I might mention about ten species common to
Guatemala and Mexico); but until we are better acquainted with the habitats of species
described as Mexican, a more detailed comparison of the two countries would be of but
little advantage. The freshwater fish-fauna of Central America may be shortly thus
characterized :—A part of the Chromides (Heros, &c.) and the Cyprinodontes generally
have attained to their greatest development ; neotropical types extending northwards
prevail over nearctic extending southwards, the latter being represented by a few extreme
branches.

§ 8. An Attempt to Subdivide this Fauna into Provinces.

We may subdivide this part of the freshwater fauna into the following provinces :—

A. The fresh waters north of the Lakes of Managua and Nigaraqua, emptying into the
Pacific.—To this province belong the fishes collected at Chiapam [Ch.], Huamuchal [H. ],
San José[J.], in the Rio Guacalate (Duefias) [G. ], San Salvador [S.], and Libertad [L.];
also the fishes from the DES of Amatitlan [Am.] and Atitlan [At.] may be referred to
the same province,

[The species printed in italics in the following lists are found also in one or more
other provinces, and in Atlantic rivers. |

Heros macracanthus 9:7: - . . . Ch:

H.
iramaculapusie es wees eee oe chy a) el. = = = =e =
nigrofaseiatus . .. +) 5. +. +. — —

eutiulatns: vs cuetas! ue st eee eee = =

Avius guatemalensis, . . . .. .Ch — =

Agonostoma microps ......— a — G. = = —
——platypogon «5. 2)... ke J

cerulescensi; a.gos) sume eet

H
Pimelodus guatemalensis . . . . . — H. — — = — =
Anacyrtus guatemalensis . . . . . — H
Tetragonopt. microphthalmus. . . . — — — — — An —
UNTER | eget gt, bal Syd Fe Baath love = — — — — An. —

Fundulus guatemalensis . . . . . — — — G, — An, —

pachyecephalusie) ey) se) ct — — — — — SSN
PUnclabush Aye tae ett eet.) tr Ohi |e — — = = —
Anablepsidoyiiemes mk eatin er oS Chy 7 = ==
Peciliamexicona. . . . . % . . Ch. 4. _ G. — An. —
CREF MAUS 20) GROEN — — =

Cy AED onion tuk O MEE? 13, era eee — — — — An. —
Girardinus pleurospilus. . . . . . — — — G: — — —
Clipeatlibertatises =) eset 20) — — a L. — —
Lepidosteus tropicus. . . . . . 2. — H. — — — — —

404 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

B. The fresh waters north of the Lakes of Manaqua and Nicaragua, emptying into the
Atlantic.—To this province belong the fishes collected in the Rio Usumacinta [U.] (and
in its tributaries Rio de Santa Isabel, Rio Chisoy, and Rio San Geronimo), in the Rio
Motagua [M.], and in the Rio Cahabon (Yzabal) [Y.].

IMlechsidormitdinie 1G) oss bees ie i) Gea ee M. Ve
Agonosioma nasutum 6: . . . . +» - » » e« « WU, M. —_
LET OSVPURIMU Leta get te es epee oho ee ores a ae
Spilurussies! APR EN. re TIN SOR Send cree

UL CUS Fe Fa RT ie Mash | NUTR WUE coe eae MADRE Se) Ge dey te

M

M

M
== motamuénsisinyid Gal Cie Piel ede ES M. —
—— microphthalmus:) 2.0 5 0. Se M
== oblongus: 8 6" Re ais. aS Re On er M
—angulifer . =

—— salvini .

: U
ATCC H ULATIS 2d Gat Tye SURMISE. ci Sled pet) Pee, doa (Ul. _

godmanni .
ATION CASSIS Ha) tse bectip Roads take ct Ate ame baave |S a huslbeidace

melanopus =
AMTUTUSeMeridionalisi; 2). ees Hea we cee ei
iPimelodusizodmanniy- meer eee: aha vo Us
MOLAPTIENSIS pseu Nee eye Lees shy” eee M. —

Salvini erg ot See aE Meee Tce Tego eee LS — me
INOKEWIC xg. alo OR) anor Gl oe Orem mete LUIs —_—

Tetragonopterus panumensis. . . . . 6 ee ee —_
brevimanus >. =O! Cpu temas ae si
@halemopsistdentex” 3) 2" 5) eo | tat eae tee ve UE
Fundulus labialis U.
Belonesox belizanus’”.) Fos Foe Pe ey aes Honduras, Belize.

Posciliaxchisoyensis# et 's74) ets. “eens alse Ue — —

| 3 |
dd i x |

Xu phophorusthell crite eae tetee wt oe eee ee Ue = _—

Sclerognathusimeridionalis=. 2-5 . 2°. 2. sas Jee Ue

= |

Carapusifasciatisistg.re tee fad boo Se en See

C. Lake Peten.—The fish-fauna of this limited district is so peculiarly developed,
that we cannot hesitate to describe it as a separate province.

Heros margaritifer, Petenia splendida.
—— melanurus. Pimelodus petenensis.
urophthalmus. Tetragonopterus petenensis.
affinis. Belonesox (im common with province
—-— friedrichsthalii. B).
salvini (in common with pro- Peecilia petenensis.
vince B). Mollienesia petenensis.
intermedius. Cheetoéssus petenensis.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA, 405

D. Lake of Managua.—Although the number of species known from this lake is
small, the forms are quite peculiar; we find here those species of Heros which are
distinguished by the extraordinary development of the lips, or by incisor-like teeth,
which render the separation into a distinct genus necessary. The development of
these Chromides is the more remarkable, as this lake occupies a space which is
supposed to have been a portion of a marine channel.

Heros erythreeus. | Heros lobochilus.
managuensis. | multispinis.
labiatus. | Neetroplus nematopus.

E. Lake of Nicaragua.—Also the fishes of this lake are, with two exceptions, peculiar ;
like Lake Managua, it appears to have been part of a marine channel.

Fleotris longiceps. Heros labiatus (Lake of Managua).

Heros longimanus. Pimelodus nicaraguensis.

|
citrinellus. | Gambusia nicaraguensis. -
|
|
|
|

dovii. Pecilia dovit (m common with Lake
nicaraguensis. Amatitlan).

F. The fresh waters south of the Lakes of Managua and Nicaragua to the Isthmus of
Darien.—We are obliged, at present, to unite into one province the fish-fauna of Costa
Rica, Veragua, Panama, and Darien, as our knowledge of the fishes of Costa Rica and
also of Veragua is too incomplete to admit of a comparison with those of the more
southern part of the isthmus. This is the more to be regretted, as a former separation
of these two parts and of their faunas is, as we have explained above, a matter of great
probability. The fishes of the Chagres River show a decidedly South-American cha-
racter. The identity of the freshwater fish-faunas of the Pacific and Atlantic sides is
here easily explained by the narrowness of the isthmus.

Hleotrusypictagy else eee — — R. Bayano.
Agonostoma nasutum . . . + . = Panama. =
TLOTULCOL Ge en — Panama. —
[Eerosy paca a eee OLACTeS: = —
altthrOnS? se, se wei = Western Veragua.
Acara ceruleomaculata . . . . Chagres. = =
Arius multiradiatus . . 2. . . — — R. Bayano.
/Blurichthys dorsalis . . . . . —_ Panama. —
jeMNI 5 5G 5 oe = Panama. —
Pimelodus wagner. . » . . . — Panama. —_—
modestus) +. woe. a ee) =) Chagres, — —
Plecostomus, sp. . . . - - = Chagres. —- —
Cheetostomus aspidolepis. . . . _- Veragua. _—
Chetostomus ?cirrhosus . . . . Chagres. = a

Loricarialima . . . . . . . Chagres. _— —

406 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

Loricaria uracantha . . . . . Chagres. — ==
Macrodon microlepis . . . . . Chagres. _ —
Tetragonopterus panamensis . . . — Panama. —

Ey ey ep —_— Panama. —
Chalcinopsis striatulus . .. . a * Panama. —

ChaeTeneisie eeu an ere. LCharres. — _—
Anacyrtus guatemalensis . . . . Chagres. — —_
Haplochilusidovit sei. < - = — Costa Rica. —
Poecilia elongata. -:-.-. . . . _ Panama. —

cilngi. Mewes. ieee. =. Chagres. — --

§ 9. Descriptive Part.

In the following descriptive part of this Memoir I have admitted full descriptions of
those species only which are not described elsewhere ; secondly, descriptive diagnoses of
those of which figures are given; and, finally, notes on some known species, if they
appeared to contribute to their better knowledge. For the descriptions of all the other
species (the insertion of which would be a repetition of matter already published), I
must refer the student to my general work on Fishes.

1. CENTROPOMUS APPENDICULATUS.
Poey, Mem. Cub. i. p. 119.
D. 85. A-j.. L. lat. 70-72.

Nine longitudinal series of scales between the origin of the second dorsal fin and the
lateral line. The height of the body is contained four times in the total length (with-
out caudal), the length of the head twice and two-thirds. Preorbital indistinctly
serrated ; suboperculum produced into a short flap, which extends to or nearly to the
vertical from the origin of the dorsal fin, The intermaxillary extends to below the
middle of the orbit. Dorsal spines of moderate strength ; the third is the longest, and
about half as long asthe head. The second anal spine is generally longer than the third ;
but sometimes they are equal in length, and even shorter than the third dorsal spine. ‘The
length of the ventral fin is more than one-half of its distance from the anal. Air-bladder
with a pair of appendages anteriorly, Silvery; dorsal fins blackish; lateral line black.

We kave received this species (which was originally described from Cuban examples)
from Surinam and Mexico, Mr. Salvin and Capt, Dow obtained a specimen from the
Chagres River, 10 inches long.

2. CENTROPOMUS MEDIUS.
Giimth. Proc. Zool. Soc. 1864, p. 144.
D.8\ 0S L. lat 67,
Eight longitudinal series of scales between the origin of the second dorsal fin and the
lateral line. ‘The height of the body is contained thrice and three-fourths in the total

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 407

length (without caudal), the length of the head twice and four-fifths. Preeorbital
finely serrated ; suboperculum produced into a flap, which does not extend to the
vertical from the origin of the dorsal fin. The intermaxillary extends somewhat beyond
the anterior margin of the orbit. Dorsal spines strong; the third is longer than the
fourth, and half as long as the head. The second anal spine long, but a little shorter
than the third, and equal in length to the distance between the extremity of the upper
jaw and the preeopercular margin. The length of the ventral fin is much more than
one-half of its distance from the anal fin. Lateral line black.
Two specimens, 13 inches long, from Chiapam.

3. CENTROPOMUS NIGRESCENS.
Giinth. Proce. Zool. Soc. 1864, p. 144.
D. 8| 7 A-g L.lat. 70.

Ten longitudinal series of scales between the origin of the second dorsal fin and the
lateral line. ‘The height of the body is contained four times and a half in the total
length (without caudal), the length of the head twice and four-fifths. Preorbital not
serrated; suboperculum produced into a short flap, which does not extend to the
vertical from the origin of the dorsal fin. The intermaxillary extends a little beyond
the middle of the orbit. Dorsal spines rather feeble; the third and fourth are equal
in length, two-fifths of the length of the head. The second and third anal spines also
are equal in length, and not longer than the dorsal spines mentioned. The length of
the ventral fin is scarcely more than one-half of the distance of its base from the anal.
Air-bladder without appendages anteriorly. Silvery; upper parts and fins blackish ;
lateral line black.

One specimen, 14 inches long, from Chiapam.

This species is allied to C. appendiculatus (Poey), but differs externally in its con-
siderably more feeble and shorter fin-spines.

4, CENTROPOMUS PARALLELUS.
Poey, Mem. Cuba, ii. p. 120.
D. 8|7 A. L. lat. 85-90.

Twelve longitudinal series of scales between the origin of the second dorsal fin and
the lateral line. The height of the body is contained thrice and three-fourths in the
total length (without caudal), the length of the head twice and a half. Preorbital
distinctly serrated ; suboperculum produced into a flap, which extends to the vertical
from the origin of the dorsal fin. The intermaxillary extends a little beyond the
middle of the orbit. Dorsal spines rather feeble; the third is the longest, half as long
as the head. The second anal spine is exceedingly strong, longer than the third and the
third dorsal spine. The length of the ventral fin is considerably more than one-half of

408 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

the distance of its base from the anal. Air-bladder without appendages anteriorly.
Silvery; upper parts and fins greenish; lateral line not black.

This species occurs in Cuba; we have received it from San Domingo, Jamaica, and
Bahia. Messrs. Dow and Salvin collected a specimen in the Chagres River.

5. CENTROPOMUS ARMATUS.
Gill, Proce. Ac. Nat. Se. Philad. 1863, p. 163.

2

D. 8|5- A.2 L.lat.51. L. transv. 7/14.

6

Six longitudinal series of scales between the origin of the second dorsal fin and the
lateral line. The height of the body is contained from thrice and two-fifths to thrice
and three-fourths in the total length (without caudal); the length of the head twice
and a half. Preeorbital serrated in its hinder half; suboperculum produced into a long
flap, which extends beyond the vertical from the origin of the dorsal fin. The inter-
maxillary extends scarcely to below the middle of the orbit. Dorsal spines of moderate
strength ; the third is the longest, and half as long as the head. The second anal spine
is exceedingly strong, much stronger than the third, and longer than the third dorsal
spine. The length of the ventral fin is scarcely more than one-half of the distance of
its base from the anal. Silvery; dorsal fins, a blotch on the opercle, and the membrane
between the anal spines blackish. Lateral line not black.

Several specimens, 12 inches long, were collected by My. Salvin at Chiapam.

6. CENTROPOMUS ENSIFERUS.
Poey, Mem. Cub. ii. p. 122, pl. 12. fig. 1.
D. 8|5, Ag, 1. lat. 53.

Seven longitudinal series of scales between the origin of the second dorsal fin and the
lateral line. The height of the body is one-fourth of the total length (without caudal),
the length of the head two-fifths. Prorbital coarsely serrated; suboperculum pro-
duced into a flap, which extends to the vertical from the origin of the dorsal fin. The
intermaxillary extends scarcely to below the middle of the orbit. Dorsal spines of
moderate strength; the third and fourth are the longest, and two-fifths as long as the
head. The second anal spine is exceedingly strong, much stronger than the third, and
much longer than the dorsal spines. The length of the ventral fin is somewhat more
than one-half of the distance of its base from the anal. Silvery; dorsal fin, a blotch on
the opercle, and the membrane between the anal spines blackish. Lateral line not
black.

This species occurs in Cuba; we have received it from Jamaica and from the Guyanas.
Mr. Godman collected a specimen, 12 inches long, at Belize.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 409

7. CENTROPRISTIS MACROPOMA. (PI. LXV. fig. 1.)
Giinth. Proc. Zool. Soc. 1864, p. 145.
De AS. Tat: 52. "L trausy. 6/16,

Closely allied to C. radialis, Q. & G.; but whilst that species has a notch above the
spiniferous angle, the present has its preopercular margin not interrupted, the long
spines of the angle gradually passing into the finer serrature. There are six series of
scales between the eye and the angle of the preoperculum. The maxillary extends
nearly to the vertical from the posterior margin of the orbit. Dorsal fin with a notch,
the ninth spine being considerably shorter than the tenth. A series of rather small
brownish spots above and below the lateral line.

Three specimens, 43 inches long, were collected by Messrs. Dow and Salvin on the
Pacific coast of Panama.

8. SERRANUS CREOLUS, CO. & V.

I have examined specimens from the Atlantic coasts only; but Mr. Gill has found it
in a collection of fishes from Lower California, the specimens being undistinguishable
from those of the West Indies and South America (Proc. Ac. Nat. Sc. Philad. 1862,
p- 249).

12. SERRANUS SELLICAUDA.
Epinephelus sellicauda, Gill, Proc. Acad. Nat. Se. Philad. 1862, p. 250.
De Are, To tat 100:

Caudal fin with the posterior margin convex. ‘The height of the body is rather more
than three-fourths of the length of the head, and one-fourth of the total (caudal
included). The diameter of the eye is one-fourth of the length of the head. Praoper-
culum finely serrated behind, with some coarser teeth at the angle, lower limb entire ;
sub- and interoperculum entire. Ventrals three-fourths of the length of pectorals, and
reaching two-thirds of the distance between their insertion and the commencement of
the anal. Brownish, with olive-coloured spots of larger and smaller size on the body
and opercles. All the fins with a narrow white margin. A square black blotch across
the back of the tail.

Description.—Body not very elevated; its greatest height is below the third spine of
the dorsal fin, rather more than three-fourths of the length of the head, and one-fourth
of the total. The distance between the end of the dorsal and the commencement of
the caudal is nearly one-sixth of the length of the base of the dorsal, is contained once
and two-thirds in the base of the anal, is one-fourth of the distance between the dorsal
fin and the snout, and equals the least depth of the tail. ‘The distance between the
eyes is one-half of the diameter of the eye, and covered with very minute scales, which
are found also on the preorbital around the nostrils. The length of the snout is two-
thirds of the diameter of the eye. The maxillary bone reaches the vertical from
the posterior margin of the eye. he mandibulary is one-half of the length of the

VOL. VI.—PART VII. 31

410 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

head. ‘The lips are not very thick. Posterior limb of preoperculum very convex,
minutely serrated, with three coarser teeth at the angle; lower limb toothless. Sub-
and interoperculum entire. Operculum terminating in three not very strong teeth, the
upper of which is somewhat more remote than the others, the middle one being the
more prominent. Suprascapular concealed by the scales.

The membrane of the dorsal fin is scaly for about half the height between the spines
and rays; the spinous portion scarcely lower but longer than the soft, with its upper
margin convex, and a small membranaceous appendage behind the tip of each spine.
The first spine is the shortest, rather more than half the length of the second, which is
one-fifth shorter than the third; from the third to the seventh the spines are equal,
becoming slightly shorter at the eighth; the last two spines are of equal length. ‘The
rays increase slightly from the first to the sixth, after which the upper margin is
straight, becoming again rounded posteriorly. ‘The first ray is one-fifth longer than
the preceding spine. Caudal with posterior margin convex. The commencement of
the anal is on a line with that of the soft dorsal, and it ends before the termination of
the dorsal; the first spine is short, not half the length of the second, which is long and
strong, longer than any of the dorsal spines; the third is slenderer, and equal to the
third dorsal spine: the margin of the soft part of the fin is nearly straight, sharply
rounded off posteriorly. The pectoral consists of eighteen rays, is rounded, and longer
than the ventral, and covered with very minute scales to one-third of the length. The
ventrals reach the vent; the second ray is the longest, the spine being equal to the
second of the dorsal. Canine teeth of moderate size, those of the lower jaw rather
small. Coloration as described above.

A single specimen, 4 inches long, was sent by Capt. Dow from the Pacific coast of
Panama. The specimen in the collection of the Smithsonian Institution is from the
coast of Lower California; a statement of its size, which would have been of some
importance, is omitted.

13. SERRANUS ANALOGUS.
Epinephelus analogus, Gill, Proc. Acad. Nat. Sc. Philad. 1863, p. 163.
D. 7 A.% L. lat. ca, 100.

Adult—The height is contained thrice in the total length (without caudal), the
length of the head twice and two-thirds. The preoperculum is finely serrated behind,
and towards the angle armed with three or four strong teeth. ‘The diameter of the
eye equals a sixth of the head’s length, and equals the interorbital space as well as
the snout behind the intermaxillaries. The third, fourth, and fifth spines are equal,
and contained twice and two-thirds in the length of the head; the tenth thrice and a
half. The caudal fin enters five times and a half in the length, the height of the
dorsal twice and three-fourths in the head. ‘The anal is deeper; its third spine is
longest, and enters four times and three-fourths in the head’s length; the pectoral is
at least half as long as the head; the ventral shorter, but coterminal with it.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 411

The colour is purplish grey, with numerous dark spots about as large as the pupil;
those of the pectoral and caudal fins are smaller and more crowded, of the dorsal, anal,
and ventral more like those of the body.

The specimens are from 11 to 15 inches long, and were found by Capt. Dow at
Panama.

We have received also a sma/ler example, 5 inches long, from the same gentleman. It
differs from those described above in being provided with five cross bands, paler in colour
than the spots, which are one-third the size of the eye. ‘The dorsal fin is scarcely
notched, the tenth spine being but little shorter than the third or fourth, the length of
which is contained twice and two-thirds in that of the head. The example being
young, its eye is comparatively larger.

14, Piecrropoma arrum. (PI. LXVII. fig. 3.)
Epinephelus afer, Bloch, Vat. 327 (fide Peters, Monatsber. Ak. Wiss. Berlin, 1865, p. 105).
Alphestes afer, Bl. Schn. p. 236.
Plectropoma chloropterum, Cuvy. & Val. 11. p. 398. Poey, Mem. Cub. i. p. 73, lam. 9. fig. 3.
monacanthus, Mull. & Trosch. in Schomb. Hist. Barb. p. 605. Giinth. Fish. 1. p. 164.
— multiguttatum, Giinth. Proc. Zool. Soc. 1866, p. 600.

ee Nee ate To.

is20"

Caudal rounded. The height of the body is equal to the length of the head, and
contained twice and three-fourths in the total (without caudal). The diameter of the
eye is one-fifth of the length of the head, and a little less than that of the snout.
Preoperculum with a strong spinous tooth below the angle, pointing forwards. Olive-
brown, head and body with numerous spots.

Description —Body somewhat elevated ; its greatest height is below the fourth spine
of the dorsal, and equal to the length of the head, which is contained thrice and one-
third in the total (the caudal included). The distance between the dorsal and the
caudal is contained seven times and one-third in the length of the base of the
dorsal fin, twice in that of the base of the anal, four times in the distance between the
dorsal fin and the snout, and is considerably less than the least depth of the tail. The
distance between the eyes is about two-thirds of the diameter of the eye, and covered with
scales which extend forward beyond the nostrils on the preorbital, and in a narrow
band on the upper maxillary. The length of the snout equals the diameter of the
eye, which is one-fifth of the length of the head. The maxillary reaches a little beyond
the level of the posterior margin of the eye. The mandibulary is covered with minute
scales, and is equal to one-half the length of the head. ‘The lips are thick and fleshy.
The posterior limb of the preoperculum slants obliquely backwards, and is minutely
serrated, the denticulations becoming coarser at the angle; and beneath on the lower
limb at some distance from the other teeth there is a single strong tooth pointing
downwards, and nearly concealed by the skin; sub- and interoperculum not serrated.

3L2

412 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

The operculum terminates in three, flat, triangular teeth, the upper of which is the
more distant and somewhat more obtuse than the others, the middle one being the
longest, but not very prominent, and the lower one the shortest and weakest. ‘The
suprascapula is concealed by the scales.

Base of dorsal fin covered with very small scales, a tapering band of scales runs up
between each pair of the spines and rays to about half the height of the fin. Spinous
portion rather lower but longer than the soft, its upper margin convex; the membrane
between each spine is deeply notched, and there is a small membranaceous appendage
behind the top of each spine. ‘The first spine is the shortest, half the size of the
second ; the second is five-sixths of the length of the third; the third, fourth, and fifth
are the longest, and of nearly equal length; the spines then become gradually shorter
to the last one, which is scarcely longer than the preceding. ‘The soft portion exhibits
an entirely rounded upper margin, the rays becoming longer from the first to the sixth
or seventh, and shorter from the fourteenth to the last; the first ray is one-fourth
longer than the preceding spine. Caudal with the posterior margin convex. Anal
commencing a little behind the commencement of the soft dorsal, and terminating in
advance of the end of the same; the first spine is not very strong, and short; the
second long, thick, and strong; the third more slender and shorter, being but little
longer than the second dorsal spine; the margin of the fin is rounded throughout, the
third ray being the longest, and the subsequent ones becoming progressively shorter.
The pectoral is composed of eighteen rays, rounded, one-fourth longer than the ventral,
and covered with minute scales for about one-third of its length. The ventral reaches
to the vertical from the origin of the eighth spine of the dorsal, but not to the vent;
the spine is a little less than two-thirds the length of the first ray; the first and second
rays are the longest, the others diminishing gradually in length; the length of the
spine is somewhat less than that of the second dorsal spine. Canine teeth small in
both jaws.

This species varies somewhat in coloration, as most of its congeners; the spots are
numerous aud small, either of a uniform dark-brown colour, or of a light colour and
mixed with large brown spots. Pectoral fins with narrow blackish cross bands.

One example, 10 inches long, and three smaller ones have been collected by Capt.
Dow on the Pacific coast of Panama. The latter have the spots somewhat larger and
less conspicuous than the adult. This species cccurs also in the West Indies and at
the Falkland Islands.

15, Ruypricus DECORATUS.
Rhypticus nigripinnis, Gill, Proc. Ac. Nat. Se. Philad. 1861, p. 53.
Promicropterus decoratus, Gill, 1. c. 1863, p. 164.
Dew A. 16,

26°
The two dorsal spines are continuous with the soft portion, Body generally with
more or less numerous round whitish spots, many of which have a brown centre.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 413

Messrs. Dow and Salvin have collected several examples, from 3 to 8 inches long, on
the Pacific coast of Panama.

The species described by Holbrook as F. maculatus, and said to have the dorsal
spines separated from the soft portion, may eventually prove to be identical with the
Pacific fish.

19. MESOPRION ARATUS.
Giunth. Proc. Zool. Soc. 1864, p. 145.
Dig Ag 4. lat 45. L. transv. 44/12.

The height of the body equals the length of the head, and is contained thrice and
two-fifths, or thrice and one-fifth in the total (without caudal). The maxillary does not
extend backwards to the vertical from the centre of the eye. Praoperculum finely
serrated, with scarcely a trace of a posterior notch. Dorsal spines of moderate strength ;
the third and fourth are the longest, two-fifths of the length of the head; the eleventh
is scarcely longer than the tenth, which is rather more than half as long as the fourth.
Caudal fin emarginate, two-thirds scaly; anal spines short, rather feeble, the third
longer than the second, and equal in length to the last dorsal spine. Upper and
lateral parts brownish-olive, each scale with a pearl-coloured spot, the spots forming
together very distinct longitudinal stripes; no black lateral spot ; hind part of the root
of the pectoral brown. Lower parts salmon-coloured..

We have six examples: two, 15 inches long, were collected by Mr. Salvin at Chiapam ;
and four others were sent by Capt. Dow from the Pacific coast of Panama.

21. APOGON Dovu.
Giinth. Proc. Zool. Soc. 1861, p. 371.
D.6|;, AZ L. lat. 25. L. transv. 3/9.

A roundish black spot on each side of the root of the caudal; the spinous dorsal
colourless, transparent; uniform olive (in spirits). Head densely punctulated with
brown. Only the hind margin of the posterior preopercular ridge is serrated. Dorsal
fins nearly equal in height.

The height of the body is one-third of the total length (without caudal); the length
of the head two-fifths; eye large, its diameter being more than one-third of the length
of the head. Palatine and vomerine teeth present. The upper jaw overlaps slightly
the lower; maxillary extending backwards to below the posterior third of the orbit.
Operculum with an upper flexible point, and with a lower stiff spine. The third dorsal
spine is a little longer than the second, one-half the length of the head. Caudal fin
slightly emarginate, with the angles rounded.

Total length 26 lines.

This species is so closely allied to A. inermis from the Mediterranean, that perhaps

414 DR. GUNTHER ON THE FISHES OF. CENTRAL AMERICA.

it would be better not to separate it; the only difference which I can find is the form
of the soft dorsal fin, which is considerably higher than the spinous in the Mediterranean
species.

22. PRISTIPOMA MELANOPTERUM.
Pristipoma melanopterum, Cuy. & Val. v. 1880, p. 273.
bilineatum, Cuv. & Val. v. 1830, p. 271, pl. 122.
Hemulon melanopterum, Ranzani, Comm. Bonon. y. 1842, p. 343, tab. 30.

Pristipoma melanopterum, Giinth. Fish. i. 1859, p. 287.
Var. Genytremus interruptus, Gill, Proc. Acad. Nat. Sc. Philad. 1862, p. 256.
Pristipoma melanopterum, Giinth. Proc. Zool. Soc. 1864, pp. 23 & 27.

This species occurs on both sides of Central America, Capt. Dow having collected
specimens at Panama and Colon. Mr. Gill has found it also in a collection of fishes
from Lower California. He describes his Pacific specimen as a distinct species ; but the
distinctive characters are, according to my views, not of specific value. He mentions it
in the following terms :—

“The species is so closely allied to dilineatus, that it might be even considered as a
variety, but it appears to differ by the steel-blue colour of the back, and the discon-
tinuance of the lateral band a short distance before the spot on the tail'; at its end the
band is bounded below by the lateral line. In. other respects, the two species are so
similar, that a detailed description would be only a repetition of that of bilineatus.”

23. PRISTIPOMA VIRGINICUM.

We have examined specimens of this species from the West Indies, from the Atlantic
coasts of Central America, and from Bahia. Mr. Gill has described an example from
Panama under the denomination of Anisotremus teniatus, Proc. Ac. Nat. Sc. 1861,
p- 107. Although six or seven is the normal number of longitudinal bands, it is some-
times increased by a more or less complete division of one or several bands. It appears
to be more natural to consider the golden colour the ground-colour than the blue, as
after death it fades into the same colour as that of the space between the black vertical
bands. In al/ specimens, I have found the bluish bands edged with purplish. Mr. Gill,
in describing his A. teniatus, has taken the blue colour as ornamental, whilst in his
description of A. virginicus the character assigned to the colours is reversed, and the
blue colour regarded as ground-colour. There is no specific difference between these
fishes.

24, PRISTIPOMA DOVIL.
Ginth. Proc. Zool. Soc. 1864, p. 23, pl. 3. fig. 1.

D. %. A.3.. L. lat. 48. L. transv..8/15.
The height of the body is one-half of the total length (without caudal); the length
of the head one-third. Snout obtuse, not much longer than the eye; cleft of the

? This is also the case in some Atlantic specimens.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 415

mouth small, the maxillary extending to the vertical from the anterior margin of the
orbit. Lips thick; a pair of pores on the symphysis of the lower jaw, a central groove
behind it. Snout naked, the remainder of the head being scaly. The width of the
interorbital space is much less than that of the orbit. Dorsal and anal spines exceed-
ingly strong; the third of the dorsal fin is the longest, and nearly two-thirds as long as
the head. ‘The second anal spine is much longer than the third, and a little shorter
(but stronger) than the third of the dorsal fin. Each ray of the soft fins is accompanied
by a series of minute scales, but only on the caudal fin are these scales dense enough
to cover the rays. Caudal fin slightly emarginate. Silvery, with four black cross
bands; the first runs from the occiput, through the eye, to behind the angle of the
mouth, the second from before the dorsal fin to below the base of the pectoral, the
third from the base of the sixth, seventh, and eighth dorsal spines to the vent; the
fourth descends from the origin of the soft dorsal to that of the soft anal. Fins
blackish. The cross bands appear to become fainter in old age.
‘Two specimens, 83 and 9 inches long, in the collection from Panama.

25. PRISTIPOMA CHALCEUM.
Giinth. Proc. Zool. Soc. 1864, p. 146.
De sae Lilet oe. 0. transy. 11/19,

The height of the body is contained twice and two-thirds in the total length (without
caudal), the length of the head thrice anda third. ‘The diameter of the eye is nearly
equal to the width of the interorbital space, and two-thirds of the extent of the snout.
The maxillary does not extend backwards to the vertical from the anterior margin of
the orbit. Praeoperculum minutely serrated behind, with the angle rounded, but not
produced. ‘There is no notch between the spinous and soft portions of the dorsal fin,
the hinder spines being only a little shorter than the anterior rays; dorsal spines of
moderate strength, the fourth being the longest, not quite half as long as the head ;
anal spines short, the second being only a little longer than the third, two-sevenths of
the length of the head. Caudal fin subtruncated, scarcely emarginate. Dorsal and
anal perfectly scaleless. The pectoral fin extends to the vertical from the vent.
Bronze-coloured, shining silvery, perfectly immaculate; vertical fins blackish, with an
indistinct light band along the base.

One specimen, 8 inches long, was discovered by Messrs. Dow and Salvin on the
Pacific coast of Panama.

26. PRISTIPOMA HUMILE.
Kner & Steindachner, Sitzgsber. Ak. Wiss. Miinch. 1863, p. 222; and Abhandl. bayer. Ak. Wiss. x.
p. 3, tab. 1. fig. 1.
D. 3 Asg. L. late 56. L. transv.. ig-ay
The height of the body is contained thrice and two-thirds in the total length (without

Cee. pyl. 3.

416 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

caudal), the length of the head thrice. The diameter of the eye equals the width of
the interorbital space, is one-fifth of the length of the head, and two-thirds of the
extent of the snout. Snout produced; cleft of the mouth wide; the maxillary extend-
ing beyond the front margin of the eye. Praoperculum with the hind margin vertical
and finely serrated. The spinous and soft portions of the dorsal fin are separated by a
notch; dorsal spines moderately strong, the fourth being the longest, its length being
contained twice and three-fourths in that of the head. Second anal spine exceedingly
strong, more than half as long as the head. Caudal fin slightly emarginate ; pectorals
terminating at some distance before the vent. Scales ctenoid. _Coloration uniform.

‘This species is known from a single example (size not stated) from the Rio Bayano
(Panama).

27. PrisTipoMA MACRACANTHUM. (PI. LXIV. fig. 1.)
Ginth. Proc. Zool. Soc. 1864, p. 146.
D.11\4. A. 3/8. L. lat. 47. L. transv. 6/13.

The height of the body equals the length of the head, and is one-third of the total
(without caudal). The diameter of the eye equals the width of the interorbital space,
and is two-thirds, or somewhat less than two-thirds, of the extent of the snout. Hind
margin of the anterior nostril with a broad flap. Snout somewhat produced; the
maxillary does not extend to below the anterior margin of the eye. Praoperculum
with the hind margin rather concave, and with stronger teeth at the angle, which is
rounded. ‘The spinous and soft portions of the dorsal fin are separated by a deep
notch, the spine of the soft portion being much longer than the preceding, which is
somewhat longer than the second. Dorsal and anal spines exceedingly strong; the
fourth dorsal spine is the longest, its length being contained twice and a third in that
of the head. ‘The second anal spine much longer and stronger than the third, and even
than the fourth dorsal spine. Candal fin truncated. Each soft ray of the vertical fins
is accompanied by a series of minute scales. The pectoral fin extends to the vent.
Scales smooth. Silvery. with several very indistinct dark cross bands on the back,
which appear to be arranged as in P. leuciscus.

Two specimens, 11 and 14 inches long, were collected by Mr. Salvin at Chiapam.

29. Pristipoma LEuciscus. (Pl. LXVI. fig. 3.)
Giinth. Proc. Zool. Soc. 1864, p. 147.

D.11|4. A.3/7-8. L. lat. 51. L. transv. 2%.

The height of the body is contained thrice or thrice and a third in the total length
(without caudal), the length of the head thrice and a fourth. The diameter of the eye
is equal to, or more than, the width of the interorbital space, but is less than the extent
of the snout. The maxillary does not quite extend backwards to the vertical from the
anterior margin of the orbit. Preeoperculum finely serrated behind, with the angle

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 417

rounded, and with the hind margin slightly concave. The spinous and soft portions of
the dorsal fin are separated by a deep notch, the spine of the soft portion being nearly
twice as long as the preceding. Dorsal spines long, of moderate strength: the third is
the longest, and one-half, or more than one-half, as long as the head. -Anal spines
rather strong: the third is a little longer than the second, equal to the seventh dorsal
spine, and more than one-third of the length of the head. Caudal fin emarginate.
Each soft ray of the vertical fins is accompanied by a series of minute scales, The
pectoral fin extends to the vertical from the origin of the anal in the younger example,
but is shorter im adult ones. Scales smooth, bright silvery; young specimens with
several very indistinct dark cross bands on the back, the first from the nape of the neck
to the gill-opening, the second below the seventh dorsal spine, the third below the last
dorsal spine ; old specimens with the marginal membrane of the operculum black.

One specimen, 74 inches long, was found by Mr. Salvin at San José. Three others,
from 11 to 12 inches long, are from Chiapam ; and Capt. Dow found it also at Panama,
where it does not appear to be rare.

30. Conopon pacirici. (Pl. LXIV. fig. 3.)
Giinth. Proc. Zool. Soc. 1864, p. 147.
D.11|%,° A. 3. Lelat. 47. L. transv. 7/13.

Diagnosis.—The spinous teeth at the angle of the preoperculum are not much
stronger than the others. The height of the body is contained twice and two-fifths in
the total length (without caudal).

One specimen, 123 inches long, was collected by Mr. Salvin at Chiapam.

Description.—The body is compressed, and considerably elevated ; its greatest height,
which is below the fifth dorsal spine, is contained twice and three-fourths in the total
length. Upper profile rounded from the first dorsal spine to the nape, concave over
the eyes, whence it descends abruptly over the snout. The upper surface of the head
is very broad, the space between the eyes being nearly twice the width of the orbit.
The snout is thick and obtuse; the lips thick and fleshy. Teeth in a villiform band in
both jaws, with an outer series of conical teeth. Chin with a median groove and a
pair of pores. Posterior limb of preoperculum straight, regularly and distinctly
serrated, the teeth becoming gradually a little larger at the angle, and continued on
the lower limb; the entire surface of the preoperculum is covered with scales, which
are smaller than those of the operculum, and reach to the margin of the bone. The
operculum has a notch behind, between two obtuse and feeble points. Suprascapular
margin indistinctly toothed or roughened. The origin of the dorsal is in the vertical
from the root of the pectoral, and its termination is vertically opposite to that of the anal;
the base of the spinous portion is nearly twice as long as that of the soft. The spines.
are strong, broader alternately on one side than on the other; the first is small, not
quite one-half the length of the second, which is rather more than half that of the

VOL. VI.—PART VII. 3M

418 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA,

third; the third spine is three-fourths the length of the fourth; the fifth is the
longest, its length being contained twice and two-thirds in the height of the body;
the sixth and fourth spines are equal in height, and the subsequent spines decrease
gradually in length; the twelfth, which must be considered part of the soft dorsal, is
slightly longer than the preceding spine, and equal to the tenth. The soft portion has
a rounded margin; the third or highest ray is not quite equal to the fifth spine, and is
twice as long as the last. The spinous portion as well as the soft can be received into
a scaly sheath. The caudal fin is very slightly emarginate, scaly to within a short
distance from its tip, and one of its longest rays is nearly one-fifth of the total length,
The distance between the caudal and anal fins is less than the base of the latter; the
first anal spine is opposite to the third ray of the dorsal, it is strong, broader on the right
side, and excavated posteriorly, and is one-half the length of the second, which is very
long and strong, equal in length to the fifth dorsal spine, and broader on the left side ;
the third anal spine is equal to the third of the dorsal, and little more than half the
height of the first ray; the first and second rays are the longest, and the margin of the
soft portion is vertical. The pectoral is moderately long, its length being contained
four times and a half in the total. Root of ventral immediately behind that of
pectoral ; the spine is of moderate size and strength, a little more than half the length
of the first ray, which is produced about one-eighth of an inch at its tip; the other rays
decrease gradually in height. The scales are of moderate size, very finely crenated,
with the margin convex. ‘The lateral line is parallel with the curve of the back. Scales
silvery, with purple reflexions ; membrane between the scales brown; fins blackish.

34. H@MULON BREVIROSTRUM.
D. 2% Ag L. lat. 50. LL, transy. 5/14.

This species is closely allied to i. chromis and H. canna, differing from both oY its
much shorter and more convex snout.

The height of the body is contained twice and
two-thirds in the total length (without caudal),
the length of the head thrice and one-fourth.
The snout is short, not much longer than the
diameter of the eye, which is more than one-
fourth of the length of the head. Cleft of the
mouth rather wide, the maxillary extending
beyond the vertical from the front margin of
the eye. Hind margin of the preoperculum
slightly emarginate, its angle with more con-
spicuous denticulations. Dorsal fin notched,
with strong spines; the fourth is the longest,
half as long as the head. Caudal: fin forked.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 419

The second anal spine is strong, and somewhat longer than the third, but not quite as
long as the fourth of the dorsal fin. Scales above the pectoral fin not conspicuously
larger than the others. More or less conspicuous oblique brown streaks run along the
series of scales, and are broken up into series of spots in larger examples. A vertical
black spot covered by the angle of the preoperculum.

We possess four examples of this species: three were collected by Capt. Dow at
Panama; and the fourth is from Puerto Cabello. The largest is 8 inches long.

35. H#MULON MARGARITIFERUM. (Pl. LXV. fig. 2.)

Giinth. Proc. Zool. Soc. 1864, p. 147.
D.% A. 3. L. lat. 55, L. transv. 6/15.

The height of the body is one-third of the total length (without caudal), the length
of the head two-sevenths. The diameter of the eye is two-sevenths of the latter, and
equal to the extent of the snout and to the width of the interorbital space, which is
very convex. The maxillary extends beyond the vertical from the anterior margin of
the eye. Preoperculum emarginate behind. Dorsal fin scarcely notched, with the
soft portion very low; its spines are moderately strong, the fourth is the longest, not
quite half as long as the head. Anal spines strong; the second is longer and stronger
than the third, and equal to the eighth of the dorsal. The soft vertical fins enveloped
in scales; caudal forked, with the upper lobe longest. The pectoral fin does not extend
to the vent. Greenish olive above, each scale with a pearl-coloured centre; sides
silvery; a blackish spot above the axil.

One specimen, 12 inches long, was obtained by Messrs. Dow and Salvin on the
Pacific coast of Panama.

39. CuzTopon HUMERALIS. (PI. LXV. fig. 3.)
Giinth. Fish. 1. p. 19.

I have given a full description of this species (/.¢.). The Pacific coast of Central
America appears to be its true home. Messrs. Salvin and Dow collected three speci-
mens at Panama; and our other specimens, which we received from the Haslar Collec-
tion, are probably from Guatemala, from which country Sir J. Richardson, as we
know, obtained a collection of fishes. J have no doubt that the statement of this
species extending to the Sandwich Islands is correct. The Panama examples differ
from the typical specimens only in having an additional black cross band near the
hind margin of the caudal fin.

41. POMACANTHUS ZONIPECTUS.
Pomacanthodes zonipectus, Gill, Proc. Ac. Nat. Sc. Philad. 1862, p. 244.
1 3
D. 93-24° A. 30°
“The form much resembles that of Pomacanthus. The greatest height equals three-
; 3M 2

420 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

fifths of the length. The head forms about a quarter, and the caudal fin about a sixth of

the total length. ..... The dorsal is considerably produced at the sixth ray, which passes

behind the rounded posterior margin. . . .__ Brownish, margined with light on each scale.

A very dark brown band girdles the breast behind the pectoral and ventral fins.” . .
Obtained by Capt. Dow at San Salvador.

43. UPENEUS TETRASPILUS. (PI. LXVI. fig. 1.)
Giinth. Proc. Zool. Soc. 1864, p. 148.

D. 8|9. “A. 7. L. lat. 33. L. transv. 2/6.

The height of the body equals the length of the head, and is contained thrice and
two-fifths in the total (without caudal); the width of the interorbital space is two-
thirds of the length of the snout. Teeth in both jaws in two series, the outer series of
the upper jaw being formed by very obtuse and partly confluent teeth. The maxillary
is dilated and rounded behind, and bent upwards into a sort of hook; the barbels
extend to the vertical from the root of the pectoral. The third and fourth dorsal
spines are subequal in length, longer than the second, and nearly three-fourths of the
length of the head. Greenish olive above, each scale above and below the lateral line
with a large pearl-coloured spot; sides yellow; a rose-coloured band on each side of the
belly. A large blackish blotch on the lateral line, behind the hind part of the spinous
dorsal fin. A second smaller blackish spot behind the orbit; the latter is sometimes
very indistinct.

