Se

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Hrbrary of the Museum

OF

COMPARATIVE ZOOLOGY,

AT HARVARD COLLEGE, CAMBRIDGE, MASS,

Founded by private subscription, in 1861.

ANATOMY OF THE INDIAN ELEPHANT

STUDIES IN COMPARATIVE ANATOMY,.—No. IL.

ANATOMY OF THE INDIAN ELEPHANT

BY

L. C. MIALL

PROFESSOR OF BIOLOGY IN THE YORKSHIRE COLLEGE AND
CURATOR OF THE LEEDS MUSEUM

AND

F. GREENWOOD

CURATOR TO THE LEEDS SCHOOL OF MEDICINE

H ond ow :
MACMILLAN AND CO.
“™1878

[The Right of Translation and Reproduction is Reserved. |

STUDIES IN COMPARATIVE ANATOMY.—No. IL

ANATOMY OF THE INDIAN ELEPHANT,

INTRODUCTION.

WHENEVER the comparative anatomy of mammalia shall be
exhaustively treated, the structure of the existing species of
elephants will claim a principal share of attention. They are so
distinctly separated from other quadrupeds that an order is
required for their sole reception. Yet, though isolated from all
other existing species, they bear marks of affinity with more than
one group, particularly with Ungulata, Sirenia, and Rodentia.
This fact of recent zoology suggests a view which the history of
extinct mammalia seems to justify, viz. that the elephants
constitute a comparatively primitive type, representing, though
perhaps not directly, a now extinct central group, from which the
orders above mentioned, and possibly some others, may have been
derived. The African and Indian elephants are further remark-
able as the largest land quadrupeds now living; nor can the
paleontologist, reviewing the land quadrupeds of all past times,
so far as they are yet made known, find a trace of any larger
form which is not also an elephant. Hence arise some curious
physiological and mechanical questions to the student of elephant
anatomy. What are the special modifications implied by a
weight of perhaps three tons? How nearly do the existing
species approach the limits of size fixed by mechanical laws and
by the physical properties of animal tissues ?

It would be an interesting task to investigate such problems
as these, and to discuss the many points of anatomy, morphology,
physiology, and palzontology suggested by the examination of a
dead elephant. Our immediate purpose is less ambitious.
Leaving the greater enterprise to other times, and probably to
other hands, we here offer a condensed statement of facts re-
specting those parts of the structure of the elephant which are at

6 STUDIES IN COMPARATIVE ANATOMY.

present least perfectly known. When the time for comparison
and inference shall have come, such facts will be found of use.

For the convenience of future students, a list of the principal
memoirs on the anatomy of the elephant is subjoined.!_ Some of these
require special mention.

1Biarr, PaTRicK.—The Anatomy and Osteology of an Elephant, being an exact
Description of all the Bones of the Elephant which died near Dundee, April
the 27th, 1706, with their several dimensions. Phil. Trans. vol. xxvii.
p. 53, fig. (1710).

On the Organ of Hearing in the Elephant. Phil. Trans. xxx. p. 885,

fig. (1718).

Camper, Prerre.—Description Anatomique d’un Eléphant male. Publiée par son

fils, A. G. Camper. Paris, fol. fig. (1802).

Kort berigt van de ontleding eens jongen elephants. Vaterlandsche
letteroeffeningen (1774). Translated in Camper’s Kleinere Schriften, pt.
i. p. 51, and Giuvres, tom. i. and il. p. 152.

CorsE, JouN.—Observations on the Different Species of Asiatic Elephants and
their Modeof Dentition. Phil. Trans. vol. [xxxix. pp. 205-236, fig. (1799).

CuvirrR, GeorcEs.—Recherches sur les Ossemens Fossiles. Paris, 4th ed. tom. i.
pp. 468-582, fig. (1834).

CuviER ET LAURILLARD.—Recueil de Planches de Myologie, tom. iii.

DavBENTON.—In Buffon’s Histoire Naturelle, vol. xi. Paris, 4to (1764).

Goopsir, Joun.—Notes on the Myology of the Elephant. Appendix to
Anatomical Memoirs, vol. i. p. 446. Edinburgh, 8vo (1868).

Harrison, RoBERT.—On the Anatomy of the Elephant. Proc. Irish Acad. vol.

iii. pp. 385-398 (1847).

On the Larynx, Trachea, and Cisophagus of the Elephant. 0. vol. iv.

pp. 1382-135 (1850).

On the Anatomy of the Lachrymal Apparatus of the Elephant. 0. vol.
iv. pp. 158-165 (1850).

Homer, EVERARD.—Some Observations on the Structure of the Teeth of
Graminivorous Quadrupeds, particularly those of the Elephant and Sus
Athiopicus. Phil. Trans. vol. \xxxix. pp. 237-258, fig. (1799).

Houvet, J. P. L. L.—Histoire Naturelle des deux Hléphans, male et femelle, du
Muséum de Paris. Paris, 4to, fig. (1808).

[External parts only. These were the animals afterwards dissected by
Cuvier. ]

Hunter, JoHN.—Essays and Observations, edited by Prof. Owen, vol. ii. pp.
170-176, London, 8vo (1861).

LEURET ET GRATIOLET.—Anatomie comparée du systéme nerveux. Paris, fol.
fig. (1839-1857). [Brain. ]

Mayer, C.—Beitrige zur Anatomie des Elephanten und der iibrigen Pachy-
dermen.—Akad. Ces. Leop. vol. xxii. 4to, fig. (1847).

Movuins, ALLEN.—Anatomical Account of an Elephant, accidentally burnt in

Dublin, 4to, 72 pp. 2 pl. (1682).

OwEN, RicHArD.—Description of the Foetal Membranes and Placeuta of the
Elephant (Ziephas Indicus, Cuv.), with remarks on the value of placentary
characters in the classification of the mammalia. Phil, Trans, exlvii. pp.
347-353, pl. xvi. (1857).

THE ANATOMY OF THE INDIAN ELEPHANT. 7

Pierre Camper dissected an elephant in the winter of 1774. The
work lasted less than three weeks. Camper had subsequently several
opportunities of examining elephants of both sexes, and it was his
intention to prepare a full anatomical description. He died, however,
in 1789, leaving only the plates, an index to the figures, and the titles
of the chapters which were to have composed the text. The published
description, which is of some value, though by no means full, was
drawn up by his son, A. G. Camper. The plates are free and taste-
ful, without much detail ; they contain useful matter respecting the
chief viscera, but only the most meagre notes of myology.

Hunter’s notes are excellent, especially the account of the female
generative organs.

Cuvier dissected a male elephant in 1805,! assisted by his pupils
and by the painter Maréchal. He subsequently dissected two other
specimens, a male and a female. The results appear in his post-
humous Recueil de Planches de Myologie, published by Laurillard.
The plates are not good. We find drawings only, without descriptive
text, and the letters of designation are not always rightly affixed.
Everywhere the want of the master’s eye seems apparent. Though
most of the details are correct, it is necessary to warn future students
that hardly anything can be taken on trust from these plates. In
the present memoir a systematic comparison with our own dissections
has been made.

Mayer’s dissection is given very briefly. Many of the muscles are
merely named. We have found several mistakes of importance in this
short account.

Owen, R.—Odontography, pp. 625-655, pl. 146-150. London (1840-1845).
PERRAULT, CL.—Mémoires de 1 Académie Royale des Sciences, tom. iii. part iii.
p. 161, pl. xxiii. (1734).
Describes the African Elephant in Mémoires pour servir a U Histoire
Naturelle des Animauzx, vol. ii. Amsterdam (1758).

Sresa, ALBERT.—Locupletissimi rerum naturalium Thesauri accurata descriptio,
&e., vol. i. tab. iii. Amst. fol. (1734-1765). [Fcetus of Elephant. ]

SERRAO.—Opuscoli di vario Argomento. Napoli, 4to, fig. (1766).

STUKELEY.—KEssay towards the Anatomy of the Elephant (1723).

On the Spleen and Anatomy of an Elephant. London, fol. (1733).

Supty.—Remarques anatomiques sur un éléphant ouvert au fort Saint-Georges
(1715).