Two specimens, 8} inches long, were collected by Messrs. Dow and Salvin on the
Pacific coast of Panama.

This species would belong to the division which has been called Mulloides.

44. UPENEUS GRANDISQUAMIS.
Gill, Proc. Acad. Nat, Se. Philad. 1863, p. 168.
This species, which belongs to Bleeker’s division Upeneus, is described thus :—
D. 8}. A. 7... L, lat. 30. L, transy, 22/5.

The greatest height is contained four times in the length to the end of the median
caudal rays, and four times and a half in the total. The head equals the height, and is
itself longer than high, the profile in front of the eyes rapidly declines downwards, and
is nearly rectilinear. The diameter of the eye enters thrice and a half in the head’s
length, and the height of the preorbitar twice and three-fourths, The supramaxillar
ends at the vertical from the front of the eye. The teeth in front of the upper jaw are
biserial ; bélow uniserial. The first dorsal fin is highest at the third spine, and there
equals the head in front of the preopercular margin; the first is exceedingly short, and
the second and fourth nearly equal, little shorter than the third; all the spines are very
slender towards the ends, The distance of the second from the first dorsal enters once

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 42]

and three-fourths in the base of the former, and in that interval are three scales; its
length is less than that of the first. The ventral equals the distance of the hinder
margin of the orbit from the snout. The tubes of the lateral line have slender branches
diverging from them, generally directed obliquely upwards. The larger scales have six
radiating strie. The colour is light greenish brown above, with an indistinct silvery
spot at the centre of each scale. Below the lateral line, especially between it and the
anal fin, the colour is rose. The dorsal fins covered with spots of the colour of the
back. The others are immaculate.

Two specimens, the longest of which is 7} inches long, were collected by Capt. Dow
on the Pacific coast of Central America.

47, CHRYSOPHRYS CALAMUS.
A fine example, 16 inches long, has lately been sent by Capt. Dow from Panama.

49, Crrruiticutuys RivuLATUS. (Plate LXX XVI. fig. 4.)
Cirrhites rivulatus, Valenc. Voy. Vénus, Poiss. p. 309, pl. 3. fig. 1 (bad).
DW. A.2.. L. lat. 47. L. transv. 6/14.

The height of the body is contained thrice in the total length (without caudal), the
length of the head twice and two-thirds. The snout is of moderate extent, compressed
and rather elevated; the maxillary extends beyond the front margin of the eye. Inter-
orbital space deeply concave, and half as wide as the orbit ; a low longitudinal median
crest on the crown of the head, Preoperculum finely serrated behind. The fourth,
fifth, and sixth dorsal spines are the longest, two-sevenths of the length of the head, all
are of moderate strength. Seven simple pectoral rays, none of which extend so far
backwards as the ventral fin. ‘The second anal spine is longer, but scarcely stronger,
than the third. Brownish, with transverse dark brown bands and spots, all of which
are edged with light blue. There are two of these bands on the head crossing the
preoperculum ; five on the body and tail, composed of large, more or less confluent,
round spots; especially the third and fourth terminate above each in a pair of large
spots, the first pair occupying the end of the spinous and commencement of the soft
dorsal, the second the basal portion of the end of the soft dorsal. Caudal and anal fins
with similar ocellated spots; a brown band across the inner side of the root of the
pectoral. :

A single example of this beautiful species, 5 inches long, was obtained by Capt. Dow
at Panama. The typical specimen was obtained at the Galapagos Islands.

51, PoLYNEMUS MELANOPOMA,
Giunth. Proc. Zool. Soc. 1864, p, 148,
Ter a ES PA
Nine free pectoral appendages, the longest of which extends to the vent. Preoper-

422 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

culum finely serrated, with a small spine above the angle. The vomerine teeth form a
rounded patch; the band of the palatine teeth is as broad anteriorly as the front part
of the intermaxillary band. Operculum black.

A single specimen, 15 inches long, was obtained by Mr. Salvin at San José.

Description.—This fish is elongated in form, its greatest height being contained five
times and a half in the total length, with the caudal, and four times and one-sixth
without it. The tail is compressed, its height above the end of the anal fin being half
the length of the head. The head is much longer than high, and is contained four
times and two-thirds in the total length with the caudal, and thrice and one-third with-
out it; its width between the eyes is two-ninths of its length. Snout produced beyond
the mouth, obtusely conical, and shorter than the diameter of the eye, which is con-
tained five times and a half in the length of the head. The cleft of the mouth is
situated on the inferior side of the head, it is extremely wide, the maxillary being more
than half the length of the head. The posterior margin of the preoperculum is finely
serrated ; the angle is produced, forming a rounded membranaceous lobe. The posterior
margin of the opercular apparatus is membranaceous, rounded, and formed by the oper-
culum and suboperculum. ‘The origin of the first dorsal is in the vertical from the ninth
scale of the lateral line, or from a point about midway between the pectoral and ventral
fins. ‘The first spine is minute, the second is the strongest, all the others being flexible;
the third is the longest, contained once and two-thirds in the length of the head; the
fourth is longer than the second, and the following rapidly decrease in length. A series
of scales ascends behind the second, third, and fourth spines, but disappears at the fifth;
the distance between the two dorsals equals the length of the base of the second, which
is entirely covered with scales and has the upper edge strongly emarginate; the second
ray is the longest, nearly as high as the spinous dorsal, and twice the height of the last
ray. The distance between this fin and the caudal is one-fourth of the total length
(without caudal). ‘The caudal fin is completely covered with scales, deeply forked, with
the lobes pointed, the upper one being slightly the longer, and one-fourth of the total
length. The distance between the anal and caudal fins is less than that between the
caudal and dorsal, as the termination of the anal falls behind that of the dorsal, and in
the vertical from the 52nd scale of the lateral line. It is entirely covered with scales ;
and its origin corresponds to that of the seventh ray of the dorsal; its lower edge is
emarginate; the first spine is very small, the second being only one-third the length of
the first ray; the first and second rays are the longest, and about thrice the length of
the thirteenth or final ray, which, however, is rather longer than the one which pre-
cedes it. The pectoral is nearly one-sixth of the total length; its root is covered with
minute transparent scales. ‘The free pectoral appendages are long, the third and fourth
being the longest, considerably longer than the pectoral fin, and reaching to the vent;
the fourth is one-eighth of an inch jonger than the head. The root of the ventral fin
falls behind that of the pectoral, and in a vertical from the twelfth scale of the lateral

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 423

line; it is short, one-eighth of the total length, and does not quite reach the vent ; its
spine is about one-half the length of the adjacent ray. The scales are of moderate size,
longer than high, and have the posterior margin minutely crenulated. Lateral line
straight, very slightly bifurcated between the lobes of the caudal. The teeth are
minute and villiform, those of the yvomer form a rounded or nearly square patch; the
band on the palatines cuneiform and elongated, broadest anteriorly. The body is
uniform silvery, greenish grey, darker on the back; the fins are minutely dotted with
black, the dorsals becoming blackish at their margins. Operculum black.

52. PoLYNEMUS APPROXIMANS.

Polynemus approximans, Lay & Benn. in Beechey’s Voy. Zool. Fish. p. 57.
Trichidion approximans, Gill, Proc. Ac. Nat. Se. Philad. 1863, p. 169.

D.7\g Ag L. lat. 60.

15"

Six pectoral appendages, the longest of which reaches to the commencement of the
anal fin. The length of the caudal lobes is rather more than one-fourth of the total
length, Pectoral fins blackish.

Description.—This fish is moderately elongate in form; its greatest height, which is
between the root of the second dorsal and anal fins, is contained four times and one-third
in the total length with the caudal, and thrice and one-fourth in the same without
caudal. ‘The tail is compressed, its height above the end of the anal being contained
seven times and one-third in the total length. The head is much longer than high ; its
length is about four times and a half in the total with, and thrice and a half without
caudal; its width between the eyes is nearly one fourth of its length. The snout is
produced, obtusely conical, and shorter than the diameter of the eye, which is one-fifth
of the length of the head. The cleft of the mouth is situated at the inferior side of
the head, as usual; it is wide; the maxillary reaching considerably behind the orbit,
but the length of the bone is only two-fifths of that of the head. The posterior margin
of the preoperculum is armed with a distinct serrature, and one or two more distinct
teeth above the projecting membranaceous lobe of the angle. The posterior extremity
of the opercular apparatus is angular, membranaceous, and formed by the operculum
and suboperculum. ‘The origin of the first dorsal is opposite to the eighth scale of the
lateral line, and in the vertical between the roots of the pectoral and ventral fins. The
first spine is minute, the second shorter than the third, which is the longest, and con-
tained about once and one-third in the length of the head; the fourth is longer than the
second; and the subsequent spines rapidly decrease in length, rendering the upper
margin almost vertical. ‘There is a series of scales behind each spine almost to the top.
The distance between the two dorsals is more than the length of the base of the
second, which is entirely covered with scales and has the upper margin emarginate ;
the first and second rays are the longest, not so high as the spinous dorsal, more than
twice as long as the hindmost rays. The distance between this fin and the caudal is

424 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

one-fifth of the total length. The caudal fin is completely covered with scales, deeply
forked, with the lobes pointed, the upper one being rather the longer. The distance
between the caudal and anal fins is less than that between the dorsal and caudal, as the
extremity of the anal falls behind that of the dorsal, or in the vertical from the forty-
third scale of the lateral line. Its origin corresponds to that of the dorsal; and it has
the lower edge straight or very slightly emarginate; it is entirely covered with scales.
The first two spines are very small, and the third not half the length of the first ray ;
the first and second rays are the longest, but not twice as long as the fifteeenth or
terminal ray. The length of the pectoral is not one-fourth of the total; it has minute
scales towards the base. The free pectoral appendages are six in number; the upper
one is the longest, reaching to the anal fin, and is not quite one-third of the total
length. The root of the ventral falls a little behind the middle of the pectoral, and in
the vertical from the eleventh scale of the lateral line; it is short, one-eighth of the
total length, reaching to the vent; its spine is more than half the length of the
adjacent ray. ‘The scales are of moderate size, scarcely higher than long, and minutely
ciliated on the posterior margin. ‘The lateral line is straight, bifurcating between the
lobes of the caudal. Teeth on the vomer in a narrow transverse patch.

Two specimens, 12 inches long, are in the Collection, one found by Mr. Salvin at
Chiapam, the other by Capt. Dow at Panama.

Mr. Gill first recognized this species, which is not identical with P. xanthonemus, as
suggested in the ‘ Catal. of Fishes.’

53. PoLYNEMUS OPERCULARIS.
Trichidion opercularis, Gill, Proc. Acad. Nat. Sc, Philad. 1863, p. 169.

This fish is described thus :—

D.8\f. A.G L. lat. 69-70. L, transv, 8/14,

The greatest height equals a fourth of the length to the fork of the caudal fin, and
more than a fifth of the extreme, while the head enters four times and a half in the
latter. The outline from the dorsal to the snout is nearly rectilinear and little declined.
The distance of the anal from the outer axil of the ventral equals that of the posterior
nostril from the margin of operculum. The first dorsal, when bent backwards, rests on
the fourth scale, in front of the second. The second commences nearly above the
twentieth scale of the lateral line. The pectoral is as long as the head behind the
pupil. There are eight pectoral filaments, the longest of which extends rather beyond
the front of the second dorsal. The colour is greenish brown above and yellowish green
below. The operculum is blackish. The first dorsal and the pectorals, except below,
are also blackish, as is likewise the margin of the caudal. The anal is tinged with
orange. j

A single specimen, 11 inches long, was collected by Capt. Dow at Panama.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 425

54. LARIMUS BREVICEPS.
Larimus breviceps, Cuv. & Val. v. p. 146, pl. 111. Giinth. Fish. ii. p. 268.
Amblyscion argenteus, Gill, Proc. Ac. Nat. Sc. Philad. 1864, p. 165.
Having recently received a fine example of this fish from Panama through Capt. Dow,
I have convinced myself that the Pacific examples are not specifically, much less
generically, distinct from West-Indian ones.

06. MICROPOGON ALTIPINNIS.
Giinth. Proc. Zool. Soc. 1864, p. 149.
D.10|5. A.2/7. L. lat. 48-50. L. transy. 7/15.

The height of the body is contained thrice and two-thirds in the total length (without
caudal), the length of the head thrice and a half. The maxillary extends scarcely
beyond the vertical from the anterior margin of the eye. A series of five minute
barbels along each side of the mental groove. Two short, strong, divergent spines at
the angle of the preoperculum. The third and fourth dorsal spines are long, their
length being three-fifths of that of the head; anal spine of moderate strength, not quite
one-fourth of the length of the head. Nearly uniform silvery.

Two specimens were procured by Mr. Salvin—one, 17 inches long, at Chiapam, and
another, 14 inches long, at San José; a third specimen, 43 inches long, was found by
Capt. Dow at Panama: this agrees in every other respect with the older examples, but
of the minute barbels only a trace of the anterior (longest) pair is visible; so that it
appears that this generic (!) character is developed with age.

57. UmBRINA ELoNGATA. (PI. LXIV. fig. 2.)
Gunth. Proc. Zool. Soc. 1864, p. 148.
D.10|4 A. 1/7. L. lat. 70. L. transv. 7/22.

The height of the body is contained four times and a third in the total length (without
caudal), and five times if the caudal is included ; the length of the head is two-sevenths
of the total, or one-fourth if the caudal is included. The depth of the head is contained
once and three-fourths in its length. Snout long; the diameter of the eye is two-fifths
of the length of the snout, and one-fourth of the postorbital part of the head. Sym
physial barbel very short, as long as the posterior nostril. Preeoperculum without
distinct serrature. The length of the second dorsal spine is one-half of that of the
head. Posterior margin of the caudal f-shaped, the upper lobe being pointed, the
lower rounded; anal spine very feeble. The maxillary extends to the vertical from the
anterior margin of the orbit. Upper parts blackish, shining silvery, the lower white.

One specimen, 17 inches long, was found by Mr. Salvin at Chiapam.

VOL. VI.—PART VII. 3N

426 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

58. UMBRINA NASUS.
D10|5. A.1/8. L. lat: 54. L, transv. 6/14.

The height of the body is contained four times in the total length (without caudal),
the length of the head thrice and one-fourth. Snout much produced beyond the

mouth, which is quite at the lower side of the snout. ‘Ihe diameter of the eye is two-
thirds of the length of the snout, and two-fifths of that of the postorbital portion of
the head. Symphysial barbel very short, scarcely as long as the posterior nostril.
Preoperculum distinctly serrated. The second and third dorsal spines are as long as
the head, without snout. Posterior margin of the caudal fin f-shaped, the upper lobe
being pointed, the lower rounded; anal spine very feeble. ‘The maxillary extends to
below the centre of the orbit. Silvery, fins blackish.
One specimen, 10 inches long, was found by Capt. Dow at Panama.

99. UMBRINA ANALIS.
D. 10|J, A. 2/6. L. lat. 46-48. L. transy. 6/15.

The height of the body is one-third of the total length (without caudal), the length
of the head two-sevenths. Snout compressed, rather deep, of moderate extent, longer
than the eye, which is two-ninths of the length of the head, and equal to the width of
the interorbital space. Snout overlapping the mouth, but not much protruding beyond
it. Barbel very short, scarcely as long as the posterior nostril. Preeoperculum
distinctly serrated. The second and third dorsal spines are not quite as long as the
head without snout. Caudal fin subtruncate. Anal spine very strong, more than half
as long as the head. The maxillary extends beyond the front margin of the eye. An
oblique dark streak runs along each series of scales. The spinous dorsal fin blackish.

One specimen, 11 inches long, was found by Capt. Dow at Panama.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 427

I thought it possible for some time that this fish might be identical with Umbrina
undulata of Girard ; however, as this writer states that the anal spines of U. undulata
are feeble, and gives 1/9 for the number of anal rays, we are not justified in identifying
these two species.

61. CorvINA cHRYSOLEUCA. (Pl. LXVII. fig. 1.)

Allied to C. ronchus.

D. 10|s555, A. 5. L. lat. 55-56. L. transv. 35.

The height of the body is contained thrice in the total length (without caudal), the
length of the head thrice and one-third. Head thick ; snout obtuse, with the upper
jaw slightly overlapping the lower, as long as the diameter of the eye, which is con-
tained four times and two-thirds in the length of the head. The maxillary is nearly
entirely hidden by the preorbital, and extends beyond the vertical from the centre of
the orbit. ‘Teeth of the outer series of the upper jaw rather stronger than the others.
Interorbital space slightly convex, only one-third wider than the orbit, its width being
two-sevenths of the length of the head. Preeoperculum with spinous teeth round its
margin, three on and below the angle being much stronger than the others. Supra-
scapular denticulated. The second dorsal spine is the strongest, and the third the
longest, being as long as the postorbital portion of the head. ‘The second anal spine is
very strong, as long as the longest of the spinous dorsal, and not much shorter than the
first anal ray. Caudal fin irregularly rounded. Silvery, irregularly mottled with large
brownish patches shining golden. A young specimen (5 inches long) is more uniform
silvery.

Two specimens, 9 inches long, were collected by Capt. Dow at Panama.

I have observed in this species a most extraordinary variation in the size of the scales
above the lateral line, such as I do not recollect having seen in other Acanthopterygian
fishes. The two larger specimens are of nearly the same size; yet the dorsal scales of
one are only half the size of those of the other. The lateral line is composed of nearly
the same number of scales in both, and also the scales below the lateral are of nearly
the same size.

62. CorviNA veRMicuLaRIS. (Pl. LXVII. fig. 2.)
D. 10 = A, = L. transv. aa

The height of the body is a little more than one-third of the total length (without
caudal); the length of the head two-sevenths. Head moderately compressed, snout
obtuse, with the upper jaw overlapping the lower, a little longer than the diameter of
the eye, which is one-fifth of the length of the head. ‘The maxillary is entirely hidden
by the preorbital, and extends somewhat beyond the vertical from the centre of the
orbit. Teeth of the outer series of the upper jaw rather stronger than the others.
Interorbital space convex, only one-fourth wider than the orbit, its width being one-

3.N 2

428 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

fourth of the length of the head. Praeoperculum with spinous teeth round its margin ;
they are rather widely set and of equally small size. Suprascapular scarcely denticu-
lated. The second dorsal spine is scarcely stronger than, and but half as long as, the
third, the length of which exceeds somewhat that of the postorbital portion of the
head. ‘The second anal spine is very strong, rather shorter than the succeeding ray,
and equal in length to the postorbital portion of the head. Caudal fin rounded, with
the upper lobe slightly produced. Scales irregularly arranged. Purplish shining
silvery; a purplish brown streak, obliquely ascending backwards, follows the middle of
each series of scales. Fins brown.
A single specimen, 8 inches long, was found by Capt. Dow at Panama.

63. CoRVINA ARMATA.
Bairdiella armata, Gill, Proc. Acad. Nat. Se. Philad. 1863, p. 164.
This species, which is evidently allied to C. ronchus, is described thus :—
D.10\5. Aj L. lat. 51. L. transv. 7/10.

The height equals a fourth of the total length, of which the head forms a fourth.
The caudal fin equals the head behind the front margin of the eye. The diameter of
the eye enters four times and a half in the head’s length, somewhat exceeds the inter-
orbital area, which is scarcely convex, and equals the snout. ‘The fourth dorsal spine
is longest, and nearly equals half the head’s length; all are stout and robust. The
second dorsal commences nearly above the twentieth scale of the lateral line, or tip of
pectoral. The second anal spine is very strong, longer than the: first ray, and nearly
equals the interval between the front of orbit and opercular flap ; the soft fin behind is
incurved. The pectoral equals the interval between the middle of the pupil and the
opercular flap, and the ventral that between the front of the pupil and the same. The
colour is hoary above, silvery below; the fins yellowish; the vertical, especially the
first dorsal, clouded with darker.

Found by Capt. Dow at Panama.

64. CoRVINA OPHIOSCION.
Ophioscion typicus, Gill, Proc. Acad. Nat. Se. Philad. 1863, p. 164.
D. 10) 5555 AG. L. dat. 49. L, transv.

The height of the body is nearly equal to the length of the head, and two-sevenths of
the total (without caudal). Head rather low, snout obtuse, but prominent, with the
upper jaw projecting beyond the lower, the cleft of the mouth being quite at the lower
side of the snout. The diameter of the eye equals the extent of-the snout, and is two-
ninths of the length of the head. The maxillary is entirely hidden by the preorbital,
and extends to below the middle of the orbit. Teeth of the outer series of the upper
jaw rather stronger than the others. Interorbital space scarcely convex, only one-third

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 429

wider than the orbit, its width being two-sevenths of the length of the head. Pre-
operculum with straight, widely-set, spinous teeth round its margin, those on or near
the angle being slightly the strongest. ‘The second dorsal spine is the strongest, the
third the longest, its length being more than that of the postorbital portion of the
head. The second anal spine is exceedingly strong, about as long as the third dorsal
spine, or as the first anal ray. Caudal fin irregularly rounded. Uniform silvery; top
of the spinous dorsal blackish.

This species appears to be scarce at Panama, Capt. Dow having collected only two
examples, of 8 and 6 inches in length.

65, OTOLITHUS SQUAMIPINNIS.
DS op Aaa Lalat, 88s

Scales rather irregularly arranged; there are nine longitudinal series between the
origin of the first dorsal fin and the lateral line, and five or six between the end of the
second dorsal fin and the lateral line. The height of the body is contained four times
and one-sixth in the total length (without caudal), the length of the head thrice and
one-fourth. Lower jaw very prominent, the extent of the snout being contained thrice
and one-third in the length of the head. ‘The width of the interorbital space is more
than the diameter of the eye, and equals the extent of the upper jaw from the orbit.
The maxillary extends to the vertical from the hind margin of the orbit. Praopercular
angle slightly produced, dilated into a membranaceous margin which is faintly striated.
The spinous dorsal is longer than high ; the spines are feeble, the length of the third
being two-fifths of that of the head. Caudal fin rounded, the middle rays being the
longest. ‘The membrane of the soft dorsal and anal fins is covered with small, transpa-
rent scales, which form a thickish cover on the base of these fins. The length of the
pectoral is three-fifths of that of the head. Body uniformly coloured, scales on the
sides minutely punctulated with brown ; hinder side of the axil of the pectoral brown.
Inner membrane of the gill-cover black. Ventral yellowish.

Two specimens, 10 & 11 inches long, were collected by Capt. Dow at Panama.

66. OTOLITHUS ALBUS.
Gunth. Proc. Zool. Soc. 1864, p. 149.
D. 10) 57. A. 2/9.

Scales rather irregularly arranged ; there are seven series between the origin of the
dorsal fin and the lateral line. The height of the body is one-fourth of the total length
(without caudal), the length of the head two-sevenths. The extent of the snout is one-
fourth of the length of the head; the maxillary extends somewhat beyond the vertical
from the posterior margin of the eye. Preopercular angle not produced behind. ‘The
spinous dorsal is much longer than high ; its spines are feeble, the length of the fourth

450 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

being two-fifths of that of the head. Caudal fin rounded, with the middle rays pro-
duced. ‘The second anal spine is truly spinous, not flexible, two-fifths of the length of
the first soft ray. Membrane of the dorsal and anal fins not scaly. The pectoral fin
extends as far backwards as the ventral, being more than half as long as the head.
Immaculate, silvery, back greenish. (Pseudobranchiz present.)

One specimen, 14} inches long, was obtained by Mr. Salvin at Chiapam.

67. OTOLITHUS RETICULATUS.
Giinth. Proc. Zool. Soc. 1864, p. 149.
D.10| sm. A. 11(2/9).

Closely allied to 0. carolinensis. Scales rather irregularly arranged; there are nine
series between the origin of the dorsal fin and the lateral line. The height of the body
is contained four times and a third in the total length (without caudal); the length of
the head thrice anda third. The extent of the snout is two-sevenths of the length of
the head; the maxillary does not extend backwards to the vertical from the posterior
margin of the eye; praopercular angle somewhat produced behind, membranaceous,
striated; the posterior margin of the preeoperculum obliquely descending backwards.
The spinous dorsal is much longer than high ; its spines are feeble, the fourth being
the longest, two-fifths of the length of the head. Caudal fin subtruncated, the middle
rays somewhat produced. ‘The first anal ray is quite rudimentary; the second as long
as the eye, flexible, scarcely spinous. The pectoral fin extends as far backwards as the
ventral, being more than half as long as the head. Back and sides with an irregular
network of brown undulated streaks ; fins immaculate.

Two specimens were collected by Mr. Salvin—one, 15 inches long, at San José, the
other, 13 inches long, at Chiapam.

71. CaRaNxX LEUCURUS.
Giinth. Proce. Zool. Soc. 1864, p. 24.
Very closely allied to C. bicolor.
158) Aa
The first dorsal fin is composed of short, stoutish spines, the fourth of which is the
longest, but scarcely longer than the eye. The soft dorsal and anal are rather elevated ;
the caudal is emarginate, and has the lobes rounded. ‘Teeth very small, forming a
single series in both jaws; palate smooth. The height of the body is one-half of the
total length (without caudal), the length of the head one-third. Snout rather obtuse,
the jaws being equal in front when the mouth is closed; the maxillary extends to
below the anterior margin of the orbit. The lateral line makes anteriorly a subsemi-
circular curve, the width of which is contained from once and two-thirds to once and
four-fifths in the length of the straight portion ; it becomes straight behind the vertical

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 451

from the origin of the second dorsal, and is armed with about fifty small and low
shields, only a few of which terminate in a depressed spine. The pectoral fin extends to
the anal spines. Brownish grey, body with six dark brown vertical bands; the first
crosses the body behind the base of the pectoral, and the fourth descends from the
middle of the soft dorsal fin. Operculum with a large black spot. Dorsal, anal, and
ventral black ; pectoral and caudal whitish.

Only two examples, three inches long, were found by Capt. Dow at Panama.

72. Caranx speciosus (Forsk.).
Having examined specimens from Panama, collected by Mr. Salvin, and compared
them with others from Borneo, Madras, Zanzibar, &c., I have convinced myself that C.
panamensis, Gill, Proc. Acad. Nat. Se. Philad. 1863, p. 166, is identical with C. speciosus.

74. Caranx uippos, L.

We have received two examples from the Pacific coast of Panama from Capt. Dow.
‘The younger one, which is 5 inches long, agrees in every point, especially in the height
of the body, with Atlantic examples of this species, whilst the older, 10 inches, is
identical with that remarkable form described by Mr. Gill as Carangus marginatus
(Proc. Acad. Nat. Se. Philad. 1863, p. 166). This is considerably lower in form than
the type, the height of the body being only two-sevenths of the total length; but having
had an opportunity of comparing the example first mentioned, I do not think it entitled
to specific rank, but regard it merely as a variety. The formula of fin rays in our

example is D. 7 = A. 2 lz

»

75. CARANX CABALLUS.
‘rachurus boops, Girard, U. S. Pac. R. R. Exped. Fish. p. 108; Giinth. Fish. nu. p. 422 (not
C. boops, C. & V.).
Tai eee | ee baer
The teeth of the upper jaw form a villiform band, those of the outer series being a
little the larger; those of the lower are in a single series; teeth on the vomer, the
palatines, and the tongue. The height of the body is two-sevenths of the total length
(without caudal), the length of the head rather more than one-fourth. Eye with a
broad adipose membrane in front and behind. Breast scaly. ‘The lateral line is curved,
the width of the arch being one-half of the length of the straight portion ; the latter
commences in the vertical from the third dorsal ray; the plates commence from the
beginning of the straight portion of the lateral line, and are well developed. Lower
jaw projecting beyond the upper; maxillary extending to below the anterior rim of the
pupil. Pectoral reaching beyond the anterior anal rays. A black opercular spot.
Two specimens were collected by Capt. Dow at Panama; the species extends north-
wards to the coast of California.

432 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

76. CARANX CANINUS.
Dis | Are |e Lat 4:

The teeth in the upper jaw form a villiform band, those of the outer series being
much the stronger, and widely set. Lower jaw with a single series of rather strong,
closely set teeth, and with the two anterior ones somewhat enlarged, canine-like ; teeth
on the vomer, the palatines, and the tongue. The height of the body is a little more
than the length of the head, and one-third of the total (without caudal). Snout
obtuse, as long as the diameter of the eye; eye with an adipose eyelid in front and
behind. Preorbital much narrower than the orbit. The maxillary extends beyond
the vertical from the centre of the eye. Breast naked; lateral line slightly bent, the
width of the arch being contained once and one-third in the length of the straight
portion; the latter commences in the vertical from the fifth dorsal ray; the plates do
not reach forward to the end of the arched portion, and are well developed. Lower
jaw scarcely projecting beyond the upper. Dorsal spines rather stont and short; the
fourth is the longest, and one-third of the length of the head. The pectoral extends to
the fifth anal ray. A black opercular spot. Membrane of the soft dorsal and anal
blackish.

One specimen, 7} inches long, was discovered by Capt. Dow at Panama.

77. CARANX DORSALIS.
Carangoides dorsalis, Gill, Proc. Acad. Nat. Se. Philad. 1863, p. 166.
D.4-5|7. A.2|7g L. Jat. 25*.

The teeth in both jaws form villiform bands; teeth on the vomer, the palatine bones,
and on the tongue. The height of the body is contained once and four-fifths in the
total length (without caudal), the length of the head thrice and one-fourth. The first
dorsal fin is but little developed, the spines being short, feeble, and flexible. Anterior
rays of the dorsal and anal fins prolonged into a very long filament, sometimes longer
than the whole body. Jaws equal in length, the maxillary extends to the vertical from
the front margin of the orbit. Lateral line bent, the width of its arch being as long as
the straight portion; the latter commences below the middle of the second dorsal fin.
The plates are moderately developed, and commence at some distance from the bend of
the lateral line. Gill-membrane above the pectoral blackish; posterior half of the
ventrals black.

Panama. We have received two examples from Capt. Dow, one 19 inches long.

* Mr. Gill counted 44; this is either a mistake, or he has counted small scales not deserving the name of

plates.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 433,

82. CHORINEMUS ALTUS,
Dye ae
The height of the body is contained thrice in the total length (without caudal), the
length of the head four times and one-fourth.
Eye rather large, its diameter being equal to
the length of the snout, and one-fourth of
that of the head. Lower jaw projecting be-
yond the upper. “Maxillary very narrow
posteriorly, scarcely extending to the ver-
tical from the hind margin of the eye; the
length of the intermaxillary is contained
once and three-fourths in the length of the
head. The infraorbital, situated above the
maxillary, is as broad as the bone next above
it; none of these bones reach to the anterior
preopercular ridge. Pectoral fin longer than
the ventral, nearly as long as the head (with-
out snout). Coloration uniform.
One example, 11 inches long, has been recently sent by Capt. Dow from Panama.

83, CHORINEMUS INORNATUS.
Oligoplites inornatus, Gill, Proc. Acad. Nat. Sc. Philad. 1863, p. 166.
D: 5|a° A. 2 lap.

“The height of the body enters four times and two-thirds in the total length; the
length of the head five times and two-thirds. The upper maxillary reaches nearly to
the vertical from the hinder margin of the orbit; the intermaxillary enters twice and
one-third in the head’s length. The snout is a little longer than the diameter of the
eye; the latter equals a quarter of the head’s length. ‘The infraorbital bones do not
extend to the preoperculum; the one above the maxillary bones is wider than the one
above itself, and as wide as that behind the eye. ‘The opercular apparatus is vertical in
front of the pectoral, and for an equal space above. The preoperculum is nearly
vertical, and its angle obliquely rounded. The width of the operculum and suboper-
culum in front of the lower axilla of the pectoral equals the diameter of the eye and
the interval between suboperculum and axil. ‘lhe pectoral equals the interval between
its axis and the hinder border of the pupil; the ventral is rather shorter, but its end
almost or quite reaches to the anus. The colour is uniform, tinged with blue above.”

One adult specimen was collected by Capt. Dow on the Pacific coast. of Central
America.

VOL. VI.—PART VII. 30

434 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

85. TRAcHYNoTUS FasclATUS. (Plate LXIX. fig. 4.)
Trachynotus fasciatus, Gill, Proc. Ac. Nat. Se. Philad. 1863, p. 86.
glaucoides, Giinth, Proc. Zool. Soc. 1864, p. 150.
D. 6p. A. le

Closely allied to 7. glaucus, but with the body more elevated. The height of the
body is one-half of the total length (without caudal); the length of the head two-
sevenths. The maxillary extends to below the middle of the eye. Anterior dorsal and
anterior anal rays, and the caudal lobes, much prolonged, the length of the latter being
two-sevenths of the total. The ventral fin does not extend to the vent. Five narrow
blackish vertical bars across the lateral line. ‘

One specimen, 7 inches long, was obtained by Mr. Salvin at San José; two others,
11 inches long, were obtained by Capt. Dow at Panama.

Description.—This species has the body (without caudal) of a rhomboidal form, its
greatest height being between the last spine of the dorsal and the vent, and one-half of
the total length (without caudal) ; the upper profile between the dorsal and the snout is
oblique, feebly convex over the eye. ‘The length of the head is contained thrice and one-
half in the total (without caudal). ‘The diameter of the eye is rather more than the length
of the snout, and contained thrice and two-thirds in that of the head. The cleft of the
mouth is narrow; the maxillary reaches nearly to the level of the centre of the diameter
of the eye; its length is a little more than one-third of that of the head. The width of
the space between the eyes is more than one-third of the length of the head, or equal to
the distance from the tip of the snout to the centre of the eye. Praoperculum with
the hinder margin straight, and at a right angle with the lower border, which is also
straight and parallel with the axis of the body. Operculum small, narrow, about two-
thirds as long as high; the hinder border of the opercular apparatus is formed almost

entirely of the sub- and interoperculum ; it is rounded and membranaceous: the line of
the separation between the operculum and suboperculum is at right angles with that
between the sub- and interoperculum. There is a recumbent spine before the com-
mencement of the first dorsal, and in a line with the posterior part of the axil of the
pectoral ; the dorsal spines, seven in number, are short; the first is minute, but erect,
and not attached by any apparent membrane to the second; the others show a slight
progression in dimensions, and are united by a low membrane. ‘The base of the soft
dorsal is not twice as long as that of the spinous; the first two rays, which are the
longest, project considerably beyond any of the others, and are equal to half the length
of the body (without the caudal); the following rays diminish very rapidly in length,
and from the eighth ray to the last the fin is scarcely higher than the spinous dorsal,
and its upper edge almost straight. The distance between the dorsal fin and the
caudal is equal to that between the anal and caudal. ‘The anal fin is preceded by three
short spines about equal to the fourth, fifth, and sixth of the dorsal. The base of the

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 435

soft portion of the anal is about as long as that of the soft dorsal: it consists of
eighteen rays, and perfectly resembles the soft dorsal in shape, having the first two rays
much longer than the others, equal to the corresponding rays in the dorsal, and the
following rays rapidly decreasing in length to the sixth, from which the margin of the
fin is almost straight. The pectoral fin is pointed, of moderate size, its length being
three-fourths that of the head. The ventrals are short, more than half the length of
the pectoral, and not reaching to the vent. The tail behind the dorsal and anal is
compressed and narrow. The caudal is deeply forked; the lobes are equal, and con-
tained thrice and a half in the total length; it is covered with small scales.

The body is covered with very minute scales ; those at the base of the vertical fins and
near the lateral line are a little larger. The head and opercular bones are entirely
naked. The lateral line shows a somewhat irregular sinuosity slightly above the median
axis of the body for the first half of its length, after which it is perfectly straight, termi-
nating between the two lobes of the caudal. Teeth small, villiform; a small central
patch on the vomer, and a narrow one on each of the palatines.

Bluish green above, silvery beneath. Five vertical brown stripes down the sides of
the body across the lateral line, the first two being nearer together than the others,
which are at almost equal distances: the first behind the axil of the pectoral, the
second below the third dorsal spine, the third below the sixth, the fourth below the
seventh dorsal ray, and the fifth below the seventeenth. However, the second and
third of these bands are placed sometimes more backwards, which is evidently the case
in the example described by Mr. Gill, and named by him 7. fasciatus. Having recently
obtained two examples from Capt. Dow, one of which shows the arrangement of the
bands as in 7. glaucoides, on one side, and that of 7. fasciatus on the other, I cannot
entertain any doubt as to the specific identity of these fishes.

86. PrLamys sarpa, BI.

We may mention this species here, although it is not contained in any of the collec-
tions forming the material for this Memoir, because Dr. Ayres alludes to it in the following
manner :—* A species of Pelamys brought to the markets of San Francisco is without ques-
tion the P. sarda. The closest examination fails to distinguish it from the Atlantic form.
Previous to this time we had no positive knowledge of any fish in the low latitudes which
inhabits Californian waters and those of the Atlantic.”—Proc. Calif. Acad. 1855, p. 74.

90. Barracnus pactrict (Gthr.).

In other specimens recently collected by Capt. Dow at Panama, I find the membrane
at the bottom of the pouch of the axil of the pectoral fin (described in Fish. iii. p. 173)
folded and wrinkled, with a great quantity of coagulated mucus between the folds.
The same species appears to occur also on the coast of West Africa, a specimen having
been lately obtained by Dr. Steindachner, who describes it as B. liberiensis (Sitzgsber.
Ak. Wiss. Wien, 1867, lv. p. 525, Taf. 1. figs. 2 & 3).

5302

436 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

THALASSOPHRYNE'.
Thalassophryne, Giinth. Fish. iii. p. 174.

Head broad, depressed ; body subcylindrical anteriorly, and compressed posteriorly ;
skin naked. Canine teeth none. Operculum with a single spine. The spinous dorsal
formed by two spines of moderate length. ‘The opercular and dorsal spines with a
canal conducting a poisonous fluid from a sac situated at their base. Gill-opening not
very narrow, not extending to the isthmus.

Atlantic coasts of Tropical America.

92. THALASSOPHRYNE MACULOSA. (Plate LX VIII. fig. 1.)
Ginth. Fish. i. p. 175.
Dy) 19. A. V8. Ve TZ:

Brown, marbled with darker; some round black spots on the pectoral and the side
of the body.

The general habitus is that of a Batrachus. The head is somewhat longer than
broad, its length being contained thrice and one-third in the total; it is moderately
depressed. The snout is short, obtuse, with the cleft of the mouth ascending obliquely
upwards, and with the chin prominent. The maxillary extends to the vertical from the
posterior margin of the orbit. . The teeth are obtusely conical, standing in single series,
except anteriorly in the lower jaw, where they form two series, and in the upper, where
they are cardiform, in a narrow band. ‘The eyes are directed upwards and very small,
their width being one-half of that of the bony bridge between the orbits. Gill-covers
with a single spine; it is long, slender, cylindrical, like one of the dorsal spines, and
has the operculum for its base. Gill-opening not very narrow; it extends from the
upper base of the pectoral obliquely downwards and forwards to the level of the inferior
base of the pectoral. The two dorsal spines are slender, pungent, about one-third of
the length of the head. Dorsal and anal fins terminate immediately before the root of
the caudal, the length of which is one-seventh of the total. Pectoral obliquely rounded,
extending to the origin of the anal; ventral rather short, not quite one-half the length
of the head, extending to the base of the pectoral. Skin perfectly smooth, with some
very short tentacles at the lower jaw. Two short horizontal muciferous channels on
the cheek and the lateral line are very distinct; they are not, as usually, composed of
a series of distant pores, but the pores are confluent, forming one continuous groove of
a white colour. Other muciferous channels, as for instance along the base of the anal,
are composed of separate indistinct pores. Colour brown, marbled with darker ;
pectoral fins and sides of the body with some round black spots; chin and ventrals
brownish ; belly white.

A single specimen from Puerto Cabello is known.

2 Greek denomination for Sea-toad.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 437

lines

Win falelentoblig $54 ccs avaceoraeactne sare as ndstceiaokgcseteer 54
dient of tienhends yeh sccasmee ny sated sSac 6 spicisnisto ae 16
Wad thi ofeth eshead ins csas quench ot sn ete sceio tenes wsilsiastie 14
Weptheotithe sh cad rrs scat pecs teas sea edeolasalaxsiese cs 10
Miameterioftheve ye). scctctins fe citsel ost aceieriesh oabscie skemeise 1
Wenothvotethe eau dalle tile aeeeeaactsmeseeeeeereae imei. 8
oletheyventrall fim is. teeneus eta k wanes ase atecist 7

93. THALASSOPHRYNE RETICULATA. (Plate LXVIII. fig. 2.)
Giinth. Proc. Zool. Soc. 1864, pp. 150, 155.
Waa|24er BAS 2a OV ad/22. Bal6;

The length of the head is two-sevenths of the total length (without caudal). ‘The
teeth on the palate are in a single series, very short, obtuse, incisor-like. Pectoral
very large, extending backwards to the sixth anal ray. Head, body, and fins brown.
with a network of yellowish lines; vertical and pectoral fins with a white margin.

In other respects this species agrees with 7. maculosa; so that we may refer to the
description of that species given above. .

Three specimens were found by Messrs. Dow and Salvin on the Pacific coast of
Panama : the largest is 13 inches long.

In this species I first observed and closely examined the poison-organ with which
the fishes of this genus are provided. Its structure is as follows :—

1. The opercular part.—The operculum is very narrow, vertically styliform, and very
mobile; it is armed behind with a spine, 8 lines long in a specimen of 103 inches, and
of the same form as the venom-fang of a snake; it is, however, somewhat less curved,
being only slightly bent upwards; it has a longish slit at the outer side of its extremity,
which leads into a canal perfectly closed, and running along the whole length of its

Fig. 1. Hinder half of the head, with the
venom-sac of the opercular apparatus in _
situ. * Place where the small opening
in the sac has been observed. a. La-
teral line and its branches. 6. Gill-
opening. c¢. Ventral fin. d. Base of
pectoral fin. e. Base of dorsal fin.

Fig. 2. Operculum, with the perforated
spine.

interior; a bristle introduced into the canal reappears through another opening at the

438 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

base of the spine, entering into a sac situated on the opercle and along the basal half
of the spine; the sac is of an oblong-ovate shape, and about double the size of an oat-
erain, Though the specimen had been ‘preserved in spirits for about nine months, it
still contained a whitish substance of the consistency of thick cream, which on the
slightest pressure freely flowed from the open:ng in the extremity of the spine. On
the other hand, the sac could be easily filled with air or fluid from the foramen of the
spine.

No gland could be discovered in the immediate neighbourhood of the sac; but on a
more careful inspection I found a minute tube floating free in the sac, whilst on the
left-hand side there is only a small opening instead of the tube. The attempts to
introduce a bristle into this opening for any distance failed, as it appears to lead into
the interior of the basal portion of the operculum, to which the sac firmly adheres at
this spot.

2. The dorsal part is composed of the two dorsal spines, each of which is 10 lines
long, The whole arrangement is the same as in the opercular spines; their slit is at
the front side of the point; each has a separate sac, which occupies the front of the
basal portion; the contents were the same as in the opercular sacs, but in somewhat
greater quantity. A strong branch of the lateral line ascends to the immediate
neighbourhood of their base.

Thus we have four poison-spines, each with a sac at its base; the walls of the sacs
are thin, composed of a fibrous membrane, the interior of which is coated over with
mucosa. ‘There are no secretory glands imbedded between these membranes, and these
sacs are probably merely the reservoirs in which the fluid secreted accumulates. ‘The
absence of a secretory organ in the immediate neighbourhood of the reservoirs (an organ
the size of which would be in accordance with the quantity of the fluid secreted), the
diversity of the osseous spines which have been modified into poison-organs, and the
actual communication indicated by the foramen in the sac, lead me to the opinion that
the organ of secretion is either that system of muciferous channels which is found in
nearly the whole class of fishes, and the secretion of which has poisonous qualities in a
few of them, or at least an independent portion of it.

This description was made from the first example; through the kindness of Capt. Dow
I received two other specimens; and in the hope of proving the connexion of the poison-
bags with the lateral-line system, Tasked Dr. Pettigrew, of the Royal College of Surgeons,
a gentleman whose great skill has enriched that collection with a series of the most
admirable anatomical preparations, to lend me his assistance in injecting the canals.
The injection of the bags through the opening of the spine was easily accomplished ;
but we failed to drive the fluid beyond the bag, or to fill with it any other part of
the system of muciferous channels. This, however, does not disprove the connexion of
the poison-bags with that system, inasmuch as it became apparent that, if’there be
minute openings, they are so contracted by the action of the spirit in which the speci-

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 439

mens were preserved, as to be impassable to the fluid of injection. A great part of the
lateral-line system consists of open canals; however, on some parts of the body, these
canals are entirely covered by the skin: thus, for instance, the open lateral line ceases
apparently in the suprascapular region, being continued again in the parietal region.
We could not discover any trace of an opening by which the open canal leads to below
the skin; yet we could distinctly trace the existence of the continuation of the canal
by a depressed line, so that it is quite evident that such openings do exist, although
they may be passable only in fresh specimens. ‘Thus, likewise, the existence of openings
in the bags, as I believe to have found in the first specimen dissected, may be proved by
examination of fresh examples.

The sacs are without an external muscular layer, and situated immediately below the
loose thick skin which envelopes their spines to their extremity; the ejection of the
poison into a living animal, therefore, can only be effected by the pressure to which the
sac is subjected the moment the spine enters another body.

Nobody will suppose that a complicated apparatus like the one described can be
intended for conveying an innocuous substance; and therefore I have not hesitated to
designate it as poisonous; and, Capt. Dow informs me in a letter lately received, “ the
natives of Panama seemed quite familiar with the existence of the spines and of the
emission from them of a poison which, when introduced into a wound, caused fever, an
effect somewhat similar to that produced by the sting of a Scorpion; but in no case wags
a wound caused by one of them known to result seriously. The slightest pressure of
the finger at the base of the spine caused the poison to jet a foot or more from the
opening of the spine.” The greatest importance must be attached to this fact, imas-
much as it assists us in our inquiries into the nature of the functions of the muciferous
system, the idea of its being a secretory organ having lately been superseded by the
notion that it serves merely as a stratum for the distribution of peripheric nerves.
Also the objection that the Sting-Rays and many Siluroid fishes are not poisonous,
because they have no poison-organ, cannot be maintained, although the organs conveying
their poison are neither so well adapted for this purpose nor in such a perfect connexion
with the secretory mucous system as in Thalassophryne.

The poison-organ serves merely as a weapon of defence. All the Batrachoids with
obtuse teeth on the palate and in the lower jaw feed on Mollusca and Crustaceans.

95, ANTENNARIUS LEOPARDINUS. (Plate LXIX. fig. 3.)
Giinth. Proc. Zool. Soc. 1864, p. 151.
IDE BGS ATG dee IU
Skin very rough, covered with minute spines; anterior dorsal spine (tentacle) not
longer than the second, terminating in a small, flat disk ; the third is separate from the
soft dorsal. . Brownish grey, marbled with rose-colour, and with brown dots on the

440 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

sides ; a black ocellus edged with rosy in the middle of the side, another larger one on
the base of the ninth and tenth dorsal rays, and one or two small ones on the side of
the tail. Belly covered with round brown spots; caudal with ovate black spots,
arranged in three transverse series ; all the other fins with similar spots.

One specimen, 24 inches long, was found by Capt. Dow on the Pacific coast of Panama.

96. ANTENNARIUS TENUIFILIS.
DPD, 3/12. <A. 7. P-. 10:

Allied to A. digibbus. Skin rough, the spines being exceedingly fine; anterior dorsal
spine (tentacle) much longer than the second, and tapering into a fine point; second
quite free, conical; third entirely covered by the skin, forming a slight protuberance.
Rose-coloured, with black markings which are most crowded and confluent on the
middle parts of the length of the fish, less so on the head and thorax, leaving the nape
and back of the trunk nearly immaculate; the markings form irregular concentric
streaks on the thorax, and larger patches on the body; a deep-black band across the
caudal and anal fins.

One specimen, 23 inches long, was found by Dr. Seemann, walking on the reefs outside
the city of Panama.

103. Exxorris Macunata (BL.).

This species attains to a large size; Mr. Salvin collected specimens 11 inches long
at Huamuchal. Such large examples have, of course, the eye comparatively smaller
than younger ones; and having also a deeper body, the number of series of scales
between the origin of the second dorsal and the anal is increased by the addition of
smaller scales. On such a large example Mr. Gill has founded his Dormitator microph-
thalmus, Proc. Ac. Nat. Sc. Philad. 1865, p. 170.

106. ELEOTRIS LONGICEPS.
Giinth. Proc. Zool. Soc. 1864, p. 151.
1.6 |59:. An aga 0 Le: lat, 66.

Vomerine teeth in a broad subcrescentic band, which is more than half as broad as
that of the intermaxillaries. Thirty-six series of scales between the occiput and the
anterior dorsal fin; twenty between the origin of the posterior and the anal. The
height of the body is nearly one-half of the length of the head, which is more than one-
third of the total (without caudal). ‘The maxillary extends to below the middle of the
eye; teeth cardiform. Caudal fin obtusely rounded, one-sixth of the total length.
Brownish black, marbled with brown and black; fins with roundish blackish spots.

This species differs from the others (which have been referred to the division of
Philypnus) in having a comparatively longer head. One specimen, 8 inches long, was
given to Mr. Salvin by Capt. Dow, who found it in the Lake of Nicaragua.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 441

107, ELEorris PICTA.
Kner, Sitzgsber. bayer. Ak. Wiss. 1863, p. 223, and Abhandl. bayer. Ak. Wiss. 1865, p.18, tab. 3. fig.1.
Di 6\25. Al5e? I. lat. 60.

This fish is closely allied to E. gyrinus, but is said to be distinguished by having the
height of the body only one-sixth or one-seventh of the total length’; lower parts of
the body with numerous whitish spots and streaks.

From the Pacific side of the Isthmus.

108. ELEOTRIS SEMINUDA.
Giinth. Proc. Zool. Soc. 1864, p. 24.
1 Diol [SUlais Bee ate

The head and the trunk are naked; the tail is covered with small scales; head
depressed, broader than high, flat above, its length being two-sevenths of the total.
Snout rather obtuse, longer than the eye, with the lower jaw somewhat prominent; the
cleft of the mouth extends to below the anterior margin of the orbit. Teeth in the
upper jaw in a narrow band; the lower has four somewhat larger and recurved teeth in
front; they appear to form a single series; palate toothless. None of the fin-rays are
prolonged; the pectoral does not quite extend to the origin of the second dorsal ;
ventral much shorter than pectoral, its inner ray is the longest, the others gradually
decreasing in length outwards; caudal fin rounded. Brown, with numerous well-
defined white cross stripes on the head as well as on the body; vertical fins black.

Although there is only a single example, 20 lines long, in the collection, the charac-
ters of this species are so well marked that I have not hesitated to describe it.

109. AMBLYOPUS BREVIS.

Giinth. Proc. Zool. Soc. 1864, p. 151.
De Gilo, As Ts.

The height of the body is one-eighth of the total length (without caudal); the length
of the head two-ninths. Eyes minute. Jaws with a series of longish, widely set teeth.
Caudal fin black.

One specimen, 3 inches long, was found on the Pacific coast of Panama by Messrs.
Dow and Salvin.

The specimen being young, I abstain from giving a detailed description. In its
dentition it agrees with A. sagitta from California, and therefore it would belong to the
subgenus Tyntlastes (Proc. Zool. Soc. 1862, p. 194), The scales must have been very
thin and deciduous, and do not appear to be very small, at least not on the hinder part
of the body. The ventral is much longer than the pectoral, aud the caudal longer than

the head.
1 According to the figure it is higher.

VOL. VI.—PART VII. 3P

449 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

114. Crinus MacrocepHaLus. (Plate LXIX. fig. 2.)
Giunth. Fish. ii. p. 266.
Dy oe Age at C122 BL 19. 1iVe Ys:

The height of the body is contained seven times and a half in the total length, the
length of the head five times. The head is depressed, rather short, nearly as broad as
long; crown of the head broad and flat ; interorbital space concave, narrower than the
orbit. Snout very short, obtuse, rounded; the maxillary does not extend to behind
the posterior margin of the orbit; lips thick. The teeth in the jaws form a band with

an outer series of stronger ones; vomerine teeth in a narrow band; palatine teeth
none. No orbital tentacles, those at the nostril and on the neck very small. Gill-
openings wide, the gill-membranes being united at the throat.. Head naked; scales on
the body not very small, cycloid. The dorsal fin commences at the occiput, and termi-
nates near the base of the caudal; the spines are flexible, and much lower than the
soft rays; the three anterior ones are rather more remote from one another than the
following: none of the rays of this or of the other fins are branched. Caudal rounded.
The anal is higher posteriorly than anteriorly, about as high as the spinous dorsal.
Pectoral rounded, with the middle rays longest, shorter than the head. Ventrals
jugular, half as long as the pectoral, with the spine and the outer ray enveloped in a
common thick membrane. Dark greyish olive; head and fins blackish; head, base of
the pectoral, anterior part of the body, and dorsal dotted with white.
Several examples, of the size of the figure, were collected by Capt. Dow at Panama.

CREMNOBATES.
Cremnobates, Giinth. Proc. Zool. Soc. 1861, p. 374.

Body moderately elongate, with the scales small or of moderate size. Snout rather
short, with the cleft of the mouth of moderate width. A band of small teeth in the
jaws; teeth on the vomer. ‘Two separate dorsals, composed of spines only; the
anterior short, formed of three spines. Ventrals jugular, composed of three rays.
Head with tentacles ; gill-opening wide.

115. CREMNOBATES MONOPHTHALMUS. (Plate LXIX. fig. 1.)
Auchenopterus monophthalmus, Giinth. Fish. ui. p. 275.
Cremnobates monophthalmus, Giinth. Proc. Zool. Soc. 1861, p. 374.
D. 3/27. A. V.B. 1. lat. 38.

A fimbriated superciliary tentacle; a small one at the nostril and on each side of
the nape, both multifid. A black ocellus, edged with white, on the posterior quarter
of the dorsal fin.

Specimens, 2 inches long, were collected by Capt. Dow at Panama.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 448

118. ATHERINICHTHYS PACHYLEPIS.
Giinth. Proc. Zool. Soc. 1864, p. 25.
D. 4 a A. = L. lat. 41. L. transv. 7.

The height of the body is nearly equal to the length of the head, and contained five
times and a half or five times and a third in the total length (without caudal). The
snout is short, not longer than the diameter of the eye ; and the cleft of the mouth does
not extend backwards to below the anterior margin of the eye. The anterior dorsal is
composed of short, feeble spines; and its origin is opposite to the fourth or fifth anal
ray. The pectoral fin is much longer than the head. The silvery streak occupies the
adjoining halves of the third and fourth series of scales.

Two specimens, 6 inches long, were collected by Capt. Dow at Panama.

119. ATHERINICHTHYS GUATEMALENSIS.
Giinth. Proc. Zool. Soc. 1864, p. 151.
DA eee ee lee lte 36.0 Ue, trenay a7.

Anterior dorsal fin very small, inserted behind the vertical from the commencement
of the anal fin. The height of the body is contained five times in the total length (with-
out caudal), the length of the head four times and a fourth. ‘The silvery band occupies
the third upper series of scales. The lower caudal lobe rather longer than the upper.

Several examples, from 2 to 24 inches long, were collected by Mr. Salvin in the Lakes
of Huamuchal.

120. Mucit BRASILIENSIS (Agass. ).

Messrs. Dow, Godman, and Salvin have collected numerous examples of Te sizes at
Belize, Chiapam, and Panama. I have no doubt that MZ. giimtheri, Gill, Proc. Ac. Nat.
Sc. Philad. 1863, p. 169, is founded on a specimen of this species; it is described as
having all the fins scaleless ; but, as all our specimens of MW. brasiliensis and M. incilis
have the dorsal and anal more or less covered with minute scales, I suppose that these
scales have either been lost in the example of the Smithsonian Institution, or overlooked.

The first dorsal spine, in this species, is either longer than or as long as the second.
L. lat. 36-38.

121. Mvein INcILIs.

Mugil incilis, Hancock in pene Quart. Journ. Se. 1830, p. 127.
Dy 4a, AS palat APA 2 Ws itraney, 15:

Closely allied to MZ. br silts but with smaller scales, and with the second dorsal
spine rather longer than the first.

The height of the body equals the length of the head, which is two-ninths of the
total (without caudal). The snout is moderately broad, scarcely convex, with the lower
profile ascending in the same degree as the upper descends; the interorbital space is
slightly convex, its width being contained twice and two-thirds in the length of the
head. Upper lip rather thin. The angle made by the two mandibulary bones is a

3P 2

444 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

right one; the preorbital tapers posteriorly, has the anterior margin finely serrated,
and-covers the maxillary, so that only a very narrow portion of it is visible on the side
of the snout. Eyes hidden anteriorly and posteriorly by a broad adipose membrane.
Nostrils rather distant from each other, the posterior situated nearer to the orbit than
to the extremity of the snout. The space of the chin, between the mandibles and the
interopercula, is elongate cuneiform. The second dorsal and the anal are enveloped
in small scales. There are twenty-three scales between the snout and the anterior
dorsal. The second dorsal spine is longer than the first, and more than half as long as
the head, The tenth, eleventh, and twenty-fifth scales of the lateral line correspond to
the extremity of the pectoral fin and to the origins of the two dorsal fins. The root of
the pectoral is above the middle of the body; and the ventral is inserted somewhat
nearer to the pectoral than to the spinous dorsal; pectoral shorter than the head ;
caudal deeply emarginate. Silvery, axil of the pectoral blackish.

We have received examples of this species from Dutch and British Guiana; Mr.
Salvin collected two fine examples in the Chagres River.

I formerly considered it possible that the fish described by Hancock might be identical
with IZ. brasiliensis; but having now received examples, I have convinced myself that it
is a distinct species.

123. AGonosToMA Microprs. (Plate LXX. fig. 1.)
Giinth. Fish. i. p. 462.
D.4\5. A. Ls lat. 43, L. transy, 12.

Broad bands of villiform teeth in the jaws, on the vomer, and the palatine and
pterygoid bones. The height of the body is contained five times in the total length,
the length of the head four times and a half; the latter is more than the distance
between the origins of the two dorsal fins; snout much longer than the eye. Upper
lip thick, protruding anteriorly. The maxillary extends to, or beyond, the vertical
from the anterior margin of the eye. ‘The interorbital space is convex. The anterior
dorsal commences midway between the snout and the base of the caudal fin.

Mr. Salvin has collected specimens of this species (of which we have given a detailed
description, /. ¢.) in the Rio Guacalate.

124. AgonostoMa NasuTuM. (Plate LXX. fig. 2.) '

Agonostoma nasutum, Giinth. Fish. ii. p. 463.

Dajaus' elongatus, Kner & Steindachner, Sitzgsber. bayer. Ak. Wiss. 1863, p. 222; and Abhandl.
bayer. Ak. Wiss. 1865, p. 6, Taf. 1. fig. 2.

nasutus, Kner & Steindachner, /.c. p. 8.

D.4|%. A. LL. lat. 42. LL. transy. 12.

Rather narrow bands of villiform teeth in the jaws, on the vomer, and palatine bones.
The height of the body equals the length of the head, and is one-fifth of the total.
‘ See Zool. Record, ii. p. 192.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 445

Upper lip thick. The maxillary extends to the vertical from the anterior margin of
the eye. The interorbital space is convex. ‘The anterior dorsal commences nearer to
the end of the snout than to the base of the caudal fin.

This species, of which we have given a full description (J. c.), occurs in rivers of both
sides of the Isthmus and of Guatemala. The specimens named Dajaus elongatus are
probably emaciated, caught after spawning-time.

125. AGoNosToMA MONTICOLA (Bancroft).

This species, which is indigenous in Jamaica and Barbadoes, has been found by
M. Sallé in Mexico; and Messrs. Kner & Steindachner enumerate it in a collection
from Panama, where it is found in rivers of both sides of the Isthmus. Abhandl. bayer.
Ak. Wiss. 1865, p. 8.

129. GoBIESOX RHODOSPILUS.
Giinth. Proc. Zool. Soc. 1864, p. 25.
eG A One C80. bs li.

A vertical fold of the skin along the lower half of the base of the pectoral; the
coracoid is scarcely below the level of the upper margin of the pectoral. The distance
of the origin of the dorsal fin from the caudal is contained twice and two-thirds in its
distance from the snout; the anal commences below the third dorsal ray. A very
narrow band of short conical teeth in the upper jaw—one of the lateral teeth being
somewhat larger than the others, recurved, canine-like. The lower jaw with a single
series of teeth, the anterior being narrow incisors, whilst the outermost on each side is
distinctly a canine tooth, corresponding to that in the upper jaw. Rose-coloured, with
dark-rose transverse spots, each spot having an edge of deep-red dots.

Two specimens, 18 lines long, were collected by Capt. Dow at Panama.

132. POMACENTRUS RECTIFRENUM.

A species from California and the western coast of Central America was described
by Mr. Gill under this name in the year 1862, and easily recognized by myself when
I gave an account of this genus in the fourth volume of my ‘ Fishes,’ having received,
beside numerous examples from Panama and the Island of Cardon, a typical specimen
from the collection of the Smithsonian Institution. At the same time a specimen was
received from the same establishment, numbered as the other, but marked in the
accompanying list as Pomacentrus bairdii, another name proposed by Mr. Gill in the same
year (Proc. Ac. Nat. Sc. Philad. 1862, p. 149). It agreed perfectly with the descrip-
tion given there; but as a comparison with our other examples of P. rectifrenum did
not reveal distinctive specific characters, I did not hesitate to regard it not only as a
typical specimen of Mr. Gill’s P. bairdii, but also as specifically identical with reeti-
frenum ; about this I had no doubt.

However, during the publication of my account of the Pomacentride, My. Gill
kindly sent me a MS. communication on the same subject, in which he pointed out the

446 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

characters of these two and of three other allied species (see ‘ Fishes,’ iv. p. 27); I had
no faith whatever in these distinctions; but, to allow others interested in the subject to
judge for themselves, I admitted into my account those MS. notes in the form in which
they were sent to me.

I was rather surprised to find in an article written by Mr. Gill a short time after-
wards (Proc. Ac. Nat. Sc. Philad. 1863, p. 218) the assertion that the specimen sent as
“ P. bairdii” is not this species, but the young of P. rectifranum, “the name, under
which the P. dairdii was sent, having doubtless by some accident been shifted to the
young of P. rectifrenum, and the specimen of the former lost.” If this be so, it is
certainly curious that the “P. dairdii” sent agrees so well with the “ P. bairdii”
described; and the shifting must have happened to the sender, inasmuch as the
number corresponding to ‘“‘ P. rectifranum in the accompanying list” is still tied, by a
thread drawn through the tail, to one of the specimens; and two specimens of Poma-
centrus only having been then sent, it is fair to assume that the other example was
meant for P. bairdit.

But in the same paper the author goes a step further: P. dairdit is not only men-
tioned as distinct, but it becomes one of the types of a new genus, Pomataprion, distin-
guished from Pomacentrus by its entire’ preoperculum. In our series of examples,
varying in size from 10 lines to 4} inches, I find that the youngest have the preeoper-
culum entire, without exception, that in specimens 2} inches long the preoperculum
is as frequently entire as but very slightly denticulated, and that the serrature even in
the oldest examples is fine and thin. Therefore I continue to regard those specific
and generic distinctions of Mr. Gill as utterly valueless, and the names of P. recti-
frenun, P. flavilatus, P. analigutta, and P. bairdii as referring to specimens of one
and the same species.

The synonymy stands now thus :—

Pomacentrus rectifrenum, Gill, l.c. p. 148.
flavilatus, Gill, ibid.
bairdii, Gill, 1. c. p. 149.

| analigutta, Gill, im Ginth. Fish. iv. p. 27.
Gunther, Pomacentrus rectifrenum, Giinth, Fish. iv. p. 26.

Gill,
1862.

1862. Helastes insolatus, young (10 inches long), Giinth. ibid. p. 61.
Pomacentrus rectifrenum, Gill, 1. c. p. 214.

) —— flavilatus, Gill, ibid.
Pomataprion bairdii, Gill, 1. c. p. 217.

Guill,
1863.

137. Heniastes MARGINATUS (Castelnau).

Chromis atrilobata (Gill) is, as already stated in Fish. iv. p. 64, identical with this
species. Mr. Gill thinks that it ought to be kept distinct on account of a “sulphur
spot behind the dorsal fin,” Proc. Ac. Nat. Sc. Philad. 1863, p. 220. In two specimens

1 «The praopercular serrature is almost obsolete,” Gill, 1862, p. 149.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 447

from Bahia, which I have examined, the larger is without any trace of such a spot;
whilst the smaller one shows one, of about the size of the scale, on each side of the base
of the last dorsal ray.

141. CossypHus PECTORALIS (Gill).

This species occurs also in the Atlantic, as I have lately received fine examples from
St. Helena. Cossyphus pulchellus (Poey) is perhaps identical with it.

143, Piatyeiossus Dispitus. (Plate LX XIV. fig. 1.)
Giinth. Proc. Zool. Soc. 1864, p. 25.
Dette = L. lat. 28. L. transv. 2/9.

The height of the body equals the length of the head, and is contained four times
and one-fourth in the total. Caudal fin rounded, with the lobes very slightly produced.
Greenish olive, with a roundish black spot edged with silvery on the lateral line,
below the fifth and sixth dorsal spines; the side of the head with five or six pearl-
coloured streaks, a part of which are continued on the body, forming a series of round
spots. An oblong variegated blotch behind the pectoral fin: it is composed of three
pearl-coloured stripes, enclosing two yellow bands, each of which has an undulated
purple edge. No spot in the axil of the pectoral. A short oblique yellowish streak
behind the base of each soft dorsal ray; these streaks form a continuous band on the
spinous portion. Anal fin with two or three whitish lines; caudal with several
irregular reddish longitudinal bands, which are convergent behind.

Young specimens are much more plain-coloured ; the black spot on the lateral line,
however, is very distinct, and there is another at the root of the caudal.

Capt. Dow’s Panama collection contains a single young specimen; but Mr. Salvin has
brought others, one, apparently adult, being 53 inches long.

144. PsrvposuLis NorospiLus. (Plate LXVI. fig. 2.

Giinth. Proc. Zool. Soc. 1864, p. 26.

D. a A. =. L. lat. 25. L. transy. =

The height of the body is rather less than the length of the head, and contained four
times and a quarter in the total. Dorsal spines pungent; caudal fin slightly rounded.
Brownish or yellowish olive; young specimens with a silvery band along each side of
the trunk, above the pectoral fin. Back with four or five indistinct broad brown cross
bars; a series of blotches on the dorsal fin corresponds to these cross bands, one of
them, on the first three dorsal rays, being the largest and most distinct ; it is of a deep
black colour, and of an ovate form. The corners of the caudal fin are white; ventral
whitish, with a broad blackish outer margin.

One adult specimen, 4 inches long, and several young ones were collected by Capt.
Dow at Panama.

448 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

150. GERRES AXILLARIS.
Giinth. Proc. Zool. Soc. 1864, p. 102.
D. 9/10. A. 3/8. L. lat. 34. L. transv. 5/10.

Allied to G. plumieri, but with considerably shorter fin-spines. The height of the
body is contained twice and a fourth in the total length (without caudal). Preorbital
finely serrated. Snout as long as the eye; the groove for the intermaxillary processes
is very broad, scaleless, extending backwards to the vertical from the centre of the eye.
Dorsal fin notched, the last spine being not much longer than the eye; dorsal spines
strong, the second as Jong as the head without snout; the second anal spine stronger,
but scarcely longer than the second of the dorsal fin. The. pectoral extends to the
vertical from the third anal spine. Caudal deeply forked, with the lobes equal in
length to each other and to the pectoral. A blackish streak along each series of scales ;
the hinder side of the axil, and sometimes the anterior, blackish.

Three specimens, from 8 to 9 inches long, were collected by Mr. Salvin at Chiapam,

151. GERRES BREVIMANUS,
Ginth. Proc. Zool. Soc. 1864, p. 152.
D.9|10. A. 3/8. L. lat. 39. L. transy. 6/11.

Preorbital minutely serrated. The height of the body is contained twice and two-
fifths in the total length (without caudal), the length of the head twice and a half.
Snout as long as the eye; the groove for the intermaxillary processes is broad, scaleless,
not extending backwards to the vertical from the centre of the eye. Dorsal fin notched,
the last spine being longer than the eye; dorsal spines strong, the length of the second
equals the distance between the end of the operculum and the anterior nostril; the
second anal spine stronger, but much shorter, than the second of the dorsal fin. The
scaly sheath of the anal fin leaves the outer half of the last ray uncovered. The
pectoral extends scarcely to the vertical from the vent. Caudal scaly, deeply forked,
with the lobes equal in length, each being one-fourth of the total. Three or four
blackish streaks along the series of scales below the lateral line; the spinous dorsal
fin black.

One specimen, 10 inches long, was found by Mr. Salvin at Chiapam.

155. GERRES DOVIL.
Diapterus dowii, Gill, Proc. Ac. Nat. Sc. Philad. 1863, p. 162.

This species is characterized thus :—

Dye Age Ae lat: 47D bransye/5/10:

‘
The greatest height is contained thrice and a half in the extreme length; the head
four times and a quarter; the diameter of the eye twice and three-fourths in the head ;
the snout equals four-fifths of the eye. The profile is rectilinear, andthe interorbital

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA, 449

space nearly flat, but convex above the eyes, and nearly as wide as the eye. The
maxillary groove is linear, and extends backwards to a vertical midway between the
front of the orbit and pupil, while the scales on each side extend to the vertical from
the front of the orbits. The exposed surface of the supermaxillary bones is at. first
triangular, and thence oblong, the whole twice and a half as long as wide. The height
of the constricted portion of the caudal peduncle equals two-thirds of its length and the
diameter of the eye. The lateral line is scarcely bent behind. The second and third
dorsal spines are slender, and nearly equal half the height of the body beneath; the
last is little more than half as long as the first branched ray. The third anal spine is
as long as the snout, and longer and more slender than the second. ‘The colour is
silvery; the spinous dorsal blackish at margin; the axilla of pectoral blackish.
Discovered by Capt. Dow on the Pacific coast of Central America.

156. ACARA CHRULEOPUNCTATA.
Kner & Steindachner, Sitzgsber. bayer. Akad. 1863, p. 222; and Abhandl. bayer. Akad. Wiss. x.
p- 16, tab. 2. fig. 3.
Dei. Az L. lat. 27. -L. transy. 24/9.

Three series of scales on the cheek. Przorbital scarcely wider than the orbit. The
greatest breadth of the head is two-thirds of its length. Cleft of the mouth oblique.
Body with four or five indistinct cross bands. A large black blotch on the middle of
the sides, and traces of a second on the root of the caudal. Each scale on the side of
the head and chest with a bluish spot.

Two specimens, 5 inches long, were collected by Mr. Salvin in the Rio Chagres.

Description.—Vhe height of the body is contained twice and a half in the total length
(without caudal), the length of the head rather more than thrice. Nape curved, the
profile of the snout straight. Width of the interorbital space two-fifths of the length
of the head, and more than that of the snout. Snout broad, moderately elevated, the
width of the preorbital being scarcely more than the diameter of the eye. Cleft of the
mouth slightly oblique, not reaching the vertical from the orbit. The fold of the lower
lip is interrupted in the middle. Lower limb of preoperculum more than half the
length of the posterior limb. There are only eight series of scales between the throat
and the root of the ventral. Dorsal spines of moderate strength, gradually increasing
in length posteriorly; the length of the ninth is more than one-third of that of the
head. The. middle of the soft dorsal’and anal produced, and extending beyond the
middle of the caudal, which is rounded. Pectoral as long as the head, reaching only
to the origin of the anal. Ventral filament rather long. Colours as described above.

157. Heros parma (Gthr.).
This species varies considerably in coloration and in form of the body. The cross
bands may be entirely absent, and replaced by a large diffuse black blotch on the end
VOL. VI.—PART VII. 3 Q

450 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

of the tail; the height of the body is contained from once and three-fifths to twice and
one-fifth in the total length. Guatemalan specimens have generally the black caudal
blotch, but they vary much in the depth of the body. The numbers of the fin-rays

(D. ao A. — appear to be very constant.

158. Heros MARGARITIFER. (Plate LXXI. fig. 2.)

Giinth. Fish. iv. p. 287.

Dd. ALL L. lat. 31. L. transv. 5/13.

The fold of the lower lip is slightly interrupted in the middle; five or six series of
scales on the cheek. The height of the body is rather less than one-half of the total
length (without caudal). Brownish olive, with seven black cross bands, each band with
pearl-coloured spots.

One specimen, 64 inches long, was found by Mr. Salvin at Lake Peten.

159. HEROS MELANOPOGON,
Steindachner, Denkschr. Ak. Wiss. Wien, xxiii. p. 72, Taf. 1. fig. 3.
D: 4 A. 8 Le lat. 30." L. transv. 63/13.

The fold of the lower lip is interrupted in the middle; five or six series of scales on
the cheek. The height of the body is four-ninths of the total length (without caudal).
Body with five irregular blackish cross bands interrupted in the middle, so as to repre-
sent two series of irregular blotches ; a large blackish blotch on the root of the caudal
fin. Small pearl-coloured spots surround the lower blotches, and are scattered over the
caudal blotch.

Known from a specimen 4} inches long.

This fish may represent merely the younger state of H. margaritifer; it is stated to
be from Central America. It formed part of a collection made by Friedrichsthal, who,
to judge from other specimens collected by him, appears to have visited Lake Peten,
which is inhabited by H. margaritifer.

160. Heros MELANURUS. (Plate LXXII. fig. 3.)
Ginth. Fish. iv. p. 288.
Dv, ALS LL. lat. 33. L. transv. 5/13.

The fold of the lower lip is subinterrupted in the middle’; five series of scales on the
cheek. The height of the body is contained twice and a third or twice and a half in the
total length (without caudal). A deep-black band along the middle of the tail; the
lower parts black in adult specimens.

Several examples, from 3 to 10 inches long, were collected by Mr. Salvin at Lake
Peten.

’ The fold is distinetly interrupted in specimens from 6 to 10 inches long, whilst it appears to be slightly
continuous in young individuals of 3 to 4 inches long.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 451

161. HEROS MACRACANTHUS.
Giinth. Proe. Zool. Soc. 1864, p. 153.
D. = A. ae L. lat. 31. L. transv. 3

The lower lip is interrupted in the middle. The height of the body is two-thirds of
the total length (without caudal) in adult specimens, but only one-half in immature ;
the length of the head is one-third of the total. Upper profile of the head very steep,
not concave. Scales on the cheek in five series. ‘The first dorsal spine is a little before
the vertical from the upper end of the gill-opening. Dorsal and anal spines strong, the
tenth of the dorsal fin being two-fifths of the length of the head. Pectoral as long as
the head. Dark greenish, many scales with a pearl-coloured spot in the upper or lower
angle. Vertical and ventral fins black.

About a dozen specimens, from 3 to 9 inches long, were collected by Mr. Salvin at
Chiapam and Huamuchal.

Description.—The height of the body is two-thirds of the total length Gaia
caudal), and nearly one-half of the entire length of the fish. The length of the head
is one-third, or slightly more than one-third of the total (without caudal). Head
rather higher than long, the nape convex, but the upper profile showing a slight
concavity above the snout. The snout is of rather considerable extent, the height of
the preorbital being one-half more than the width of the orbit. The cleft of the
mouth is slightly oblique; the preorbital almost covering the posterior end of the
maxillary, which does not attain the line of the front margin of the eye. Jaws rather
protractile, armed with a broad band of villiform teeth, those of the outer series being
enlarged. Interorbital space convex, nearly twice the width of the orbit. Eye some-
what nearer to the end of the operculum than to that of the snout. Base of soft
dorsal and anal with a few small scales. Dorsal spines strong; the twelfth is a little
less than one-half of the length of the head in adult specimens; the fifteenth is the
longest, and more than half the length of the head. Soft dorsal and anal much
elevated; the middle rays produced; caudal rounded. Pectoral rounded, about as
long as the head. First ventral ray slightly prolonged. The free portion of the tail
is nearly twice as deep as long. Greenish or brownish olive; fins black; a more or
less distinct black spot on the root of the caudal fin, above the lateral line. Immature
specimens with six very indistinct dark cross bands, the third of which has a blackish
blotch below the lateral line; an indistinct blackish spot at the root of the caudal fin.

162. Heros spinurus. (Plate LX XIII. fig. 1.)
Ginth. Fish. iv. p. 289.

18 8-9 44
D. in’ A. — L. lat. 29. LL. transv. i

The fold of the lower lip is interrupted in the middle; four series of scales on the
cheek. The height of the body is one-half of the total length (without caudal), the
3Q2

452 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

length of the head nearly one-third. Head a little higher than long; snout of
moderate extent, its length being two-fifths of that of the head. The diameter of the
eye is two-sevenths of the length of the head, two-thirds of that of the snout, and less
than the width of the interorbital space, which is convex; the eye is situated below
the upper profile, a little nearer to the extremity of the operculum than to that of the
snout, Preeorbital as wide as the orbit. Dorsal spines of moderate length and strength,
the length of the twelfth being contained twice and a third in that of the head. The
distance between dorsal and caudal is less than the depth of the tail. Greenish olive,
with nine dark yertical bands; a large, roundish black spot on the middle of the root
of the caudal; no spot on the temple ; caudal. and the posterior part of the dorsal and
anal with whitish spots.

Three examples were collected by Messrs. Salvin and Godman at Yzabal and in the
Rio Motagua. Length 33 inches.

163. Heros nicrorasciatus. (Plate LXXIV. fig. 3.)
D. 2. A. 2. L. Jat. 29. L. transy. 4/11.

The lower lip is interrupted in the middle. Scales on the cheek in four or five series.
Dark blackish brown, with nine deep-black cross bands.

Numerous examples, from 2 to 3} inches long, were collected by Mr. Salvin in the
Lakes of Amatitlan and Atitlan.

Description.—Vhe height of the body is contained twice and one-sixth in the total
length (without caudal), the length of the head thrice; the free portion of the tail is
considerably deeper than long. Head as high as long, with the upper profile convex
to the snout, where it is straight. Snout of moderate extent, the width of the pre-
orbital being equal to that of the orbit. The eye is somewhat nearer to the end of the
snout than to that of the operculum ; its diameter is considerably less than the width
of the interorbital space, and one-fourth of the length of the head. Jaws equal in
length. The soft dorsal and anal fins have scarcely any scales on their base, and are
more or less produced in the middle, the longest rays reaching to the middle of the
caudal. ‘The dorsal fin commences in the vertical from the humerus; its spines are of
moderate strength, rather short, the length of the twelfth being somewhat less than
one-third of that of the head. Anal spines as long as, but rather stronger than those
of the dorsal fin. Caudal rounded, two-ninths of the total length. Pectoral as long
as the head, without snout, extending to the second or third anal spine. Ventral but
slightly produced.

This spécies is very dark-coloured. The ground-colour is a dark blackish purplish
brown. An arched black band runs ‘from the nape of the neck round the opercular
margin to the interoperculum. A second is nearly concentric with the first, running
from the nape to behind the pectoral and ventral. ‘The third is short, like a spot,

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 455

between the anterior dorsal spines and the lateral line. The following are subvertical,
slightly inclined. backwards, and broader than the imterspaces between them. The
penultimate connects the ends of the dorsal and anal fins; the last across the root of
the caudal. Fins black.

This species appears to remain within small dimensions.

164. Heros MULTISPINosuS. (Plate LX XIV. fig. 2.
D. 7. Az L.lat. 29. LL, tansy: 4/12.

‘The lower lip is interrupted in the middle. Three series of scales on the cheek. A
blackish band, interrupted on the tail, runs from the eye to the caudal; a round black
spot in the middle of the length of the band.

A single specimen, 34 inches long, was obtained by Capt. Dow in the Lake of
Managua.

Description—tThe height of the body is contained twice and one-seventh in the total
length (without caudal), the length of the head thrice. The free portion of the tail is
twice as deep as long. Head as deep as long, with the upper profile nearly straight.
Snout rather short; the width of the preorbital being considerably less than that of the
orbit. The eye is situated immediately below the upper profile, nearer to the end of
the snout than to that of the operculum; its diameter is a little less than one-third of
the length of the head, and much less than the width of the interorbital space, which
is flat. Mouth with the jaws equal in length, small, the maxillary not nearly reaching
the vertical from the orbit. Suboperculum with two series of scales. The soft dorsal
and anal fins are scaly at the base, they are scarcely prolonged, and not extending to
the middle of the caudal. The dorsal fin commences above the humerus ; its spines are
of moderate strength, and rather long, the length of the eighth to the last spine being
not much less than one-half of that of the head. Anal spines stronger, and even a
little longer than those of the dorsal. Caudal fin rounded, two-ninths of the total
length. Pectoral shorter than the head, extending to the fifth anal spine. The outer
ventral ray produced into a. short filament. Brownish olive, each scale somewhat
darker at the base. A blackish band, as broad’as a scale, runs from the eye to a round
black spot situated before and below the termination of the upper part of the lateral
line; thence it is continued to the root of the caudal as a series of four or five irregular
spots. Fins blackish, apparently immaculate.

165, Heros toncimanus. (Plate LXXII. fig. 2.)
D. jg Aug: L, lat. 28... L.-transv. 43/12.
The fold of the lower lip is interrupted in the middle. Three or four series of scales
on the cheek. Pectoral very long, extending nearly to the end of the anal. Greenish

454 Dk. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

olive, with an indistinct blackish band running from the orbit to a large black spot on
the middle of the side. Dorsal fin with numerous round whitish spots.

One specimen, 5} inches long, was found by Capt. Dow in the Lake of Nicaragua.

Description—The height of the body is contained twice and one-sixth in the total
length (without caudal), the length of the head twice and three-fifths. Upper profile
of head straight. Head rather longer than high; cleft of the mouth slightly oblique,
with the lower jaw rather prominent. Jaws moderately protractile; the maxillary does
not extend to the vertical from the front margin of the eye. Preorbital as wide as the
diameter of the eye, which is somewhat less than the width of the interorbital space,
and more than one-fourth of the length of the head. The eye is situated immediately -
beneath the upper profile of the head, and a little nearer to the end of the operculum
than to that of the snout. Scales on the cheek in three or four series; scales on the
opercles large. The dorsal commences vertically above the scapula; the spinous portion
has its upper margin convex; the spines are slender and long, the fifth and sixth being
the longest, one-half of the length of the head. The soft dorsal and anal have the middle
rays somewhat longer than the others, and reaching to about the middle of the caudal ;
the soft anal is slightly scaly at the base, the soft dorsal scarcely or not at all scaly.
Anal spines shorter but somewhat stronger than those of the dorsal. Caudal slightly
emarginate. Pectoral very long, slightly longer than the head, and extending nearly to
the end of the anal. Ventral with the outer ray produced into afilament. The distance
between the vent and the root of the ventrals is equal to one-third of the length of the
head. Teeth in the jaws small, cardiform, forming a band, those of the outer series
being somewhat larger than the others.

This species is similar to the Mexican //. helleri, but has a considerably longer
pectoral fin, and also less anal rays.

166. Heros vroputuatmus. (Plate LXXII. fig. 1.)
Ginth. Fish. iv. p. 291.
Dei. A. 3. -L'lat: 28," Li transv. 5/12:

The fold of the lower lip is continuous in the middle. Scales on the cheek in six
series. The height of the body is contained twice and a half or twice and a quarter in
the total length (without caudal), the length of the head nearly three times. Head as
high as long; snout rather elevated, with the cleft of the mouth oblique and with the
lower jaw prominent. Preorbital as wide as the orbit; interorbital space flat, wider
than the orbit. The eye is nearer to the extremity of the snout than to that of the
operculum. Dorsal spines of moderate length and strength, the length of the twelfth
being two-fifths of that of the head. The free portion of the tail is higher than long.
Anal spines strong and long. ‘The distance between the vent and the root of the ventral
is three-fifths of the length of the head. Brownish- or greenish-olive, with seven blackish
cross bands, as broad as the interspaces between: the first descending - obliquely back-

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 455

wards across the nape; the. second, third, and fourth below the dorsal spines; the
seventh across the free portion of the tail. A large, black, white-edged ocellus on the
root of the caudal. Fins blackish; pectoral yellowish towards the base.

Mr. Salvin obtained three examples, 7 inches long, at Lake Peten.

167. Heros aurgus. (Plate LX XIII. fig. 2.
Giinth. Fish. iv, p. 292.
Di tees Mudate33:1 LL. trausy..6/13.

The fold of the lower lip is continuous in the middle. Scales on the cheek in five
series. Base of the dorsal scaleless. The height of the body is contained twice and a
third in the total length (without caudal), the length of the head three times. Head
as high as long; snout somewhat elevated, with the cleft of the mouth oblique and
with the jaws equal anteriorly ; preorbital as wide as the orbit. The eye is a little
nearer to the extremity of the operculum than to that of the snout. Dorsal spines
rather slender, the length of the twelfth being a little less than one-half of that of the
head. The distance between the dorsal and caudal is somewhat less than the greatest
depth of the free portion of the tail. Caudal slightly emarginate. The distance
between the vent and the root of the ventral is one-third of the length of the head.
Yellowish-olive, with six dark cross bands, extending downwards to a yellow longi-
tudinal band running from above the pectoral to the lower half of the base of the
caudal. The third cross band terminates in a large black lateral spot; sides of the
head with several bluish dots, and with a blackish spot on the operculum and suboper-
culum, darkest on the latter bone. Fins light-coloured, immaculate.

Two specimens, 4 and 5 inches long, were collected by Messrs. Salvin and Godman
at Yzabal and in the Rio Motagua.

168. Heros arrinis. (Plate LXXIX. fig. 1.)
Ginth. Fish. iv. p. 292.
D. ge Av Ge lat. 29. L. transv. 5/12.

The fold of the lower lip is continuous im the middle. Scales on the cheek in four
series. The height of the body is contained twice and two-fifths in the total length
(without caudal), the length of the head twice and three-fourths. Head as high as
long; snout compressed, elevated, with the cleft of the mouth oblique and with the
lower jaw prominent. Preorbital wider than the orbit (in the larger individuals) ; the
eye is considerably nearer to the extremity of the operculum than to that of the snout.
Dorsal and anal fins entirely scaleless; dorsal spines rather strong and long, the length
of the twelfth being two-fifths of that of the head. Anal spines very strong. ‘The free
portion of the tail is a little higher than long. Caudal slightly emarginate, two-ninths
of the total length. The distance between the vent and the root of the ventral is one-

456 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

third of the length of the head. Olive, with five or six dark cross bands, the middle one
of which has a deep-black spot where it passes the lateral line; a more or less distinct
black spot on the suboperculum ; sides of the head and vertical fins with bluish dark-
edged ocelli.