TurNER, W.—Lectures on the Comparative Anatomy of the Placenta, 1st series,
Edinburgh (1876). [Relates some Observations on the Structure of the
Placenta in the Elephant. ]

VULPIAN ET PuHitIppAux.—Notes sur le cceur, le foie et les poumons d’un
éléphant (femelle). Ann. Sci. Nat. Zoologie, vol. v. (1856),

Watson, Morrison, M.D.—Contributions to the Anatomy of the Indian
Elephant. Journal of Anatomy and Physiology, 1871-74. Observations
in Human and Comparative Anatomy. Hdinburgh (1874).

ZIMMERMANN, E. A. W.—Beschreibung und Abbildung eines Neugebornen
Elephanten. Erlangen, 4to (1783).

1 Deleuze, ‘‘ History and Description of the Royal Museum of Natural History,”

ting. Trans. vol. i. p. 91.

8 STUDIES IN COMPARATIVE ANATOMY.

Vulpian and Philippaux vive an excellent description of the heart,
lungs, and liver. We have not thought it needful to describe these
viscera afresh.

In 1856 the late Professor Goodsir purchased an Indian elephant, and
dissected its fore and hind limbs, and the notes of his dissections, un-
fortunately very imperfect, are published in the Appendix to his
Anatomical Memoirs, vol. i. p. 446. As the head and several of
the viscera of this specimen had been preserved in spirit in the stores
of the Anatomical Museum of the University of Edinburgh, Dr
Morrison Watson was permitted by Professor Turner to examine them,
and the results of his dissections were published in this Journal.

Our dissections, for the most part, agree closely with Dr Watson’s, |
and had he been able to complete his researches in the same manner,
our treatise would have proved superfluous.

A few words may usefully be said upon the mode of dissection.
When this particularly awkward subject was offered to us we had
no experience or instructions to guide us as to the best way of pro-
ceeding. We have, nevertheless, succeeded in preserving all the parts
in a nearly unaltered state for three years without immersion in fluid.
Any future dissector of a very large animal will do well to inject
a preservative solution repeatedly before dismembering the carcase.
Afterwards, it is merely necessary to keep the parts in a close-fitting
box, lined with lead, and to wrap them in cloths wetted with dilute
spirit, or other antiseptic. Occasional change of position is desirable.

Our example was a’ young female, about five feet high. .It was
purchased in December 1874 by the Council of the Leeds Philoso-
phical and Literary Society, to whom our thanks are due, not only
for permission to dissect the animal, but also for the readiness with
which they have provided the appliances of all kinds necessary for
such a piece of work.

MUSCLES OF THE FoRE-LIMB.

Pectoralis major consists of two distinct portions. The super-
ficial arises from the forepart of the sternum, and passes nearly
transversely outwards, to be inserted about the middle of the
shaft of the humerus. It is separated at its insertion by a small
interval from the other part of the muscle. A few of the lowest
fibres are continued into the fascia of the arm. Pectoralis major
is overlaid by masto-humeralis and deltoid, with which some of
its lower fibres are blended. The remaining portion of pectoralis
major arises from the posterior two-thirds of the sternum, and
passes forwards and outwards beneath the superficial part of the
muscle, to be inserted into the humerus along the outer border
of the bicipital groove, reaching upwards nearly to the apex of
the greater tuberosity.

THE ANATOMY OF THE INDIAN ELEPHANT. 9

C. and L.1—274-5, 7; 283, figs. 1,2, 7,71; 284, fie. 1, 7,7 (should
be j,) ; 285-6, 7,7,; 287-8, 7,7,; 292-3, 7,7? (should be /,).

Pectoralis minor.—Cuvier and Laurillard figure (292-3, j*) a
small muscle arismg beneath the anterior portion of pectoralis
major, and widening considerably towards its insertion into the
upper end of the humerus. This is apparently the pectoralis
minor of their index. We have not found this muscle.?

Serratus magnus arises by digitations from the 7th, 8th, 9th,
and 10th ribs, and from the fascia above them. It is inserted
into the inner surface of the apex of the scapula.

C. and L., 276-7 (g), 282, upper figure (9). Represented as
arising from five posterior cervical vertebre, as well as from
ribs, 283, figs. 1, 2 (9).

Masto-humeralis arises by two distinct heads, the larger of
which is attached by a strong rounded tendon to the basilar
process of the occipital bone. The second and thinner head
arises by a flat tendon from the mastoid process between the
stylo-mastoid foramen and the origin of digastricus. It is inserted
into the top and outer side of the tuberosity of the humerus, blend-
ing with the anterior part of the deltoid.

C. and L.—274-5 (61), “cleido-mastoidien ou trachélien ;”
276-7 (6) ; 287-8 (6') Gncorrectly shown as inserted into the first
rib); 290-1 (61), 292-3 (61).

The plates indicate an insertion much lower down upon the
humerus than that given above. We cannot reconcile the rela-
tions of b' to & (masto-humeralis to deltoid) in the different plates.

Delioideus is comparatively thin. It arises from the under
surface of a strong fascia, which is attached to the spine of the
scapula, and stretches over the shoulder-joint. Part of the
muscular fibres pass back to the angle formed by the spine and
the unciform process. The remaining fibres arise from the fascia,
along a line which extends from the tip of the unciform process
to the posterior -border of the scapula. The fascia is continued
upwards and backwards over the infraspinatus to the borders of
the fossa. The deltoid passes downwards, winding over the

1 Cuvier and Laurillard, Recueil de Planches de Myologie.

* Our notes of the sterno-humeral muscles are, unfortunately, not quite com-
plete, and certain details cannot now be recovered. The parts were cut through,
when little beyond a post-mortem examination was thought possible.

10 STUDIES IN COMPARATIVE ANATOMY.

tuberosity of the humerus, and is inserted into the outer and
anterior surface of the humerus, immediately below the root of
the tuberosity. Some of its fibres run into the teres minor, q.v.

C. and L.—274-5, k! (we have found nothing exactly corre-
sponding to k); 276-7 (k'); 283, fig. 1 (£2); 284, fig. 1 (4);
292-3 (k 2).

Subscapularis arises from the whole internal surface of the
scapula, except at the angles. Serratus magnus is inserted into
the superior angle; teres major arises from the inferior angle ;
while above the glenoid cavity is a surface unoccupied by
muscle, over which the subscapularis plays. This muscle takes
origin also from the suprascapular ligament. It is inserted by a
thin flat tendon into the internal surface of the head of the
humerus close to the articulation. The insertion is overlaid by
the coraco-brachialis.

C. and L.—283, figs. 1, 2 (n); 284, fig. 1 (n).

Supraspinatus.—A thick muscular mass, fillmg the supra-
spinous fossa, and taking origin also above from the fascia which
covers the muscle. Inserted into the upper border of the tuber-
osity of the humerus. Fleshy throughout.

—C. and L.— 276-7; 283, figs. 1, 2; 284, fig. 1 (2).

Infraspinatus arises from the posterior surface of the spine of
the scapula, and from the under surface of the unciform process.
Its origin from the bone is defined below by a line drawn from
the tuberosity of the humerus to the junction of the upper and
middle thirds of the posterior border of the scapula. It arises
also from the intermuscular septum between it and teres minor,
and from the fascia which covers in the infraspinous fossa above
the level of the unciform process. The muscle converges to a
strong flat tendon, which is inserted into an oblique line along
the outer surface of the base of the tuberosity of the humerus.
Above this line a large and distinct bursa separates it from the
tuberosity. |

C. and L., 276-7 (mm).

Teres minor arises from the outer surface of the scapula along
the middle third of the posterior margin, and from the intermus-
cular septum beneath the muscle. It is inserted into the neck
of the humerus behind and below infraspinatus, with the lower
border of which some of its fibres are connected. Its tendon is

THE ANATOMY OF THE INDIAN ELEPHANT, 11

also closely connected with a very strong bundle of ligamentous
fibres from the deep surface of the deltoid.

C. and L., 276-7 (0, the letter indicates teres major, according
to the index).

Teres major arises from a small triangular surface on the
internal aspect of the inferior angle of the scapula. It is in-
serted into the neck of the humerus along about two inches of a
line connecting the inner side of the head of the humerus with
the internal condyle. The lower fibres of the tendon are con-
nected with part of coraco-brachialis. Immediately behind it is
a bundle of muscular fibres, forming part of the triceps. The
tendon of teres major is embraced by the two portions of the
tendon of latissimus dorsi, of which the more superficial is
inserted into the humerus a little above teres major, while the
deeper and smaller portion is inserted immediately behind it.