This species is very closely allied to H. aureus, but may be distinguished from it by
larger scales, by a more backward position of the eyes, by stronger spines, &c.

Four examples, from 3} to 5} inches long, were obtained by Mr. Salvin at Lake Peten.

169. Heros LABIATUS.
Giinth. Proc. Zool. Soc. 1864, p. 27, pl. 4. fig. 1.

D.v Alt. L lati 32... ‘L. transv. 6/13.

The anterior portions of the upper and lower lips are much enlarged, each forming
a moveable subtriangular flap (probably in old males only). The height of the body is
somewhat more than the length of the head, and two-fifths of the total (without caudal) ;
the eye occupies the middle of the length of the head. Scales on the cheek in four
series. Base of the dorsal fin almost scaleless. The length of the eighth dorsal spine ts
less than one-third of that of the head. The depth of the free portion of the tail is
scarcely more than its length. Uniform red, or red irregularly marbled with black, or
nearly entirely black.

Two specimens, 63 and 7 inches long, were collected by Capt. Dow in the Lake of
Managua; three others were lately sent by the same gentleman from the Lake of
Nicaragua. We do not yet know the female sex and the young state of this species.

Description —Head rather longer than high; snout somewhat elevated; cleft of the
mouth slightly oblique, with the lower jaw a little prominent. Teeth in narrow bands,
those of the outer series enlarged, with brown tips. The maxillary does not nearly
attain the vertical from the front of the eye. Preeorbital as wide as the orbit, the
diameter of which is less than the extent of the snout, and one-fourth of the length of
the head. Interorbital space somewhat convex, wider than the orbit. The eye is
situated not quite immediately beneath the upper profile of the head, and midway
between the end of the snout and that of the operculum. Opercles scaly, the scales
being larger than those on the cheek; suboperculum with two series of scales. Soft
portions of dorsal and anal fins with minute scales between the rays at their base.
Dorsal spines of moderate length and strength, the length of the eighth dorsal spine
being less than one-third of the length of the head. Points of the produced aiddle
rays of the soft dorsal and anal reaching to the middle of the caudal fin. Caudal
rounded, its length being contained rather more than five times in the total. Anal
spines of nearly the same length and strength as those of the dorsal fin. Pectoral
rounded, reaching to the fourth or fifth spine of the anal; ventral filament produced.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 457

The distance between the vent and the root of the ventral is less than one-half of the
length of the head.

170. Heros ERyTHREUS. (Plate LXXV. fig. 2.)
D4. AL Eilat. 31. 1. trafisv. 64/14.

Lips thick, with broad free margin in their entire circumference. The height of the
body is contained twice and a third in the total length (without caudal), the length of
the head twice and two-thirds; the eyes occupy the middle of the length of the head.
Scales on the cheek in four or five series. Base of the soft dorsal fin with very small
scales. The length of the eighth dorsal spine is less than one-third of that of the head.
The depth of the free portion of the tail is conspicuously more than its length. Of a
deep orange-colour; many of the scales of the tail with a blackish spot on the base.

One specimen, 7 inches long, was collected by Capt. Dow in the Lake of Managua.

I was for some time inclined to regard this fish as a variety of sex or age of 7.

labiatus. This, however, is not the case, all the specimens being males, and the speci-
men of H. erythrwus larger than one of the two of H. labiatus. Besides, it appears to
be sufficiently distinguished by its much shorter and deeper tail.
. Description—Head as high as long ; snout rather elevated, with the cleft of the mouth
slightly oblique, and the lower jaw scarcely prominent. ‘Teeth in narrow bands, those
of the outer series enlarged, with brown tips. The maxillary does not reach the
vertical from the front margin of the eye; preorbital wider than the orbit. The
diameter of the eye is contained nearly five times in the length of the head. Inter-
orbital space slightly convex, much wider than the orbit. Eye situated near the upper
profile of the head, and equidistant from the end of the snout and that of the oper-
culum. Opercles scaly, the scales being larger than those on the cheek ; suboperculum
with two series of scales. The soft dorsal and anal fins with a few minute scales
running up between the bases of the rays; dorsal spines of moderate strength. Soft
dorsal and anal slightly produced, not reaching to the middle of the caudal. Caudal
rounded, one-fifth of the total length. Anal spines stronger but not longer than those
of the dorsal fin. Pectoral rounded, extending to the fourth anal spine, somewhat
shorter than the head. The outer ventral ray produced; the distance between the
yentral and the vent is one-half of the length of the head.

171, Heros Losocuinus. (Plate LXXV. fig. 1.)
Dior WA gan Eaileti32sy Tiitransy,.6/14.

Old males with the anterior portions of the upper and lower lips much enlarged,
each forming a moveable subtriangular flap; in young males the lips are simple, the
fold of the lower being continuous. The height of the body is contained twice and a
third in the total length (without caudal), the length of the head twice and three-

VOL. VI.—PART VII. aR

458 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

fourths. The eye occupies the middle or nearly the middle of the length of the head.
Scales on the cheek in four or five series. Base of the dorsal fin scaly. The length of
the eighth dorsal spine is more than one-third of that of the head. The depth of the free
portion of the tail is scarcely more than its length. Greenish or yellowish, with about
six indistinct dark cross bands; that below the fourteenth dorsal spine with a large
black blotch below the lateral line; sometimes a black spot on the upper half of the
base of the caudal.

Two male specimens were collected by Capt. Dow in the Lake of Managua; the
larger, 8 inches long, has the labial lobes and black caudal spot; the smaller is 7 inches
long, and, without doubt, of the same species. The female sex is unknown.

Description —Head as high as long; snout rather elevated, with the cleft of the
mouth oblique, and the lower jaw rather prominent. Upper profile very concave.
Teeth in narrow bands, those of the outer series enlarged, with brown tips. The
maxillary does not reach nearly to the vertical from the front of the orbit; preeorbital
as wide as the orbit, being contained very slightly more than four times in the length
of the head. Interorbital space flat, much wider than the orbit. The eye is situated
immediately below the upper profile, slightly nearer to the extremity of the snout than
to that of the operculum. Opercles scaly, the scales being larger than those on the
cheeks; suboperculum with one series of scales. ‘The soft portions of the anal and
dorsal fins with a series of small scales between the rays at their base. Dorsal spines
of moderate strength, the length of the eighth to twelfth being more than one-third of
that of the head. Points of the soft anal and dorsal reaching to the middle of the
caudal. The free portion of the tail is scarcely higher than long. Caudal rounded, its
length being one-fifth of the total. Anal spines strong and long. Pectoral rounded,
reaching to the fourth anal spine; outer ventral ray produced. The distance between
the vent and the root of the ventral is three-sevenths of the length of the head.
Coloration as described above.

172. Heros citRINELLUS. (Plate LXXI. fig. 1.)
Giinth. Proc. Zool. Soc. 1864, p. 153.
p. $3" A. ZL. Jat. 30. L. transv. 6/13.

The fold of the lower lip is continuous in the middle. ‘The height of the body is
contained twice and a fifth in the total length (without caudal), the length of the head
twice and seven-eighths; the free portion of the tail is conspicuously deeper than long ;
nape of the neck very convex ; interorbital space broad, its width being two-fifths of the
length of the head. Snout not obtuse; scales on the cheek in four series. The first
dorsal spine is inserted above the upper end of the gill-opening. Dorsal and anal
spines slender, the eighth or tenth of the dorsal fin being two-fifths of the length of
the head. Pectoral nearly as long as the head. Lemon-coloured, either nearly uni-

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 459

form or with the back black, which colour sometimes forms irregular blotches on the
vertical fins.

Three specimens, two males and one female, from 7 to 8 inches long, were collected
by Capt. Dow in the Lake of Nicaragua.

Description.—Head as high as long; snout rather elevated, the cleft of the mouth
almost horizontal, the lower jaw scarcely prominent. Teeth in narrow bands, those of
the outer series enlarged, with brown tips. ‘The maxillary does not reach the vertical
from the front margin of the eye. Prezorbital wider than the orbit. The eye is
situated close to the upper profile, and a little nearer to the end of the snout than to
the opercular margin; its diameter is one-fourth of the length of the head. Inter-
orbital space flattish, twice the width of the orbit. Opercles scaly, the scales being
larger than those on the cheeks; suboperculum with two series of scales. Soft anal
and dorsal fins slightly scaly at the base. The points of the soft dorsal and anal con-
siderably produced, and extending beyond the middle of the caudal fin, sometimes to
its extremity. Caudal rounded, its length being contained four times and a half in the
total. Pectoral long and rounded, extending to the fifth anal spine. Outer ventral
ray produced. ‘The distance between the vent and the root of the ventral is nearly one-
third of the length of the head.

173. HEROS ALTIFRONS.
Kner & Steindachner, Sitzgsber. bayer. Akad. 1863, p. 223; and Abhandl. bayer. Akad. x. p. 11,
Taf. 2. fig. 1.
ia
The lower lip is dilated into a lobe on each side, which is broadest behind. Scales
on the cheek in four or five series. The height of the body is contained twice and two-
fifths in the total length (without caudal), the length of the head twice and four-fifths.
Snout rather high, the width of the preorbital being more than that of the orbit. Jaws
equal in length. Eye considerably nearer to the end of the operculum than to that of
the snout. Dorsal spines of moderate length and strength. Body with four or five dark
vertical bands’, each band with a darker‘blotch. Scattered pearl-coloured dots all over
the body; a dark spot on the middle of the root of the caudal fin.
Southern (Pacific) rivers of the district Chiriqui (Western Veragua).

174. Heros FRIEDRICHSTHALU.

Heckel, Flussfische Brasil. p. 381. Giinth. Fish. iv. p. 294.
18 8-9

D. 9qp- A- gy. 4. lat. 31. L. transv. 4/12.
The fold of the lower lip continuous in the middle. Scales on the check in seven
series. Antero-inferior margin of preorbital concave, the greatest width of this bone
' The authors describe them as “ teenie,” instead of * fascie.”

R2

460 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA,

being only two-thirds of that of the orbit. The length of the twelfth dorsal spine is
two-sevenths of that of the head. Yellowish-olive, with six or seven blackish cross
bands; a black band from the eye to the upper part of the root of the caudal, inter-
rupted by the interspaces between the cross bands; the origin and end of this band are
edged with yellow; suboperculum with a black ocellus; an oblique black streak from
the eye towards the ocellus.

Lake Peten. Several examples, 5 inches long, were collected by Mr, Salvin.

175. Heros satvint. (Plate LXXIIL fig. 3.)

Heros salvini, Gunth. Fish. iv. p. 294.
triagramma, Steindachner, Denkschr. Akad. Wiss. Wien, xxiii. p. 70, tab. 3. fig. 2.

Dee OMe SE Pe BOOK: trainsy.6/10.

‘

Fold of the lower lip continuous in the middle; scales of the cheek in five series.
Preorbital a little narrower than the orbit, with the antero-inferior margin concave,
Base of the soft dorsal scaly. The height of the body is contained twice and a fourth
in the total length (without caudal), and the length of the head twice and three-fourths.
Head: somewhat longer than high; snout of moderate extent, longer than the eye,
pointed, with the cleft of the mouth very oblique, and with the lower jaw projecting ;
the maxillary does not quite extend to the vertical from the anterior margin of the
orbit. The eye is situated immediately below the upper profile, in the middle of the
length of the head. Suboperculum of moderate width, with one series of scales. The
length of the twelfth dorsal spine is two-fifths of that of the head in specimens from
Lake Peten, and one-third in those from Santa Isabel. The distance between dorsal and
caudal is considerably less than the depth of the free portion of the tail. ‘The distance
between the vent and the root of the ventrals is two-fifths of the length of the head.
Dark greenish olive, with a black band, edged with yellow, running from the snout,
through the eye, to the root of the caudal; it is most distinct on the head, but inter-
rupted on the tail by lighter interspaces ; it passes a black lateral spot, and, in young
specimens, terminates in another black spot. An irregular black band along the back,
below the base of the dorsal fin. Sometimes ‘three bands across the upper surface of
the head. A blue horizontal line below the orbit; a more or less distinct black
ocellus on the suboperculum is sometimes entirely absent. Fins blackish, immacu-
late, or with faint dots only in small number. The sides below the black band are
sanguineous in mature specimens.

The largest specimen is 43 inches long.

This species occurs in Lake Peten as well as in the Rio Santa Isabel ; specimens from
the former locality are distinguished by somewhat longer dorsal spines. H.. triagramma
appears to have been founded on a Lake-Peten example.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 461

176. Heros rrmacunatus. (Plate LX XVI.)
D. =. A. = L. lat. 31. L. transv. 5/14.

Allied to H. salvini. Fold of the lower lip continuous in the middle; scales of the
cheek in five series. Lower jaw prominent. Preorbital as wide as the orbit, with the
antero-inferior margin concave. The length of the twelfth dorsal spine is rather less
than one-third of that of the head. Dark greenish olive, with three black spots; the
first above the origin of the lateral line, the second in the middle of the side, and the
third above the end of the lateral line. Fins black.

Three adult examples, from 8 to 11 inches long, and one of 2} inches, were collected
by Mr. Salvin at Chiapam and Huamuchal.

Description—The height of the body is contained twice in the total length (without
caudal), the length of the head twice and two-thirds. Head nearly as high as long;
snout rather pointed, much longer than the eye, with the cleft of the mouth very
oblique, and the lower jaw prominent; the maxillary extends nearly to the vertical
from the front margin of the orbit. Praorbital as wide as the orbit, with the antero-
inferior margin concave. The width of the orbit is one-fifth of the length of the head,
but only two-thirds of that of the interorbital space. ‘The eye is situated immediately
below the concavity of the upper profile of the head, and is very slightly nearer to the
tip of the snout than to the opercular margin. Opercles scaly; suboperculum with two
series of scales. The vertical fins are scaly at the base. Dorsal spines of moderate
strength and length, the twelfth being rather less than one-third of the length of the
head. The points of the soft dorsal and anal extend beyond the middle of the caudal.
Caudal much rounded. The distance between the caudal and the dorsal is considerably
less than the depth of the free portion of the tail. Pectoral much shorter than the
head, extending only to the second anal spine; ventrals with the outer ray produced.
The distance between the vent and the root of the ventrals is nearly half the length of
the head. The coloration of the young example is exactly the same as that of the adult.

177. Heros povit. (Plate LXXIII. fig. 4.)
Giinth. Proc. Zool. Soc. 1864, p. 154. '
D. a A. — L. lat. 35. _L. transv. =
The fold of the lower lip is continuous in the middle. The height of the body is
contained thrice in the total length (without caudal); the length of the head twice and
three-fifths. Snout pointed, with the lower jaw very prominent. Preeorbital with the
antero-inferior margin but slightly concave, its greatest width being three-fourths of
that of the orbit. Both jaws with a pair of fangs, those of the upper pair being close
together in the middle of the jaw, whilst the lower are separate. Scales on the cheek
small, rather irregularly arranged, in about eight series. The first dorsal spine is
inserted behind the vertical from the upper end of the gill-opening. Dorsal and anal

462 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

spines slender, the length of the twelfth of the dorsal fin being one-fourth of that of the
head. Pectoral three-fifths as long as the head. Brown, irregularly marbled with
darker; fins black; an indistinct black band along the operculum and the side of the
trunk; an oblique blackish band descends from the eye towards the root of the pectoral ;
a black spot behind the angle of the mouth.

‘This species is allied to H. friedrichsthalii, H. salvini, &c. ‘Two specimens, 6 inches
long, were collected by Capt. Dow in the Lake of Nicaragua.

Description.—Head much longer than high. Snout rather elongate, much longer than
the eye, pointed, with the cleft of the mouth oblique, and the lower jaw very prominent.
‘lhe maxillary reaches the vertical from the anterior margin of the orbit. The width of
the orbit is contained four times and a half in the length of the head, and equal to that
of the interorbital space. The eye is situated immediately below the upper profile, but
is considerably nearer to the end of the snout than to that of the operculum. Opercles
scaly, the scales on the operculum larger than those on the cheek; suboperculum with
two series of scales. The soft portions of the dorsal and anal fins are scaly at the base,
and do not reach much beyond the origin of the caudal. Caudal rounded. The
pectoral is about two-thirds as long as the head, and scarcely reaches the vertical from
the origin of the anal. Ventral pointed, slightly produced, reaching only to the vent.
The distance between the vent and the root of the ventral is two-fifths of the length of
the head.

178. Heros Moracuensis. (Plate LX XVII. fig. 2.)

De Awe. Lu. lat. 32." T. transy. 5/13.

The fold of the lower lip is continuous in the middle. Snout pointed, with the lower
jaw prominent. Preorbital with the antero-inferior margin but slightly concave, its
greatest width being equal to that of the orbit. Dentition as in H. dow. Scales on
the cheek small, in eight series. The first dorsal spine is inserted behind the vertical
from the upper end of the gill-opening. Dorsal and anal spines short, the length of
the twelfth of the dorsal fin being two-ninths of that of the head. Brownish, a black
interrupted band runs from the eye to a spot on the root of the caudal, this spot being
situated above the lateral line. An oblique short black streak runs from the lower
posterior angle of the orbit towards a spot situated on the suture between operculum
and suboperculum, close to the interoperculum, the band being not continuous with the
spot. Back with traces of irregular cross bands, more distinct in young than in old
individuals. Vertical fins with numerous brown dots.

Five examples, from 4 to 10 inches long, were obtained by Mr. Godman from the Rio
Motagua. This species is closely allied to H. friedrichsthalii.

Description of an example 10 inches long.—The height of the body is nearly equal to
the length of the head, and is contained thrice in the total length (without caudal); the
length of the head is contained twice and five-sixths in the same. Head longer than

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 465

high; snout of moderate extent, much longer than the eye, pointed, with the cleft of
the mouth very oblique, and the lower jaw very prominent. The maxillary reaches
nearly to the vertical from the anterior margin of the orbit. The width of the orbit is
not quite one-fifth of the length of the head, and less than the width of the interorbital
space. ‘The eye is situated near the upper profile of the head, nearer to the end of the
snout than to that of the operculum. Opercles scaly; suboperculum with two series of
scales. Vertical fins scaly at the base, their points do not reach the middle of the
caudal. Caudal rounded. ‘The distance between the dorsal and caudal is somewhat
less than the depth of the free portion of the tail. Pectoral short, less than two-thirds
of the length of the head, and scarcely reaching to the vent; ventral short, pointed, with
the outer ray produced. The distance between the vent and the root of the ventral is
more than half the length of the head.

179. Heros MANAGUENSIS. (Plate LX XVII. fig. 3.)

D. i A.j. L. lat.32. ‘L. transy. 43/13.

The fold of the lower lip is continuous in the middle. Snout somewhat pointed, with
lower jaw prominent. Preorbital with the antero-inferior margin concave, narrow, its
greatest width being scarcely more than one-half of that of the orbit. Dentition as in
H. dovii. Scales on the cheek small, rather irregularly arranged, in eight or nine
series. The first dorsal spine is inserted behind the vertical from the upper end of the
gill-opening. Dorsal and anal spines of moderate length and strength, the length of the
twelfth of the dorsal fin being contained thrice and two-thirds in that of the head.
Greenish brown, shining golden, and irregularly marbled with dark brown. A series
of quadrangular black spots (probably a band in young examples) runs from the eye to
a black spot on the root of the caudal, this spot being situated above the lateral line;
a brown band descends obliquely from the lower posterior angle of the orbit to the
lower posterior angle of the operculum. Vertical fins with black spots, each spot
being half as large as a scale.

This species is allied to H. friedrichsthalit, salvini, &c.; a single specimen, 74 inches
long, was found by Capt. Dow in the Lake of Managua.

Description —tThe height of the body is nearly equal to the length of the head, and
two-fifths of the total length (without caudal). Head longer than high; snout of
moderate length, somewhat pointed, with the lower jaw prominent, and the cleft of
the mouth oblique. The maxillary reaches beyond the anterior margin of the eye.
The width of the orbit is one-fifth of the length of the head, and three-fourths of the
width of the interorbital space. The eye is situated immediately below the upper
profile ; its distance from the end of the snout is a little more than half of that from
the hinder margin of the operculum. Opercles scaly, the scales on the operculum
larger than those on the cheek; suboperculum with two series of scales. Vertical fins

464 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

slightly scaly at the base. The soft dorsal and anal do not reach to the middle of the
caudal. Caudal rounded. The distance between the dorsal and caudal is much: less
than the depth of the free portion of the tail. Pectoral short, more than half the
length of the head, and extending only to the origin of the anal; ventral, with the
outer ray slightly produced, reaching beyond the vent. The distance between the vent
and the root of the ventral is not quite half the length of the head.

180. Heros MICROPHTHALMUS.
Ginth, Fish. iv. p. 295.
5-6

D. a A. jap LL. lat. 34. L. transv. 5/14.

The fold of the lower lip is continuous in the middle; six series of scales on the
cheek. The height of the body is contained twice and a third in the total length
(without caudal), the length of the head thrice and a third. Head as high as long ;
snout of moderate extent; preorbital wider than the eye. Cleft of the mouth rather
narrow, horizontal, with the jaws equal anteriorly. Interorbital space very convex,
twice as broad as the- orbit; -the eye is a little nearer. to the extremity of the snout
than to that of the opercle. Vertical fins scaly at the base; the spinous dorsal is low,
the length of the twelfth spine being one-third or rather less than one-third of that of
the head. The free portion of the tail is rather higher than long. Pectoral much
shorter than the head, equal in length to the ventral, which does not extend on to the
vent. Brownish, with indistinct dark cross bands, and with a dark band along the sides
and tail, terminating at a black spot in the middle of the root of the caudal. Each
scale on the lateral and lower parts with a purple spot at the base. The soft portions
of the vertical fins with series of blackish dots; axil of the pectoral orange-coloured.

Numerous examples, from 4 to 8 inches long, were collected by Mr. Godman in the
Rio Motagua.

. 181, Heros oBLonevs.
D. gus A-gg LL. Jat. 38. LL. trangv/63/15,

Closely allied to H. microphthalmus, but with the body considerably more elongate.

The fold of the lower lip is continuous in the middle; five series of scales on the
cheek. The height of the body is one-third of the total length (without caudal), the
length ofthe head two-sevenths. Mouth small, horizontal, with the jaws equal in
length. The length of the twelfth dorsal spine is less than one-third of that of the
head. Coloration as in H. microphthalmus.

Two examples, 7 and.8 inches long, were obtained by Mr. Godman from the Rio
Motagua. be

Description. Head a little longer than high; snout of moderate extent; preorbital
wider than the eye, the diameter of which is one-fifth of the length of the head. Cleft
of the mouth rather narrow, horizontal, with the jaws equal anteriorly, and the maxil-

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 465

lary not extending backwards to the vertical from the front margin of the eye. Teeth
in a band, those of the outer series much larger and stronger than the others, and with
brown tips. Interorbital space convex, not quite twice as broad as the orbit. Hye
about equidistant from the end of the snout and that of the opercle. Vertical fins
scaly at the base. The spinous dorsal is rather low, the length of the twelfth spine
being less than one-third of that of the head. Soft dorsal and anal somewhat produced,
the former reaching to the middle of the caudal. The free portion of the tail is rather
‘longer than high. Caudal subtruncated, its length being a little less than one-fifth of
the total. Pectoral shorter than the head, about equal in length to the ventral, the
outer ray of which reaches to the vent. Brownish, with about five very indistinct
broad darker cross bands, descending from the back to a not less indistinct longitudinal
band which runs from above the pectoral to a black spot in the middle of the root of
the ventral. Vertical fins with transverse series of round whitish spots, separated by a
network of dark lines. Pectoral yellowish.

182. Heros nicaracuEnsis. (Plate LX XVII. fig. 1.)
Ginth. Proc. Zool. Soc. 1864, p. 153.
D. a A. sa L, lat. 35. L. transv. 5/13.

The fold of lower lip is interrupted in the middle, The height of the body is con-
tained twice and two-fifths in the total length (without caudal); the length of the head
thrice and one-fifth. Head much higher than long, in consequence of an adipose
swelling above the eye, which renders the shape of the head Coryphzna-like. Scales
on the cheek in six series, rather irregularly arranged. The first dorsal spine is inserted
aboye the upper end of the gill-opening. Dorsal and anal spines long, the sixteenth of
the dorsal fin being one-half the length of the head, Pectoral not quite as long as
the head, Brownish olive above, yellowish below; back with five or six blackish cross
bands, not extending downwards to beyond the middle of the side; many scales with a
brown vertical marginal streak. The soft vertical fins with brown spots, each half as
large as a scale.

Two specimens, 64 and 7 inches long, were collected by Capt. Dow in the Lake of
Nicaragua.

Description.—Snout elevated; preorbital wider than the orbit, the diameter of which
is more than one-fourth of the length of the head. Cleft of the mouth rather narrow,
horizontal, the jaws equal in front, and the maxillary not extending back to the vertical
from the front of the orbit. Teeth in a band, those of the outer series being somewhat
enlarged, and with brown tips. Interorbital space very convex, not quite twice as broad
as the orbit. The eye is about equally distant from the end of the snout and that of
the opercle, and far below the upper profile of the head. Vertical fins scaly at the
base. Spinous dorsal not very low, the sixteenth spine being half as long as the head ;

VOL. VI.—PART VII. 38

466 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

the soft portions of the dorsal and anal slightly produced, the former extending nearly
to the middle of the caudal. Free portion of the tail as high as long. Caudal slightly
emarginate, its length being considerably more than one-fifth of the total. The ventral
has the outer ray much produced, and reaches to the sixth anal spine.

183. Heros copMAnni. (Plate LXXIV. fig. 5.)
Giinth. Fish. iv. p. 296.
D. ee A. 2. L. lat. 33. L. transyv. 6/13.

The fold of the lower lip is interrupted in the middle; six or seven series of scales on
the cheek. The height of the body is contained twice and three-fourths in the total
length (without caudal), the length of the head thrice or thrice and a third. The
profile of the nape is much curved. Head rather longer than high; snout rather
elevated, the preorbital being wider than the orbit. Cleft of the mouth rather narrow,
horizontal, with the jaws equal anteriorly, and with the maxillary not extending
backwards to the vertical from the front margin of the eye. The nape is elevated, and
the orbit considerably below the upper profile of the head. Dorsal and anal fins very
slightly scaly at the base; the spinous dorsal is low, the length of the twelfth spine
being one-fourth of that of the head. The free portion of the tail is a little longer
than high. Head greyish olive ; cheeks and body reddish olive; an irregular blackish
band proceeds from above the pectoral to a black spot in the middle of the root of the
caudal. A black spot above the origin of the lateral band. Opercles, back, and vertical
fins with black dots.

Two specimens, 7 inches long, were collected by Mr. Salvin in the River of Cahabon.

184. Heros sieBo.pil.
Kner & Stemdachner, Abhandl. bayer. Ak. Wiss. x. (1864) p. 13, Taf. 2. fig. 2.

This fish is probably not sufficiently distinct from H. godmanni; it is from New Gra-
nada, and the dark markings are arranged in irregular cross bands.

185. Heros currunatus. (Plate LX XVIII. fig. 3.)
Ginth. Proc. Zool. Soc. 1864, p. 152.

D. 7 A.5- L. lat. 33. L. transy.

Very closely allied to H. godmanni.

The fold of the lower lip is interrupted in the middle. The height of the body is
contained twice and three-fifths in the total length (without caudal), the length of the
head thrice and a fifth. Head as high as long. The upper profile of the head descend-
ing in a gentle curve. Scales on the cheek in four or five series. The first dorsal spine

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 467

is inserted behind the vertical from the upper end of the gill-opening. Dorsal spines
rather feeble, the length of the twelfth being two-sevenths of that of the head. Pectoral
two-thirds as long as the head. Upper parts blackish, each scale with a black base;
lower parts reddish, with a broad blackish band from behind the pectoral to the base
of the caudal; many scales within or below the band with a black spot in the upper
or lower angle; each scale on the side of the head with a black spot; chin and throat
violet. The spinous dorsal black, with yellowish margin; the soft parts of the vertical
fins with blackish spots.

This species inhabits the Lake of Amatitlan, where Mr. Salvin obtained numerous
examples up to 7 inches in length.

Description,—The profile of the head and nape forms a curve. Head as high as long ;
snout rather elevated; przorbital wider than the eye, the diameter of which is a little
more than one-fifth of the length of the head, and about three-fifths of the width of the
interorbital space. Cleft of the mouth narrow, horizontal, the upper jaw slightly over-
lapping the lower, and the maxillary not extending backwards as far as the anterior
margin of the orbit. The six front teeth of the outer series are the longest, deep
brown. ‘The orbit is considerably below the upper profile of the head, and somewhat
nearer to the end of the opercle than to that of the snout. Opercles scaly, the scales
on the cheek in four or five series, and smaller than those on the opercle. Vertical fins
not scaly at the base; the soft dorsal and anal do not reach far beyond the root of the
caudal. Free portion of tail a little higher than long. Caudal subtruncated, two-
ninths of the total length. Pectoral three-fourths as long as the head; ventral with
the outer ray slightly produced, rather longer than the pectoral, and reaching nearly to
the vent.

186. Heros rrrecutaris. (Plate LX XVIII. fig. 2.)
Theraps irregularis, Giinth, Fish. iv. p. 284.
Die Al SP Rlates5: . Le tranev 4/14.

The fold of the lower lip is interrupted in the middle. The height of the body is
nearly equal to the length of the head, which is two-sevenths of the total (the caudal fin
not included). Head longer than high, with the snout compressed and prominent; the
length of the snout is two-fifths of that of the head, and twice or more than twice the
width of the orbit. The cleft of the mouth is small, extending backwards somewhat
behind the vertical from the nostril; upper jaw slightly overlapping the lower; teeth
in a narrow band, those of the outer series largest. Praorbital wider than the orbit,
its width being equal to that of the interorbital space, which is rather convex. he eye
is situated immediately below the upper profile, its centre being a little behind the
middle of the length of the head. Scales on the cheek small, in six oblique series.
Scales on the opercles as large as those on the neck; those near the base of the dorsal

“and on the abdomen very small. Scales finely serrated. The dorsal fin commences
382

468 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA,

above the root of the ventral, and is not scaly. ‘The spines are of moderate length and
strength, the length of the fifteenth being two-sevenths, or in old examples one-third of
that of the head. The soft portion does not extend to the caudal, if laid backwards. The
free portion of the tail much longer than high. Caudal rounded. Pectoral shorter than
the head. The ventral does not extend on to the vent. Reddish olive, marbled with
blackish; the latter colour forming seven rather irregular transverse bands, some of
which extend on the dorsal fin. Belly silvery, marbled with blackish; opercles and
some scales on the body with blue dots. ‘The inner half of the soft vertical fins blackish
violet, the outer yellow; spinous dorsal with yellow margin. Lower side of head
blackish violet.

I have now before me numerous examples of this species from the Rivers Chisoy,
San Geronimo, and Santa Isabel; and finding that the anal spines are normally five in
number, the number four of the typical specimen being merely accidental, I do not
hesitate to reunite the genus Theraps with Heros. The largest example in the collection
is 8 inches long.

187. Heros interMepivus. (Plate LXXVIII. fig. 1.)
Ginth. Fish. iv. p. 298.
Dee MANES) igalaty BOle- Ma tramev5 ABs

The fold of the lower lip is interrupted in the middle; five or six series of scales on
the cheek. The height of the body is contained twice and three-fifths in the total
length (without caudal), the length of the head thrice and a fourth. The eye is not
very remote from the profile of the nape, which is curved. Head as high as long;
preorbital rather wider than the orbit. Cleft of the mouth rather narrow, horizontal,
with the jaws equal anteriorly. Base of the soft dorsal and anal with scarcely any
scales; dorsal spines of moderate length and strength, the length of the twelfth being
one-third or nearly one-third of that of the head’. ‘The soft dorsal and anal extend
slightly beyond the root of the caudal. The free portion of the tail is not quite so long
as high. Caudal subtruncated, its length being one-fifth of the total. Pectoral shorter
than the head, but rather longer than the ventral, which extends nearly to the vent.
Brownish, lower parts red in adult specimens; a broad angular brown band on the
trunk, its horizontal branch extending from the gill-opening to the vertical from the
first anal spine, whilst its vertical branch ascends to the hinder dorsal spines. Each
scale within this band with a black vertical streak. A rather narrow brown band runs
from the angular band to a blackish spot at the root of the caudal. Vertical fins with
whitish ocelli, enclosed by reddish streaks.

This species is closely allied to H. nebulifer and H. angulifer, from which it may be
distinguished by its colours, and by the size of its scales. It inhabits Lake Peten, where
specimens 5 and 6 inches long were collected by Mr. Salvin.

‘ These spines are represented a little too short in the figure.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 469

188. Heros ancunirer. (Plate LXXXYV. fig. 1.)
Ginth. Fish. iv. p. 298.

D. — Ahs 2. L. lat. 38. L. transv. 4/12.

The fold of the lower lip is interrupted in the middle; four series of scales on the
cheek. The height of the body is two-fifths of the total length (without caudal),
the length of the head three-tenths. The eye is not very remote from the profil of
the nape, which is slightly curved. Head as high as long; preorbital scarcely wider
than the orbit. Cleft of the mouth rather narrow, horizontal, with the jaws equal
anteriorly, and with the maxillary not extending backwards to the vertical from the
front margin of the eye. Dorsal and anal fins not scaly; dorsal spines of moderate
length and strength, the length of the twelfth being one-third of that of the head; the
soft dorsal and anal extend to the root of the caudal. The free portion of the tail is as
long as high. Caudal subtruncated, its length being not quite one-fifth of the total.
Pectoral shorter than the head, but longer than the ventral, which does not extend to
the vent. Brownish olive, with a broad angular black band on the trunk, its horizontal
branch extending from the eye to the vertical from the first anal spine, whilst its
vertical branch ascends to the hinder dorsal spines. Some scales within the band
and on the opercles with a black dot. A round blackish blotch on the root of the
caudal fin.

‘Two examples, 4 inches long, were collected by Messrs. Godman and Salvin at Yzabal.

PETENIA.
Ginth. Fish. iv. p. 301.

Body compressed, oblong, covered with ctenoid scales of moderate size. Dorsal
spines numerous, anal spines more than four; the soft dorsal scaleless. Teeth in a
band, small, conical. Anterior prominences of the first branchial arch short, com-
pressed, distant. Cleft of the mouth wide; jaws very protractile. Scales on the cheeks
small. ‘The origin of the ventral falls vertically below that of the dorsal.

189. PETENIA SPLENDIDA. (Plate LX XIX. fig. 2.)
Ginth. /. ec.

1843, 1D) = A. = L. lat. 41. L. transv. 6/17.

Scales on the cheek in about seven series. Greenish shining golden; head, body,
and vertical fins with black dots. A series of six or seven large round black spots along
the middle of the side, the last spot being edged with white, and situated on the upper
half of the root of the caudal.

Three examples were collected by Mr. Salvin in Lake Peten, the largest being 16
inches long.

NEETROPLUS.

This genus differs from Heros in having a front series of flat incisor-like teeth. It is

470 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

also closely allied to Etroplus, which genus, however, has but a rudimentary lateral line,
whilst in Neetroplus it. is as much developed as in Heros.

190. NrETROPLUS NEMATOPUS. (Plate LXXIV. fig. 4.)

Dc. A. iL, lat. 34... L, transv. 54/12,

. 10°
The fold of the lower jaw interrupted in the middle; five series of scales on the

cheek. Incisors = The outer ventral ray produced into a filament as long as the fin.

One specimen, 44 inches long, was discovered by Capt. Dow in Lake Managua.

Description —The height of the body is contained twice and three-fifths in the total
length (without caudal), the length of the head thrice and two-fifths. Head as high as
long, with an adipose prominence over the eye, which renders the profile of the fore-
head somewhat abrupt; snout rather compressed and prominent, the length of the
snout is two-fifths of that of the head, and more than the width of the orbit, which is
nearly one-third of the length of the head. Cleft of the mouth small, extending back-
wards somewhat behind the vertical from the nostril; jaws equal in front; teeth in a
band, those of the outer series being genuine incisors, which appear to be replaced by
smaller ones, standing behindin a band. Preeorbital wider than the eye, equal in width
to the interorbital space, which is convex. ‘The eye is situated at some distance from
the upper profile, nearer to the end of the opercle than to that of the snout. Scales on
the cheek small, in about five oblique series. Vosterior limb of preeoperculum about
twice as long as the inferior, and descending obliquely forwards. Scales on the
opercles as large as those on the neck; those near the base of the dorsal and on the
abdomen very small. The dorsal fin commences above the vertical from the hinder
margin of the operculum. Dorsal and anal scaly at the base. Spines rather strong,
the sixteenth dorsal spine being nearly one-half of the length of the head. The soft
portions of both fins are produced, and reach beyond the middle of the caudal. Caudal
truncated. Pectoral shorter than the head. Outer ray of ventral produced into a
filament as long as the fin. Brownish-olive, with irregular darker clouds.

MICRODESMUS.
Ginth. Proc. Zool. Soc. 1864, p. 26. :

Body much elongate, eel-like, covered with rudimentary scales. Head rather short,
with snout obtuse, cleft of the mouth narrow, and lower jaw prominent. Eyes minute.
Teeth in both jaws minute; palate toothless. ‘The gill-opening is reduced to a small
slit in front of the pectoral fin. Vertical fins united by a membrane; but the caudal
can be easily distinguished from the two other fins. Dorsal fin very long, composed of
flexible, undivided rays, like the anal. Pectorals short; ventrals thoracic, each reduced
toa single ray. Vent in the middle of the total length.

Iam not able at present to add anything to the knowledge of this fish which would
elucidate its natural affinities and indicate its systematic position.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 471

191. MicropESMUS DIPUS.
Gunth. /. ¢., pl. 3. fig. 2.

D. 55. A. 34. C.16. P.12. V.1.

The depth of the body is about one-eighteenth of the total length ; the length of the
head one-eleventh. The head is rather compressed, the snout short, the mouth very
narrow, and the lower jaw very prominent. The minute eye is lateral, and in the
anterior third of the length of the head. The dorsal fin commences at a distance from
the occiput which is somewhat less than the length of the head; it is nearly even, and
the rays are very distinct, the interradial membrane being thin and transparent. The
anal fin commences immediately behind the vent. The caudal rays are much more
slender and more closely set than those of the dorsal and anal; the caudal fin is
rounded, two-thirds of the length of the head. Pectorals as long as the ventrals, and
half as long as the head; the latter fins are close together, and inserted a little behind
the root of the pectoral. Upper parts uniform brownish olive.

‘The single specimen is 44 inches long, it was found by Capt. Dow at Panama.

192. Broruta MuLTIBARBATA (Schleg.).

Mr. Salvin has found on the Pacific coast of Guatemala a small fish, which I am
unable to distinguish from the Indian or Japanese species. However, having only small
examples for examination, I am not prepared to maintain the specific identity of these
fishes.

195. CitHARICHTHYS SPILOPTERUS. (Plate LX XX. fig. 2'.)
Giinth. Fish. iv. p. 421.
D. 76-78. <A. 60-63. L. lat. 47-80.

The height of the body is two-fifths of the total length (without caudal), the length
of the head two-sevenths. Scales of the lateral line subquadrangular; lateral line
nearly straight, gently descending anteriorly. Snout with the jaws equal in front,
rather longer than the eye, the diameter of which is one-sixth of the length of the
head. The maxillary, the length of which is contained twice and two-thirds in that of
the head, extends beyond the middle of the orbit. Anterior teeth of the upper jaw
widely set, much larger than the posterior, which are close together and very small ;
the lower jaw with seven or eight distant teeth of moderate size on each side. Eyes
separated by a very narrow scaleless ridge, their front margins being nearly on the same
level. Fin-rays scaly. The dorsal commences a little before the upper eye, and
terminates close by the caudal ; its longest rays are behind the middle, and one-half of
the length of the head. Anal spine none. Caudal rounded ; its length is one-sixth of
the total. ‘The pectoral is rather longer than half the length of the head; ventral
much shorter, extending beyond the origin of the anal. Gill-rakers lanceolate, pointed,

' The artist has unfortunately omitted to reverse the view of this figure.

472 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

one-third as long as the eye. Greenish olive (in spirits); a series of distant blackish
spots along the basal portions of the anal and dorsal fins.

This species occurs on the shores of the tropical parts of the Atlantic, and has been
found by Mr. Salvin also on the Pacific coast, at Chiapam.

196. CrrHARICHTHYS GUATEMALENSIS.
Bleeker, Nederl. Tydschr. Dierk. 1864, p. 73.
D. 77. ~A. 57. “L. Tat. ca. 45.

“(Citharichthys corpore ovali, altitudine 23 circiter in ejus longitudine; capite 43
circiter in longitudine corporis, eque alto circiter ac longo; oculis sinistris, minus
diametro + distantibus, superiore quam inferiore vix majore 5 circiter in longitudine
capitis, paulo ante inferiorem prominente; linea frontonuchali declivi rectiuscula;
naribus non tubulatis, utroque latere approximatis; rictu curvato; maxillis subzquali-
bus, superiore usque ante oculi inferioris marginem superiorem adscendente, sub oculi
inferioris partem posteriorem desinente, 22 circiter in longitudine capitis; dentibus
maxillis conicis acutis parvis, caninis nullis, utroque latere maxilla superiore numerosis
postrorsum magnitudine decrescentibus, maxilla inferiore parcioribus, intermaxillaribus
majoribus subequalibus; praoperculo obtusangulo angulo rotundato; squamis pre-
operculo in series verticales 8 circiter, operculo in series verticales 6 vel 7 dispositis,
linea laterali corpore antice parum declivi; capite regione oculo-temporali conspicua
obliqua; prima dorsali ante medium oculum superiorem incipiente et vix ante pinnam
caudalem desinente radiis longissimis corpore plus triplo humilioribus; pinnis pecto-
ralibus et ventralibus (ex parte abruptis) ; anali dorsali vix humiliore; caudali postice
angulata 53 ad 5$ in longitudine corporis; colore corpore latere oculari viridescente (‘),
latere anophthalmo albido, pinnis flavescente (?).”

The specimen, which is 145 mm. long, and in the Leyden Museum, is stated to be
from Guatemala.

197. Hemrruomeus ovauis. (Plate LXXX. fig. 1")
Giinth. Proc. Zool. Soc. 1864, p. 154.
D. 86. A. 69, L. lat. 58.

The dorsal commences before the eye. Teeth of the upper jaw in a double series,
with one or two pairs of small canine teeth in front; those of the lower jaw closely set,
conical, in a single series. Scales largest in the middle of the body, adherent, ciliated ;
lateral line ascending gently over the pectoral fin, straight for the rest of its length.
The height of the body is contained twice and two-thirds in the total length; the length
of the head four times and two-thirds. Snout rather shorter than the eye, the diameter
of which is two-ninths of the length of the head. Jaws equal anteriorly; the length of

* The artist has unfortunately omitted to reverse the view of this figure.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 473

the maxillary is a little more than one-third of the length of the head; maxillary scaly.
Interorbital space very narrow, concave, one-third of the vertical width of the orbit;
the concavity is produced by two ridges convergent posteriorly. Head nearly entirely
covered with ciliated scales. Rays of the vertical fins scaly, the distance between the
dorsal and caudal fins is one-half of the depth of the free portion of the tail. The
longest dorsal rays are somewhat behind the middle of the fin, and four-ninths of the
length of the head. Pectoral rays not prolonged. Body nearly uniform reddish olive ;
some of the dorsal, anal, and caudal rays with elongate dark-brown spots.

One specimen, 7 inches long, was collected by Messrs. Dow and Salvin on the Pacific
coast uf Panama.