C. and L.—283, figs. 1, 2 (0). In the second figure the origin
would agree better with that of teres minor.

Coraco-brachialis arises thin and flat, by tendinous fibres from
the tip of the coracoid tubercle, and from the upper part of the
capsule of the shoulder-joint, It is inserted by fleshy fibres into
the anterior surface of the humerus, between the insertion of
latissimus dorsi and the internal condyle.

C. and L.—283, fies. 1, 2; 284, fig. 1 (q).

Biceps arises within the capsule of the shoulder-joint, from
which it emerges as a strong round tendon: There is no trace of
a coracoid head. The muscle gradually increases in thickness to
the elbow-joint. Some of the deeper fibres are inserted into the
anterior part of the capsular ligament, the rest pass over the
joint to be inserted into the front and inner side of the ulna,
close to its upper extremity. A large vein separates the inser-
tion of the biceps from brachialis anticus.

C. and L.—283, figs. 1, 2 (r); 284, fig. 1 (71). The drawings
are correct; but the index mentions a coracoid head which does
not exist.

Brachialis anticus arises from the humerus just below the
articular end, and external to the greater tuberosity. The
muscle winds round the shaft of the bone to reach the front of
the external condyloid ridge, from the commencement of which
it receives some of its lower fibres. Inserted into the front of

14 STUDIES IN COMPARATIVE ANATOMY.

the ulna, immediately below the biceps. The musculo-spiral
nerve lies along its posterior border, while the extensor carpi radi-
alis longior crosses it superficially. Beneath it les a plexus of
large veins, just above the elbow-joint, and a large horizontal
branch of the brachial artery.

C. and L.—276-7 (s), 283, figs. 1, 2 (s); 284, figs. 1, 2 (s)?

Triceps—The middle or long head (C. and L., 274-5, ¢, 276-7
t, 283 fies. 1, 2, 2, origin only 284 fig. 1717), arises from the
whole axillary border of the scapula by a thick and strong
fleshy mass. It expands as it passes down, blends towards the
elbow with the external head, and is inserted with it into the
posterior surface of the end of the olecranon, lying superficially
to the remainder of the muscle, from which it is separated indis-
tinctly by connective tissue. The external head (C. and L.,
274-5, 1, 276-7, #1), the stronger of the remaining two, arises
from the posterior surface of the humerus, reaching as high as
the greater tuberosity. It bridges over the musculo-spiral nerve
just above the external condyloid ridge, and blends with the
deeper surface of the middle head. The imner head (C. and L.,
283, fig. 1, #3, 283, fig. 2, ¢) has a small pointed origin close
below the insertion of latissimus dorsi. It passes downwards
and widens out so as to occupy the whole posterior surface of
the bone, between the condyloid ridges, a strong bundle taking
origin from the external condyloid ridge. The inner head is
inserted anterior to the rest of the muscle into the olecranon
process.

Dorso-epitrochlearis—A thin and flat muscle, which arises
close to the posterior angle of the scapula on its ventral surface,
and from aponeurotic fibres superficial to triceps. The lower
part of the muscle is connected, close to its origin, with strong
aponeurotic fibres derived partly from Jatissimus dorsi, partly from
panniculus, and passing into the posterior border of the middle
head of triceps. Dorso-epitrochlearis passes down as a thin flat
muscle, extensively tendinous on its deeper surface, and is
inserted by a small rounded tendon into the olecranon on its
inner side, close to triceps, with which it partly blends.

C. and L., 283, fig. 1 (@), fig. 2 (@ 7).

Pronator radit teres—A strong ligament, which may repre-
sent this muscle, springs from the inner side of the internal con-

THE ANATOMY OF THE INDIAN ELEPHANT. 13

dyle of the humerus, about on a level with flexor carpi radials,
but anterior to it. This ligament passes obliquely over the
muscles and vessels, and is inserted into the middle of the
radius for about two inches, expanding greatly over the bone.
Beneath it, towards the radial extremity, are some muscular
fibres, which spread out towards their insertion as a small trian-
gular muscle, continuous with the ligament, and covered by it
superficially. .

C. and L., 283, figs. 1, 2 (@).

Fleaor carpi radialis arises by a flat tendon from the back of
the internal condyle of the humerus, passes directly along the
inner side of the fore-arm, and ends in a round tendon, which
occupies a distinct sheath connected with the annular liga-
ment. The tendon is finally inserted rather deeply into the
carpus. This muscle is remarkably elastic. When cut, it is
retracted 3 inches, and is seen to be composed of regular alter-
nate layers of muscular and elastic fibres in about equal propor-

‘tions. The muscular fibres predominate towards the deeper
surface, and also towards the origin of the muscle. This unex-
pected structural peculiarity would give rigidity to the limb,
when it is extended in a perfectly straight line. When the
animal stands, the toes are, so to speak, over-extended, so that
the flexor carpi radialis, stretched like the chord across an are,
tends to draw the carpus upwards and forwards, and opposes the
flexors. When, on the other hand, flexion has carried the limb
in the least beyond the neutral line, the elastic band assists the
flexors, drawing the carpus upwards and backwards.

C. and L—283, fies. 1, 2 (vy); 284, fig. 2 (y).

Elastic ligament of outer side of fore-arm.—Connected with the
external condyle of the humerus is a very strong flattened band,
composed partly of yellow elastic, but chiefly of white fibrous
tissue. This passes down along the outer side of the fore-arm,
and blends on its anterior and posterior borders with the strong
fascia of that part of the limb. Opposite the upper fourth of the
bones of the fore-arm, this ligamentous band spreads out laterally,
becomes greatly thickened, and is converted almost wholly
into yellow elastic tissue. It continues to widen as far as 2
inches above the lower end of the ulna, where it divides into
two portions; of which the anterior and stronger is inserted into

14 STUDIES IN COMPARATIVE ANATOMY.

the front of the end of the ulna, the posterior into the outer sur-
face of the end of the same bone. The tendon of extensor minimi
digitt separates the two portions. The insertion consists of white
fibres chiefly.

Palmaris longus arises from the back of the internal condyle
of the humerus, and from the strong intermuscular septa which
divide it on the radial side from flexor communis digitorum, and
on the ulnar side from flexor carpi ulnaris. Palmaris is a flat
muscle, becoming tendinous at the commencement of the carpus,
where it spreads out over the palm. The strongest portion of
the tendon passes to the ulnar side, and is inserted into the
sesamoid bone of the 5th digit. A median portion is much
slighter, and is lost m the palm. On the radial side, part of the
expansion blends with that of the extensor tendon. Lastly, a
strong tendinous bundle of palmaris is attached by its deep
surface to the annular ligament, nearly over the tendon of flexor
carpt radialis, and superficial to the origin of flexor brevis.

C. and L.—283, figs. 1, 2 (a); 284, fig. 2 (a).

Flexor carpi ulnaris arises by two heads, one, thin and pointed,
from the internal condyle, the common aponeurosis, and exten-
sively from the intermuscular septum between this muscle and
flexor communis digitorum. The second head, on the ulnar side,
arises thick and fleshy from the back and inner side of the ole-
cranon, immediately below the insertion of triceps. The ulnar
nerve passes between the two heads of origin. The united
muscle is inserted into the pisiform bone, and continued thence
by strong fibres to the ulnar side of the manus.

C. and L.—274-5 (@); 276--7 (8); 283, figs. 1, 2 (6); 284,
figs. 2, 4 (8).

Flexor communis digitorum arises by a common tendon with
palmaris and flexor carpi ulnaris from the back of the internal
condyle of the humerus. Its deep surface is tendinous, and
separated by loose cellular tissue from the bones of the fore-arm.
It ends in a very strong, flat tendon, which divides into five
opposite the base of the radial sesamoid bone. Of these tendons,
those to the Ist and 5th digits are only half the size of any of
the others. The middle tendon is rather the strongest of the
remaining three. Inserted as usual into the terminal phalanges.
A small muscle, which may correspond to jlexor longus pollicis,

THE ANATOMY OF THE INDIAN ELEPHANT. 15

arises, close to the common tendon, from the internal condyle of
the humerus, but somewhat deeper. It lies beneath flexor com-
munis digitorum, and to its radial side. At the lower fourth of
the fore-arm it ends in a round tendon, which joins the deep
surface of the common flexor. Another muscle arises by fleshy
fibres from the ulna, beneath the ulnar origin of flexor carpr
wlnaris, and to its inner side. It consists of a small conical
bundle of muscular fibres, ending in a thin round tendon near
the upper extremity of the fore-arm. The tendon joins the deep
surface of the tendon of the common flexor on its ulnar side.

C. and L.—283, fig. 2 (uw, w+); 284, fic. 2 (ut, w, the letters of
the preceding figure are reversed here, pu”), fig. 4 (w).

Hlexor brevis digitorum arises from the annular ligament of the
wrist, where it forms a sheath for flexor carpr radialis. It con-
sists mainly of fleshy fibres. Opposite the base of the radial
sesamoid it divides into two nearly equal masses, of which that
to the radial side is inserted into the sheath of the long flexor
tendon of the 4th digit. The short flexor crosses the other
superficially, and is attached to its ulnar side. The second mass,
shorter and smaller than the other, is similarly inserted into the
flexor tendon of the 5th digit, blending also with the palmar
fascia at its attachment to the sesamoid bone of the 5th digit.

C. and L.— 283, fig. 2 (€); 284, figs. 2. 3, 4 (&).

Extensor carpi radialis longior (supinator longus?) arises
chiefly from the lower part of the tendon of insertion of the
deltoid on its outer side. It extends a little further upwards,
and is connected with the lower and anterior border of the ex-
ternal head of the ¢riceps. Between these poimts a thin slip
arises from the humerus. The muscle passes down in its usual
position, bridging over the brachialis anticus. It ends at the
lower third of the fore-arm in a small tendon, which passes
together with extensor carpi radialis beneath extensor proprius
pollicis, then through a sheath upon the radius, having a separate
compartment, but lying in the same groove with eatensor carpi
radialis. It lies rather deeper than that muscle, and to its inner
side. It is inserted into the semilunar bone of the carpus.

C. and L.—274-5 (6) ; 276-7 (6); 283, figs. 1,2 (6); 284, fig. 1 (6).

Extensor carpi radialis brevior arises from the front of the
external condyloid ridge. It ends in a strong tendon which

16 STUDIES IN COMPARATIVE ANATOMY.

passes beneath extensor pollicis, and has a distinct sheath upon
the lower end of the radius; crosses the carpus, and is inserted
principally into the base of the metacarpal of the 3d digit, and
partly into the metacarpal of the 2d.

C. and L.—274-5 (61); 276-7 (61); 283, figs. 1, 2 (6'); 284,
fig. 1 (64).

Hxtensor communis digitorum arises by fleshy and tendinous
fibres from the lower part of the external condyloid ridge, on the
outer side of extensor carpi radialis, and also from the outer side
of the external condyle, the two portions being indistinctly sepa-
rated by a septum. The muscle passes down in a special sheath
over the end of the radius, and becomes tendinous opposite the
wrist-jomt. The tendon spreads over the carpus, and is finally
inserted into the upper surface of the 2d, 3d, 4th, and 5th digits.
The divisions of the tendon of insertion form arches beneath
which the vessels appear, as in the pes.

C. and L.—274-5 (e); 276-7 (e); 284, fig. 1 (©).

Extensor minim digiti arises by two heads—one muscular,
and the stronger of the two, from the outer surface of the ex-
ternal condyle; the other, which is smaller and tendinous, from
the front and outer side of the head and shaft of the ulna, and
from an intermuscular septum between it and extensor carpi
ulnaris. The muscle passes through a distinct sheath on the
back of the ulna to the outer side of the manus, crossing the
insertion of extensor carpi ulnaris, and is inserted into the base
of the 5th digit. A tendinous expansion passes from the fore-
part of the tendon to the outer side of the base of the Ist
phalanx of the 4th digit, and blends with the deep surface of
extensor communis.

C. and L.—274-5 (e!); 276-7 (1) ; 284, fig. 1 (e"). The addi-
tional slip of insertion is shown as passing to the 3d and not to
the 4th digit.

Extensor carpi ulnaris, a strong muscle arising by a round
tendon from the external condyle of the humerus, behind and a
little above eatensor minim digiti. It lies along the outer sur-
face of the ulna, passes through a distinct sheath on the outer
side of the lower extremity of that bone, and is inserted by a
strong round tendon into the metacarpal of the 5th digit.

C. and L.—274-5 ; 276-7; 284, fig. 2 (6).

THE ANATOMY OF THE INDIAN ELEPHANT. 17

Anconcus arises chiefly from the back of the external condy-
loid ridge; some of the upper fibres taking origin from the
posterior border of the elastic ligament of the fore-arm, and some
of the deeper fibres from the superior tendon of extensor carpi
ulnaris, From this origin the muscle spreads out, and is inserted
into a triangular surface on the outer side of the ulna, imme-
diately beneath the olecranon. The apex of this triangle is
situated nearly half way down the shaft. Below, the muscle is
overlaid by extensor carpi ulnaris, and above by a dense fascia.

C. and L.—274-5 ; 276-7; 283, fig. 2; 284, fig. 2 (w).

Extensor pollicis arises from nearly the whole anterior surface
of the radius, with the exception of a small part towards the
lower end of the bone, from the interosseous membrane and
from the adjacent surface of the ulna. At the lower part of the
radius the muscle is twisted upon itself, so that the external
fibres become anterior, and ultimately internal. The tendon
passes through a special sheath on the inner side of the end of
the radius, and is inserted into the base of the Ist digit. The
tendon is connected by a lateral expansion with that of the
extensor indicis. A large vein and the posterior interosseous
nerve pass between the radial and ulnar origins of the muscle.
Several large veins, communicating with the plexus in front of
the elbow-joint, lie in the interosseous space beneath the muscle.

C. and L—274-5; 276-7; 283, figs. 1,2; 284, fig. 1 (0).

Extensor indicis—Small and pointed above, where it arises
from the front of the ulna close to the head of the radius, and
below this from the intermuscular septum on the outer side
between it and extensor minimi digiti, and on the inner side
between it and extensor pollicis. The extensor indicis passes over
the wrist in a separate synovial sheath, but in the same groove
with extensor communis digitorum, and to its inner side. The
tendon passes obliquely to the inner side of the manus, crossing
the tendon of extensor carpi radials. It then spreads out, and
is inserted into the inner (radial) side and front surface of the
1st phalanx of the 2d digit. An expansion connects this tendon
with that of extensor pollicis, close to their insertion.

C. and L.—274-5 ; 276-7; 284, fig. 1 (&). Shown as con-
nected with the inner side of the tendon of extensor communis
digitorum, not with extensor pollicis.

B

18 STUDIES IN COMPARATIVE ANATOMY.

Abductor minim digiti arises from the annular ligament and
pisiform bone. It is inserted into the sesamoid bone of the 5th
digit to the ulnar side of flexor brevis digitorum.

C. and L.—283, fig. 2; 284, figs. 2, 3,4 (v). In 283, fig. 2, a
small muscle (“abductor”—adductor?—* minimi digiti”) is shown
internal to the short flexor of the toes. We have not found this.

The palmar fascia consists of strong fibrous bands crossing i
various directions, with large intermediate spaces filled with fat,
especially between the digits. An exceedingly strong semi-
circular band, concave towards the wrist, stretches across the
heads of the metacarpal bones, and forms sheaths in the usual
way for the flexor tendons. By dissection these sheaths can be
resolved into a superficial or proper transverse ligament, and a
deeper layer, which, when divided, presents on its deep surface
two tendinous slips passing to be inserted into the base of the
second phalanx on each side of the flexor tendon. This dis-
position resembles the ordinary arrangement of the /lexor
sublumis.

C. and L., 284, fig. 3.

Lumbricales arise all together from the superficial surface of
the flexor tendon, just above its division. They are four in
number, of which the 1st and 2d (from the radial side) are
respectively inserted into the bases of the 1st and 2d digits on
the ulnar side. The 3d lumbrical is inserted into the deep side
of the sheath of the flexor of the 3d digit, and is continuous
with one of the lateral tendinous bands of the flexor sheath.
The 4th lumbrical passes deeply between the 3d and 4th digits,
and is connected with an aponeurosis from the extensor tendon.
The 1st and 2d lwmbricales appear to have a similar attachment
on the extensor side of the manus.