198. PsEUDORHOMBUS BRASILIENSIS.

Hippoglossus brasiliensis, Ranzani, Comm. Bonon. v. p. 10, tab. 3.
Rhombus aramaca, Casteln., not Cuv.
Pseudorhombus vorax, Giinth. Fish. iv. p. 429.

This species is known to occur on the coast of Brazil; however, there is a specimen
in the British Museum, which formed part of a collection containing numerous fishes
from Guatemala; and I mention it, therefore, in this list to draw attention to this species.

200. APHORISTIA ORNATA, Var. ELONGATA.

Two examples, 5 inches long, collected by Mr. Salvin on the Pacific coast of Panama,
differ, from specimens from the West Indies, only in having the body more elongate, its
depth being contained four times and two-thirds in the total length (with the caudal).
The number of fin-rays is the same, viz. D. 97, A. 82; L. lat. 98.

201. AmiuRUS MERIDIONALIS. (Plate LX XXL. fig. 1.)

Ginth. Fish. v. p. 102.
D. 1/6. A, 28-29. P.1/9.

Head one-half or one-third longer than broad; the maxillary barbels extend to the
end of the head. The length of the dorsal spine is somewhat less than that of the head
without snout, and nearly equal to that of the pectoral spine. Adipose fin short. The
height of the body is one-fifth of the total length (without caudal), the length of the
head one-fourth or two-ninths. Snout obtusely rounded, with the upper jaw longer
than the lower. The diameter of the eye is one-half or two-fifths of the extent of the
snout, and one-third or two-sevenths of the length of the postorbital portion of the
head. ‘The band of maxillary teeth is five or six times as broad as long. The outer
mandibulary barbels extend to the posterior margin of the gill-‘membrane. The distance
of the dorsal spine from the snout is a little more than one-half of its distance from the
caudal fin; it is finely serrated behind. The length of the base of the adipose fin equals
that of the dorsal. Caudal fin deeply forked; the upper lobe is somewhat the longer,

VOL. VI.—PART VII. 3T

474 DR, GUNTHER ON THE FISHES OF CENTRAL AMERICA.

its length being equal to, or a little less than, that of the head. The anal fin terminates
behind the adipose fin, and its last rays do not extend to the base of the caudal. Axil
of the pectoral with a very distinct porus mucosus. ‘The pectoral spine is serrated
interiorly, sometimes a little longer, sometimes a little shorter, but always rather stronger
than that of the dorsal fin. Pectoral fin longer than ventral, two-thirds or three-fifths
ot the length of the head. The ventral extends to the origin of the anal. Upper parts
brownish, with steel-blue reflexions; lower parts silvery, with a reddish tinge.

Three examples were collected by Mr. Salvin in the Rio Usumacinta; the largest is
15 inches long.

205. PIMELODUS WAGNERI.
Pimelodus cinerascens, Kner & Steindachner, Abhandl. bayer. Akad. x. p. 49 (not Gthr.).
D. 1/6. <A. 11-12.

Head covered with soft skin above. Adipose fin one-third of the total length (without
caudal). ‘The maxillary barbels extend beyond the root of the yentrals; the outer ones
of the mandible do not quite reach the base of the pectorals. ‘The length of the head
is contained from five times and two-fifths to five times and seven-eighths in the total
(with the caudal), the height of the body seven times and one-third, or seven times and
one-half. Upper jaw projecting beyond the lower, ‘The eye is equidistant from the
end of the snout and from that of the gill-cover; its diameter is one-seventh or one-
eighth of the length of the head, and contained twice and two-thirds or twice and
three-fourths in the width of the interorbital space. Dorsal fin with the spine very
feeble, as high as long. Pectoral fin two-thirds as long as head. Porus axillaris
small. Coloration uniform, a darker streak along the lateral line; dorsal fin with the
usual whitish cross band, and sometimes with a large round black spot between the
last two rays.

Pacific and Atlantic rivers of Panama.

The complete diagnosis of P. cinerascens (Gthr.), accompanied by a most accurate
figure, proves at once that the species discovered by Hr. Wagner is distinct from it. It
appears to be nearest to P. godmanni.

206. PIMELODUS MANAGUENSIS. ,
D. 1/6. ; A, 14-15, P.1/9.

Head covered with soft skin above; occipital process styliform, not extending to the
basal bone of the dorsal spine, Adipose fin very long, rather more than one-third of
the total length (without caudal); its distance from the dorsal is equal to the length of
the base of the latter. ‘The maxillary barbels are rather short, extending nearly to the
base of the dorsal fin, the outer ones of the mandibles reach beyond the root of the
pectoral. The height of the body is contained six times in the total length (without

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. AT5

caudal); the length of the head five times. The lower jaw is slightly shorter than the
upper. Interorbital space flat, its width being less than twice the width of the eye.
Dorsal fin with the spine very feeble, somewhat higher than long. Pectoral fin rather
short, as long as the head, without snout; its spine about double the length of the
humeral spine. Porus axillaris distinct. Ventral rather longer than the pectoral.
Anal fin with the base longer than that of the dorsal; its rays do not extend nearly to
the end of the adipose fin if laid backwards. Caudal cleft to the base ; its upper lobe
less rounded and narrower than the lower one, which is one-seventh of the total length.
Coloration uniform, dorsal fin with a whitish cross band.
One specimen, 9 inches long, was obtained by Capt. Dow in the Lake of Managua.

214. ARIUS ASSIMILIS.
Gunth. Fish. v. p. 146.
ID); Wt 2A WO), TR IVAN

The height of the body is contained four times and two-thirds in the total length
(without caudal), the length of the head thrice and three-fifths; head much broader
than high, its greatest width being three-fourths of its length. Eyes rather small,
situated nearer to the end of the snout than to that of the operculum; the length of
the snout is three-fifths of the width of the interorbital space. The median longitudinal
fonticulus on the upper side of the head does not extend to the base of the occipital pro-
cess. Teeth onthe vomer but slightly separated in the middle, forming 4
a pair of onl transverse patches, which are confluent il those on LES
the palatine bones; the latter are short, club-shaped. ‘The band of iD
intermaxillary teeth is five times as broad as long. All the teeth villi- i A
form. ‘The maxillary barbels extend nearly to the end of the head; the
length of the outer ones of the mandible is one-half or two-thirds of that of the head.
Crown of the head granular, the granulations being arranged in radiating streaks.
Occipital process broader than long, triangular, with its hinder end concave. ‘The
basal bone of the dorsal spine of moderate size, crescent-shaped. Dorsal spine of
moderate strength, more than half as long as the head, granulated in front and slightly
* serrated behind; the first soft ray is longer than the spine and as high as the body.
Adipose fin shorter than the dorsal, Caudal deeply forked, with the upper lobe
longest, its length being contained five times and a half in the total. Pectoral spine
serrated along its inner edge and on the extremity of the outer edge. Ventral fin
shorter than pectoral. Sides of the body silvery; vertical fins greyish; basal half of
the inner side of the paired fins black.

One example, 13 inches long, was obtained by Messrs. Godman and Salvin in the
Lake of Yzabal.

Hexanematichthys hymenorrhinos, Bleek. Vers]. & Mededeel. Akad. Wetensch. Amsterd.
1862, xiv. p. 377, appears to be closely allied to the above species; and we should not

P
a7 2

476 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

hesitate to refer our specimens to it, if the barbels of Bleeker’s species were not much
longer, those of the maxillaries extending on to the base of the ventral fin, and the
outer ones of the mandible to the base of the pectoral. ‘The specimen in the Leyden
Museum is 8} inches long.

219. ARIUS DOVII.

Mr. Gill (Proc. Acad. Nat. Sc. Philad. 1863, p. 170) describes a species discovered
by Capt. Dow on the Pacific coast of Central America, under the name of Leptarius
dowii. Leptarius is a distinct genus, according to Mr. Gill, characterized by having
the band of teeth quadripartite, the head granulated and without lateral fontanelles, a
slender body, and a very slender caudal peduncle, the anal fin rather low and oblong,
the thin adipose fin extending behind the anal, and the fins little developed.—The
species is not described; but detailed comparative measurements of the single example
(which is 5? inches long) are given.

222. ARLURICHTHYS NUCHALIS. (Plate LXXXI. fig. 2.)
Giinth. Fish. v. p. 179.
ID Wee alg AAO LEE A

The height of the body is rather less than the length of the head, which is two-ninths
of the total (without caudal); the greatest width of the head is three-fourths of its
length ; snout longer than the eye, the diameter of which is rather less than one-fourth
of the length of the head. The vomerine band of teeth is separated in the middle by a
short interspace, each half being as broad, and long as the palatine band, with which it
is subcontinuous. The maxillary barbels extend to the root of the ventral, those of the
mandible nearly to the pectoral. The dorsal buckler is as broad behind as in front,
with rounded lateral margins, each half being bent downwards on the side. Dorsal fin
narrow and elevated, the first ray being considerably longer than the spine, which is as
long as the head without snout; pectoral spine equal to the dorsal spine. The origin
of the anal fin is much nearer to the base of the caudal than to that of the pectoral.
The first pectoral ray is produced into a long filament reaching beyond the origin of the
anal. Ventrals extending beyond the vent, their length being three-fifths of that of the
head. Indescent blue above, silvery below.

One example, 11 inches long, was obtained by Messrs. Salvin and Dow on the Pacific
coast of Panama.

223. ASLURICHTHYS PANAMENSIS.
Gill, Proc. Ac. Nat. Se. Philad. 1863, p. 172.
This species is described thus :—
Dislvii.o A.c27./gbsal /l3.

The greatest height is contained five times in the length to the base of the caudal fin,

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. AT7

and six times and a half in the total. The height of the caudal peduncle equals half
the interorbital area, and is half its length behind the anal fin. The smooth head
enters four times in the length to the middle of the central caudal ray, and nearly five
times in the total. The width of the head enters once and one-third in its length, and
the width of the interorbital area once and two-thirds. The eye is elliptical, its diameter
equals a fourth of the head’s length, and the distance from the anterior nostril is equal
to it. The maxillary barbels extend backwards nearly to the anus, and the mental to
the bases of the pectoral fins. The dorsal buckler is rather longer than wide, with its
anterior margin concealed, and its lateral and posterior very conspicuous, rounded
towards the posterior angles, and emarginated behind; the sides slope and form a
rectangle, and the surface is filled with deep oblong pits. ‘he anal fin is situated
midway between, or scarcely in advance of, the central point between the bases of the
pectoral and caudal fins; it is oblong, and equals or nearly equals the width of the
head. The pectoral filaments extend about to the middle of the anal fin; the ventrals
are inserted midway between the lower jaw and base of caudal, and extend backwards
to the anus, entering twice and a half in the head’s length. The colour above is
plumbeous; the pectorals thickly dotted with black on their inner faces, and the anal
less so.

One specimen, 8 inches long, has been collected by Capt. Dow on the Pacific coast ot
Panama.

224. PLECOSTOMUS BICIRRHOSUS (Gronoyv.).

Messrs. Kner & Steindachner (Abhandl. bayer. Ak. x. p. 60) mention a species of this
genus from Pacific and Atlantic rivers of Panama, which they regard as a variety of
Plecostomus bicirrhosus, but which differs in several respects.

225. CH&TOSTOMUS ASPIDOLEPIS.
D.1/7. A.5. P.1/6. L. lat. 26.

Head large, depressed, a little longer than broad, its length being contained thrice
and one-third in the total (without caudal); snout very broad, rounded in front. Inter-
orbital space nearly flat, with a very slight rising along the middle. Orbit small, its
diameter being one-third of the width of the interorbital space. Margin of the snout
granulated. Interoperculum with very few, and for the greater part short, setiform
spines, the longest of which is about half as long as the orbit. Thorax and belly
granulated, with naked patches. Dorsal fin higher than long, the length of its anterior
rays being nearly equal to that of the head; the length of its base equals its distance
from the hinder axil of the adipose fin. There are seven scutes between the two
dorsal fins. The pectoral spine is strong, rather longer than the head, covered behind
with setiform spinules. The ventral fins extend somewhat behind the anal. Twelve
scutes between anal and caudal. Scutes of the body with a prominent keel, each keel

478 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

with from four to seven short setiform spines. Posthumeral ridge rather distinct. Each
scute is variegated with dirty yellow and dark brown.
I have received of this species only a single skinned example, 123 inches long; it is
from Veragua.
226. CumTostomus cirRHOsUS (Val.).
Messrs. Kner & Steindachner (/. ¢. p. 61) mention this species from the Rio Chagres ;
but their species is probably distinct from it.

227. LORICARIA URACANTHA.
Kner & Steindachner, Abhandl. bayer, Akad. x. p. 56, Taf. 6. fig. 3.

Snout broad, of moderate length; eye of moderate size, with a notch in its posterior
margin, its horizontal diameter is one-half of the width of the interorbital space, which
is slightly concave, owing to the raised supraorbitals. Eight or ten rather large bifid
teeth in each jaw. Labial folds broad, with numerous papille, and a small lateral
barbel. ‘The lower side of the head naked; scutes of the neck but very indistinctly
bicarinate. L.lat.27. There are seven lateral scutes between the pectoral and ventral
fins. Thorax and belly with numerous smali irregular scutes. The origin of the dorsal
is opposite to that of the ventrals. ‘The length of the outer pectoral ray is contained
six times and a half in the total (without caudal). The upper caudal ray very thick and
strong. Rays of all the fins spotted.

Pacific and Atlantic rivers of Panama.

229. Macropon MIcROLEPIS (Gthr.).

The fish described by Messrs. Kner and Steindachner (/. ¢. p. 28) under the name of
M. tareira belongs to this species.

256, TETRAGONOPTERUS NEUS (Gthr.).
This species has been recognized in a collection from Panama by Messrs. Kner and
Steindachner (/. c. p. 46).

237. CHALCINOPSIS DENTEX. (Plate LXAXXII. fig. 1, nat. size.)
Brycon dentex, Giinth. Proc. Zool. Soc. 1860, p. 240.
Chalcinopsis dentex, Giinth, Fish. y. p. 337.

D.11. <A. 38-36, L. lat. 48-55. LL, transv. Ea Vert. 23/22.

‘The height of the body is contained thrice and one-fourth or thrice and one-third in
the total length (without caudal), the length of the head four times and one-third or
four times and two-thirds, The maxillary does not quite extend to below the centre of
the eye. Snout as long as the eye in young examples, but much longer in adult ones,
Interorbital space convex, its width being much more than the diameter of the eye in

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 479

old specimens. ‘The origin of the dorsal fin is nearer to the root of the caudal than to
the extremity of the snout; its hinder rays are vertically above the anterior anal rays.
The free portion of the tail is considerably longer than high. Caudal deeply forked.
The pectoral extends to, or nearly to, or a little beyond, the ventral. Silvery, sometimes
with a reddish hue; a part of the scales have sometimes a black margin, or are spotted
with black; humeral part of the gill-opening black; sometimes a black spot at the root
of the caudal. Anal fin generally with a black margin.

Specimens, up to 16 inches in length, were collected by Messrs. Salvin and Godman in
the Rio Motagua and Usumacinta, and at Yzabal. The species occurs also in Ecuador.

240. ANACYRTUS GUATEMALENSIS. (Plate LXXXII. fig. 4.)

Anacyrtus (Reboides) guatemalensis, Giinth. Fish. v. p. 347.
D.11. A. dl. V.8. L. lat. 80. L. transv. 19/22.

Upper and lower jaw anteriorly on each side with a short, conical, tooth-like process
directed forwards; teeth in the intermaxillary, maxillary, and mandible in a single,
rather irregular series; no canine teeth in the upper jaw, those in the lower small and
short. Back elevated, the upper profile of the head and nape forming an S-shaped
curve. The height of the body is contained twice and three-fourths in the total length
(without caudal), the length of the head four times. The lower jaw is rather shorter
than the upper; the maxillary extends to the vertical from the centre of the eye. ‘The
width of the interorbital space is a little less than the diameter of the eye, which is
two-sevenths of the length of the head. The humeral process in front of the pectoral
terminates in a point anteriorly and posteriorly. The origin of the dorsal fin is a little
nearer to the extremity of the snout than to the root of the caudal, above the fifth or
sixth anal ray; caudal deeply forked; the ventral is inserted below the middle of the
pectoral, which extends nearly to the origin of the anal. Light reddish olive with a
silvery lateral band.

Specimens, up to 6 inches in length, were collected by Mr. Salvin at Huamuchal.
and in the Chagres River.

244, Exocamtus cALLopTeRus. (Plate LX XXIII.)
Ginth. Fish. vi. p. 292.
D, 11-12. A. 8. . L. lat. 46.

Body stout, its height being one-fifth of the total length (without caudal), the length of
the head being somewhat less than one-fourth. The depth of the head equals the distance
between the extremity of the snout and the hind margin of the preoperculum, Snout
obtuse and depressed, three-fifths of the length of the diameter of the eye, which is
one-third of the length of the head, and less than the width of the interorbital space,
which is slightly concave. The pectoral fin extends to the end of the dorsal. Ventral

480 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

fins midway between preoperculum and root of the caudal, extending nearly to the end
of the base of the anal. The dorsal commences far in advance of the anal, its anterior
rays being half as long as the head. The distance between the first dorsal ray and the
Jirst rudimentary caudal ray equals the length of the head. here are thirty-four scales
between the occiput and origin of the dorsal, and nine longitudinal series of scales
between the origin of the dorsal and the lateral line. Pectoral with numerous small
roundish blackish-brown spots and with the lower and upper rays whitish. Ventral
white, the middle rays greyish.

Two examples, 10 inches long, were obtained by Capt. Dow on the Pacific coast of
Panama.

CHARACODON.
Ginth. Fish. vi. p. 308.

Cleft of the mouth small, developed laterally and horizontally ; mandible short, with
the bones of each side firmly united. Snout short. ‘Teeth rather small, bicuspid, in a
single series; but there is a narrow band of villiform teeth behind the series of incisors.
Scales of moderate size. Origin of the anal fin opposite, or nearly opposite, to that of
the dorsal. Sexes not differentiated. Intestinal tract but slightly convoluted.

247. CHARACODON LATERALIS. (Plate LX XXII. fig. 2, fem.)
D. 10-11. <A. 13 in fem., 15-16 in male. L. lat. 35. L. transv. 12.

In general habits very similar to a Cyprinodon. Body rather elevated, with the neck
somewhat arched, its greatest depth being rather more than the length of the head,
and one-third of the total (without caudal). Head thick and broad, with the snout
obtuse, as long as, or rather longer than, the diameter of the eye, which is one-fourth
or two-ninths of the length of the head. The mandible ascends obliquely, and is longer
than the eye. There are about twenty smallish teeth in each jaw, their apex is indistinctly
notched. Interorbital space flat, its width being two-fifths of the length of the head.
The origin of the dorsal fin is a little nearer to the end of the caudal than to the
occiput, and a little behind that of the anal. Both fins are small and rounded. In the
male the six anterior rays of the anal are of nearly equal length, but considerably
shorter than the following, forming a very distinct portion of the fin; all these rays
are very closely set. Caudal fin small, truncate or slightly convex. The distance
between dorsal and caudal is somewhat more than the least depth of the tail, and equal
to the distance between eye and gill-opening. Brownish olive (in spirits), with a darker
band running from the eye to the root of the caudal: this band is sometimes broken up
into a more or less regular series of brownish-black spots.

There are several examples, from 13 to 2} inches long, in the British Museum; they
are from Dr. Seemann’s collection, who obtained them in Southern Central America.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 481

248. Hapiocuitus povit. (Plate LX XXII. fig. 5.)
Ginth. Fish, vi. p. 315.
D. 8. A. 14. V.6. L. lat. 31. L. transv. 8.

The height of the body is contained five times in the total length (without caudal),
the length of the head thrice and two-thirds ; head elongate, low, and depressed, with
the snout much produced and the upper jaw somewhat longer than the lower; the eye
occupies exactly the middle of the length of the head, its diameter being two-ninths of it,
and more than one-half of the width of the interorbital space, which is flat. The origin
of the dorsal fin is a little nearer to the extremity of the caudal than to the gill-opening,
and corresponds to the twenty-third scale of the lateral line. Anal fin entirely before
the dorsal ; pectoral extending to ventral, which reaches the vent; caudal rounded; all
the fins well developed. Light brownish olive; posterior half of the dorsal and anal fins
with black cross bands; basal half of the caudal with round light spots.

Two specimens, 6 inches long, probably males, were collected by Capt. Dow at Punta
Arenas, Costa Rica.

249. FUNDULUS LABIALIS. (Plate LX XXIV. figs. 1 & 2.)
Ginth. Fish. vi. p. 319.

B. 6. D. 13-14. A. 16-17. V.6. L. lat. 37-39. L. transy. 15.

The height of the body, taken on the level of the base of the pectoral, is two-ninths
of the total length (without caudal). Head rather depressed, its length being contained
four times or four times and a third in the total. Interorbital space broad, slightly
convex, its width being less than one-half of the length of the head. Snout broad,
obtuse, depressed, with the jaws perfectly equal in front; mandible very short, not
longer than the eye. Upper lip well developed, broad, extending to the angle of the
mouth. The diameter of-the eye is less than the length of the snout, or than one-
fourth of that of the head, and, in females, one-half of the width of the interorbital
space, whilst in males the forehead is somewhat narrower. The origin of the dorsal
fin is midway between the extremity of the caudal and the orbit, and corresponds to the
first ray of the anal. Dorsal fin as high as long in both sexes; anal fin rounded in the
male, scarcely higher than long; much elevated in the female, the length of its base
being two-thirds only of its depth. Genital opening of the female immediately in
front of, but disconnected from, the anal fin. Basal third of the caudal fin (which is
subtruncate) scaly. Body uniform brownish olive, paler below ; sometimes irregular
cloudy markings on the tail. Fins immaculate; the anal fin of the male is black at the
base, and bright yellow on its marginal half; also the upper margin of the dorsal fin of
the same sex is yellowish.

Numerous examples, up to 45 inches long, were collected by Messrs. Salyin and
Godman in the Rio San Geronimo and at Yzabal. Figure 1 represents the male, and
fig. 2 the female. !

VOL. VI.—PART VII. 30

482 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

250. Funputus puncratus. (Plate LX XXIV. fig. 5.)
Ginth. Fish. vi. p. 320.
D.12. A. 13. V.6. L. lat. 34. L. transv. 12.

The height of the body, taken on the level of the base of the pectoral, is two-ninths
of the total length (without caudal). Head depressed, its length being one-fourth of
the total. Interorbital space very broad, slightly convex, its width being one-half of
the length of the head. Snout broad, obtuse, much depressed, with the lower jaw
scarcely projecting beyond the upper; mandible longer than the eye. Upper lip of
moderate breadth, not extending to the angle of the mouth. The diameter of the eye
is less than the length of the snout, two-ninths of that of the head, and less than one-
half of the width of the interorbital space. The origin of the dorsal fin is somewhat
nearer to the extremity of the caudal than to the orbit, and corresponds to the nine-
teenth scale of the lateral line. The first anal ray is opposite to the third of the dorsal.
Dorsal and anal fins subquadrangular, with the outer margins convex: both are a little
longer than high. Caudal fin subtruncate, scaly on its basal half. Pectoral fins shorter
than the head without snout, not extending to the base of the ventrals. Brownish
olive, paler below, each scale, especially those on the tail, with a vertical dark purplish
violet spot on the centre. Dorsal with three or four series of blackish dots, anal with
a whitish margin.

A single male, 34 inches long, was obtained by Mr. Salvin at Chiapam.

251. FUNDULUS GUATEMALENSIS. (Plate LXXXLV. figs. 3 & 4.)
Ginth. Fish. vi. p. 321.
D.12 (13). A. 14-15 (16). L. lat. 32-35. LL. transv. 12.

The height of the body, taken on the level of the base of the pectoral, equals the
length of the head, and is one-fourth or rather more than one-fourth of the total length —
(without caudal). Head thick and broad; interorbital space broad, slightly convex, its
width being a little less than one-half of the length of the head. Snout broad, obtuse,
with the lower jaw slightly projecting beyond the upper; mandible longer than the eye.
The diameter of the eye is equal to, or, in the larger specimens, less than the length of
the snout, one-fourth of that of the head, and one-half of the width of the interorbital
space. ‘The origin of the dorsal fin is midway between the extremity of the caudal and
the posterior margin of the orbit, and corresponds to the nineteenth scale of the lateral
line. The first anal ray corresponds to the second of the dorsal. Dorsal and anal fins
snbquadrangular, rather low, longer than high in the male, and as long as high in the
female. Two-thirds of caudal covered with small scales. Brown above and on the
sides, pale below; females with a very indistinct dark band along the side. Fins
immaculate; anal with a light margin.

The sexual opening of the female is not attached to the anterior anal rays

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 483

Numerous examples, up to 34 inches long, were collected by Mr. Salvin in the Lakes
of Duefias and Amatitlan, and in the Rio Guacalate. This species occurs also in
Western Ecuador. Figure 3 represents the male, and fig. 4 the female.

252 FuNpULUS PACHYCEPHALUS. (Plate LX XXIV. fig. 6.)
Giinth. Fish. vi. p. 321.

This species is closely allied to F. guatemalensis, but has a thicker head and
smaller eye.

DBA TAS tbe VeOy I lat. 35. I: transy. 12.

The height of the body, taken on the level of the base of the pectoral, is contained
thrice and one-fifth or thrice and four-fifths in the total length (without caudal). Head
very thick and broad, its length being contained thrice and one-third in the total.
Interorbital space very broad, slightly convex, its width being one-half of the length
of the head. Snout broad, obtuse, with the lower jaw slightly projecting beyond the
upper; mandible longer than the eye. The diameter of the eye is less than the length
of the snout, one-fourth of that of the head, and one-half of the width of the inter-
orbital space. The origin of the dorsal fin is midway between the extremity of the
caudal and the anterior or posterior margin of the orbit, and corresponds to the six-
teenth scale of the lateral line. The first anal ray corresponds to the third of the
dorsal. Dorsal and anal fins subquadrangular, of moderate height, the latter fin being
scarcely higher than long. Caudal fin subtruncate. Brownish above and on the sides,
each scale darker on the tip; an indistinct dark band along the middle of the tail.
Fins immaculate, anal with the lower margin whitish.

Three males, 24 inches long, were obtained by Mr. Salvin in the Lake of Atitlan.

254. GAMBUSIA NICARAGUENSIS. (Plate LX XXII, fig. 3, fem.)
Giinth. Fish. vi. p. 336.
D. 8. A.10. L. lat. 29. L. transv. 8.

The height of the body is contained thrice and a third in the total length (without
caudal), the length of the head thrice and two-thirds. Snout broad, subspatulate, with
the lower jaw projecting beyond the upper. The diameter of the eye is a little more
than the length of snout, one-third of that of the head, and three-fifths of the width of
the interorbital space. In the female the origin of the dorsal fin is somewhat nearer to
the extremity of the caudal than to the end of the snout, and opposite to the last ray
of the anal fin. Pectoral fins not quite reaching as far backwards as the ventrals,
which terminate immediately in front of the anal fin. Free portion of the tail rather
short, the length of the base of the anal fin being one-half of its distance from the
caudal fin. Brownish olive above, sometimes with series of black dots along the rows

aU 2

484 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

of scales. Dorsal and caudal fins crossed by series of black dots; middle of anal
blackish.

Several females, 14 or 2 inches long, were obtained by Capt. Dow in the Lake of
Nicaragua.

259. PaciLia ELONGATA. (Plate LXXXYV. fig. 2, fem.)
Ginth. Fish. vi. p. 342.
D.9. A.8. L. lat. 30-32. L. transv. 9.

The height of the body is contained four times in the total length (without caudal),
the length of the head four times and a third. The free portion of the tail is elevated,
its least depth being contained once and two-thirds in its own length, and nearly equal
to the length of the head without snout. The diameter of the eye equals the length
of the snout, is two-sevenths of that of the head, and more than one-half of the width
of the interorbital space. Snout much depressed. Lower jaw with a single series of
very small teeth only; and also in the upper the posterior band of villiform teeth is
very indistinct. Origin of the dorsal fin nearer to the root of the caudal than to the
occiput, a little behind that of the anal, above the fourteenth scale of the lateral line.
Dorsal fin higher than long, its longest ray being as long as the head without snout.
Anal small. There are eight longitudinal series of scales on each side of the tail.
Caudal rounded, its base only covered with scales. Uniform greenish ; the membrane
of the pouches of scales with a blackish margin. Fins immaculate.

This species is known from a female only, 5 inches long; it was obtained by Capt.
Dow at Panama.

260. Pacinia PeTENENSIS. (Plate LXXXV. fig. 3, male; fig. 4, fem.)
Ginth. Fish. vi. p. 342.
D.11.° A. 8. L. lat. 29-30. L. transv. 8-9. Vert. 16/14.

‘The height of the body (measured below the anterior dorsal rays) is contained four
times and one-fifth in the total length (without caudal) in females, and thrice and one-
fourth in males, the males having the body much higher and shorter. ‘The length of
the head is one-fifth of the same length in the female, and one-fourth in the male. ‘The
diameter of the eye is a little less than the length of the snout, two-sevenths or one-
fourth of that of the head, and somewhat more than one-half of the interorbital space.
The origin of the dorsal fin is further distant from the root of the caudal than from the
occiput, and corresponds to the eleventh or twelfth scale of the lateral line. Origin of
the anal opposite to the fourth ray of the dorsal (in the female). Dorsal tin of moderate
size; anal rather small, but pointed; caudal scaly in its basal third. ‘The free portion
of the tail is compressed, rather high, its least depth being one-half of its length, and
equal to the length of the the head without snout. ‘here are seven longitudinal

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 485

series of scales on each side of the tail. Lateral line rather indistinct. Greenish, each
scale with a black vertical spot. Dorsal and basal half of the anal irregularly and finely
marbled with brown.

The male has the dorsal fin somewhat elevated, the longest ray being rather longer
than the head. Anal fin modified into an intromittent organ, and advanced to between
the ventrals, in front of the dorsal. ‘Tail strongly compressed, and much higher than
in the female, its least depth being equal to the length of the head.

Five examples, up to 6 inches in length, were collected by Mr. Salvin in Lake Peten.

262. Paci.ia GILLI.

Messrs. Kner & Steindachner (Abhandl. bayer. Akad. Wiss. x. p. 25, Taf. 4. fig. 1) have
described a species of this genus from the Rio Chagres under the name of Xiphophorus
gilli. It would appear to be most closely allied to P. dovit; but there are some appa-
rently slight differences, which have induced me to keep the two species distinct until I
shall have had an opportunity of comparing specimens from the Rio Chagres with the
typical examples of P. dovit.

264. MOLLIENESIA PETENENSIS. (Plate LXXXVI. figs. 1-3.)
Giinth. Fish. vi. p. 348.

B.6. D.15. A. 8-9. L. lat. 30. L. transv. 10. Vert. 17/13.

The height of the body is one-third of the total length (without caudal), the length
of the head one-fourth or two-ninths. ‘The diameter of the eye is equal to the length of
the snout, two-sevenths of that of the head, and rather less than one-half of the width
of the interorbital space. The length of the dorsal fin of the male is one-half of the
distance between eye and root of the caudal, in the female two-fifths; caudal rounded,
with scales at the base only. The free portion of the tail is as high as long, and
covered by nine longitudinal series of scales on each side. Lateral line very indistinct.
Greenish, or brownish green, silvery below ; a dark spot to each scale of the upper and
middle caudal series and the lower part of the trunk. Dorsal fin of the adult male with
small irregular brown lines or spots, and with a row of large rounded spots along the
middle of its height. Interradial membrane of the caudal with numerous black dots;
the lower part of the hind margin black. Females and immature males have the
dorsal fin simply ornamented with small irregularly curved brown spots.

Three examples, up to 5 inches in length, were collected by Mr. Salvin in Lake Peten.

Figure 1 represents the adult male, fig. 2 the immature male, and fig. 3 the adult
female, all of the natural size.

265. XIPHOPHORUS HELLERI (Heck.). (Plate LXX XVII. figs. 2-6.)
This species varies considerably in coloration. Two varieties occur in the river
Chisoy—one with two yellowish green bands along the side, separated, and bordered

486 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

above and below by a blue band; the second, without bands, has the body covered all
over with irregular black spots.

‘The figures are of the natural size. The specimens are from an affluent of the
Chisoy River,—fig. 4 representing an adult male of a variety, fig. 2 an adult female,
fig. 5 a male approaching to maturity, fig. 3 an adult female of a variety; finally, fig. 6
represents a Mexican example, half-grown male.

266. GIRARDINUS PLEUROSPILUS. (Plate LXXXVII. fig. 1.)
Ginth. Fish. vi. p. 353.
D.8. A. 9. V. 6. L. lat. 28. L. transv. 8.

The height of the body is somewhat more than the length of the head, which is one-
fourth of the total (without caudal); the diameter of the eye is more than the length
of the snout, one-third of that of the head, and two-thirds of the width of the inter-
orbital space, which is slightly concave. In the female the origin of the dorsal fin is in
the middle of the total length, and conspicuously behind that of the anal fin. Caudal
fin large, longer than the head, subtruncate behind; the free portion of the tail is
somewhat elongate, the length of the base of the anal being one-third of its distance
from the caudal. Pectoral fin not quite as long as the head, and not extending as far
backwards as the ventral fins, which reach the vent.

In the male the origin of the dorsal is somewhat nearer the extremity of the caudal
than that of the snout; the anal process is quite straight, nearly twice as long as the
head, and terminating in a simple tapering point. Caudal very short. Reddish olive ;
a series of six or seven round blackish spots, each about the size of the eye, runs along
the middle of the side, a black line along the base of the anal fin and the lower and
upper margins of the tail. Caudal fin with two indistinct dark cross bands.

Mr. Salvin has discovered this species in the Lake of Duenas. Females attain toa
length of 2 inches, males to half that size only.

267. SCLEROGNATHUS MERIDIONALIS.
Ginth. Fish. vu. p. 23.
D..29-30. A. 10. _L. lat. 38. .L. transv. 74/73.

Mouth small, inferior, slightly corrugated. The height of the body is contained
thrice and one-half or thrice and one-fourth in the total length (without caudal), the
length of the head four times or four times and one-half. Head not much longer than
high. Eye rather small, one-fifth of the length of the head, and two-thirds of that of
the snout. Suborbitals narrow. The anterior dorsal rays are not much produced,
being shorter than the head. Caudal fin forked. ‘The origin of the ventral fin is
vertically below the fourth dorsal ray. Pectoral fin not extending to ventral. . There
are five longitudinal series of scales between the lateral line and the root of the ventral.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 487

Coloration uniform. Pharyngeal teeth very numerous and small, increasing somewhat
in size downwards.
Four examples, from 9 to 10 inches long, were obtained by Mr. Salvin in the Rio
Usumacinta.
271. PRISTIGASTER MACROPS.
Ginth. Proc. Zool. Soc. 1866, p. 603 ; and Fish. vil. p. 461.
p.13. A. 61. L. lat. 53.

Abdominal profile but slightly convex, the greatest depth of the body being one-third
of the total length (without cauda)); the length of the head is contained four times and
two-thirds in the same length ; eye very large, its diameter being more than one-third
of the length of the head, and nearly equal to that of the postorbital portion of the
head. ‘There are thirteen scales in the transverse series ascending from the origin of
the anal fin to that of the dorsal, four of the series being above the lateral line. Origin
of the dorsal fin midway between the root of the caudal and the scapula ; origin of the
anal nearer to the end of the snout than to the root of the caudal. A round black spot
on the scapula.

A specimen, 8 inches long, was found by Messrs. Dow and Salvin on the Pacific coast
of Panama.

272. PRISTIGASTER DOVII.
Pristigaster aryenteus, Giinth. Proc. Zool. Soc. 1866, p. 603 (not Cuv.).
dovii, Giinth. Fish. vii. p. 461.
D. 11. A. 56. . 1. lat. 51.

Abdominal profile but slightly convex, the greatest depth of the body being two-
sevenths of the total length (without caudal); the length of the head is nearly one-fifth
of the same. Eye large, its diameter being two-sevenths of the length of the head, and
two-thirds of that of the postorbital portion of the head. There are eleven or twelve
scales in the transverse series ascending from the origin of the anal fin to that of the
dorsal, four of the series being above the lateral line. Origin of the dorsal fin much
nearer to the root of the caudal than to the scapula; origin of the anal midway between
the end of the snout and the root of the caudal. Scapula with an indistinct blackish
spot.

A specimen, 8} inches long, was found by Capt. Dow at Panama.

273. CLUPEA LIBERTATIS,

Meletta libertatis, Giinth. Proc. Zool. Soc. 1866, p. 603.
Clupea libertatis, Giinth. Fish. vii. p. 433.

Dyhie®, ACeLOM I. lat. 48.
Closely allied to C. thrissa. ‘The length of the head is contained thrice and two-

488 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

thirds in the total (without caudal), the height of the body thrice and a half. The
origin of the dorsal fin is much nearer to the end of the snout than to the root of the
caudal. ‘The dorsal filament does not extend on to the caudal. Uniform silvery, with-
out humeral spot.

A single example, 24 inches long, was obtained by Messrs. Salvin and Dow at Libertad.

274, CHATORSSUS PETENENSIS.
Meletta petenensis, Giinth. Proc. Zool. Soc. 1866, p. 603.
Chatoéssus petenensus, Giinth. Fish. vu. p. 408.

D. 14-15. A. 20-23. L. lat. 40.

‘The length of the head is two-sevenths of the total (without caudal); the height of
the body is contained thrice or twice and three-fourths in the same. ‘The origin of the
dorsal fin is nearer to the end of the snout than to the root of the caudal, and in advance
of the ventrals. The dorsal filament does not extend on to the caudal. A small black
round spot on the shoulder.

Four examples, from 3 to 4 inches long, were obtained by Mr. Salvin in Lake Peten.

276. ENGRAULIS POEYI.
Kner & Steindachner, Abhandl. bayer. Akad. x. p. 23, Taf. 3. fig. 3.
D.16. A. 24. L. lat. 42.

The length of the head is nearly equal to the height of the body, which is two-ninths
of the total (without caudal); snout very short and rather obtuse; eye rather larger
than one-fourth of the length of the head. The origin of the dorsal fin is nearer to the
root of the caudal than to the end of the snout; origin of the anal fin opposite to the
middle of the dorsal. Pectoral fin reaching a little beyond the root of the ventral.
Upper and lower jaws with small teeth.

Rio Bayano.

277. ENGRAULIS MACROLEPIDOTA.
Kner & Steindachner, /. c. p. 21, taf. 3. fig. 2.
B. 12-13... D. 12. A. 29.. L. lat. 35. — L. transv. 9.

The length of the head is two-sevenths of the total (without caudal), the height of
the body one-third. Snout pointed, very short. The diameter of the eye is one-fourth
of the length of the head. ‘The origin of the dorsal fin is a little nearer to the root of
the caudal than to the end of the snout ; origin of the anal fin immediately behind the
end of the dorsal. Maxillary edentulous, extending to the angle of the preoperculum.

Rio Bayano.

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 489

278. CETENGRAULIS MYSTICETUS.
Engraulis mysticetus, Giinth. Proc. Zool. Soc. 1866, p. 604.
Cetengraulis mysticetus, Giinth. Fish. vii. p. 383.
D.17. A. 20. L. lat. 42. L. transy. 14.

Head exceedingly large, its length being contained twice and four-fifths in the total
(without caudal); the height of the body is contained thrice and two-thirds in the
same; the depth of the head is two-thirds of its length; snout compressed, pointed,
considerably shorter than the eye, the diameter of which is contained five times and a
half in the length of the head. The origin of the dorsal fin is nearer to the root of the
caudal than to the end of the snout; origin of the anal somewhat in advance of the end
of the dorsal. Pectoral fin reaching a little beyond the root of the ventral. Scales
adherent. Silvery, back greenish.

Three examples, the largest 6 inches long, were obtained by Messrs. Dow and Salvin
on the Pacific coast of Panama.

279. CARAPUS FASCIATUS (Pall.).

Two examples from the Rio Motagua are of a uniform brown coloration, but do not
differ structurally from South-American specimens.

284. Trrropon po.itus (Gir.).
Dns: AS ie

Nasal cavity with a short, imperforated papilla. Body smooth, except in the inter-
pectoral region, which is provided with minute spines. Head as broad as high, its
greatest depth being equal to the distance between the gill-opening and the front
margin of the orbit. Eye rather nearer to the gill-opening than to the end of the
snout. Upper parts blackish brown, with numerous black dots; belly and lower part
of the sides white. Dorsal and caudal fins brown; axil of the pectoral blackish.

One specimen, 13 inches long, was obtained by Mr. Salvin at San José.

285. TETRODON GEOMETRICUS.
D. 8. A. 7.

Nasal cavity with a short, imperforated papilla. Body covered with minute spines,
except on the snout and tail. Belly pendent, very extensible. Head nearly as high
as broad, its depth being equal to its length without snout. The eye occupies the
middle of the length of the head. Upper parts blackish, with bluish transverse lines,
curved on the sides; sides with some scattered black spots, lower parts white. Caudal
fin white in its basal, and black in its outer half; the other fins whitish.

One example, 3 inches long, was obtained by Messrs. Dow and Salvin at Panama.
VOL. VI.—PART VII. 3X

490 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

289. BALIsTEs FRENATUS (Lacép.).

A specimen, 8 inches long, obtained by Capt. Dow at Gonzalez Island, differs in
being of a more uniform coloration, the yellow band on the head being but slightly
indicated.

292, LEPIDOSTEUS TROPICUS.
Atractosteus tropicus, Gill, Proce. Ac. Nat. Sc. Philad. 1863, p. 172.
D.10-11. A.11. LE. lat. 54. L. transv. 8/12.

The length of the head is nearly one-fourth of the total; the width of the inter-
orbital space is two-fifths of the length of the snout, which equals the distance of the
front margin of the orbit from the fifth scale of the lateral line. The root of the
ventral fin is nearer to the base of the caudal than to the end of the snout.

Two examples, 18 inches long, were obtained by Mr. Salvin at Huamuchal.

295. MUSTELUS DORSALIS.
Gill, Proc. Ac. Nat. Se. Philad. 1864, p. 149.

“Teeth unicuspid. The posterior angle of the first dorsal fin projects to the vertical
of the origin of the ventrals, although the anterior fourth of the base of the fin is above
the pectoral. The caudal fin equals the distance between the snout and third branchial
aperture; and its terminal lobe nearly equals a third of the length, and is obliquely
truncated behind.”

Panama.

295. CARCHARIAS MACULIPINNIS.
Isogomphodon maculipinnis, Poey, Repertor. Fis. Nat. Cub. 1865, p. 191.

This species belongs to the subgenus Prionodon. Teeth with the terminal portion
much constricted—the serrature being very fine, and only in a few distinct to the point ; -
there are twelve on each side of the upper jaw; teeth of the lower jaw without any
denticulations. The length of the snout, from the front margin of the mouth, is not
much less than the width of the cleft of the mouth; the latter very deep, forming
nearly a semicircular arch. ‘The dorsal fin commences opposite to the inner posterior
angle of the pectoral; pectoral pointed, not twice as broad as long. Coloration uniform
grey, tips of most of the fins black.

One example, 4 feet long, was obtained by Mr. Salvin at Chiapam; the species was
first described from a Cuban specimen.

297. RHINOBATUS LEUCORHYNCHUS.
The anterior nasal valve is not prolonged to the inner angle of the nostril. Disk
longer than broad; the prenasal part of the snout is not so long as broad at the base,
but longer than the distance between the front extremities of the nostrils. Skin very

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 49]

finely granular; a series of very small, distant, smooth, oblong tubercles along the
median line of the back. Nostrils longer than the space between their posterior
extremities, but shorter than the mouth. Upper parts uniform ashy brown, the pre-
ocular part of the snout yellowish white.