C. and L.—284, fig. 4. The 1st lumbrical is not seen in this
drawing.

Flexor brevis pollicis—The only short muscle of the first digit.
Arises by a pointed origin from the os magnum, and is inserted
into the sesamoid bone on the head of the first metacarpal. This
muscle corresponds most nearly with the inner head of flexor
brevis pollicis.

Not in C. and L.

Flexor brevis mininu digitc consists of two slips, which arise

THE ANATOMY OF THE INDIAN ELEPHANT. 19

from the unciform bone, and from a ligamentous band uniting
the unciform and pisiform bones. The slips run side by side to
their insertion into the ulnar side of the sesamoid on the head
of the fifth metacarpal.

Not in C. and L.

Opponens minima digiti arises from a prominence upon the
unciform bone, opposite the articulation of the fourth meta-
carpal, and is inserted into the radial side of the sesamoid of the
fifth digit.

Not in C. and L.

Interosset (manus).—There are three palmar interossei, which
arise near together from the carpus, opposite the articulation of
the third metacarpal. They diverge to be inserted—the first
into the expansion of the extensor communis digitorum on the
ulnar side of the second digit ; the other two in the same manner,
but on the radial sides of the fourth and fifth digits. The inser-
tion corresponds, therefore, with that in the human subject. The
dorsal interossei are also three in number, and form short, thick
muscles. That of the index digit consists of two distinct por-
tions, one arising from the base of the first metacarpal and the
adjacent surface of the trapezium; the other from the base of
the second metacarpal. They are inserted together into the
sesamoid of the index. The inéterosseus of the third digit arises
from the base of the third metacarpal, and is inserted like the
preceding. The interosseus of the fourth digit is quite similar to
that of the third. Each of the dorsal interosse: shows a distinct
separation into lateral halves, one to each division of the sesa-
moid. The inéerosset are invisible from the dorsal side of the
manus.

Not in C. and L.

Of the foot as a whole, and this remark applies to both
fore and hind extremities, the separate mobility of the parts
is greater than would be suspected from an external inspection,
and much greater than in most Ungulates. The palmar and plantar
soles, though thick and tough, are not rigid boxes like hoofs, but
may be made to bend even by human fingers. The large develop-
ment of muscles acting upon the carpus and tarsus, and the
separate existence of flexors and extensors of individual digits, is
further proof that the elephant’s foot is far from being a solid,

20 STUDIES IN COMPARATIVE ANATOMY.

unalterable mass. There are, as has been pointed out, tendinous
or ligamentous attachments which restrain the independent
action of some of these muscles; but anatomical examination
would lead us to suppose that the living animal could, at all
events, accurately direct any part of the circumference of the
foot by itself to the ground.

The metacarpal and metatarsal bones form a considerable
angle with the surface of the sole; while the digits, when sup-
porting the weight of the body, are nearly horizontal.

MUSCLES OF THE HIND Limp.

Psoas magnus arises chiefly from the sides of the bodies of the
four lower dorsal vertebrae. It receives slips from the 15th,
16th, 17th, 18th, and 19th ribs. Inserted, together with zliacus,
into the inner side of the femur below the neck of the bone.
Another slip arises from the brim of the pelvis and the capsular
ligament, passing to the front of the femur, 3 inches below the
capsular ligament.

C. and L., 290-1, h.

Psoas parvus arises from the sides of the bodies of the lumbar
vertebre, internal to psoas magnus. It ends in a tendon about
4 inches long, which is inserted into the brim of the pelvis for
about the middle third of the lateral margin. The external iliac
vessels lie adjacent to the inner side of this muscle.

C. and L., 290-1, g.

Ilacus arises from the whole of the iliac fossa, except a small
space adjacent to the brim of the pelvis, over which psoas magnus
passes. It is inserted together with that muscle. The anterior
crural nerve emerges between psoas and iacus in the usual
position.

C, and L.—290-1, 2; 292-3, 7; 294, fig. 1,7. In 290-1 the
muscle is shown as a rounded tapering mass, lying upon the
inferior surface of the ilium, and projecting considerably from it.
The full width of the muscle (which occupies the whole of the
ilium) and its conformity to the curves of the iliac fossa do not
appear in the drawing.

Tensor fascie femoris, a thick strong muscle, arises from the
outer surface of the anterior spine of the ilium, and from the

THE ANATOMY OF THE INDIAN ELEPHANT. 21

bone immediately adjacent. It passes downwards on the outer
side of the thigh, and is lost in the fascia at the junction of the
lower third with the remainder of the thigh. Here it is closely
united with the anterior fibres of gluteus maximus.

C. and L.— 274-5 «, 276-7, 287-8, 290-1, 292-3; 294, fig. 1 a.
Omitted by Mayer.

Gracilis arises by a very thin flat tendon from the posterior
three-fourths of the middle line of the pubic symphysis. The
tendon has a straight anterior border, and terminates about half-
way between the origin and insertion, where muscular fibres take
its place. On the posterior border the muscular fibres run up to
the origin. Most of the fibres pass backwards and downwards,
and are inserted into the fascia of the thigh. The anterior
fibres, which form a separable band about two inches wide, are
inserted into the inner side of the tibia, an inch and a half below
the tuberosity.

C. and L.—285-6 uw. In this drawing the posterior margin is
too distinct, and the muscle far too rounded and solid. 290-1;
292-3 ; 294, fig. 2 wu.

Sartorius.—A. broad, flat muscle, which arises from the fascia
on the front and inner part of the thigh below Poupart’s
ligament. The muscle consists of loosely coherent muscular
bundles which thin out below. The most prolonged are inserted
by several tendinous strings into the thick yellow fascia of the
leg immediately below the knee-joint on its inner side.

Not figured by Cuvier and Laurillard.

Gluteus maximus occupies, superficially, the inner two-thirds
of the buttock. It is sheathed in fascia, which, on the outer
side of the muscle, is strengthened by a considerable thickness
of yellow elastic tissue, but thins away internally. The muscle
arises from the crest of the ium by means of a thin fascia,
from the dense fascia which covers glutwus medius, from the side
of the sacrum and upper caudal vertebrae, and from the great
sacro-sciatic ligament. The fibres are inserted as follows :—The
anterior or external part into the upper part of the fascia lata,
the internal fibres into the inner edge of the same, while a strong
bundle of the deep inner fibres passes to the middle fifth of the
femur.

C, and L.—274-5 ; 276-7; 287-8; 294, fig. 2 (a, g part),

22 STUDIES IN COMPARATIVE ANATOMY.

We cannot understand Pl. 294, fig. 2. The foreshortening is
probably wrong. What are described above as the internal
fibres of gluteus maximus, Cuvier and Laurillard call biceps,
while the deep inner fibres correspond to their “ biceps—portion
fémorale.” Mayer assigns to gluteus maximus an insertion into
the great trochanter; he speaks further of its tendon as blending
with a portion of the “ caput breve des M. occipitis.”

Gluteus medius arises from the side of the upper part of the
sacrum beneath gluteus maximus, from the upper half of the
back of the ilium, from the dense fascia covering the muscle,
and from the lower and anterior part of the crest of the ilium as
far as the anterior spine. It is separated from gluteus minimus
by vessels, and the superior gluteal nerve. From this origin the
fibres converge and are inserted as follows:—The anterior two-
thirds into a line which extends obliquely downwards and for-
wards from the superior and posterior part of the great trochanter,
the posterior third into the apex of the trochanter and into a
ligament which passes thence to the acetabulum. The most
posterior fibres pass down to the posterior root of the trochanter
about 8 inches from its apex.

C. and L.—276-7 ; 287-8; 294 (a+).

Gluteus minimus arises from the middle third of the back of
the ilium beneath glutwus medius. The fibres converge to a
small tendon which is inserted into a depression on the outer
aspect of the root of the trochanter.

C. and L.—287-8 (a,); 294, fig. 1 (a,). The f of the last-
mentioned figures, “pyramidal (sacro-trochantérien),” we have
not found as a separate muscle. In the absence of a, this would
agree with our G. minimus.

Biceps femoris arises by a thin round tendon from the tuber-
osity of the ischium beneath and external to the other hamstring
muscles. It is narrow above, but widens greatly below. The
more anterior fibres are continuous with the lower fibres of
gluteus maximus, and are inserted into the outer side of the
femur for several inches above the knee. A separable strap-like
bundle of the most posterior fibres proceeds from the belly of
the biceps to the fascia of the leg. The peroneal nerve separ-
ates the anterior from the posterior division of the muscle.

C. and L. (biceps—portion fémorale).—274-5; 276-7; 287-8;

THE ANATOMY OF THE INDIAN ELEPHANT. 23

294, fig. 2 (part of g, and in 294, fig. 2 g). The biceps of Cuvier
and Laurillard includes also what we have described as the pos-
terior part of gluteus maaxinvus.

A strong band of fascia is continued from the back of the
leg to the caleaneum ; biceps passes into the oblique upper edge
of this fascia, and the tendon of gastrocnemius externus blends
with its deep surface.

Semitendinosus arises by fleshy fibres from the ramus of the
ischium, and ends about the middle of the thigh in a thin tendon,
which is lost in the fascia of the inner and back part of the
thigh.

C. and L.—274-5 ; 285-6 ; 287-8; 289, figs. 1, 3, 4; 294 (7).
Drawn as passing to the inner side of the tibia, and ending in a
distinct tendon.

Semi-membranosus arises by a double tendon. The external
head, long, round, and slender, is attached to the great sacro-
sciatic ligament and the side of the sacrum. The other, square
in section, and tendinous upon its deep surface, springs from the
tuberosity of the ischium. The muscle ends in a flattened
tendon about three inches below the knee-joint. This tendon
divides into two parts, of which the anterior, broad and flat, is
inserted into the anterior and internal surface of the tibia, about
the middle of the bone; the posterior portion is continued to the
internal malleolus, and is connected along its posterior border
with the deep fascia of the lee.

C. and L.—274-5 ; 285-6; 287-8; 289, fig. 4; 294 (s).

Obturator externus arises from the outer surface of the obturator
membrane, and from the bone beneath. It is inserted into the
digital fossa of the femur, anterior to obtwrator internus, and
somewhat below it. i

Not given in Cuvier and Laurillard.

Obturator internus arises from the inner margin of the obtura-
tor foramen, and is inserted into the digital fossa, posterior to
obturator externus, and by a narrower tendon.

Not given in Cuvier and Laurillard.

Adductor magnus takes origin from the os pubis and ramus of
the ischium. It is inserted into the back and inner side of the
femur for about the middle two-thirds of its length, and by a
separate slip into the internal condyle of the same bone.

24 - STUDIES IN COMPARATIVE ANATOMY.

Adductor magnus is pierced, as usual, by the femoral vessels,
which intervene between the main part of the muscle and the
slip which passes to the internal condyle. It is pierced also by
the obturator nerve.

C. and L.—285-6 (1); 289, figs. 3, 4 (2) ; 290-1 (21); 292-3
().

Adductor longus is closely connected above with pectineus, but
has a distinct rounded tendon of origin, attached to the front of
the pubes beneath pectineus, and nearer to the symphysis. Ad-
ductor longus is inserted into the inner side of the femur, reach-
ing to a point just above the internal condyle.

C. and L.—(“ 2° Pectiné”), 290-1; 292-3 (}).

Adductor minimus.—This large and distinct quadrilateral
muscle arises from the ramus of the ischium, beneath the ham-
string muscles, and is inserted into the posterior surface of the
femur for about the second fourth of the bone, counting from
above. Towards its insertion it is partly enveloped by adductor
magnus, a large vein separating the two muscles. Their fibres
ultimately unite, and are inserted together.

C. and L., 290-1 (d?).

Pectineus takes origin between the iliopectineal eminence and
the symphysis pubis. Some of its fibres are closely connected
with the tendon of origin of adductor longus. It is inserted
into the middle of the femur on its inner side, above and
external to adductor longus.

C. and L.—285-6 ; 290-1; 292-3 (&).

Gastrocnemius externus arises by a small round tendon from
the external condyle of the femur in front of plantaris and
external to it; also from the intermuscular septum between it
and the peronez. It is a thin, flat muscle, ending by a tendon
in the deep surface of the common tendon of the os calcis.

C. and L.—(“ Plantaire gréle ”), 276-7, 285-6, 287-8, 289, figs.
3,4 (y). Mayer's figure of the gastrocnemu (Act. Acad. Ces,
Leop. vol, xxii. t. ix. fig. 4), is very indistinct and not accurate.

Gastrocnenvius internus takes origin as a thin flat muscle from
the internal condyle of the femur, and partly from the capsule
of the knee-joint. It passes downwards and outwards, ending a
little below the middle of the leg in a small flattened tendon.
The lower four inches are sheathed in a strong aponeurosis

THE ANATOMY OF THE INDIAN ELEPHANT, 25

derived chiefly from biceps femoris. Gastrocnemius externus and.
internus, semimembranosus, semitendinosus, and the aponeurosis
from biceps femoris unite to form the common tendon of the os
calcis.

C. and L.—276-7 ; 285-6 ; 289, figs. 3,4; 292 (a1).

Popliteus arises by a round tendon from the external condyle
of the femur, beneath the external lateral ligament. The muscle
occupies a deep groove, lined with cartilage and synovial mem-
brane, which les obliquely above the head of the fibula, and is
inserted into a triangular surface of considerable extent on the
posterior aspect of the upper half of the tibia. Below and to
its outer side, popliteus is bounded by a strong ligamentous
band, which stretches from the tibia to the head of the fibula.
Part of the muscle is inserted into the deep surface of this
band.

C. and L., 289, fig. 4 (v).

Plantaris arises from the posterior surface of the external
condyle of the femur by a roundish mass of muscular fibres.
About the junction of the upper and middle thirds of the leg it
passes into a tendon, very similar to that of gastrocnemius inter-
nus, but more slender. It passes beneath that muscle towards
the middle of the leg, and receives an investment from the
fascia. On reaching the os calcis, plantaris spreads out into a
thin fascia, separated from the heel by a bursa. Externally,
it is connected with a strong annular ligament which surrounds
the tarsus. Plantaris is finally inserted into the plantar fascia, q.v.

C. and L. (“ Gastrocnemien externe”)—274-5; 276-7; 285-6;
287-8 ; 289, figs. 3, 4 (a).

Soleus arises by a thin, flat tendon from the posterior and
external surface of the head of the fibula, and from the inter-
muscular septum between it and the perone. It is inserted by
fleshy fibres into the upper part of the tuberosity of the os calcis,
beneath the common tendon, but closely blended with it.
Soleus is channelled by several large veins, which course longi-
tudinally through it.

C. and L.—274-5; 276-7; 287-8; 289, fies. 3, 4; 292-3
(p).

Peroneus longus arises from the head of the tibia, and from
strong fibres which pass over the capsule of the knee-joint. It

26 STUDIES IN COMPARATIVE ANATOMY.

becomes tendinous, and passes down into a groove behind the
external malleolus, external to Peronewus brevis. The muscle
winds round the outer edge of the foot, to be inserted deeply,
beneath the calcaneo-cuboid ligament, into the posterior border
of the cuboid. A small part of the tendon passes further into
the sole, towards the bases of the metatarsals.

C. and L—274-5 ; 276-7; 287-8; 289, figs. 1, 2, 3, 4 (€).
Mayer states (incorrectly), that peroncus longus is pierced by
the peroneal nerve.

Peroneus brevis arises from nearly the whole of the outer
surface of the fibula. It becomes tendinous along its anterior
border immediately above the external malleolus; but the
muscular fibres are prolonged nearly to the insertion along the
posterior border. The tendon lies in the groove behind the
external malleolus, and is separated from that of peroncus longus
by a division of the external annular ligament. It then crosses
the base of the 5th metatarsal, and is inserted principally into
the fibular side of the metatarsal of the 4th digit; also, by a
small slip, into the proximal phalanx of the 5th. Peroncwus brevis
underlies P. Jongus, and is much broader than that muscle.
The peroneal nerve passes between them.