One male, 21 inches long, was obtained by Capt. Dow on the Pacific coast of Panama.

299. UROLOPHUS MUNDUS.
Urotrygon mundus, Gill, Proc. Ac. Nat. Se. Philad. 1863, p. 173.

Mr. Gill proposes the generic name of Urotrygon for U. torpedinus and the present
species, the new genus being distinguished by the rounded and not angular outline, the
longer tail and posterior insertion of the spine, and especially the acute teeth. The
new species is thus characterized :—

The disk is orbicular, with a slight linguiform projection in front, and with the
pectoral fins behind broadly rounded. The distance of the snout from the hinder
margin of the pectorals equals the width of the disk. The tail (behind the anus) is
rather longer than the body (in front). The spine is inserted behind the middle of the
tail, and is about as long as the distance between the snout and the nostrils. The
ventral fins extend outwards, the rectilinear anterior margin tending little backwards ;
and the external margins are on a line with and complete the outline of the disk. The
posterior margin in the male is nearly rectilinear, while in the female it is slightly
convex, especially towards the inner angles. The upper velum is very sinuous and
fimbriated. The teeth are pointed and pyramidal. ‘The spiracles are oval, interrupted
at the intero-anterior angle by the eyes; and the margins are entire and well defined.
The skin is beset with numerous small stelliform tubercles, larger on the dorsal region.
The colour is a uniform dark brown above.

Two small specimens, male and female, were collected by Capt~Dow on the Pacific
coast of Central America.

300. A#TOBATIS LATIROSTRIS (A. Dum.).

This species was known from one very young example only, from the west coast of
Africa; Messrs. Dow and Salvin have rediscovered it in the Bay of Panama. The
specimen, which to the root of the tail is 12 inches long, and has a tail of 44 inches,
does not differ in anything from the Atlantic example. I may remark here that the
soft rostral appendage is naturally bent upwards, like the nose-leaf of certain Chiroptera,
and is not horizontally stretched forward as represented by M. A. Duméril.

3) eB

492 DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

EXPLANATION OF THE PLATES.

PLATE LXIII.

Map of the States of Central America, exhibiting localities mentioned in the paper.
The areas less than fifteen hundred feet above the sea-level are coloured green.

PLATE LXIV.

Fig. 1. Pristipoma macracanthum, Fig. 2. Umbrina elongata, p. 425.
p- 416. Fig. 3. Conodon pacifici, p. 417.
PLATE LXV.

Fig. 1. Centropristis macropoma, p. 409. Fig. 3. Chetodon humeralis, p. 419
Fig. 2. Hemulon margaritiferum, p. 419.

PLATE LXVI.

‘ig. 1. Upeneus tetraspilus, p. 420. Fig. 3. Pristipoma leuciscus, p. 416.
Fig. 2. Pseudojulis notospilus, p. 4477.

PLATE LXVII.

Fig. 1. Corvina chrysoleuca, p. 427. Fig. 3. Plectropoma afrum, p. 411.
vermicularis, p. 427.

PLATE LXVIII.
Fig. 1. Thalassophryne maculosa, p. 436. Fig. 2. Thalassophryne reticulata, p. 437.

PLATE LXIX.
Fig. 1. Cremnobates monophthalmus, Fig. 3. Antennarius leopardinus, p. 439..
p- 442. Fig. 4. Trachynotus fasciatus, p. 434.

Fig. 2. Clinus macrocephalus, p. 442.

PLATE LXX.
Fig. 1. Agonostoma microps, p. 444. Fig. 2. Agonostoma nasutum, p. 444.

PLATE LXXI.
Fig. 1. Heros citrinellus, p. 458. Fig. 2. Heros margaritifer, p. 450.

a
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(SU oy

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA. 493

PLATE LXXII.

9

. Heros urophthalmus, p. 404. Fig. 5. Heros melanurus, p. 450.
. Heros longimanus, p. 453.

PLATE LXXIITI.

. Heros spilurus, p. 451. Fig. 3. Heros salvini, p. 460.
. Heros aureus, p. 455. Fig. 4. Heros dovii, p. 461.

PLATE LXXIV.

. Platyglossus dispilus, p. 447. Fig. 4. Neetroplus nematopus, p. 470.
. Heros multispinosus, p. 453. Fig. 5. Heros godmanni, p. 466.
. Heros nigrofasciatus, p. 452.

PLATE LXXV.
. Heros lobochilus, p. 457. Fig. 2. Heros erythreus, p. 457.

PLATE LXXVI.

Heros trimaculatus, p. 461.

PLATE LXXVII.

. Heros nicaraquensis, p. 465. Fig. 3. Heros manaquensis, p. 463.
. Heros motaquensis, p. 462.

PLATE LXXVIII.
. Heros intermedius. p. 468. Fig. 3. Heros guttulatus, p. 466.

. Leros irregularis, p. 467.
PLATE LXXIX.

. Heros affinis, p. 455. Fig. 2. Petenia splendida, p. 469.
PLATE LXXX}.

. Hemirhombus ovalis, p. 472. Fig. 2. Citharichthys spilopterus, p. 471.
PLATE. LXXXI.

- Amiurus meridionalis, p. 473. Fig. 2. Mlurichthys nuchalis, p. 476.

' The figures on this plate ought to have been reversed.

. VI.—PART VII. 3

494

Ss:

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. Heros angulifer, p. 469, Fig.
. Pecilia elongata, fem., p. 484. Fig,

DR. GUNTHER ON THE FISHES OF CENTRAL AMERICA.

PLATE LXXXII.

. Chalcinopsis dentex, p. 478. Fig. 4. Anacyrtus quatemalensis, p. 479.
. Characodon lateralis, p. 480. Fig. 5. Haplochilus dovit, p. 481.
. Gambusia nicaraquensis, p. 485.

PLATE LXXXIII.

Exocetus callopterus, p. 479.

PLATE LXXXIV.

. Fundulus labialis, male, p. 481. Fig. 4. Fundulus guatemalensis, fem.,p.482.
. Fundulus labialis, fem., p. 481. Fig. 6. Fundulus punctatus, male, p. 482.
. Fundulus guatemalensis, male, Fig. 6. Fundulus pachycephalus, male,

p. 482. p. 483.

PLATE LXXXYV.
3. Pecilia petenensis, male, p. 484.
4. Pecilia petenensis, female, p. 484.

PLATE LXXXVI.

. Mollienesia petenensis, male, Fig. 3. Mollienesia petenensis, female,
adult, p. 485. adult, p. 485.
. Mollienesia petenensis, male, Fig. 4. Cirrhitichthys rivulatus, p. 421.

immature, p. 485.

PLATE LXXXVII.

. Girardinus pleurospilus, p. 486. Fig. 5. Xiphophorus hellerii, male approach-
. Xiphophorus hellerii, female,

ing to maturity, Chisoy River,
Chisoy River, p. 485. p. 485.

. Xiphophorus hellerit, vay., female, Fig. 6. Xtphophorus hellerti, half-grown

Chisoy River, p. 485. male, Mexico, p. 489.

. Xiphophorus hellerii, var., male,

Chisoy River, p. 485.

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[ 495 j

XV. On Divornis (Part XI.): containing a Description of the Intequment of the Sole,
and Tendons of a Toe, of the Foot of Dinornis robustus, Ow. By Professor
OwEN, F.R.S., F.ZS., &c.

Read November 28th, 1867.
[Puate LX XXVIII. }

THROUGH the liberality of the President and Council of the Philosophical Society
of York I have been favoured by the transmission, for examination and description, of
the portion of the foot, with the adherent tendons and integument, of the skeleton of
the Moa, referred to in the letter of Dr. Hector, F.G.S., quoted in a former Memoir
(Part IX.)', which skeleton, having been transmitted to the Museum of the Philoso-
phical Society at York, I determined, and referred, in that Memoir, to the Dinornis
robustus.

The toe, so preserved, with the tendons, sesamoids, and tegumentary sole-pads
(Pl. LXXXVIII.), is the inner one (#) of the right foot, and includes the three pha-
langes, corresponding in size and character with those figured in the Memoir
(Part IV.)?.

To the outer side of the proximal end of the first phalanx the capsule of the joint
adheres (Pl. LXX XVIII. fig. 1, ¢), whence it is continued upon the same side of that
part enclosing the metatarsal trochlea, and is extended upon the upper part of the
sheath of the tendons and sesamoids (s, ib. a, 6, c) beneath the joint between that
phalanx and the corresponding (innermost) trochlea of the tarso-metatarsal. Part of
the dried cartilage also adheres (at f) to the proximal articular surface of the phalanx.

The terminal portions of the tendon (a), inserted by means of the sesamoids and
their ligament into the base of the first phalanx, part of the perforating tendon (4),
similarly inserted into that of the second phalanx, and part of the upper perforating
tendon (c), which is continued to that of the last phalanx, are here preserved.

The largest portion of the sole-skin includes the pad (ib. fig. 2, dd’) beneath the
trochlear division of the metatarse answering to the above toe, with part of that which
extended beneath the middle trochlea, but wanting the outer portion. The under
surface of so much as remains of this ‘‘ heel-pad” measures 3 inches 8 lines by 3 inches
6 lines.

The margin (d' d') towards the missing outer heel of the pad shows abrasion; but
where the under surface is entire, it is beset with large papille, having a tendency to a
» Transactions of the Zoological Society, vol. y. part v. p. 340.

* Ib. vol. iv. part i. pl. 1. fig. 1, 11.1, 11.2, 113,
VOL. VI.—PART VIII. 3Z

496 PROFESSOR OWEN ON THE GENUS DINORNIS.

circular arrangement about a smoother space, which seems to have received the chief
pressure of the inner trochlea. On the inner or tibial side of the pad (d) the papill
increase in breadth and number, and are aggregated in the form of penta- or hexagonal
oblongish scales, about 23 lines in long diameter, diminishing in size (and chiefly in
breadth) at the inner margin of the pad.

The skin becomes thinner and smooth at the part yielding to the bend of the toe
upon the metatarse (ib. fig. 1, 4), and that to an extent and with a degree of infolding
in the dry integument which indicate great flexibility of the toe.

In advance of this, beneath the expanded ends of the first and second phalanges, the
skin thickens and spreads into a second broad flat pad (7) beset with coarse scattered
papille.

A short smooth tract (4) below the middle of the second phalanx intervenes between
the second (¢) and the third (7) papillose pad, which latter is beneath the joint between
the second and third phalanges; the papille are here longer and more close-set, the
transverse extent prevailing in most.

All the papillz are formed, or covered, by thick epidermal matter as hard as horn.
It is, however, together with the more gelatinous matter of the tendons and ten-
dinous sheaths, soluble; and, unless the remains of the bird had been buried in very
dry sand, it can hardly be supposed that it would have resisted for many years the —
action of moisture.

DESCRIPTION OF THE PLATE.

PLATE LXXXVIIL

. 1. Inner-side view of the inner toe (¢) of the right foot, Dinornis robustus, Ow.

. 2. Under view of the same toe.

. 3. Proximal end of first phalanx of the same toe, with appended integument,
tendons, &c. Natural size.

2 ete
18 fe

Les Le iL |
ey
go dQ

M&N Hanhart imp

Dinkei del* on Stone by J Erxlebern.

DINORNIS ROBUSTUS

[ 497 ]

XVI.—On Drivornis (Part XII.): containing a Description of the Femur, Tibia, and
Metatarsus of Dinornis maximus, Owen. By Professor Ownn, F.R.S., F.Z.S., Le.

Read November 28th, 1867.

[Puares LXXXIX., XC.]

IN the letter of the date of February 15th, 1864, in which Dr. Hector, F.G.S., Pro-
vincial Geologist of Otago, New Zealand, communicated to me the particulars of the
discovery of the almost entire skeleton of the Dinornis, of which the skull and scapulo-
coracoid were described in a previous Memoir (No. IX.), he remarked that ‘‘ The
skeleton was not that of one of the largest-sized Moas, the tibia, for instance, being
only 27 inches in length, whereas I have frequently seen them as much as 36 inches.”

The tibia of the specimen in the British Museum, which is the type of my Dinornis
robustus, measures 32 inches in length; and it is probable that the difference in the
length of the tibia of this specimen and that of the skeleton at York (27 inches) indi-
cates the range of size as exemplified in individuals of different sexes of this species.

I have, however, for some years, been cognizant of a species of Dinornis from the
Middle Island of New Zealand, having a tibia rather exceeding the length stated by
Dr. Hector, and of a thickness proportionally the same as in Dinornis robustus. In
1858 the Duke of Argyll favoured me by sending for my inspection a tibia of this size,
together with a femur and metatarsus of like proportions, and purporting to be of the
same limb of a Dinornis, which bones had been transmitted to His Grace from the
Middle Island, New Zealand, by the Rev. Dr. Lillie. With the liberal permission of the
Duke, casts were taken from these bones for the British Museum, which have been
exhibited in the Paleontological Gallery as of the “ Dinornis giganteus, var. maximus.”

In 1861 I was favoured by Henry Joseph, Esq., with an inspection of a femur of a
Dinornis of the dimensions of that of D. maximus, which had been found beneath drift-
sand at Otago, New Zealand.

In 1863 Professor Tennant, F.G.S., was so kind as to bring for my inspection the
shaft of a femur of a Dinornis, from New Zealand, locality not stated, of the general
dimensions of the two above specified, but heavier from some infiltration of mineral
matter, and rather more robust. The least circumference, e. g., of the shaft of the
femur in Dr. Lillie’s and Mr. Joseph’s specimens was 8 inches 13 line; in Professor
Tennant’s specimen it was 8 inches 9 lines.

In March, 1867, I was favoured by Major J. Michael, of the Madras Staff Corps, with

' Trans, Zool. Soc. vol. vy. p. 340.

BVA 2

498 PROFESSOR OWEN ON THE GENUS DINORNIS.

the opportunity of inspecting the femur, tibia, and metatarse figured in Plates LK X XIX.
and XC., of the natural size, which had been discovered in August, 1865, on the Glen-
mark Estate of ‘“ Kermode & Co.,” about forty-five miles from Christchurch, Canter-
bury Settlement, Middle Island, New Zealand. ‘They were discovered, in the course of
running a drain across a bog or swamp, about 4 feet below the surface, in such juxta
position as to lead to the inference that they were bones of the same leg (the left); and
their dimensions a little exceed those of the bones on which I had previously founded ~
the variety or species Dinornis maximus. They are such, indeed, as to lead me to
believe that the proposed specific term may be a safe one. I can hardly conceive
that any bones as much larger than these as they are in comparison with Dinornis
yiganteus remain to be discovered in New Zealand—that land of these strange giants
of the feathered class.

To have evidence of a bird as large as the Ostrich of Africa, from so comparatively
small a tract of territory, seemed to me in 1839 the most wonderful result of the deter-
mination of the bone figured in plate u1., Volume III., of the ‘ Zoological Transactions.’
When I subsequently received a femur surpassing in length that of the struthioid
species (Dinornis struthioides') by 2 inches, I called the species Dinornis ingens? ; then
receiving a femur of the length of 15 inches, with other leg-bones to match, I proposed
for it the term Din. giganteus*. Leg-bones equalling those of Dinornis giganteus in
length, but in all cases exceeding them in thickness, and from an island where bones
of the true Dinornis giganteus have never been found, represent the species called
Dinornis robustus*; and now, having almost exhausted the vocabulary of terms expres-
sive of hugeness, I venture on the superlative for the species represented by the bones
which form the subject of the present Memoir.

Femur. (Plate LXXXIX. fig. 1.)

This presents all the generic characters of that bone in Dinornis’. The roundness of
the shaft, the thickness of the walls of the medullary cavity, the absence of pneumatic
foramina, the thickness of the shaft, and breadth of the articular extremities, especially
of the distal one, in proportion to the length of the bone, the tuberous “ line aspera ”
on the back of the shaft (Pl. LXX XIX. fig. 1), the production of the anterior inter-
muscular ridge from the lower end of the longitudinally extended thick and rugged
pretrochanterian ridge, the rough, deep, well-defined fossa at the upper and fore part
of the femoral shaft, the still deeper ecto-gastrocnemial fossa, and the very wide rotular
channel—each and all of these Dinornithic characters of the avian femur are strongly
marked in the present species. The surface, on the head of the femur, for the attach-

' Trans, Zool. Soc. vol. iii. pl. 21. fig. 3. (Length 10”.) * Ibid. fig. 1. (Length 12’.)
* Thid. p. 250. (Length 15’.) ‘ Thid. vol. iy. pl. 1. fig. 1. (Length 14" 6’”.)
® Tbid. vol. iii. p. 247.

PROFESSOR OWEN ON THE GENUS DINORNIS. 499

ment of the ligamentum teres is as if a slice of the convexity had been cut off obliquely
from its most prominent part backward and a little upward, so that no part of this sur-
face appears in a direct front view; and it is slightly, if at all, depressed: part of this
large flat surface had been broken away in the specimen figured (Pl. LXX XIX. fig. 1).

In the few femora of the general size attributable to Din. maximus there are varia-
tions in the relations of the circumference of the shaft to the length of the bone, and in
one instance (the femur from Mr. Joseph) in the proportion of the breadth of the distal
end. In the instance (Mr. Joseph’s) where this end of the bone is narrower, the back
part of the inner condyle is much narrower, and is more convex and more backwardly
produced. The form of the popliteal space also differs. In Plate LXXXIX., and the
majority of femora of Dinornis, it is a deep oblong oval pit, definitely excavated, the
larger end toward the inner posterior tuberosity of the “linea aspera,” and deepening to
the back ridge of the inner condyle, which extends toward the outer one. In the
femur from Mr. Joseph the depth of the popliteal pit is due to the backward projection
of the condyles and their uniting posterior ridge, dividing the popliteal from the in-
ferior intercondylar fossa, which is unusually deep. The modification of the distal end
of the less robust femur (from Mr. Joseph) is, indeed, such as to suggest a specific dif-
ference. The trochanter, also, rises more abruptly, is higher, and the outer ridge of the
antero-superior pit is unusually prominent.

Tibia. (Plate XC.)

Of the tibia (Pl. XC.) there need only to be given the dimensions, taken in accord-
ance with those which have been previously recorded of the Dinornis giganteus and
other species’. All the characters of the bone which distinguish it generically as of
Dinornis are closely repeated in the present specimen.

Metatarsus. (Plate LXXXIX. figs. 3 & 4.)

The same remark applies to the metatarse (P]. LXX XIX. fig. 3); but between that
bone in Major Michael’s series and the one in Dr. Lillie’s there are differences of pro-
portion (probably within the limits of individual variety), which may be mainly appre-
ciated by comparison of the outline of the latter (fig. 4) added to the Plate (LXX XIX.)
which contains the finished lithograph of Major Michael’s specimen (fig. 3).

Dr. Lillie’s specimen is longer and more slender, but with a greater transverse expan-
sion of the distal end. The back part of the middle articular trochlea at the distal end
projects more abruptly in Dr. Lillie’s specimen; but the generic characters of the meta-
tarsus of Dinornis are closely maintained in both specimens.

Subjoined are dimensions of the three chief bones of the hind limb of the present
enormous species (). maximus), together with those of the same bones in Dinornis
giganteus.

! Trans. Zool. Soc. vol. iii. p. 246,

500 PROFESSOR OWEN ON THE GENUS DINORNIS.

Dimensions of Femur.

Dinornis maximus. Din. giganteus.
Major Michael's. Dr. Lillie’s. Mr, Joseph's.
in. lines. in. lines. in. lines, in. lines.

ene this ha Sefer e ne, Pen kee 18 3 17 0 18 0 16 0
Breadth of proximal end, in the axis of

the neck, or transverse diameter... . 6 6 6 3 6 2 6 0
Breadth of antero-posterior diameter.. 6 0 5 6 4 7 5 6
Breadth of distal end, transverse diam. 7 6 7 0 6 9 6 3
Circumference of middle ............ 9 6 8 9 8 0 7 9

Dinornis maximus. Din. giganteus.
in. lines. in. lines.
era prtilie: Pea vens ee dys ci Vsk one ence cuoue tots ook Sahl akon Nees ave tien auen tint ral cays 39 0 35 0
Preadtheonproximalyend cs. ca5 eck: apne nil Page epsi eevee ere skels 8 6 G 6
Bread tvOMaisavend tract ciist nc kiciieiecse ste beter cers «nice 5 0 4 0
Gmeumferente of middle ts). 5.20. ee EME A 8 6 6 6
Tibplan zidreextends|dowa 2) ks. t- Saurtinuys Meher tanaka tates 20 0 13 0

Dimensions of Metatarse.

Dinornis maximus. Din. giganteus.
Major Michael’s. Dr. Lillie’s.
in. lines. in. lines. in. lines.

Here Gee. anon Meviar cn aarti. at lunes Rae 20 0 21 6 18 6
Circumference of middle of shaft .............. 8 4 6 4 5 6
Breadth of middle of shaft..............:...4. 3 0 2 7 uP bik
Breadth, transverse, of distal end.............. 6 9 ¥( 3 3 1
Breadth, transverse, of proximal end .......... 5 6 5 3

DESCRIPTION OF THE PLATES

PLATE LXXXIX.

. Femur, back view, Dinornis maximus: nat. size.

. Femur, back view, Turkey (Meleagris gallopavo): nat. size.
3. Metatarsus, front view, Dinornis maximus (Major Michael’s specimen): nat. size.

Fig. 4. Metatarsus, front view, Dinornis maximus (outline of Dr. Lillie’s specimen) :°
nat. size.

Fig. 5. Metatarsus, front view, Turkey (Meleagris gallopavo): nat. size.

=}
Een Se
3 de OG
bp

PLATE XC.

Fig. 1. Tibia, front view, Dinornis maximus: nat. size.
. Tibia, front view, Turkey (Meleagris gallopavo): nat. size.

les]
es
ho

f 501 ]

XVII. On the Osteology of the Kagu (Rhinochetus jubatus).
By W. XK. Parker, F.RS., F.Z.S.

Read January 9th, 1868,

[Puates XCI., XCII.]

IN the Proceedings of this Society for 1864 (pp. 70-72) there is a short account
given of my views of the zoological position of the Kagu; but no details are added as
that paper was merely intended to be an introduction to one more exhaustive and that
should contain the results of much more labour and thought. More recently, in my
memoir “On the Shoulder-girdle and Sternum” (Ray Soc. 1868, pp. 158-160), I have
spoken of the relationships of this bird; but those remarks merely have reference to
what is indicated by the parts of the skeleton which are there treated of. With regard
to the nomenclature of those parts (namely, the breast-bone and shoulder-bones), re-
ference to the memoir itself will show that there is some change of the terms used in
my older papers on these subjects; this has become a necessity on account of the
additional light obtained from severer research. Here, also, I must crave the liberty of
modifying terms used by me in time past, and also of dropping some that seem now to
be inaccurate, and of coining new words’in cases of absolute need. The splint-bones
that invest the face have cost me the most trouble in researches into the morphology
of the skull; for I have strained after an harmonious view of the facial bones in the
whole vertebrate subkingdom, and the Bird has always appeared to me to be the very
class-type that ought to show the transition from the Ovipara to the Mammal. Un-
doubtedly it does; but it presents the greatest difficulties to the anatomical student—
the process of ossification being so intense in degree, and so varied in relation to time
in this Class. Hence the morphological observer has to lie in wait for the various
osseous centres, never knowing when they may appear in the different groups, and
being equally uncertain when they shall lose their individuality. The Bird’s face has
always appeared to me to be what one might suppose that of a Fish to become, if that
low type were to undergo a series of metamorphic changes; it is this great unlikeness of
the Bird’s face to that of a Reptile or a Mammal which makes its morphology so difficult
of interpretation. When I first lighted upon an additional bony element (a bone
which I at once saw must answer to the outer alveolar plate of the mammalian max-
illary), my difficulties with regard to the Bird’s upper jaw were only becoming greater
instead of receiving their ultimate elucidation. Well knowing from embryological
researches into the structure of the skull and face in the large Serpents that their so-

502 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

called “ turbinal” was, in reality, one of the maxillary series, it seemed to me perfectly
possible that this bone, which I proposed to call the “prevomer” (see Zool. Trans.
vol. vy. p. 157), should reappear in the Bird as enormously developed as the premaxillary,
and that thus I had an explanation of those quasi-turbinal outgrowths which make the
maxillary of the Bird so unlike that of a Mammal.

Thus it appeared that the true maxillary was only exceptionally present in the Bird,
and that the huge prevomerine splint not only retained its reptilian character but also
grew largely into tracts which were left open to it by the abortion of the “maxilla
proper.”

Nothing occurred to disturb this view until the end of the year 1865, when Professor
Huxley very forcibly put it to me that the bird’s great maxilla does actually correspond
to that of the mammal, minus its outer alveolar plate; this view is given in my paper
on the Ostrich’s skull (Phil. Trans. 1866, pp. 113, 114). If that view were the true
one, then the so-called “‘turbinal”’ of the Snake and the Lizard might also be referred
to the same category. Frequently returning to this subject, being severely criticised
by Professor Huxley for using the term “ prevomer,” and receiving new ideas upon the
subject from fresh research, I have at last determined to alter my nomenclature. I
give up the obnoxious term “ prevomer,” and propose to call the pseudo-turbinal splint
the “ septo-maxillary”; the small additional maxillary of certain birds may be called the
‘‘postmaxillary”’; and the great upper jaw-bone of the bird will retain its old name—
the “ maxillary.” Nevertheless it is ess than the maxillary of the Mammal, its outer
alveolar plate being aborted by the premaxillary; and more, for it oftentimes has the
attributes of the ‘‘septo-maxillary” added to its own, although in the Bird merely as a
region, not as a distinct osseous piece. I have traced, as I think, the septo-maxillary
through a large series of Ovipara, from the Ganoid fish to the Lizard; I am quite pre-
pared to see it continuous with the maxillary of the bird, which has many splint-bones
single that are double in the Lizard. A few other changes will be spoken of in the
course of my description; but the maxillary series has presented the greatest difficulty,
and needed to be put plain at once; for my determination of the so-called “ turbinals ”
of the Lizard to belong to the maxillary splint-system, and the discovery of the attri-
butes of these bones in the maxillaries of the bird, have caused no little difficulty in the
nomenclature of those parts.

I have already ventured to classify the Kagu (see ‘Shoulder-girdle and Sternum,’
p. 158), putting it into a family with the Psophia and the Sun-Bittern, and calling this
group the “ Psophiinz,” with the qualification, however, that each of the three types
deserves to be placed at the head of a distinct subfamily. These Psophiine types are
essentially Cranes, but they are very aberrant.

Notwithstanding the essential relationship of these three ‘‘ Geranomorphs,” the first
view of their skulls would not suggest so near an affinity. If the Kagu’s skull be
placed in the midst of a series of skulls of the ‘‘ Grallie,” the Night-Heron’s skull and

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. 503

that of the Norfolk Plover (@dicnemus) would arrest attention first, as coming nearest ;
and if that of the Kagu were placed between these two, it would at once be seen to be
a perfectly intermediate form.

This need excite no surprise ; for the Cranes are certainly Pluvialine birds, having no
little affinity for the Herons; and, moreover, the Kagu, being a generalized type, comes
much nearer to the true Ardeine than do the typical Cranes.

In the Psophia’s skull there is a very strong resemblance, in general outline, to that
of a Pheasant; but this vanishes on closer inspection, and then its exceeding nearness
to the Rails is at once evident: moreover it has the superorbital chain of the Timamou.
The skull of the Ewrypyga is seen at once to come quite as close to that of a Stilt-
Plover (Himantopus) as to that of a Bittern, and, indeed, closer; more minute observa-
tion shows that it is essentially the same as that of the Kagu, but feebler and having a
more elongated face.

The face and skull of the Kagu are of equal length (Pl. XCII. figs. 1-3); the orbits,
as in @dicnemus, are extremely large, the postfrontal region being unusually wide and
overhanging. The skull is very broad behind (PJ. XCII. fig. 4); the only related bird
that makes any approach to the Kagu in this respect is the Night-Heron; and here the
Kagu departs furthest from the Pluvialine, and comes nearest to the Ardeine type; for
the temporal fossz are extremely large, and are bounded by strong ridges in front (the
coronal), at the mid line (the sagittal), and behind (the lambdoid). Compare with this
state of things the low, small, widely separated temporal fossee of Anthropotdes, Balea-
rica, Psophia, Eurypyga, Ocydromus, and Cdicnemus. In Nycticorax the postorbital
angle is very distinct from the postfrontal process; in the Kagu they are coincident
(Pl. XCIL. fig. 3). But the Kagu and Nycticoragr are very similar in the bounding of the
temporal fossa below, this valley being in both very deep between the postfrontal and
the descending spur of the squamosal (zygomatic process of the Mammal). This spur,
however, is much the largest in the Kagu; and in this respect this bird shows an affinity
to the Struthionid on one hand, and to the Cariama on the other. The foramen mag-
num (fig. 2) is large; and the condyle is a little wider in the transverse than in the lon-
gitudinal direction. The tympanic wings of the exoccipital (figs. 2, 3,4) are very much
like those of the Night-Heron, but are larger and deeper. The symmetrical bosses of
the basitemporal are very mammillate and distinct ; they agree with those of the Gruiniwe
proper; but the anterior lip of the basitemporal plate is delicately thin, as in the
Ardeine (Pl. XCII. fig. 2). The large, trumpet-like anterior extension of the tympanic
cavity, on each side, is like what is seen in all the congeners of the Kagu; they are
formed in front by the posterior pterygoid processes of the basisphenoid. In front of
these, the “anterior pterygoid processes” are seen as small, aborted, angular projections
(Pl. XCII. fig. 2); and in front of these the “rostrum” (formed by the grafting of the
parasphenoid on to the true basisphenoidal beam) is narrow, subcarinate, and com-
pletely fused with the base of the orbito-septal plate. ‘These two parts are somewhat

VOL, VI.—PART VIII. 4A

504 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU,

separated by a notch anteriorly (Pl. XCII. fig. 3). The under surface of the superorbital
plate of the frontal forms one continuous plate of bone with the alisphenoid, which
has no fenestra, and which has completely coalesced with all the surrounding bones
(Pl. XCII. figs. 2 & 3)".

The Kagu approaches both the Herons and the Rails in the structure of the inter-
orbital septum; for the orbito-presphenoid (Pl. XCII. fig. 3) is feeble and oblique; but
it is feebler and more obiique than in either of those types, the posterior process of the
perpendicular ethmoid and the alar part of the anterior sphenoid being very feeble at
their junction. This leads towards Eurypyga, which has complete abortion of the
connecting bar, as in Himantopus and Phalacrocorax. Two small fenestre are seen
on each side of the feeble four-winged orbito-sphenoid; below, the presphenoid is a
minute separately ossified spur (fig. 5). As in the Stanley Crane, and unlike that in the
Night-Heron and Rail, the great ethmo-basisphenoidal bar, bounding the interorbital
space below, is very deep (fig. 3), and thickens as it approaches the parasphenoidal
beam. ‘The cleft between the orbital and the nasal septum is incomplete (fig. 3), and
the latter is partly ossified, as in the Ardeine: it is much deeper than in Gralle
generally. There are three septal ossifications—two upper (the small one foremost), and
one lower; this lower bony plate answers to the antero-inferior bone of the Rapacious
bird—that which sends out the vestibular bars to join the maxillary on each side. The
nasal septum, like the al nasi, only occupies the hinder half of the bony nasal opening ;
the rest is filled up by fibrous tissue. The septum nasi is alate below, and each semilan-
ceolate cartilaginous wing is attached to the septal process of the maxillo-palatine plate.

The prefrontal region (Pl. XCII. fig. 3) is only partly ossified, the antorbital plate
being cartilaginous above and externally, and the perpendicular ethmoid sending very
little bony matter into the upper ale. The aliseptal region (inferior turbinal and its
root) and the alinasal are quite soft in the adult; the alinasal flaps are obliquely oblong,
lie low down, bulge but little, and only reach halfway along the bony nasal opening;
the anterior nostril is a low-lying valvular slit.

As the bony nostrils are extremely open, the body of the premaxillary only reaches
one-third of the way to the angle of the bone. ‘The nasal processes (Pl. XCII. fig. 1)
are completely fused, as in the Cranes, the Psophia, the Ewrypyga, and the Rail, but
they keep very distinct from the nasals. The nasal process is strong, flat, even some-
what concave at the middle of the bone, and is very unlike what is seen in the related
types. Below (fig. 2) there is seen to be a degree of filling-in by bony matter that
approaches what we find in the true Herons, and the fusion of the parts is complete.
This narrow, gently concave anterior part of the bony palate is composed of the inner
part of the dentary plates and the palatine bars of the premaxillaries. Joined to them are

* I must refer the reader to the plates accompanying my paper on the Gallinaceous and Struthious skulls for

the lettering of especial parts, with this proviso, namely, that p.v. (prevomer) should be read as mx. (maxillary),
and mx. as pt.mx. (postmaxillary).

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. 505

the sharp anterior spurs of the palatines and of the maxillaries. As the maxillary of the
bird has cost me more trouble than any other part, I shall take some pains to describe
it, especially as the Kagu and its relatives have this part in a very instructive condition.
Here the strictest watch has to be kept, lest the mind should be deceived by external
resemblances, and by actual function of the parts; nothing but a study of the develop-
ment of the face can give any solution of this difficulty. In the first place let it be
noted that the maxillary of the Bird is extremely like that of the typical Fish, being
secondary in so great a degree to the premaxillary, and lying within and behind it; the
manner, also, in which the palatine stretches forwards to the front of the face and
articulates with it is very ichthyic. But the maxillary of the Bird for a long while
seemed to me to be merely a very inordinately developed septo-maxillary (the so-called
turbinal of the Lizard); so that the Lacertian relationship appeared to be that which was
most evident. Now, however, I have no doubt about its being truly the maxillary of the
other Vertebrata; but I do see in it also the Lacertain septo-maxillary in a high state
of development in many cases, but continuous with the true maxillary. But comparison
of the Bird’s maxillary with that of the cold-blooded Vertebrata (excluding the Croco-
diles) fails to give any explanation of its meaning; and it is only by comparing it,
also, with a series of Mammalian counterparts that it can fully be understood. Perhaps
the first thing that strikes the eye in the anterior part of a bird’s palate is the large
azygous ‘anterior palatine foramen;” this is quite unlike what is seen in Mammals
generally. In the Bird the space between the dentary and palatine plates of the pre-
maxillary is filled up by the sharp wedge-like fore end of the maxillary; whilst the
‘anterior palatine foramen” is seen to be merely the space between the palatine plates
of the premaxillaries. The Pangolin (J/anis) and, especially, the Hedgehog (Erinaceus)
explain this, the symmetrical passages being almost filled up by the wedge-like anterior
ends of the maxillary palatine plates. But in the Pangolin, and also in the Coati
(Nasua fusca), the palatine plates of the premaxillaries are bowed out in some degree,
forming a third opening in their fore-palate. This structure is precisely the morpho-
logical link wanting to make the Bird and the Mammal hang together here. In the
Pangolin the palatine plate of the maxillary is one continuous, narrow, obliquely scooped
sheet of bone, attached by “‘harmony” to the palatine plates of the palatine behind,
and wedged in between the premaxillary forks in front. In the Hare (Lepus timidus),
however, four-fifths of this plate is absent; and behind only does the maxillary send
inwards an elegant, subquadrate, pedate plate to meet its fellow of the opposite side,
and to be underlain behind by the much feebler palatine plates of the palatines.

In the Bird the part answering to the fore end of the Pangolin’s palato-maxillary plate
is differentiated from the hinder part, is very narrow, and entirely fills up the space
between the premaxillary forks. Behind, the Bird’s maxillary developes the exact
counterpart of the palato-maxillary plate of the Hare; and this part, instead of keeping
to the palatal floor, as in the Hare, ascends into the nasal labyrinth, occluding it in

4a2

506 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

no small degree, and, being thus developed into pseudo-turbinal outgrowths, does
correspond, not numerically so far as ossific points are concerned, but truly, in a mor-
phological sense, to the pseudo-turbinal (my “ prevomer” or “ septo-maxillary ”) of the
Snake and the Lizard. At first sight, no one would doubt that the Pangolin came in
to explain the small “postmaxillary” seen in certain birds; for on the projecting
zygomatic angle of the main bone there is a small ossicle, apparently the precise coun-
terpart of the little face-bone of the bird. This likeness, however, is quite deceptive ;
for the bone in the Pangolin is articulated to the zygomatic process of the maxillary,
and is the “malar” or “ jugal;” whereas in the Bird it answers to the extreme angle
of the dentary plate, and is a curiously segmented representation of that elegant
pupiform maxillary chamber which, in the young Pig (Sus scrofa), contains the
hindermost molar tooth. The supplementary ornithic bone can only be explained by
referring to the multiple ganoid face-wall of the Lepidosteus. The ‘“ septo-maxillary ”
gets its interpretation from the same fish ; but it crops up in several other ichthyic
types—for instance, in another ganoid the (Ama), in Clarias, and in the Clupeoids.

The maxillary of the Bird may be described as a lateral facial splint, which develops
several spurs and plates, passing in various directions, namely :—the outer or dentary part,
which only appears for a small space on the outside, close below the descending crus
of the nasal; the palatine part, which is composed of two portions, an anterior and a
posterior; and the long, styliform zygomatic process. The long anterior palatine pro-
cess wedges in between the two forks of the intermaxillary, and articulates by its inner
side with the styliform fore end of the palatine. The posterior palatine plate is only
moderately broad where it is given off; it then suddenly narrows, both margins being
concave, and then expands into what Professor Huxley terms the “ maxillo-palatine
plate.” Below, this part tends to, and often does unite with, its fellow of the opposite
side, between the palatines ; this answers to the submesial sutural part of the palatine
plate of the maxillary in the Hare. Above, the plate in many birds developes an
“anterior septo-maxillary spur” to articulate with the septum; this spur answers to
the transverse bar of the septo-maxillary which partly occludes the nares in the
Cyclodont Lizards. Above and behind, we have the large, oblique “ posterior septo-
maxillary lobe;” this answers to that part of the Lizard’s and Snake’s septo-maxillary
which overlaps the vomer.

In the Kagu the maxillary has coalesced to a great extent with the surrounding
bones; but with care its boundaries can be discovered ; it is best seen from the under
and side aspect (Pl. XCI. figs. 2 & 3). This bone is a long style, pointed at both ends,
and sending inwards a broad and complex plate. The anterior style fills up the inter-
maxillary fork, and answers to the fore end of the palato-maxillary plate of the
Mammal; the posterior style is the very long and slender zygomatic process; and the
middle transverse bar answers to the palato-maxillary plate of the Zeporide, and is
developed upwards into the septo-maxillary processes. ‘The anterior style has an

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. 507

inward direction; the posterior or zygomatic style is turned obliquely outwards, and
is clamped by the equally delicate jugal. The posterior maxillo-palatine bar lies at an
exactly right angle to the cranio-facial axis: it is 3 lines broad at its root, but as it passes
inwards it becomes only half that width; and both its margins are elegantly concave.
This plate lies a considerable height above the deep palatine (figs. 2 & 3); it shows
itself very little in the interpalatine region; but the widening of the palatine on the
inner side, near the fore end of the vomer, is due to the down-growth of this part of the
maxillary. This edge does not nearly meet its fellow of the opposite side!; but it is the
exact homologue of the sutural region of the palato-maxillary plate of the Hare. ‘This
interpalatine portion of the maxillary is merely the lower edge of the large spoon-
shaped “ posterior septo-maxillary lobe” (fig. 3), which is gently convex on the inner
face, scooped on the outside, moderately thick, and slightly fenestrate. The soft
inferior turbinal is attached to the upper and inner edge of this lobe; below, the vomer
lies a small distance within it (as in its counterpart in the Lacertilia and Ophidia); and
in front it thickens where it underprops the descending crus of the nasal (fig. 3).
Inside the foot of the nasal, before the bone has narrowed to form its anterior palatine
portion, it developes a squarish mass, which is prickly in front, and is attached to the
cartilaginous wing of the septum; this is the “ anterior septo-maxillary spur,” and is
the counterpart of the transverse vestibular bar of the septo-maxillary of the Cyclodont
Lizard. This hooked plate is better developed in Eurypyga; but attains its highest
condition in the Psophia, where it is a large, transversely oval lobe, constricted round
its base, so as to be somewhat pedicellate ; it also reappears in Caprimulqus europeus
as a long, slender, sigmoid style.

In Psophia, as in Cyclodus, the nasal vestibule is formed by the facial splint (by the
distinct septo-maxillary piece in the Lizard); but in many birds, especially the Raptores,
the “anterior septo-maxillary spur” is but little produced inwards, and the vestibule is
completed by the largely developed transseptal bar: without a transverse section the
septo-maxillary of the Cyclodont might easily be mistaken for one side of an ossified
septum nasi like that of the Owl or Hawk. In the Eurypyga the maxillary comes
exceedingly close to that of the Kagu; but the plate answering to the Hare’s palato-
maxillary is extremely contracted before it expands submesially ; its ‘‘ posterior septo-
maxillary” lobe is also much more fenestrate, and developes itself behind into an
elegant, flattened thimble-like pouch: it also appears more in the interpalatine space.
In the Psophia these plates are very thick and spongy (approaching those of the
Herons, Pigeons, Owls, &c.), and they have a large, oblong interpalatine portion.
There is no “postmaxillary” bone in the Kagu; and the “ dentary” portion of the
maxillary is an extremely small angle just below the angular flap of the premaxillary
(fig. 3). The fusion of these parts, however, makes the boundaries somewhat indistinct ;

' In this respect the Kagu is ‘‘ Schizognathous ;” but in the fresh state the wings of the nasal septum are
strongly tied to the maxillaries, and thus it is, practically, “‘ Desmognathous.”

508 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

but in the Psophia and Eurypyga this angle is a very delicate style: in the latter the
premaxillary angle also is continued over the zygoma as a very slender needle of bone.
The vomer (fig. 2) is a very long, delicate style in these three aberrant types; it comes
very near to that of Talegalla, but is longer. The posterior third is bifurcate; it then
becomes deeply grooved above, and sharply carinate below, and is bowed upwards:
this middle part lies between the interpalatine portion of the maxillaries. The anterior
third then descends and becomes broader, as in the Chelonians; but what is gained in
depth is lost in height, and the fore part is a very slender needle of bone. In the
Psophia the vomer is split halfway ; and the edges of the basifacial groove are thickened
and denticulate, as in the Heron; so they are in the Hurypyga, which, however, in other
respects has a vomer almost the exact counterpart of that of the Kagu,—save that it is
still more exquisitely slender, and elongate in front. As to the upper and lateral facial
splints the Kagu and the Ewrypyga agree very closely. The nasals (fig. 1) are very
sharply split, the clean fissure between the upper and ascending ramus turning inwards
between the lacrymals above. The nasals reach half an inch further backwards than the
nasal part of the premaxillaries in the Kagu, and are as thick and cellular as the frontals
where they coalesce with those bones. The upper crus of the nasal runs forwards three-
fourths of the distance to the fore end of the nasal fossa; the descending crus is sharp
below, and is strongly interwedged between the maxillary and premaxillary. Thus this
part of the face is thoroughly Gruine in both the Eurypyga and the Kagu; the long,
open nasal fossa, so sharp above at the bifurcation of the nasals, gives a character to the
face common to large groups of Grallz and Palmipeds (see my figure of the Lapwing’s
skull, Trans. Zool. Soc. 1864, vol. v. pl. 37. fig. 8,2, pa). In the Rallidew the nasal
fossa is semioyal, and so it is in the Psophia; but in it the division of the nasal, whilst
equally obtuse and rounded, yet helps to form a much shorter nasal fossa, like that of
the young of a typical Raptorial bird, before the ossification of the alinasal cartilages:
in that aberrant 4tomorph the Cariama, this open condition of the nasal fossa is per-
sistent, but somewhat occluded by the bony “anterior alinasal bridge.” The lacrymal
(figs. 1 & 3) of the Kagu agrees with that of the Eurypyga, save that in the latter the
bifurcated descending part is slenderer and is fenestrate, whilst in the Kagu it is cellular.
They both differ from Grus in having the superorbital part’ not more developed outwards
and backwards than in the Night-Heron; these bones keep distinct, as in the Cranes and
Rails. The lacrymal of Psophia is larger and thicker, and is greatly clubbed below;
but the most remarkable thing in the Psophia is, that the lacrymal (preorbital) is followed
by a chain of 5-7 free superorbitals, like those of the Tinamous (see Trans. Zool. Soc.
vol. v. pl. 40; and Phil. Trans. vol. clvi. pl. 15). This cropping-up again of what the
‘Tinamou had adopted from the Reptile is very interesting, and is not the only character
by which these two birds may be connected. In the Psophia the anterior frontal
region is swollen and cellular, somewhat approaching to what is seen in the Coot

‘ A separate bone in the Lizard.