C. and L—274-5, but incorrectly drawn and not lettered ;
e! is put upon another muscle ; 276-7 (e+); 287-8 (¢!); 289, fig.
3 (e), figs. 1, 2, 4, not lettered; ¢! of these figures is our P.
anticus. ,

Peroneus anticus arises by a round tendon from the external
lateral ligament of the knee, and by a second head from the top
of the tibia and lower two-thirds of the front of the fibula, ad-
jacent to the tibia. It divides into two main slips. The more
superficial passes outwards, and is inserted into the outer border
of the 5th metatarsal. The deeper portion is subdivided into
two parts, of which the shorter passes to the 4th and 5th meta-
tarsals; the longer to the fibular side of the 4th. A slip from
the anterior border of the external malleolus blends with the
muscle. This accessory slip would agree with P. éertius.

C. and L.—274-5 (e,, €,); 276-7 (e,) ; 287-8 (+ «, accessory
slip) ; 287, fig. 1 (e,); fig. 2 (e, +); fig. 4 (+ «, accessory slip).
Not in Mayer.

Rectus femoris arises partly from the notch between the

THE ANATOMY OF THE INDIAN ELEPHANT. QO”

anterior spine of the ilium and the border of the acetabulum.
The tendon of attachment arches over the centre of the con-
cavity. Between it and the bone is a large branch of the
external iliac artery, and a considerable venous anastomosis.
The external head arises from the root of the great trochanter
on its anterior surface, and extensively from an intermuscular
septum between this muscle and vastus eaternus. The long
head is inserted into the front of the patella; the external
head has a slightly distinct insertion above, and to the outer
side of the other.

C. and L—285-6 (p); 287-8 (p); 290-1 (p); 292-3 (p);
294, fic. 1 (p; m, and o perhaps represent the external head).

Vastus internus arises from the lower half of the femur on its
inner surface. It becomes very thick and fleshy in its lower
part, and is inserted into the inner border of the patella and the
ligamentum patelle.

CO. and L.—285-6 (nm); 290-1 (nm); 292-3 (n); 294, fig.
1 (n).

Vastus externus arises from the outer surface of the trochanter,
and from nearly the whole length of the femur along its outer
border. The line of origin then curves out towards the external
condyle. ‘The muscle is inserted, chiefly by fleshy fibres, into
the outer side of the patella, into the heamentum patelle, and
the aponeurosis of the leg.

C. and L.—276-7 (the right-hand m) ; 287-8 (m); 289, fig. 2
(m); 290-1 (m); 294, fig. 1 (a).

Crureus arises from nearly the whole anterior surface of the
femur, and from the lower part of the external surface of the
trochanter. It sends processes of attachment between the ad-
jacent muscles, thus embracing by pointed slips the femoral
origin of rectus, and the insertion of glutwus minimus. It is
inserted into the upper border of the patella. This muscle is
much divided by vessels and tendinous septa.

C. and L.—290-1 (0); 294, fig. 1 (0). Mayer appears to con-
sider this muscle a part of vastus caternus.

Tibialis anticus is the most internal of three muscles which
occupy the front of the leg, and which are hardly separable
above. It arises from the external surface of the tibia, for the
middle three-fifths of the length of the bone, passing downwards

28 STUDIES IN COMPARATIVE ANATOMY.

and slightly inwards to its insertion into the metatarsal of
the 2d digit. It crosses the tendon of tibialis posticus, close
to its insertion. TZ%bialis anticus is fleshy almost to its ter-
mination.

C. and L.—285-6 (6); 287-8 (6); 289, figs. 1, 2 (6); 292-3.
Mayer states that this muscle is inserted into the cuboid and 1st
and 2d metatarsals.

Extensor communis digitorum arises from the tibia, external to
tibialis anticus, for nearly the whole length of the bone; also
from the intermuscular septum on its outer side. Passing down-
wards, it spreads out, and becomes tendinous over the tibio-tarsal
articulation. It finally divides into four slips, which are inserted
into the upper surface of the digits. The tendons to the third
and fourth toes are much stronger than the rest. That to the
fourth gives off a slip to the base of the proximal phalanx, an -
arrangement which does not exist in the other tendons.

C. and L—274-5 (€) ; 276-7 (&) ; 285-6 (€); 287-8 (€); 289,
figs. 1, 2 (€); 292-3 (€). The slips appear more distant from
each other in the drawings than in our dissections.

Extensor brevis digitorum has an oblique origin from the
dorsum of the tarsus, following a line drawn to connect the
external malleolus with the base of the second metatarsal. The
dorsal vessels pass between the two main divisions of the muscle,
close to its origin. The outermost and highest head of extensor
brevis is partly concealed by a strong ligamentous band con-
nected with the external lateral ligament. Opposite the heads
of the metatarsal bones the muscle blends with the under
surface of the tendinous expansion of extensor communis digit-
orum, but the deeper fibres pass into a strong ligament which
encloses the heads of the metatarsals and the bases of the first
phalanges. No part of the muscle is attached to the first or
second digits.

C. and L—276-7 (6) ; 287-8 (@); 289, figs. 1, 2 (@); 292-3
(0). Mayer’s brief account is very accurate.

We have not found the muscle named extensor of the meta-
carpus, and lettered + ¢ by Cuvier and Laurillard.

Tibialis posticus is concealed beneath flexor longus digitorum.
It arises from the posterior surface of the tibia, below what
answers to the oblique line; from the adjacent surface of the

THE ANATOMY OF THE INDIAN ELEPHANT. 29

fibula; and largely from the intermuscular septum between this
muscle and flexor longus digitorum. It ends in a tendon, which
passes through a distinct sheath on the inner malleolus, and is
inserted into the upper surface of the basis of the second and
third metatarsals, being crossed, immediately above its insertion,
by tubialis anticus.

C. and L.—239, fig. 4 (6); 292-3 (61). Mayer's description is
very incorrect; the muscle is said to pass over the outer malle-
olus, and to blend with tibialis anticus.

Flexor communis digitorum arises by tendinous and fleshy
fibres from the postero-internal surface of the head of the fibula.
An additional slip, small and tendinous, proceeds from the tibia,
internal to the insertion of popliteus, from the oblique ligament
between the tibia and fibula, and from the septum between
flewor communis and tibialis posticus. The muscle becomes
tendinous opposite the tuberosity of the os calcis, and passes
into the sole along a groove at the junction of the astragalus
with the sustentaculum tali. Flexor longus hallucis; a larger
muscle than the last, arises from the posterior and internal sur-
faces of the fibula, and from the septum between the two
muscles. It passes through a distinct sheath. Opposite the
ento-cuneiform bone flexor digitorum, which lies superficial to
the other, spreads out, and blends by its deep surface with the
tendon of flexor hallucis. The separate flexor tendons are given
off after this junction. The small tendon to the first digit, and
that. to the fifth, are furnished almost entirely by jlexor digit-
orum, while those to the middle digits proceed from the two
flexors jointly. There is no trace of a short flexor digitorum,
unless the lwmbricales from plantaris represent it. In the flexor
sheaths there is on each side a strong band, which passes to the
second phalanx, and the two bands enclose the tendon, very
much as the perforatus usually encloses the perforans. In the
case of the third digit, a lumbrical muscle is distinctly inserted
into one of these bands.

C. and L.—285-6 (c); 289, fig. 3 (x), 4 (). In this figure the
tendon is not correctly drawn—292-3 (c,). The above refer to
flexor communis digitorum; flexor longus hallucis is shown in
289, fig. 3 (u,), fig. 4 (c); 295-3 (0).

The plantar fascia is exceedingly strong on the outer side of

30 STUDIES IN COMPARATIVE ANATOMY.

the foot, forming a sheath for the short muscles of the fifth
digit. In front, it is firmly connected with a strong transverse
band, which stretches across the sole over the heads of the meta-
tarsal bones. The fascia is not so strong on the inner side of
the foot; here it blends with the other connective tissues of the
sole. Close to the inner side of the tuberosity of the heel, a
rather strong tendinous portion of plantaris passes deeply. The
first dwmbricalis is attached to this; beyond, the tendinous slip
gives off a bundle of fibres to the deep flexor tendon, and itself
ends in a small round tendon, from which the third and fourth
lumbricales take origin.