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. 509

and the Palamedea; but in the Eurypyga and the Kagu (PI. XC. fig. 1) this part is
gently concave.

There is nothing to remark upon in the slender jugal and quadrato-jugal (figs. 1-3),
save that they coalesce in some degree, as in the Stanley Crane.

The palatines of the Kagu are very noteworthy; they are very steep, forming an
acute angle with the cranio-facial axis, and are sharply kecled (Pl. XCII. figs. 2, 3).
The pointed anterior portion has coalesced with the maxillary and palatine plate of the
premaxillary ; this part is nearly horizontal. The next part, on each side of the maxillo-
palatine plates, however, is oblique, and is twice as broad. Above the junction of the
middle and hinder portion there is a very large “ orbital plate” (fig. 3); it is a low triangle
in outline, and is convex on its outer side, curling inwards towards the “ parasphenoidal
rostrum.” This orbital plate is in relation in front with the posterior margin of the
“posterior septo-maxillary lobe,” and above with the vomerine fork of the same side.
The hinder part of the palatine is very steep; it is rather thin, but strong: there isa
small submesial keel (fig. 2) growing from the basicranial edge of the bone, and a
larger outer keel: both these keels send backwards a retral process; and that of the
outer keel belongs to the “transpalatine” region. This region is also indicated by a
small “ fenestra” (fig. 2). The oblong condyle at the end of the palatine, and much of
the basicranial edge, posteriorly, is formed by the once distinct ‘“ mesopterygoid,” a
strong wedge of bone with its sharp end forwards. In the Eurypyga these characters are
repeated in a somewhat softened form; but the fenestree marking the distinction into a
palatine and a transpalatine plate are larger, and there are one or two additional spaces,
as is the wont of these arrested clefts. These fenestree have the highest development
in Tigrisoma leucolophum, but they occur in other Ardeine and in the long-billed
Pluyialines.

In Psophia the transpalatine angles are more rounded, the keels are less sharp, the
bones are less steep, and there are no fenestre; altogether it comes much nearer the
typical Gruinz, whilst in the Kagu they are very Ardeine, the nearest form being
Nycticorax.

The pterygoids (Pl. XCII. figs. 2 & 3) are typically Gruine; they are small, and rather
slender, flattened anteriorly, with a keeled outer edge, a scooped under surface, and the
posterior third compressed, with a rounded upper and lower edge. The anterior con-
dyle is subconvex and three-sided ; the posterior is a round, shallow cup, surmounted by
a small, obtuse, ‘‘ epipterygoid” process, the counterpart of the columelliform epiptery-
goid of the Lizard.

The Kagu comes very close to the typical Cranes (and also to the Herons) in its os
quadratum (Pl. XCII. figs. 2,5); the two neat rounded upper condyles are far apart, and
the inner is scarcely behind the outer. The metapterygoid process is bowed out and
broad, the free terminal part being somewhat pedate. This bone is broad at its nar-

510 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

rowest part, and is much dilated at its base, which turns inwards and forwards. There
are a deep “ quadrato-jugal cup,” a high and neatly finished semioval convex “ pterygoid
condyle,” and two reniform convexo-concave condyles for the “articulare ;” the inner of
these is the lesser, and has the “hilum” looking backwards; the outer has its concave
margin looking forwards and inwards.

The mandibles are typically Gruine, but there is a good Ardeine character even in
them ; this is the long retral process between the mandibles (Pl. XCII. fig. 1a), growing
backwards from the ankylosed symphysis’. Here, again, the Kagu and the Eurypyga
correspond, whilst in the Psophia there is the merest rudiment of this part. ‘The fore
part of the dentaries is like the fore part of the intermaxillaries; further backwards
(Pl. XCII. fig. 3@) the ramus is high, as in Anthropoides, and only lowers a little in
front of the hinge, where it becomes thick also. The fore part of the “coronoid” and
the hinder part of the “splenial” is unankylosed; the “dentary,” also, has not coa-
lesced with the combined “ angular” and “ surangular.” The fenestra which appears in
the Psophia, between the forks of the dentary behind, is filled up; so is that which is
seen in the Stanley Crane and the Eurypyga above the root of the “coronoid” and
through the substance of the “surangular;” in this the Kagu agrees with the Grey
Heron: there is a trace of this fenestra in Mycticoraz. The convexo-concave condyles
(Pl. XCII. fig. 1 @) conform exactly to those of the os quadratum, the concavities and
convexities being reversed. The posterior face of the mandible is essentially Gruine, but
most evidently Ardeine also; it is triangular, the upper and outer edge forming a right
angle (Pl. XCII. fig. 4a).

The “internal angular process” is blunt, and perforated for the passage of air; the
‘* outer angular process” is rather sharp, and projects downwards as much as backwards.
In the posterior face of the mandible the Kagu agrees with the typical Gruine, but
makes some approach to the Ardeine; in the Ewrypyga the Ardeine characters prepon-
derate, whilst the same part in the Psophia is about equal to that of the Kagu.

The ceratohyals are lost in my specimen; I suppose that they were long, slender,
and unossified. The basihyal (figs. 8 & 8a) is rather long, slender, and well ossified ;
the urohyal is small, slender, and scarcely ossified at all; and the thyrohyals are slender
and feebly ossified.

The sclerotal ring is ossified ; but the bony plates are scarcely larger than in Psophia,
with a much smaller eyeball. In Dicholophus, which has the eyeball but little larger
than that of the Kagu, the sclerotals are twice as large and twice as strong, and the
diameter of the ring is but little short of an inch (114 lines) from edge to edge. In the
Kagu this diameter is 2 lines less; and in it the sclerotals turn forwards very little at
their inner edge; in Dicholophus this Owl-character is very strongly developed.

The vertebra vary as to number in the Kagu and its relatives, as follows :—

‘In this character the Herons and Pelecanine birds agree; in the Cormorant and Gannet this process is
well developed.

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. 511

| Cervical. | Dorsal. Lumbo-sacral. | Caudal (free).

Psophia erepitans ...........- | 18 6 17 6
Rhinochetus jubatus .......... | 16 5 15 7
Eurypyga helias .. 0.00226. 0 | 18 5 13 7
Botaurus viridis .......... 000% 13 4 14 6
Anthropotdes stanleyanus ...... | 19 5 17 7
Balearica pavoninad .......-.- | 20 6 |

GRUSMONG ONC eo ere aed oe ns 19 (Cieaacataain lian ate, | a

In the Psophia there is one pair of free cervical ribs, and one perfect lumbo-sacral arch ;
behind this there is a free styloid rib occasionally on the right side, the left coalescing
with the sacrum; there is one pair of abdominal ribs. In the Kagu there are two pairs
of free cervical ribs, and one nearly perfect lumbo-sacral arch (Pl. XCI. fig. 1); there
isno abdominal rib. In the Eurypyga the cervical ribs are all ankylosed to the ver-
tebre; there is one perfect lumbo-sacral arch, small free ribs to the second of that
series, and a very small abdominal rib on each side. In Botaurus viridis there are free
ribs to the last two cervical vertebre, a nearly perfect lumbo-sacral arch, and a pair of
styloid abdominal ribs.

In the Anthropoides stanleyanus there are two pairs of free cervical ribs, three pairs
of lumbo-sacral ribs, the last of which is apt to become ankylosed to the sacrum.

In the Balearic Crane there is only one arch developed from the lumbo-sacral region,
and it is perfect; but in Grus antigone two pairs of such arches exist, and a pair of styloid
floating ribs behind them.

In the Kagu, the Psophia, and the Eurypyga the first four dorsal ribs are ankylosed
together; in Grus antigone the last cervical and first two dorsal blend together; in
Balearica pavonina the first two dorsals only, whilst the Stanley Crane agrees with the
Bittern and the Heron in having these bones free of each other—at least for some
years.

The last caudal vertebra may, at least, count for seven (it is composed of ten in the
Duck) ; and thus the number six ought to be added to the seven caudal. We may thus
make a numerical comparison of the whole series as follows :—

Total Nr. of Vertebre.

ERO DEMIRCL DRUMS teh Mee Ta peta niclste + iaciektel eters ste eh 53
TRICO CR ALUS SUB OLS eta cats eval taes-)elp eka aeeraeley «aeons 49
UP YY GE EAE ae hase a oem wore sieisy eee lenis) s wehn ais == 49
PRODI RHEE eagle le Sa etn DON OIe ar a eae oe 43
Anthropoides stanleyanus .. 0.60.00. reer teers 54

These numerical comparisons are of considerable interest, thus:—We see that the
Eurypyga and the Kagu have the same total number of vertebri, although they are
disposed of differently, that the Psophia differs but little from the typical Cranes, that
the dorsal vertebre are more confluent in the aberrant Gruine forms than in the types,
and that this character is not constant in the typical genera.

VOL. VI.—PART VIII. 4B

512 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

The Ewrypyga is very typical in having a long neck, and very curiously isomorphic
of the Anhingas (Plotus) in the unusual elongation of this part’; for whilst it has five
more joints in this region than the Green Bittern, it has the individual vertebre as long
as in that true Ardeine bird. Yet this elongation of the cervical vertebrae, a most con-
stant character in the true Ardeinz, is not coupled with the swollen, spongy condition
seen in those birds, but the bones are quite Gruine in their histological characters.

But the Kagu differs from all its congeners in the stoutness of its cervical vertebrae
(Pl. XCI. figs. 1, 2); this is a correlate of its very large ridgy head, and is similar to
what is seen in the Baleniceps. This incrassation of the cervical vertebree is coupled
with a diminution of their number; and the strength of the head and neck of this bird
is in curious contrast with the feeble condition of many other parts of the skeleton.

In the cervical vertebra, as in the rest of the skeleton, the Kagu shows a very much
feebler ossification (especially with relation to the fibrous bands that are attached to
the bones) than the Psophia and the typical Cranes; in this it agrees with the Eurypyga;
and the difference between the vertebre of these two types is merely such as arises from
the slenderness of those of the Sun-Bittern and the breadth and shortness of those of
the Kagu.

The “atlas” of the Kagu (Pl. XCI. figs. 1 & 2, and Pl. XCII. figs. 12 & 12a) is a
strong ring of bone; its upper portion projects on each side behind; its centrum is tri-
dentate; and it has the ‘‘ cup” cut away in a rounded manner for the “ odontoid process
of the axis.”

The “axis” (Pl. XCLI. fig. 1 and fig. 2, 13, and Pl. XCII. figs. 13, 13 a, 13 6) has a thick
upper and a triangular lower spine; over each of its oblique facets there is a thick ridge;
and there is a small bony bridge on each side of the odontoid process (Pl. XCII. figs. 13a,
13).

The next vertebra (Pl. XCII. figs, 14, 14 a, 14, 14 ©) is intermediate in shape between
the “axis” and the cervicals of the hind region; its spine is smaller than that of the
“axis,” it has a short, stout rib, confluent with the sides of the bone and forming a
bridge for the vertebral artery. The next ten vertebre have scarcely any spine; the
sixth approaches in character the typical form of those of the hind region (Pl. XCI.
figs. 1 & 2,15, and Pl. XCII. figs. 15, 15 a,.15.6, 15). In this bone the transverse pro-
cesses project considerably; there is no process tending to wall-in the carotid artery
below; the centrum is deeply concave beneath, and is produced into two ears behind,
the transverse convexo-concave hinge being greatly produced outwards. The next four
have paired processes, tending to embrace the carotid artery; the penultimate bone of
these (Pl. XCL. figs. 1 & 2,16, and Pl. XCII. figs. 16, 16 a, 16 6, 16 ¢) is seen to be of great
breadth, and rather short, with its oblique facets very diverging. In the next five joints,
instead of the paired “carotid processes,” there is a flat inferior spine; in these the

1 The abortion of the “ ethmo-presphenoidal” band in the Sun-Bittern is in harmony with what we see not
only in the Stilt-Plover but also in the Cormorant.

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. ol3

upper spine begins to reappear, and in the last two the ribs do not become ankylosed.
The last but three (the thirteenth)’ has more breadth than length (Pl. XCI. figs. 1 & 2,
and Pl. XCII. figs. 17, 17 a, 17 b, 17 ¢); and this great development in the transverse
direction is still greater in the three last. The spine of the last cervical (PJ. XCL.
figs. 1 & 2, and Pl. XCII. figs. 18, 18a, 180) is very high and also very thick; it
has a low inferior spine, a large transverse process that projects beyond the “ tuber-
culum ” of its styloid rib, and a neat cup on each side for the corresponding “ capitulum
cost ;” the styloid feeble ribs (Pl. XCI. fig. 2, 18) are less than an inch in length.
The five dorsal vertebre are all ankylosed together, except the last, and in the free
bone the inferior crest has died out (Pl. XCI. fig. 1,d5). The upper spine of the first
dorsal is higher than that of the second; they then increase in height until we reach the
third sacro-lumbar. As the tendons are but little ossified in the Kagu, the transverse
processes of the dorsals are permanently distinct (Pl. XCII. fig. 5d); the bodies of the
second, third, and fourth are very thin and carinate.

In the height of the dorsal and lumbo-sacral spines the Kagu comes near to Ocydro-
mus; in the Kagu the third sacro-lumbar attains its greatest height (Pl. XCI. fig. 1);
behind this the spine aborts rapidly, and the hinder two-fifths of the sacrum has no
spine. ;

The eleventh sacral has a long, unsegmented rib on each side; for this part of the
sacrum is of great width (Pl. XCII. figs. 5 & 7); behind the eleventh the component
vertebre gradually lose the distinction in rib and transverse process. The sacrum
(Pl. XCI. fig. 1) has its last third more bent downwards than in any bird I know, and
the caudal series (cd) takes the same downward direction. The caudal vertebre of the
Kagu (Pl. XCI. fig. 1, ed, and Pl. XCII. figs. 5 & 7, cd) are furnished with long
transverse processes, as in the Huwrypyga; in this they both differ from the Psophia.
The last piece (Pl. XCII. fig. 1, cd) is sharper than in either of the related types. The
ribs (P]. XCI. fig. 1, and Pl. XCII. fig. 5) are feebler than in any related type, and the
appendages are small, and only present on the first four. The upper part of the ribs is
very much enlarged in Psophia, Ocydromus, and Eurypyga, much less so in the Kagu.
The sternal pieces (s7) are also feeble, and that belonging to the sacro-lumbar rib
(s./r) is imperfect below; altogether, the ribs are very feeble and embryonic.

But the shoulder-girdle and sternum show the most remarkable embryonic characters,
and haye their only counterpart in this respect amongst the Carinate in the feeblest-
winged Rail (Brachypteryx australis). Ocydromus has a more perfect sternum.

The scapula (Pl. XCI. fig. 3, sc, and Pl. XCII. fig. 5, sc) is an extremely feeble and
very curved bone; it is only half the relative width of the scapula of Psophia and
Eurypyga; but in these types this bone is very much curved. The acromion process
(Pl. XCII. fig. 5, sc) is blunt and of a moderate size, and the suprascapular tip, of the

1 In Plate XCI. fig. 2, the penultimate vertebra is numbered 17 by mistake ; in Plate XCIL., fig. 17 repre-
sents the thirteenth vertebra.

4B2

o14 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

bone is rounded. The coracoid (Pl. XCI. fig. 5, cr, and Pl. XCII. fig. 6, mer) is an
unusually long and feeble bone; its head is hooked and tuberculate, and its meso-
coracoid process (mcr) is square; this process, as in the Ewrypyga, is continuous with a
sharp keel that runs along the inner side of the bone, and becomes considerably
developed at the angle below. At the outer angle below, the epicoracoid hook (eer)
is nearly obsolete. ‘The coracoids (Pl. XCII. fig. 6, cr) are separated below by the
width of the anterior sternal notch; these bones are most slender just above the middle,
where they are pinched to little above a line in thickness, whilst measured across the
epicoracoid region they are five lines in breadth. The fureula (fr) is of a narrow
U-shape; it is a much stronger bone than that of Brachypteryx or Ocydromus: in
Psophia and Eurypyga the bone is stouter, more V-shaped, and has a distinct inter-
clavicular process; in the Kagu this part is much aborted. ‘The whole outline is
that of three-fourths of a very elegant ellipse (Pl. XCII. fig. 6, fr); and the bone is
widest below and above; the upper outcurved parts are broad and flat; and there is a
very evident enlargement from the “ precoracoid segment ”’ where it articulates at its
side with the hooked head of the coracoid, and another addition at its tip where it joins
the acromion (mesoscapula); this additional piece, which has become confluent with
the clavicle on each side, is the ‘mesoscapular segment.” (See ‘Shoulder-girdle and
Sternum,’ Ray Society, 1868, p. 179.)

In relative size, general form, and in the degree of its morphological development,
the sternum of the adult Kagu agrees with that of the ripe chick of a typical Crane
(op. cit. pl. 14. p. 158). The curvature of the bone is so great that the lower view of
the shoulder-girdle, when fairly seen, gives a foreshortened view of the sternum (see
Pl. XCII. fig. 6, where the sternum appears too short by half an inch); the lateral views
(Pl. XCI. figs. 1 & 3, st) show the elegant curvature that the xiphoid part of the
sternum makes, passing both upwards and backwards to support the abdominal viscera.
As in the Psophia and in the typical Cranes, the sternum of the Kagu is as narrow in
front as in the Rallide, but further backwards it narrows-in still more, and then dilates
somewhat at the xiphoid end (Pl. XCII. fig. 6, 2): this part is only slightly trifid. The
general feebleness of the Kagu’s sternum will be well seen by comparing it (Pl. XCI.
figs. 1 & 3, and Pl. XCII. fig. 6) with that of the entirely unossified sternum of the
newly hatched Mantchouri Crane (see ‘ Shoulder-girdle and Sternum,’ pl. 14. figs. 6-8).
Even at that early stage the sternum of the typical Crane has a much larger keel (which
articulates with the “furcula”), a definite “rostrum” in front overlapping “ coracoid
grooves,” and has the normal Pluvialine fission of the xiphoid region of the sternum
into middle, intermediate, and external xiphoid processes.

The general curvature of the sternum is very similar in both these instances. The
costal processes (Pl. XCI. fig. 3, ¢.p) are long, triangular, and are hooked inwards
behind the posterior face of the coracoid.

The two triangular flaps in which the sternal keel terminates in front in the embryo

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. 515

of Grus (op. cit. pl. 14. figs. 6,7) have their counterpart in the thickened projecting
lobe of the sternal angle in the Kagu (Pl. XCI. figs. 1 & 3). The primordial notch
between the coracoid grooves (Pl. XCII. fig. 6) is quite similar to what is seen in the
Cassowary (op. cit. pl. 17. fig. 3), and in Brachypteryx amongst the feeble-winged Land-
Rails. The ridge on the sternal keel of the Kagu, indicating the origin of the pectoral
muscles, shows the extreme thinness of the large pectorals ; altogether, the outer face of
the sternum bespeaks a very feeble development of the muscular masses.

The wing of the Kagu (Pl. XCI. fig. 1, and Pl. XCIL. fig. 5) is much feebler than that of
either the Psophia or Eurypyga, but it is considerably stronger than that of Ocydromus.

In Eurypyga the arm is 24 lines shorter than the forearm, in Psophia it is 2 lines
longer ; in Ocydromus the arm is 7 lines longer than the forearm, but in the Kagu
the arm is almost equal to the forearm; so that the wing of the Kagu, whilst very
inferior to that of the Eurypyga, is somewhat better proportioned, although relatively
feebler, than that of Psophia; it is much stronger than that of the great Land-Rail
(Ocydromus).

The humerus of the Kagu (h) is perfectly typical; but the overhanging anterior
crest for the insertion of the “ pectoralis major” is much less sharp and outstanding
than in Ewrypyga; the condyles are quite normal; but the rounded, thick process below
the lower condyle for the ulna is very large (see Pl. XCI. fig. 1,4, wu, where it is seen
lying below the end of the ulna, on the os femoris, f). The radius (r), ulna (7),
upper and lower carpals (c), free and coalesced metacarpals (mc), and digits (d) are
feeble, but quite normal. There is some appearance of a second free digital joint to
the pollex (as is seen in Porphyrio) (Pl. XCL. fig. 1), but this is somewhat doubtful; the
whole of the “ hand” is shorter than the other two regions by about a line in extent.

The following Table will show the relative lengths of the three regions of the wing in
the Kagu and its congeners :-—

| Arm. Forearm. | Hand. |
in. lin. in. lin. in. lin |
TER ULOCILELUS oysters Rus otto yon easiest see | 2.3 2 ar al |
Hr apPy GO Heh.iera valley ote ata cede = eeeh fot let Dias 2 32 2 02
IGE nase ee OI Oe PA 12a ero o SOOT e | 3-03 2 103 Deh, |
Ohi ie Sere dao hon uma sc eda clothe lio nDP Oe 1 8 1 10

The pelvis of the Kagu is very peculiar (Pl. XCI. fig. 1); it is bent upon the spine
to a greater degree than in any bird I have examined, and the height of the iliac
crests and of the third sacro-lumbar spine is very remarkable. This condition is seen,
but in a less degree, in Psophia, Ocydromus, and Talegalla; but in Eurypyga the whole
pelvis is as straight, as broad, and as flat as in the Umber (Scopus); indeed the
resemblance of the pelves of these two types is very great. But if we come to minutie,
the pelvis of the Eurypyga and that of the Kagu agree very closely, notwithstanding the
compression and elevation of the latter and the broad outspread condition of the former.

516 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

In curious contrast with the smooth and rounded condition of the Kagu’s sternum is
the high, ridgy pelvis, the whole structure being scooped, carinate, and in every way
forming a fit foundation for most powerful muscles. The high iliac crests (Pl. XCI.
fig. 1, ¢/) culminate on each side of the third sacro-lumbar vertebra, the spine of which
they strongly clamp, like the relation of the suprascapula of the Skate to the cervical
and anterior dorsal spines (‘Shoulder-girdle and Sternum,’ pl. 1. figs. 1 & 2, ssc, v).
This junction of the sacro-lumbar spine with the iliac crests keeps unankylosed
(Pl. XCII. fig. 5. s7,70) as in Hurypyga; at any rate it is so in this specimen, the age
of which I cannot tell, but which is evidently quite adult. In Psophia, as in Grus
and Ocydromus, these parts coalesce. Here let it be remarked that the pelvis of Psophia
is more like that of Ocydromus than that of the typical Crane.

The angle formed by the iliac crest as it bends downwards towards the acetabulum
is quite a unique condition. There is as distinct a “ preacetabular spur” in the
Psophia as in Talegalla; there is scarcely a trace of this process in Grus proper, none
in the Kagu and Eurypyga, but in Ocydromus it is as large as in the typical Galline.
The canals formed by the junction of the iliac crests with the sacro-lumbar spine are
equally deep in the Kagu and Ocydromus; they are open in Eurypyga, where the crests
keep wide apart, and are filled in by bony matter in Psophia and Grus. The acetabular
fenestrae are very large in all these related Gralle, the articular surface being a mere
crescent below ; above, there is an oval, slightly concave facet for the “ trochanter major.”
The overhanging crest of the postfemoral part of the ilium (Pl. XCI. fig. 1, i, and
Pl. XCIL. fig. 5) is nearly as strong as in the Psophia. The descending ilio-ischiadic
plate (Pl. XCL. fig. 1, ¢/./se) is relatively deeper than it is even in Psophia, but in this
latter bird this plate is less notched than it is in Ocydromus: the notch is more definite
in Grus; but in the Kagu, as in Huwrypyga, this notch is very deep, and the ilium and
ischium both end in long processes. The long ‘‘ obturator notch ” (0.7) and the ovoidal
ischiadic fenestra (is. f) are quite alike in these two species. The ischiadic fenestra is
broad behind alike in Ocydromus, Psophia, the Kagu, and in Eurypyga; but in Grus the
posterior end of this space is narrow. ‘The pubis (4), which is nearly straight in Ocy-
dromus, Scopus, and Psophia, is much more curved and sigmoid in Grus, and still more
so in Eurypyga; but this curvature is greatest in the Kagu, it becomes almost suddenly
bent upwards below the angle of the ischium, and is then deflected inwards in some
considerable degree towards its fellow of the opposite side. In its rounded and very
feeble condition the pubis of the Kagu comes closest to that of Eurypyga; for in Psophia
it is a stout and even a broad bone, much like that of the Rail (Ocydromus). ‘The pubis
of Grus, making allowance for its great size, is intermediate between that of the Kagu
and Psophia.

With regard to the serial homologies of the pelvis it may be remarked that the
whole of the spinal edge of the ilium is the counterpart of the upper edge of the
suprascapula; this is best seen by comparing the ilium of the bird with the supra-

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU., 517

scapula of the Skate (‘Shoulder-girdle and Sternum,’ pl. 1. figs. 1-4, sev). The
great backward extension of the ilium in the bird is illustrated by the large, flappy
suprascapula of the Lacertians (op. cit. pls. 9-11, s.sc); in the Rhea, and in a less
degree in the Apteryx, the ilium stops short of the ischium behind. The manner in
which the three main elements of the “os innominatum ” meet in the “ acetabulum” is
beautifully illustrated in the shoulder-girdle of the Batrachia, especially in Dactylethra
(op. cit. pl. 6. fig. 11). The pelvic counterpart of the great notch between the supra-
scapula and scapula above and the coracoid below is filled up behind in most birds by a
continuous growth of cartilage, so that we have not an ischiadic “notch,” but a large
“fenestra.” In the Skate (op. cit. pl. 1. fig. 2, sc. f, cr. f) there are two fenestre in the
corresponding region of the shoulder-girdle moiety. In the Apteryx, the Emu, the
Cassowary, the African Ostrich, and the Tinamous (Trans. Zool. Soc. 1864, vol. v. part 3,
pl. 39, 2/,7sc) this space is open in the same manner as in the shoulder-girdle of the
Frog. In Dactylethra, im the Chelonians, and in the African Ostrich the space be-
tween the precoracoid and coracoid (the counterparts of the pubis and ischium in the
pelvis) is a large, deep notch (‘ Shoulder-girdle and Sternum,’ pl. 6. fig. 11, pl. 12. fig. 2,
and pl. 17. figs.5,6). In birds, generally, the “ obturator” space is a very deep notch;
in the Rhea it is a huge “fenestra” from the first; in the Buceride it becomes so by
coalescence of the pubis with the ischium; these differences are all in harmony with
what is seen in the various conditions of the shoulder-girdle. The moieties of the
hip-girdle only meet below and coalesce in the African Ostrich; in other birds they
keep apart; in the Toad (op. cit. pl. 5. figs. 15-17) the counterparts of the pubes meet
and coalesce, this being, however, an exceptional condition as to the Batrachia, and
borrowed from the Skate and Shark (op. cit. pl. 1. figs. 1-4).

In certain Fishes the scapular region becomes segmented from the coracoid (op. cit.
pl. 2. fig. 12, se, per); in certain Amphibia, as in some other fishes (e. g. Trigla, Agonus,
Gobius, op. cit. pl. 2. fig. 13), this connectivum is only partially separated; this partial
separation appears again in the rudimentary cleft seen in the fundus of the glenoid
cup in Proteus and Cryptobranchus (op. cit. pl. 3. figs. 1 & 3, gl).

In birds the pelvic counterpart of this cleft is constantly present, and its large size
and irregular shape is well illustrated in Professor Owen’s paper on Cnemiornis (Trans.
Zool. Soc. 1866, vol. v. part 5, pl. 64. fig. 7 a).

Hence it will be seen that the hip-girdle is the same in its morphology as the
shoulder-girdle, that in the Class of Birds its spinal crest is hugely developed but per-
fectly normal, and that the whole of each moiety is a cartilaginous plate superadded to
the axial skeleton and tending to undergo fission in exactly the same manner as the
shoulder-girdle moiety. Also it is evident that the hip-girdle is always of a lower
morphological type than the shoulder-girdle, its fission and general metamorphosis
being so much more arrested in the same individual type; so that, whilst the shoulder-
girdle of the Skate illustrates the hip-girdle of the Bird, the hip-girdle of the Mammal

518 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

and the shoulder-girdle of the Frog lie very nearly on the same morphological level
(op. cit. pls. 6 & 7).

With regard to the setting-on of the limb-girdle moieties, it will be seen that there
is the greatest liberty with regard to the angle these plates form with the vertebral axis.
As they are merely supplementary parts, and as they appear between the skin and the
axial cartilages and muscles, there is nothing to prevent them shooting along the sub-
cutaneous plane in any direction. In the Bird-class, where the skeleton is so marvellously
modified in correlation with the functions and habits of a flying creature, the upper
edge of the shoulder-girdle is directed very far backwards, the direction, from the narrow
suprascapular top, being forwards and a little downwards to the glenoid region, and
then suddenly downwards and backwards, the upper and lower halves lying at an acute
angle. In the hip-girdle, on the contrary, the true apex is turned forwards, but runs
also backwards by a very long upper margin, the general direction of the whole plate
being backwards and a little downwards. The Kagu is peculiar among birds for a
much more downward direction of the hip-girdle moiety than is common in its Class;
but even in it there is but little approach to the condition of these parts in the
Lacertian, where the ilium is set on to the spine at almost a right angle.

With regard to the arrested metamorphosis of the hip-girdle in the Bird, it is worthy
of remark that the pubis (the counterpart of the precoracoid bar which is the subject
in the Bird and in the Mammal of such a large amount of morphological change) does
die out at its anterior part in certain Raptores—for instance, the Falcons, Hawks, &c.
The pubis, also, like the precoracoid, is the first to become enfeebled and modified,
although in a less degree.

The hinder limbs of the Kagu (Pl. XCI. fig. 1, and Pl. XCII. figs. 9-11) are very
much like those of its immediate congeners the Psophia, the Eurypyga, the Cranes,
and the Rails. The “os femoris” (Pl. XCI. fig. 1, f, and Pl. XCII. figs. 9, 9a, 90) is
slender, more arcuate, and longer, relatively, than in Psophia and Eurypyga, and there-
fore more Ralline. A Table showing the comparative lengths of the femur, tibia,
tarsus, and middle toe will illustrate this :—

Femur. Tibia. | Tarsus. Middle toe.
in. lin. in. lin, | in. lin, in. lin,
Rhinochetus jubatus .........- 2 7 4 9 3 11 2 33
Eurypyga helias..... are eetetttonaye As 3 0 Broil Lh
Psophia crepitans .. 1.6.22 204. 210 5 10 ae fs! 2 4
Porphyrio poliocephalus........ 2 103 5 4 3.9 4 03
Ocydromus australis .... 2.146: 3 24 4 63 278 2 64

Here it is shown how extremely variable the relative proportions of these regions are
even in birds so closely related as those given in the list. Altogether, the bones of
the hinder limbs are much slenderer in the Kagu than in Psophia, Ocydromus, and

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU. 519

Porphyrio. Notwithstanding the greater slenderness and inferior size of the tibia
of the Kagu (Pl. XCI. fig. 1, ¢, and Pl. XCII. figs. 10, 10a, 106) as compared with
that of Porphyrio and Ocydromus, yet the crests at its upper part (ectocnemial, ento-
cnemial, and epicnemial) are quite equal to what is seen in those large Rails. ‘This
brings us to consider the extraordinary development of these parts in that extinct
Grallatorial bird which Professor Owen has called Cnemiornis (see Zool. Trans. 1866,
vol. v. part 5. pl. 66, p. 401). The condition of the parts in that huge type is merely
an exaggeration of what is seen in Porphyrio and Ocydromus, and especially, also, in
the Kagu.

Both extremities of the tibia of the Kagu are large as compared with the slender
shaft; the front view of the bone (Pl. XCII. fig. 10) shows this, and also how closely
this bone corresponds with that of Cxemiornis. The lower condyles are very strongly
marked (Pl. XCII. figs. 10 & 102); there is an oblique tendon-bridge in front, a cleanly
cut notch above the ectocondylar elevation, and between the two condyles there is a
large cotyloid cavity for the precalcaneal knob. ‘The slender fibula (Pl. XCI. fig. 1, fb)
is three-fifths the length of the tibia; the patella (Pl. XCI. fig. 1, p) is rather larger.
The separateness of the lower articular portion of the tibia of this bird was familiar to
me in my early days, and the drawings made by me of these parts in the young Emu
twenty-four years since “are alive to testify” to this. Afterwards, when writing upon
the osteology of the Balwniceps (Trans. Zool. Soc. 1861, vol. iv. part 7, p. 343), I strongly
doubted the merely epiphysial nature of this piece, and put this question: “Is it the
homologue of the mammalian astragalus?” This question has now been definitively
settled by Professor Gegenbaur (see his paper “ Vergleichend-anatomische Bemerkungen
iiber das Fussskelet der Vogel,” Archiv fiir Anat. u. Phys. Jahrgang 1863, p. 455).

This apparent obliteration of the tarsal bones is a remarkable feature in the pelvic
limbs of the bird; after all only two segments are found, one coalescing with the tibia
and the other with the three main metatarsals; for the sesamoid bone occasionally found
behind the joint is not an “os calcis,” but a tendon-bone. At first sight, the tarso-
metatarse of the Psophia and that of the Kagu seem to have scarcely anything, except
the superior size of the former, to distinguish them; but there are many important
differences. The whole piece is much more feebly ossified in the Kagu than in the
Psophia, and this to a degree that is very remarkable for a Bird, reminding the observer
of the Penguin and of young birds generally. The ectotarsal and entotarsal keels
behind the upper head of the bone are only united by membrane (PI. XCII. fig. 11 a);
they are connected by a bony bridge in Psophia: the Kagu and Eurypyga agree as to
the condition of these parts.

The great “lower tarsal” is but feebly molten into the heads of the main metatarsals
(Pl. XCII. fig. 11); and these three long bones keep their sutures for a long time, and
have large “fenestre” between them above—a very remarkable and interesting cha-
yacter. There is very little trace of these spaces in Psophia, Eurypyga, Ocydromus, and

VOL. VI.—PART VIII. 4c

520 MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

Porphyrio. The posterior sharp edges of the metatarsus seen in these birds have
scarcely any existence in that of the Kagu. As in these its congeners, the Kagu has
the outer lower condyle connected with the middle condyle by a bony bridge, so that
the considerable open space between these two, below, is divided into a fenestra and a
notch. The condyles themselves (Pl. XCII. figs. 11 & 114) differ in the Kagu from
those of its congeners; they are much more outspread (Reptilian) than in the Psophia,
whilst in Ewrypyga they are more compressed than even in Porphyrio, equalling what is
seen in the Coot (Fulica atra), and approaching the condition of these parts in the
Grebes (Podiceps). The small free metatarsal (Pl. XCI. fig. 1,7. m.t) is placed high
up, as in the congeners of the Kagu ; and the “ hallux”’’ is of moderate length. The rest
of the toes are slender, and rather long (Pl. XCI. fig. 1), the middle toe (d. 3) being: more
than an inch and a half shorter than the tarso-metatarse. In the Psophia the middle
toe is only half the length of the shank; in Ewrypyga the middle toe is nearly two-thirds
the length of the shank; whilst in Porphyrio it is a quarter of an inch longer than the
tarso-metatarse.

These variations are worth mentioning because of their zoological value; morpho-
logically, however, they have much less import, as they relate mainly to the correlation
of each type to its surroundings in actual life.

In summing up the affinities of the Kagu, I may say that my view of it is that it is a
generalized Crane, that it is nearer of kin to Eurypyga than to Psophia—the latter
coming near to the Balearic Crane, whilst Eurypyga, like the Kagu, makes a very near
approach to the Night-Herons amongst the typical Ardeine. The Kagu is related to the
Rails ; but so, indeed, are all the Gruinz ; and Professor Huxley has, with great sagacity,
put both these families into one group, and has called this group the “‘ Geranomorphe ”
(see Proc. Zool. Soc. 1867, p. 457).

EXPLANATION OF PLATES XCI. anp XCII.

PLATE XCI.
Fig. 1. Side view of skeleton.
Fig. 2. Upper view of cervical vertebra.
Fig. 3. Side view of sternum and shoulder-girdle.

* The hinder toe is evidently the true “hallux;” but in my former papers it was described as the second
(d. 2), on the supposition that the spur of the Cock represented the innermost digit.

Fig

Fig.

Fig

g.
Fig.
Fig.
Fig.
Fig.

MR. W. K. PARKER ON THE OSTEOLOGY OF THE KAGU.

PLATE XCII.

1. Upper view of skull.

la. Upper view of lower jaw.
2. Lower view of skull.

3. Side view of ditto.

3a. Side view of lower jaw.
4, End view of skull.

4a. End view of mandibles.

. 5. Upper view of trunk and wings.
. 6. Lower view of sternum and

shoulder-girdle.

. 7. Lower view of pelvis and tail.
. 8. Upper view of os hyoides.

g. 8a. Side view of ditto.

. 9. Front view of os femoris.

. 9a. Upper view of ditto.

. 96. Lower view of ditto.

g. 10. Front view of tibia.

ig. 10 a. Upper view of ditto.

. 106. Lower view of ditto.

. 11. Front view of tarso-metatarse.
. 11 a. Upper view of ditto.

. 116. Lower view of ditto.

. 12. Side view of atlas.

. 12 a. Front view of ditto.

g. 13. Upper view of axis.

. 13a. Front view of ditto.

Fig.
Fig.

13 5. Posterior view of axis.
14. Upper view of third cervical
tebra.

Ver=

. 14a. Lower view of ditto.

. 146. Front view of ditto.

. 14c¢. Posterior view of ditto.

. 15. Upper view of sixth cervical

ver-
tebra.

. 15 a. Lower view of ditto.

. 156. Front view of ditto.

. 15 c. Posterior view of ditto.

g. 16. Upper view of ninth cervical ver-

tebra.

. 16 a. Lower view of ditto.

. 166. Front view of ditto.

. 16 ¢. Posterior view of ditto.

. 17. Upper view of thirteenth cervical

vertebra.

. 17 a. Lower view of ditto.

. 176. Front view of ditto.

. 17 ¢. Posterior view of ditto.

. 18. Side view of sixteenth (last) cervical

vertebra.

. 18a. Posterior view of ditto.
. 18%. Anterior view of ditto.

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RHINOCHETUS JUBATUS (nat-sye-.)

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INDEX OF SPECIES, ETC., IN VOL. VI.

Acanthias vulgaris, 384, 396.
Acanthopterygii, 385.

Acanthurus chirurgus, 387.

Acara ceruleopunctata, 384, 391, 405, 449.
Acronuride, 387.

Alurichthys dorsalis, 405.

—— nuchalis, 393, 476.

panamensis, 383, 393, 405, 476.
Aipyornis, 70.

Aétobatis latirostris, 397, 491.
Agonostoma microps, 389, 403, 444.
monticola, 390, 405, 445.
nasutum, 384, 390, 404, 405, 444.
Agonus, 517.

Albula conorhynchus, 395.

Alea impennis, 70, 76.

Aleuteres monoceros, 396.

Alligator, 160, 162, 168.

cuviert, 168.

cynocephalus, 160, 163.
gibbiceps, 166.

Lucius, 128 note, 168.

—— mississippiensis, 128 note, 168.
niger, 128 note, 163.
palpebrosus, 154, 167.

—— punctulatus, 161, 165.

sclerops, 160, 162, 163.
Alligator, banded, 167.

, rough-backed, 167.
Alligatoride, 130, 160.

Alphestes afer, 411.

Amblyopus brevis, 389, 441.
Amblyscion argenteus, 383, 425. ,
America, Central, fishes of, 377-491.

, fish-provinces of, 403-406.
, list of fishes of, 385-397.
——,, topographical features of, 378-381.
Amia, 506.

Amiurus meridionalis, 393, 404, 473.
Anableps dovii, 394, 403.

VOL. VI.—PART VIII.

Anacanthini, 392.
| Anacyrtus quatemalensis, 394, 403, 406, 479.
Ancistrus cirrhosus, 384.
Auhingas, 512.
Anisotremus teniatus, 414.
| Antennarius leopardinus, 388, 439.
tenuifilis, 388, 440.
Anthropoides, 503, 510, 511.
stanleyanus, number of vertebre in,
511.
Aphoristia ornata, 392, 473.
Apogon dovit, 385, 413.
inermis, 413.
Aptenodytes, 70, 76.
Apteryx, 68, 74, 76, 517.
Aptornis, 70, 79.
| otidiformis, 78.
«« Aquez palin,” 145.
Ardeine, 503, 504.
| Argyriosus setipinnis, 388.

vomer, 388.

Arius assimilis, 393, 404, 475.
| caerulescens, 393, 403.
—— dovii, 383, 393, 476.
guatemalensis, 393, 403.
melanopus, 393, 404.
—— multiradiatus, 384, 393, 405.
—— platypogon, 393, 403.
seemanni, 393.
troscheliz, 383, 393.
Ateles, 214.
Atherinichthys pachylepis, 389, 443.
guatemalensis, 389, 443.
Atherinide, 389.
Atractosteus tropicus, 383, 490.
Auchenopterus monophthalmus, 442,
Bagrus? arioides, 384.
Bairdidla armata, 383, 428.
Balena, 44, 45, 115.
——, cervical vertebrie of, 328.

4D

524 INDEX OF SPECIES.

Balena mysticetus, accessory pelvic bones or cartilages

of, 364,

, weight of ribs of, 351.
Baleeniceps, 512, 519.

Balenide, 115.

Balenine, 115.

Balsnoidea, 110, 115.

Balenoptera, 4, 115.

Balenopteride, 115, 329.

Balenopterine, 115.

Balearica, 503, 511.

pivonina, number of vertebre in, 511.
Balistes frenatus, 396, 490.

niger, 396.

vetula, 396.

Batrachia, 517.

Batrachide, 388.

Batrachus liberiensis, 435.

pacifici, 383, 388, 435.
surinamensis, 388.

Belonesox belizanus, 394, 404.

Beluga, 96, 97, 98, 114, 115.

, atlas of, 330.

, carpus of, 361.

——,, cervical vertebre of, 329, 330, 333.
lewcas, vertebral formula of, 349.
Belugine, 115.

Bernicla brenta, 119, 123.

ruficollis, 375.

Bird, Didine, of the island of Bourbon, 373.
, raptorial, 117.

Birds, fossil, from the Zebbug Cave, Malta, 119-124.
, homologies of the maxillary bones of, 501,

502, 505, 506.

, homologies of the pelvis in, 517, 518.
Bittern, Green, 512.

Black Dolphin, 23.

Blenniide, 389.

Blennius brevipinnis, 389.

“ Bolla gadimi,” 21.

Bombifrons, 135, 136, 137, 139.

—— indicus, 128 note, 140, 149, 150.
siamensis, 128 note, 144,
trigonops, 140.

Botaurus viridis, number of vertebre in, 511.

Brachypteryx, 514, 515.
australis, 513.
Brotula multibarbata, 392, 471.

Brycon dentex, 478.
Buceride, 517.

Bustards, 72.

Cachalot, osteology of the, 309-369.
, Snub-nosed, 30.
Cachalots, 112.

Caiman, 160, 161, 166.
fissipes, 160, 161, 163.
moschifer, 167.

—— niger, 160, 162.
palpebrosus, 167.
—— punctulatus, 165.

trigonatus, 167.

Caiman 4 paupiéres osseuses, 155.
Caprimulqus europaeus, 507.
Carangide, 387.

Carangoides dorsalis, 383, 432.
Carangus amblyrhynchus, 387.
caballus, 388, 431.

caninus, 388, 432.

carangus, 388.
crumenophthalmus, 387.
—— dorsalis, 388, 432.

—— hippos, 383, 388, 431.
leucurus, 387, 430.
marginatus, 383, 431.
panamensis, 383.

speciosus, 383, 387, 431.
Carapus fasciatus, 395, 404, 489.
Carcharias maculipinnis, 396, 490.
Cariama, 72, 508.

Cassowary, 517.

Casuarius, 70, 74, 76.

Catodon australis, 32.
macrocephalus, 33,
Centropomus appendiculatus, 385, 406.
armatus, 383, 385, 408.
—— ensiferus, 385, 408.

medius, 385, 406.

nigrescens, 385, 407.
parallelus, 385, 407.
Centropristis macropoma, 385, 409.

Cepphus grylle, 375.

Cetacea, 17, 87,171.

, classification of the, 113-115.

, comparison of characters of, 44, 45.

——., progression or succession in, 45.
——,, teeth of, 44.

INDEX OF SPECIES.

Cetengraulis mysticetus, 395, 489.
Cheetodon capistratus, 386.

—— humeralis, 386, 419.

striatus, 386.

Cheetostomus aspidolepis, 393, 405, 477.
cirrhosus, 393, 405, 478.
Chalceus atrocaudatus, 384.
Chaleinopsis chagrensis, 384, 394, 406.
dentex, 394, 404, 478.
striatulus, 384, 394, 406.
Champsa, 134, 161, 168.

Jissipes, 138, 161, 163.

—— gibbiceps, 167.

nigra, 161, 162.

palpebrosa, 167.

—— punctulata, 161, 165.

sclerops, 161, 165.

—— trigonata, 167.

vallifrons, 161, 165.

Chanos salmoneus, 395.

Characinide, 393.

Characodon, 480.

lateralis, 394, 480.

Charadrius, 74,76, 77, 79.

Chatoéssus petenensis, 395, 404, 488.

Cheiromys, 214.

Chelonians, 508.

Chimpanzee, limbs of, compared with those of the
Orang, 175-218.

Chorinemus altus, 388, 433.

imornatus, 383, 388, 433.

occidentalis, 388.

saliens, 388.

Chromides, 391.

Chromis atrilobata, 446.

Chrysophrys calamus, 386, 421.

Ciconia, 76, 79.

alba, 375.

Cirrhites rivulatus, 421.

Oirrhitichthys rivulatus, 387, 421.

Cirrhitide, 387.

Citharichthys guatemalensis, 392, 472.

spilopterus, 392, 471.

Clangula glaucion, 375.

Clarias, 506.

Clinus delalandii, 389.

macrocephalus, 389, 442.

—— nuchipinnis, 389.

525

Clupea libertatis, 395, 403, 487.
Clupeide, 395.

Clupeoids, 506.

Cnemiornis, 70, 78, 79, 517, 519.
Coati, 505.

Cockatoos, 69.

Columba galeata, 77.

magnifica, 75.

palumbus, 77.

Columbide, 72, 73.

Colymbus, 76.

Conodon pacifict, 386, 417.
Coot, 508, 520.

Cormorant, 512 note.

Corvina armata, 383, 387, 428.
chrysoleuca, 387, 427.
— ophioscion, 383, 387, 428.
ronchus, 387.
vermicularis, 387, 427.

Cossyphus diplotenia, 390.
pectoralis, 390, 447.
—— pulchellus, 447.
rufus, 390.
Crane, Balearic, 511.
——, Mantchouri, 514.
, Stanley, 504, 509, 510.
Cranes, 72, 503, 504, 509, 514, 518.
Cremnobates, 442.
monophthalmus, 389, 442.
Crocodile, Black, 166.
——, Black African, 153.
——, Green, 147.
——, Narrow-beaked, 132.
——,, Olive African, 147,
——, Orinoco, 151.
——, Pondicherry, 139.
— 4 nuque cuirassée, 159.
de l’Orénoque, 151.
Crocodiles, American, 145, 150.
, Asiatic, 139.
——, Brackish-water, 137.
——, Estuarine, 137.
—., Fluviatile, 139.
——., Gavialian, 156,
——, Normal, 137,
——, River, 139.
, Saltwater, 138.
Crocodilians, geographical distribution of, 130, 131.
4p2

526 INDEX OF SPECIES.

Crocodilians, synopsis of, 125-168.
Crocodilide, 130, 132, 134.
Crocodilus, 134, 135, 136, 137, 146.

—— americanus, 150, 151, 158, 165.
arctirostris, 133.

binuensis, 147, 149.

biporcatus, 128 note, 138, 140, 150.
raninus, 138.

biscutatus, 150, 155, 157, 158.
—— bisulcatus, 157.

bombifrons, 140.

—— caiman, 165.

—— cataphractus, 127 note, 133, 147, 157.
—— chamses, 147.

—— cuvieri, 168.
dubius, 140.
frontatus, 153, 154.
galeatus, 144.

—— gangeticus, 133.

—— gravest, 133, 145.

—— intermedius, 151.

—— lacunosus, 147, 149.

—— latirostris, 163.

—— leptorhynchus, 157, 159.
longirostris, 133.

—— lucius, 168.

—— marginatus, 147, 149, 150.
—— mississippiensis, 168.

—— moreletii, 146.

niger, 153.

—— niloticus, 144, 147.

oopholis, 138.

palpebrosus, 153, 154, 155, 167.
palustris, 128 note, 140, 144, 149.
planirostris, 145.

pondicerianus, 139.

porosus, 138.

—- rhombifer, 140, 145.
— schlegelii, 133, 134, 156.
—— sclerops, 162, 163, 165.
— siamensis, 144.

—— suchus, 140, 147, 149.
tenuirostris, 133.

—— trigonatus, 153, 167.
—— trigonops, 140, 143.
vulgaris, 128 note, 134, 140, 144, 147, 149,
150.

acutus, 127 note, 128 note,138, 147, 150, 151.

Crocodilus yakare, 162, 163.
Crocodilus de St. Domingue, 150.
verd de Sénégal, 147.
Crown-Pigeon, 70.
Cryptobranchus, 517.

Cybium maculatum, 388.
Cyclodont Lizards, 506, 507.
Cyclodus, 507.

Cygnus bewickii, 123.

falconeri, 120, 227.
musicus? 120, 123.

olor, 119.

Cynogale velox, Du Chaillu, 2, 3.
Cynosuchus, 163.

Cyprinide, 395.
Cyprinodontide, 394.
Dactylethra, 517.

Dajaus elongatus, 384, 444.
monticola, 384.
nasutus, 444.

Delphinapterus, 115.

molagan, 24.

Delphinide, 98, 113, 114, 115.

, Tibs of, 354.

Delphinine, 115.

Delphinoidea, 111, 115.

Delphinoid Cetacea, vertebral formule of, 348, 349.

| Delphinus, 44, 94, 114, 115.

——,, trapezoid of, 362.

amazonicus, 87.

attenuatus, 19.

compressus, 19.

cuviert, 17, 25.

delphis, vertebral formula of, 349.
euphrosyne, 24,

Sluviatilis, 87 note.

forsteri, 23, 24.

frontatus, 19.

—— fusiformis, 20-23.

gadamu, 17-24,

geoffrensis, 88.

— geoffroyi, 88.

heavisidii, vertebral formula of, 349.
lentiginosus, 20-22.

longirostris, 23, 24,

—— maculiventer, 21.

Seep

| —— malayanus, 19.
—— pomeegra, 23, 24.

INDEX OF

Delphinus sinensis, vertebral formula of, 349.

superciliosus, vertebral formula of, 349.
—— tursio, atlas of, 330.
, pisiform bone of, 360.

, trapezoid of, 362.

, vertebral formula of, 349.
Diapterus dovii, 383, 448.

Dicholophus, sclerotal ring of, 510.

Didine bird of the Island of Bourbon, 373.
Didunculus, 67, 68, 72, 73, 75, 76, 77.
Didus ineptus, 49-82, 373.

, axis of, 56.

——, brain of, 69:

, cervical vertebre of, 54, 73.
, clavicle of, 63.

, concluding remarks on, 80.

, coracoid of, 62, 74.

, cranial cavity of, 71, 76.

, diploé of, 69.

——. __, discovery of remains of, 51-53.
, dorsal vertebra of, 53; 72.
— , femur of, 64, 78.

—_—_ ——, fibula of, 65.

, frontals of, 67.

, humerus of, 63, 78.

ee linmyof.o0:

—__— ——,, ischium of, 60.

—— , leg-bones of, 64-66.

ae mastoid ols) Ge

___ ___,, maxillaries and intermaxillaries of, 68,
71.

____ ___,, metatarsus of, 65.
, nasals of, 68.

, occipital of, 66.

, origin of, 70, 80.
——,, palatines of, 69.

, pelvis of, 57, 75.

, probable habits of, 70.
——, pubis of, 60.

, radius of, 63.

——, ribs of, 56, 72, 73.

, sacral vertebrae of, 58.

——, scapula of, 62.

, skull of, 66-72.

——,, sphenoids of, 66, 67.
——, sternum of, 60-62, 73.
, temporal fosse of, 67.

-) JalMeie ese

—__— —_, comparison of the skeleton of, 72-79.

SPECIES.

Didus ineptus, tibia of, 65.

——, tympanic bones of, 69.
—— ——,, ulna of, 63, 78.

, vertebre of, 53-56, 72, 73.
, wing-bones of, 63, 64,
Dinematichthys marginatus, 392.
Dinornis, 67-70, 77, 79.

maximus, femur of, 498.

, metatarsus of, 499.

, tibia of, 499.

—— robustus, integuments and tendons of the foot

of, 495.
Diodon sex-maculatus, 396.
© Dodeersen,” 373.
Dodo, 49-82, 373.
——,, axis of the, 56.
——,, brain of the, 69.
——,, cervical vertebre of the, 54, 73.
——, clavicle of the, 63.
——, comparison of the skeleton of the, 72-79.
——, concluding remarks on, 80.
—, coracoid of the, 62, 74.
——, cranial cavity of the, 71, 76.
——, diploé of the, 69.
——,, discovery of remains of the, 51-53.
——., dorsal vertebre of the, 53, 72.
——., femur of the, 64, 78.
——, fibula of the, 65.
——, frontals of the, 67.
——., humerus of the, 63, 78.
——,, ilium of the, 59.
——, ischium of the, 60.

| ——, leg-bones of the, 64-66.
| ——, mastoid of the, 67.
——, maxillaries and premaxillaries of the, 68,

(fle
——, metatarsus of the, 65.
——,, nasals of the, 68.
——, occipital of the, 66.
——, origin of the, 70, 80.
——,, palatines of the, 69.
——, pelvis of the, 57, 75.
, probable habits of the, 70.
——, pubis of the, 60.
, radius of the, 63.
——., ribs of the, 56, 72, 73.
, sacral vertebra of the, 58.
, scapula of the, 62.

528 INDEX OF SPECIES.

Dodo, skull of the, 66-72.

——., sphenoids of the, 66-77.

, sternum of the, 60-62, 73.
, temporal fosss of the, 67.
——,, tibia of the, 65.

——, tympanic bone of the, 69.
——, ulna of the, 63, 78.

, vertebre of the, 53-56, 72, 73.
——., wing-bones of the, 63, 64.
Dolphin, Black, 23.

, Freckled, 20.

——, “ Gadamu,” 17.

——, Gangetic, 92.

——, Pomeegra, 23.

——, Spindle-shaped, 22.

, Spot-bellied, 21.
Dolphins, 111.

Dormitator microphthalmus, 383.
Dromaius, 70, 74, 76.

Dugong, 97.

Echeneis naucrates, 388.

—— remora, 388.
Elasmobranchii, 396.

Eleotris dormitatrix, 389, 404.
longiceps, 389, 405, 440.
—— maculata, 383, 389, 440.
—— picta, 389, 405, 441.
—— pictus, 384.

seminuda, 389, 441.
somnolenta, 389.
Elephant, 69.

——, African, exoccipital bone of, 234.

——,, extinct species of, dentition of, 283-302.

. , from Malta, 227.

Elephas, sp.? 231.

, dentition of, 283-302.

——, milk-incisors of, 284.

——, milk-molars of, 286-291.

, permanent incisors of, 285.

——, true molars of, 291-302.

——,, young bones of, from Malta, 271-283.
africanus, astragalus of, 269, 306.
—— ——, atlas of, 306.

—— ——, femur of, 248-250.

, humerus of, 256, 257.
—— ——, mandible of, 237.

—— ——, milk-molars of, 301.

, obturator foramen of, 243.

Elephas africanus, osteology of uterine foetus of,

274-277.

——, scapula of, 244,

, seventh cervical vertebra of, 239, 240.
, ulna of, 246, 247, 262.
antiquus, 230.

——, milk-molars of, 298.

, molars of, 299,

falconeri, 230, 251.

, astragalus of, 268, 306.

, atlas of, 251, 306.

——, bones of the fore foot of, 263.
, bones of the hind foot of, 268.
——,, femur of, 264,

——., humerus of, 255.

——, metatarsal, fourth, of, 271.

, pelvis of, 263.

, Tibs of, 253.

, scapula of, 254.

——, ulna of, 260-263.

-——, vertebre of, 253.

, young bones of, 271.

indicus, 233.

, astragalus of, 270, 306.

~, atlas of, 306.

, femur of, 249, 250,

——, humerus of, 245, 256, 257.
——, mandible of, 237.

——, milk-molars of, 298, 301.

, obturator foramen of, 243.
——,, ribs of, 253 note.

, scapula of, 244,

——, seventh cervical yertebra of, 239, 240.
——, ulna of, 247, 262,

insignis, milk-incisors of, 284.
melitensis, 230, 235.

, astragalus of, 270, 306.
——., atlas of, 238, 306.

-——, cervical vertebrx of, 238, 240.
, cranium and face of, 236.

, dorsal vertebrae of, 240.

, femur of, 247, 265, 267.
——, humerus of, 244,

——, lumbar vertebre of, 241.
——,, pelvis of, 242.

——, rib of, 241.

, scapula of, 244.

——,, tusk of, 285.

INDEX OF SPECIES.

Elephas melitensis, ulna of, 245, 262.
—— primigenius, 232, 233.

—— ——,, humerus of, 245.

—— —— , milk-molars of, 298.
—— ——,, molars of, 299.

, scapula of, 244,

—— sumatrensis, lumbar vertebre of, 241.

, Tib of, 242.

, scapula of, 244.

, Seventh cervical vertebra of, 239,

240.

Emu, 517.

Emydosauri, 129.

Emydosaurians, geographical distribution of, 130,
131.

——, synopsis of, 125-168.

Engraulis brownii, 395.

macrolepidota, 395, 488.

macrolepidotus, 384.

—— mysticetus, 489.

poeyi, 384, 395, 488.

Ephippus faber, 386.

Epinephelus afer, 411.

analoqus, 383, 410.

sellicauda, 409.

Erinaceus, 505.

Eubalena, 115.

Euctenogobius sagittula, 389.

Eudyptes, 76.

Euelephas, milk-incisors of, 284.

, molars of, 292, 298.

Euphysetes, 30, 44.

breviceps, 41, 42.

grayi, McLeay, 34, 42.

Eurypyga, osteology of, compared in various points
with that of the Kagu, 503, 504, 507-516, 518-

520.

helias, number of vertebre in, 511.

, proportions of leg-bones in, 518.

Exocetus albidactylus, 383, 394.

bahiensis, 383, 394.

callopterus, 394, 479.

dovii, 394.

— dowii, 383.

Falcons, os pubis in some, 518.

Fishes of the Atlantic and Pacific Coasts of Central
America, comparison of the, 397-403.

of Central America, 377-491.

529

Fishes, scapular region of, 517.
Fistularia tabacaria, 390.
Fistulariide, 390.

Freckled Dolphin, 20.

Fulica atra, 520.

Fulix fuligula, 375.

Fundulus guatemalensis, 394, 403, 482.
labialis, 394, 404, 481.
pachycephalus, 394, 403, 483.
—— punctatus, 394, 402, 482.
“‘Gadamu” Dolphin, 17.

Galline, 516.

Gambusia nicaraquensis, 394, 405, 483.
Gangetic Dolphin, 92.

Ganoidei, 396.

“ Ganumu,” 25 note.

Gar-fowl, 76.

Garial, African False, 157.

Garials, 130.

Gayial, 132, 133.

—, Bornean, 134, 135.

of Senegal, 157.

Gavialia, 132.

Gavialide, 129, 132.

Gavialis, 132, 133.

gangeticus, 127 note, 132, 133.
longirostris, 133.

tenuirostris, 133.

Genytremus interruptus, 414.

Gerres aprion, 391.

axillaris, 391, 448.
brevimanus, 391, 448.

dovit, 383, 391, 448.
plumieri, 391.

rhombeus, 381.

squamipinnis, 391.

Gerride, 391.

Girardinus pleurospilus, 395, 403, 486.
Globiocephalus, 25 note, 98, 114, 115.
, atlas of, 330.

, trapezoid of, 362.

melas, vertebral formula of, 349.
Glyphidodon concolor, 390.
declinifrons, 390.

saxatilis, 390.

Gobiesocide, 390.

Gobiesox nigripinnis, 390.

nudus, 390.

530

Gobiesox rhodospilus, 390, 445.

Gobiide, 388.

Gobius, 517.

—— mewicanus, 389.

paradoxus, 388.

seminudus, 389.

soporator, 388.

Gorilla, limbs of, compared with those of the Orang,
175-218.

Goura, 67, 70, 72, 73, 75-79.

Grampus, 115.

Grebes, 520.

Gruine, 503.

Grus, 79, 508, 515, 516.

antigone, number of vertebree in, 511.

Gymnotide, 395.

Hemulon brevirostrum, 386, 418.

—— canna, 386.

chromis, 386.

—— margaritiferum, 386, 419.

—— melanopterum, 414.

wanthopterum, 386.
Halerosia, 135-137, 152.
frontata, 153.

—— nigra, 153.

palpebrosa, 166.

Halicore, 97.

Haplochilus dovii, 394, 406, 481.
Hare, 505, 507.

Hawks, pubis in some, 518.
Hedgehog, 505.

Heliastes insolatus, 446.
marginatus, 390, 446.
Hemirhamphus unifasciatus, 394.
Hemirhombus ovalis, 392, 472.
Herons, 503, 504, 508, 509.
Heros affinis, 391, 404, 455.
altifrons, 384, 391, 405, 459.
angulifer, 392, 404, 469.
aureus, 391, 404, 455.
citrinellus, 391, 405, 458.
dovit, 391, 405, 461.
erythreeus, 391, 405, 457.
—— friedrichsthalii, 382, 391, 404, 459.
godmanni, 392, 404, 466.
guttulatus, 392, 403, 466.
intermedius, 392, 404, 468.
irregularis, 392, 404, 467.

INDEX OF SPECIES.

Heros labiatus, 391, 405, 456.

lobochilus, 391, 405, 457.

longimanus, 391, 405, 453.

macracanthus, 391, 403, 451.

managuensis, 392, 405, 463.

margaritifer, 391, 404, 450.

melanopogon, 382, 391, 450.

melanurus, 391, 404, 450.

microphthalmus, 392, 404, 464.

motaguensis, 391, 404, 462.

multispinosus, 391, 405, 453.

nicaraguensis, 392, 405, 465.

nigrofasciatus, 391, 403, 452.

oblongus, 392, 404, 464.

parma, 391, 404, 405, 449.

salvini, 382, 391, 404, 460.

sieboldii, 384, 392, 466.

spilurus, 391, 404, 451.

—— triagramma, 382, 460.

—— trimaculatus, 391, 403, 461.

urophthalmus, 382, 391, 404, 454.

Hexanematichthys hymenorrhinos, 475.

Himantopus, 503, 504.

Hippopotamus, bones of, from Malta, 228.

Hylobates, 213.

Hyperoodon, 44, 98, 111, 112.

, atlas of, 330, 331.

, axis of, 333.

——.,, carpus of, 361.

, cervical vertebre of, 328, 330, 332, 333.

, comparison of skull of, with that of Physeter
macrocephalus, 314-323.

——., dorsal vertebre of, 340.

| ——, hyoid bones of, 326.

, vertebrae of, 327.
, scapula of, 357.
latifrons, pelvic bones of, 365, 366.
rostratum, pelvic bones of, 365, 366.
, vertebral formula of, 348.
Hypostomus plecostomus, 384.
Indris, 214.
Inia, cervical vertebre of, 329.
——, pisiform bone of, 360.
—— boliviensis, 87.
geoffrensis, 87-106.
, atlas of, 96.

—— ——, axis of, 96.

——-— ——, carpus of, 105.

Inia geoffrensis, caudal vertebre of, 101, 102.

, cervical vertebre of, 96, 97.

, chevron bones of, 102.

, costal cartilages of, 103.
——,, digits of, 105.

——, history of, 87, 88.
——, humerus of, 104.

—, hyoid apparatus of, 95.
——, lumbar vertebre of, 101.
——, malar of, 91.

——, mandible of, 94.

——, maxillaries of, 91.
——,, nasals of, 91.

—— —, occipital of, 90.

——, orbits of, 91.

——,, palate of, 92, 93.

——,, parietal of, 90.

——,, premaxillaries of, 91, 92.
, pterygoids of, 93.

— , radius of, 105.

——, ribs of, 98, 99, 102, 103.
, rostrum of, 92.

, scapula of, 104.

-—, skull of, 89-94.

, sphenoids of, 93.

, spinal column of, 95-99.

, squamosal of, 90.

, sternum of, 103.

, teeth of, 94, 95.

, temporal fossa of, 90.

——, thoracic vertebre of, 97, 98.

——, ulna of, 105.

, vertebre of, 95-99.
, vertebral formula of, 349.
, vomer of, 92, 93.
Iniine, 114, 115.

Insectivora, 1.

Tsogomphodon maculipimnis, 490.
Jacaré, Black, 162.

— , Brazilian, 164.

——, Dog-headed, 163.

——, Dotted-jawed, 165.

— _, Eyed, 164.

——, Long-shielded, 164.

, Rough-necked, 165.
Jacare, 160, 161, 162.

JSissipes, 163.

VOL. VI.—PART VIII.

‘elalae (cle SS Gia e s8 aie

——,, tympanic bulle and fosse of, 94.

SPECIES.

Jacare hirticollis, 165.

—— latirostris, 163.

—— longiscutata, 164.

—— multiscutata, 164.

—— nigra, 128 note, 162, 163,
ocellata, 164.

—— punctulata, 165.
sclerops, 165.

vallifrons, 165.
Jacaretinga, 160, 162.

Julis lucasana, 391.

Kagu, 501.

——,, atlas of the, 512.

, axis of the, 512.

, cervical ribs of the, 511.
—-—, coracoids of the, 514.
——,, dorsal ribs of the, 511.
——, femur of the, 518.
——, fureula of the, 514.
——, hyoid bones of the, 510.
——,, interorbital septum of the, 504.

, lacrymal of the, 508.

——, lumbo-sacral arch of the, 511.
——, mandible of the, 510.

——, maxillaries of the, 504, 506, 507.
, nasals of the, 504, 508.

, orbits of the, 503.

, 08 quadratum of the, 509.

——,, palatines of the, 509.

, pelvis of the, 515-518.

——,, pterygoids of the, 509.

, scapula of the, 513.

, sclerotal ring of the, 510.

, shoulder-girdle of the, 513-515.
, skull of the, 503-510.

, sternum of the, 514, 515.

, systematic position of the, 502.
——, tarso-metatarsus of the, 519, 520.
——,, temporal fosse of the, 503.
, toes of the, 520.

— , tibia of the, 519.

, vertebre of the, 510-513.
, vomer of the, 508.

, wing-bones of the, 515.
Kogia, 37 note, 111, 112.

, cervical vertebrae of, 328.
——,, hyoid bones of, 326.

, scapula of, 357.

531

532 INDEX OF SPECIES.

Kogia grayi, vertebral formula of, 348.

Krokodile noir du Niger, 153-155.

Labride, 390.

Lacerta gangetica, 132.

Lacertilia, 507, 517.

Lachnolemus faleatus, 390.

Lagenorhynchus, 22-24, 115.

—— albirostris, vertebral formula of, 349.

electra, 23.

—— fusiformis, 23.

leucopleurus, vertebral formula of, 349.

Larimus breviceps, 387, 425.

Lepidosteus, 506.

—— tropicus, 383, 396, 403, 490.

Leporide, 506.

Leptarius dowii, 383.

Lepus timidus, 505.

Lizards, Cyclodont, 506.

Lobotes auctorum, 386.

Loricaria lima, 384, 393, 405.

uracantha, 384, 393, 406, 478.

Loxodon, 299.

Lycodide, 392.

Macherhamphus aleinus, 118.

Macrodon brasiliensis, 384.

microlepis, 384, 393, 406, 478.

tareira, 478.

Man, limbs of, compared with those of the Orang,
175-218.

Mareca penelope, 375.

Mecistops, 134-137, 156.

bathyrhynchus, 151.

burnettiz, 157, 159. ‘

— cataphractus, 127 note, 133, 134, 150, 157.

journei, 134, 151.

Megalops thrissoides, 395.

Megaptera, 115.
Megapterine, 115.
Melanosuchus, 162.

Meletta libertatis, 487.
petenensis, 488.
Mergus castor, 375.
Mesoprion aratus, 385, 413.
chrysurus, 385.

griseus, 385.

—— uninotatus, 385.
vivanus, 385.
Microdesmus, 470.

Microdesmus dipus, 392, 471.
Microglossum aterrimum, 69.
Micropogon altipinnis, 387, 425.
undulatus, 387.

Micropteron, 111.

, cervical vertebrae of, 328.
sowerbyense, carpus of, 361.
—— ——,, dorsal vertebre of, 340.
—-—, vertebral formula of, 348.
“ Molagan”’ (Dolphin), 24.

Molinia, 135-137, 150.

americana, 127 note, 150, 151.
intermedia, 151.

Mollienesia petenensis, 395, 404, 485.
Monodon, 44, 96, 98, 115.

, atlas of, 330.

——, carpus of, 361.

, cervical vertebree of, 329.
——,, trapezoid of, 362.

—— monoceros, vertebral formula of, 349.
“ Muggar,” 140.

««___” Siamese, 144.

Mugil brasiliensis, 383, 389, 443.
guentheri, 383.

—— incilis, 389, 443.

—— proboscideus, 389.
Mugilide, 389.

Mullide, 386.

Murena lineopinnis, 396.
Murenide, 396.

Mustelus dorsalis, 396, 490.
Mycteria, 79.

Myoaxus cartei, 307.

melitensis, 228, 307.
Mystacoceti, 110, 115.
Mythomys velow, Gray, 2, 3.
Myaus harengus, 390.

Nakoo, 132.

Narwhal, 114.

Nasua fusca, 505.

Neomeris, 114, 115.
phocenoides, vertebral formula of, 349.
Neetroplus, 469.

—— nematopus, 392, 405, 470.
Night-Heron, 502.

Norfolk Plover, 503.

Notornis, 70.

Nycticebine, 214.

INDEX OF SPECIES.

Nyeticorasx, osteology of, compared in various points
with that of the Kagu, 503, 509-511.

Ocydromus, 503, 513-516, 518, 519.

—- australis, proportions of leg-bones in, 518.

Odontoceti, 111, 115.

, homologies of the carpal bones of the, 360.

Gidicnemus, 74, 76-78.

, osteology of, compared in yarious points with

that of the Kagu, 503.

—— crepitans, 79.

Oligoplites inornatus, 383, 433.

Oopholis, 135-137.

pondicherianus, 139.

porosus, 128 note, 138.
Ophidia, 507.

Ophidiide, 392.

Ophidium brevibarbe, 392.
Ophioscion typicus, 383, 428.
Ophiurus breviceps, 396.
boro, 396.

triserialis, 396,
Orang-outang, 175.

Orca, 24, 98, 114, 115.

, atlas of, 330.

——, carpus of, 361.

——, pelvic bones of, 365 note.

, trapezoid of, 362.

gladiator, vertebral formula of, 349.
Osteolemus, 152.

tetraspes, 153.

Ostracion bicaudalis, 396.

cornutus, 396.
Ostrich, African, 517.
Otis, 74, 76-79.

Otolithus albus, 387, 429.
— reticulatus, 387, 430.
—— squamipinnis, 387, 429.
Owls, 69.

Palamedea, 72, 509.
Palapteryx, 68,70.
Paleosuchus, 167.

Palinia, Cuban, 145.

, Yucatan, 146.
Palinia, 135-137, 145.

2 moreletiz, 146.
rhombifera, 145.
Pangolin, 505, 506.
Pediculati, 388.

Pelamys sarda, 388, 435.

Penguins, 72, 79.

Percida, 385.

Petenia, 469.

splendida, 382, 392, 404, 469,

Pezophaps, 70, 79, .

solitarius, 373.

Phalacrocorax, 504.

Pharyngognathi acanthopterygii, 390.

Phocena, 24, 25, 44, 92, 98, 114, 115.

, atlas of, 330.

, trapezoid of, 362.

— brevirostris, foramina in the skull of, 29,
30.

, frontals of, 27.

—, malar bone of, 28, 39.
——, mandible of, 29.

——, maxillaries of, 27,

——,, occipital bone of, 26.

, palatine bones of, 27.
, premaxillaries of, 28.
——,, pterygoid of, 28.
——, skull of, 24-30.
——, sphenoid bones of, 26.

——, squamosal of, 29.

Ce Saai se aegis

, vomer of, 27.

—— communis, vertebral formula of, 349.

globiceps, 25, 28.

Pheenicopterus, 72.

Physalus, 115.

Physeter, 44, 45, 98, 111, 112, 114, 119.

australis, characters of, 367.

breviceps, 33, 41, 42.

catodon, 369.

— macrocephalus, 32, 33, 35-41 notes, 44,
iba,

, atlas of, 328.

, axis of, 332.

—— ——, base of skull of, 319.

, carpus of, 359-364,

, caudal vertebre of, 342-347.
, cervical vertebrae of, 328-335.
—_— ——,, characters of, 367-369.

, chevron bones of, 350.

, comparison of yertebre in known spe-
cimens of, 347, 348.

, cranium of, 314-323.

—— — , digits of, 362-364.

442

554

Physeter macrocephalus,historyof the knowledge of,309.

, humerus of, 357.

—— ——, hyoid bones of, 325.

—— ——, known specimens of, 311-314.

—— —, measurements of skulls of, 321-323.

—— ——, metacarpals of, 362-364.

— — , nares of, 317.

, nasal bones of, 318.

—— — , orbits of, 319.

, osteology of the, 309-369.

, pectoral limb of, 356-364.

, pelvis of, 364-366.

——, petrotympanie of, 321.

——,, pisiform bone of, 360.

, radius of, 358.

——,, ribs of, 351-354.

, Scapula of, 356.

, Sternum of, 354-356.

——, teeth of, 320, 324,

——,, ulna of, 358.

, vertebral column of, 326-350.

——, vertebral formula of, 348.

microps, 369.

(Euphysetes) siaus, 30-43.

, comparison of skull of, 41, 42.

—— —— ——,, description and measurements of,
172-174.

, external characters of, 30-32.
, fins of, 43.

—— ——.,, frontals of, 37.

, interior of cranium of, 40.

, lower jaw of, 40.

—— ——, malars of, 39.

—— ——, maxillaries of, 38.
—— ——, nasals of, 37.
—— ——,, occipital of, 34,
—— ——,, palatines of, 37.

, pelvic bones of, 43.
——, pterygoids of, 38.
——, ribs of, 43.

, Seapula of, 43.

, Skull of, 34.

——,, sphenoids of, 35, 36.
, squamosals of, 39.
—— ——, ulna of, 43.

—— ——,, vertebre of, 42, 43,
, vomer of, 37.
tursio, 369.

Berea

INDEX OF SPECIES.

Physeteride, 30, 32, 33, 93, 103, 113, 114, 116,
366, 368.

Physeterine, 114, 115, 368.

Pimelepterus boscii, 386.

Pimelodus cinerascens, 384, 474.

godmanni, 393, 404.

—— guatemalensis, 393, 403.

managuensis, 393, 474.

—— micropterus, 393.
modestus, 384, 393, 405.
motaguensis, 393, 404.

—— nicaraquensis, 393, 405.

petenensis, 393, 404,

polycaulus, 393, 404.

salvini, 393, 404.

—— wagneri, 384, 393, 405, 474.
Platalea, 72,76.

Platanista, 44, 87, 89, 92, 94, 96, 98, 100, 102-104,
112, 115.

, atlas of, 330,

——-., cervical vertebre of, 328, 330, 333.
gangetica, vertebral formula of, 349,
Platanistide, 113, 115.

Platanistine, 114, 115.

Platyglossus bivittatus, 390.

dispilus, 390, 447.

Plecostomus, 393, 405.

bicirrhosus, 477.

Plectognathi, 396.

Plectropoma afrum, 385, 411.

— chloropterum, 411.

-— monacanthus, 411.

multiguttatum, 411,
Pleuronectide, 392.

Plotus, 512.

Plovers, 72, 75, 79.
Podargus humeralis, 74.
Podiceps, 76, 520.

Peecilia chisoyensis, 395, 404.
dovit, 395, 403, 405.
elongata, 395, 406, 484.
— gilli, 395, 406, 485.
mexicana, 395, 403.
petenensis, 395, 404, 484,
spilurus, 395.
thermalis, 395, 403.
Polynemide, 387.

| Polynemus approaimans, 383, 387, 423.

INDEX OF SPECIES.

Polynemus melanopoma, 387, 421.
opercularis, 251, 383, 387, 424, 429.
Pomacanthodes zonipectus, 383.
Pomacanthus paru, 386.
zonipectus, 386, 419,
Pomacentride, 390.

Pomacentrus analigutta, 446.
—— bairdii, 445, 446.
flavilatus, 446.

—— lewcostictus, 390.
rectifranum, 390, 445, 446.
Pomatoprion bairdii, 446,
Pomeegra Dolphin, 23.

Pontoporia, cervical vertebre of, 329.
, systematic position of, 109-115,
—— blainvillii, 106-109.

— — , history of, 106.

, mandible of, 109.

— — , skull of, 107.

, teeth of, 109.

, vertebral formula of, 349.
Porichthys porosissimus, 388.
Porphyrio, 79, 515, 519, 520.

Porpoise, 111.

, short-snouted, 24.

Potamogale, 1; characters of the genus, 15.
velow, anal glands of, 13.

, brain of, 14.

, caleaneum of, 13,

, carpus of, 12.
, cloacal character of, 13.

, ears of, 5.
—— — , eyes of, 5.

, femur of, 13.
— — , fibula of, 13.

, fore limbs of, 5.
— — , hairs of, 3.

, head of, 4.

—_— ——,, hind limbs of, 5.
—— — , humerus of, 12.

, lium of, 12.

, ischium of, 12.

, mandible of, 11.

, metatarsals of, 13.
, ossa innominata of, 9, 12.

, ovaries of, 14.
——, pelvis of, 12.

poliocephalus, proportions of leg-bones in, 518.

§35

Potamogale velox, pubic bones of, 12.
, radius of, 12.

, rectum of, 13.

, scapula of, 12.

, Skeleton of, 8.

, Skull of, 10.

, Soft parts of, 13.
, tail of, 6.

, tarsus of, 13.

, teeth of, 6.

—— —, tibia of, 13.

, toes of, 5.

— — , wna of, 12.

aaeaaals

—— ——, uterus of, 13.

, vagina and vulva of, 14.
, vertebrae of, 8, 9.
Potamogalide, 15.

Primates, 175.

Prionodon, 490.

Pristigaster argenteus, 487.

— dovii, 395, 487.

macrops, 395, 487.
Pristipoma bilineatum, 414.
chalceum, 386, 415.

crocro, 386.

dovit, 386, 414.

— humile, 382, 384, 386, 415.
— leuciscus, 386, 416.

—— macracanthum, 386, 416.

—— melanopterum, 386, 414.
virginicum, 386, 414.
Pristipomatide, 386.

Pristis antiquorum, 397.
Promicropterus decoratus, 383, 412.
Proteus, 517.

Pseudochalceus lineatus, 384.
Pseudojulis notospilus, 390, 447.
Pseudorca, 115.

, trapezoid of, 362.

crassidens, vertebral formula of, 349.
Pseudorhombus brasiliensis, 392, 473.
vorax, 473.

Pseudoscarus guacamaia, 391.

sancte crucis, 391.

Psophia, 72.

, osteology of, compared in various points with

that of the Kagu, 503, 504, 507-516,518—520.
crepitans, number of vertebre in, 511.

536 INDEX OF

Psophia crepitans, proportions of leg-bones in, 518.
Psophiine, 502.

Rails, 504, 518.

Rallide, 508.

Ramphostoma, 132.

tenuirostre, 133.

Raptores, pubis in some, 518.
Rhamphognathus, 132.

Rhea, 70, 74, 517.

Rhinobatus leucorhynchus, 396, 490.
Rhinochetus jubatus, atlas of, 512.

, axis of, 512.

, cervical ribs of, 511.

, coracoids of, 514.

—— ——,, dorsal ribs of, 511.

—— ——,, femur of, 518.

—— —— , furcula of, 514.

— —, hyoid of, 510.

—— ——,, interorbital septum of, 504.
—— ——,, lacrymal of, 508.

—— ——,, lumbo-sacral arch of, 511.
—— ——, mandible of, 510.

—— ——, maxillaries of, 504, 506, 507.
—— ——,, nasals of, 504, 508.

—— —-—,, orbits of, 503.

508 quadratum of, 509.

, palatines of, 509.

—— ——,, pelvis of, 515-518.

——. ——.,, pterygoids of, 509.

, scapula of, 513,

, Sclerotal ring of, 510.

—— ——, shoulder-girdle of, 513-515.
—— ——, skull of, 503-510.

—— ——, sternum of, 514, 515.

—— ——, systematic position of, 502.
, tarso-metatarsus of, 519, 520.
— ——,, temporal fosse of, 503.
—— ——,, tibia of, 519.

, toes of, 520.

, vertebre of, 510-513.
—— ——, vomer of, 508.

, wing-bones of, 515.
Rhombus aramaca, 473.
Rhynchosuchus, 133.

schlegelit, 134.

Rhypticus decoratus, 383, 385, 412.
—— nigripinnis, 412.

Saccodon wagneri, 384.

pz

SPECIES.

Salarias atlanticus, 389.
Salmo fario, 44.
Sargus aries, 386.

unimaculatus, 386.

Saurus fetens, 394.

myops, 394.

Scarus abildgaardii, 391.

Sciades troschelii, 383.

Scienide, 387.

Sclerognathus meridionalis, 395, 404, 486.
Scombresocide, 394.

Scombride, 388.

Scopelide, 394.

Scopus, 515, 516.

Scorpana plumieri, 387.
Scorpenide, 387.

Serpents, “ turbinal” bone of, 502.
Serranus analogus, 383, 385, 410.
coronatus, 385.

creolus, 385, 409.

—— sellicauda, 385, 409.
striatus, 385.

undulosus, 385.
Short-snouted Porpoise, 24.
Sibbaldius, 115.

Sicyases fasciatus, 390.

Sticydium plumieri, 389.
Siluride, 393.

Simia, appendicular skeleton of, 175-218.
——, carpus of, 187.

——, clavicle of, 178, 215.
——,, digits of, 198, 212.

, femur of, 198, 217.
——,, fibula of, 203.

, humerus of, 180, 215.
——, manus of, 187, 218.
——, metacarpus of, 192.
—— , metatarsus of, 209.

, os innominatum of, 194, 216,217.
——,, patella of, 201.

, pes of, 204, 218.

, radius of, 183, 216.

, scapula of, 175, 214, 215.
, tarsus of, 204,

— , tibia of, 201, 217.

, ulna of, 184, 216.
Sirenia, 87, 110.

Skate, 516, 517.

INDEX OF SPECIES. 537

Snub-nosed Cachalot, 30.
Solea scutum, 392.

Solitaire, 373.

Sparide, 386.

Sperm- Whale, osteology of the, 309-369.
Sperm- Whales, 30.
Sphyrena forstert, 389.
picuda, 389.
Sphyrenide, 389.
Spindle-shaped Dolphin, 22.
Spot-bellied Dolphin, 21.
Squamipinnes, 386.

Stanley Crane, 504, 509, 510.
Stegodon, 284.

Steno, 17-21, 23, 24.
Stilt-Plover, 503, 512 note.
Struthio, 70, 74, 76.
Sun-Bittern, 502, 512.

Sus scrofa, 506.
Symbranchide, 396.
Symbranchus immaculatus, 396.

marmoratus, 396.
Talegalla, 508, 515, 516.
Tantalus, 79.

Tarsius, 213, 214.

Temsacus, 151.

Tetragonopterus eneus, 384, 394, 399, 406, 478.
brevimanus, 394, 404.
fasciatus, 393.

gronovit, 384.

humilis, 394, 403.
microphthalmus, 394, 403.
panamensis, 394, 404, 406.
petenensis, 394, 404.
Tetrodon geometricus, 396, 489.
politus, 396, 489.
Thalassophryne, 436.

maculosa, 388, 436.
reticulata, 388, 437.

Theraps irregularis, 467.
Tigrisoma leucolophum, 509.
Tinamous, 508, 517.

Toad, 517.

Tomistoma, 132, 133.

schlegelit, 134, 135.
Toothed Whales, 110-112.
Trachynotus fasciatus, 388, 434.
—— glaucoides, 434.

—— ovatus, 388.

Trachurus boops, 431.

Trichidion approximans, 383, 423.

opercularis, 383, 424.

Trigla, 517.

Troglodytes, limbs of, compared with those of Simia,
175-218.

Tyntlastes, 441.

Umber, 515.

Umbrina analis, 387, 426.

elongata, 387, 425.

nasus, 387, 426.

undulata, 427.

Upeneus grandisquamis, 383, 386, 420.

tetraspilus, 386, 420.

Uria, 76.

Urolophus mundus, 397,491.

Urotrygon mundus, 383, 491.

Vultur, 76.

Vultures, 72, 74,76, 79.

Whalebone- Whales, 110.

, accessory pelvic bones or cartilages of 364.

« Wonga,” 30.

Xiphophorus gillii, 384, 485.

hellerii, 395, 404, 485.

Ziphiine, 114, 115.

Ziphioids, 91, 92, 111, 112.

Ziphius, 44.

ceryptodon, vertebral formula of, 348.

Zygena tiburo, 396.

END OF VOLUME VI.

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