The general disposition of the superficial ligaments of the sole
is similar to that of the fore-limb. A synovial bursa and a great
quantity of highly elastic connective tissue in each case separates
the ligamentous from the horny sole.

Lumbricales—The first, short and much stronger than the
rest, arises from the tendon of plantaris on its inner side, close
to the tuberosity of the os calcis. It is inserted by a flattened
tendon into the tibial side of the first phalanx of the first toe.
The tendon of plantaris, prolonged towards the middle digit,
gives off next, on the inner side, the second lumbrical, which
passes to ths flexor sheath of the second toe. On the other side
two lumbircales arise, close together, from the tendon of plan-
taris. One passes to the sheath of the flexor tendon of the third
digit, the other similarly to that of the fourth. The /uwmbricalis
of the third digit arises also by a small head from the deep flexor
just above its division. Another lumbrical, given off at the same
point, is inserted into the tibial side of the sheath of the third
digit, and is connected further with the extensor tendon. A
shorter and thicker lumbrical arises from the flexor tendon just
behind the preceding, and passes to the tibial side of the second
digit, where it is attached, like the rest, to the deep surface of
the sheath. There is no dwmbricalis of the fifth digit.

C. and L.—289, figs. 4, 5 (A); 292-3 (x).

Abductor minima digitt (pedis) —A short, thick muscle, arising
from the under surface of the tuberosity of the os calcis, and
inserted by means of a sesamoid into the proximal phalanx of
the fifth digit.

Not in C. and L.

THE ANATOMY OF THE INDIAN ELEPHANT. ob

Adductor minimi digiti—A smaller muscle than the last.
Arises from under surface of os calcis, in front and to the inner side
of the abductor ; also from calcaneo-cuboid ligament. Inserted
into the sesamoid just mentioned to the inner side of the abductor.

Not in C. and L.

Flexor brevis hallucis—A small, short muscle, arising from the
sustentaculum of the os calcis, beneath a strong ligamentous
band which stretches across the centre of the tarsus.

Not in C. and L.

Interossei (pedis)—Only a single plantar interosseus can be
distinguished. It arises from the tarsus, opposite the bases of
the third and fourth metatarsals, and is inserted into the fibular
side of the base of the second digit. The second, third, and
fourth digits are provided with dorsal interossei, almost exactly
resembling those of the manus.

Not in C. and L.

MUSCLES OF THE HEAD AND TRUNK—FACIAL MUSCLES.

Occipito-frontalis—A double-bellied muscle, with intermediate
tendon. The posterior belly, the larger of the two, arises from
the cranial aponeurosis along a line connecting the two auditory
apertures. The anterior belly arises from the supra-orbital
region, partly blending with the orbicularis palpebrarum.

C. and L.—274-5 (1a), anterior belly—“ surciliers.”

Platysma myoides (facial portion of Panniculus).—This muscle,
passing over the angle of the jaw, is inserted as follows :—A few
of the lower fibres pass to the outer surface of the horizontal
ramus of the mandible, a little in front of the facial artery,
where it crosses the bone ; the uppermost fibres to the facial
aponeurosis below the zygoma; the intermediate and principal
portion of the muscle converges to the angle of the mouth,
where its fibres interlace with those of orbicularis oris and
levator anguli oris.

C. and L.—272-3 (2, 4), “ facien ”—“ thoraco-facien.”

Cutaneous muscle of the occiput.—This is described by Dr Wat-
son as arising from the cranial aponeurosis of the posterior sur-
face of the skull. The fibres pass transversely outwards to be
inserted into the cranial aponeurosis immediately behind the

32 STUDIES IN COMPARATIVE ANATOMY.

origin of the posterior belly of the occipito-frontalis. This muscle
was not met with in our dissection.

Not in C. and L.

Zygomatico-labialis arises from the zygoma, and runs parallel
with the upper border of platysma, partly blending with it, to
the angle of the mouth.

Not in C. and L.

Orbicularis oris forms a tolerably distinct bundle, which passes
from the middle line of the lower lip, where it joins the corre-
sponding muscle of the opposite side, to the angle of the mouth.
Here it is partly blended with fibres of platysma and zygomatico-
labialis. The muscle is continued forwards along the margin of the
upper lip,becoming stronger and well-defined as it passes the socket
of the tusk, beyond which it is united with levator proboscidis.

C. and L.—274-5 (7) ; not im the index.

Levator labw superioris arises immediately below the origin of
orbicularis palpebrarum, slightly overlying the origin of levator
proboscidis. It passes almost directly downwards, and blends
with orbicularis oris. Dr Watson gives a similar origin to this
muscle, but describes an anterior portion as inserted into the lip;
while a posterior, weaker bundle is inserted into the facial apo-
neurosis beneath the eye.

C. and L—274-5 (not lettered); 276-7 (2), “grand zygo-
matique.”

Levator anguli oris consists of about six fasciculi, which arise
from the maxilla, below and internal to the infra-orbital foramen.
It passes thence backwards and downwards, and ends by inter-
lacing with orbicularis oris and platysma.

Not in C. and L.

Orbicularis palpebrarum.—This muscle is considerably more
developed below the aperture of the eye than above. It arises
from a tubercle upon the inner margin of the orbit. The fibres
pass downwards and outwards, spreading out into a semicircle
below the eye. The lower fibres extend over the zygomatic
process of the maxilla, becoming indistinct towards the outer
side of the orbit, and finally losing. themselves in the fascia.
Dr Watson describes a more elaborate arrangement of the fibres
than we were able to distinguish in our example.

C. and L.—274-5 (d).

pe)

THE ANATOMY OF THE INDIAN ELEPHANT.

MUSCLES OF THE PROBOSCIS.

The proboscis of the elephant has been fully and accurately
described by Cuvier, but the arrangement of its muscles is in-
cluded in the present account for the sake of completeness.
This organ in the elephant differs from the somewhat similar
snout of certain Insectivora (¢.g., Macroscelides) in its homologies
and internal structure. The imsectivorous snout derives its
form and extent from the great development of the nasal carti-
lage ; whereas the proboscis of the elephant consists wholly of
muscular and membranous tissue, the nasal cartilages simply
serving as valves to the entrance of the bony nares. The muscles
consist of a superficial set, the fibres of which are more or less
longitudinal in direction, and a deep set, which are radiate and
transverse. The longitudinal muscles are further divisible into
anterior, lateral, and posterior.

Levatores proboscidis, or anterior longitudinal, arise from the
frontal bone, along a semicircular line arching upwards above
the nasal bones and between the orbits. The muscles of the
two sides blend, and their fasciculi, running in nearly straight
lines, extend the entire length of the proboscis along its anterior
surface, and are continued into the finger-hke process at its

extremity.
C. and L.—274-5; 276-7; 278, fig. 1; 279, figs. 1, 2, (8);
280 (a). ,

Depressores proboscidis, or posterior longitudinal, arise from the
anterior surface and lower border of the premaxille, and form
two layers of oblique fasciculi along the posterior surface of the
proboscis. The fibres of the superficial set are directed down-
wards and outwards from the middle line; they do not reach the
extremity of the trunk, but disappear by curving over the sides
a littie above the end of the organ. The fibres of the deeper set .
take the reverse direction, and are attached’ to a distinct ten-
dinous raphe along the posterior median line.

C. and L.—274-5 ; 276-7 ; 278, fig. 2; 279, fig. 2 (42); 280 (c).

The lateral longitudinal muscles consist of two portions on
each side; one is derived from orbicularis oris, the other springs
from the anterior margin of the orbit. They are inserted by a

C

o4 STUDIES IN COMPARATIVE ANATOMY.

series of slips between the anterior and posterior longitudinal
muscles along the proximal half of the proboscis.

All these muscles are very intimately connected with the
integument, especially in the lower two-thirds of their extent.

C. and L.— 274-5 ; 276-7; 2179, fig. 2 (5); 280 (0).

The foregoing muscles form a sort of sheath to the deep radiat-
ing and transverse muscles, which, with the nasal canals, make up
the bulk of the proboscis. The deep muscles are best seen in a
cross section of the trunk.

The radiating muscles consist of very numerous distinct fas-
ciculi, stretching directly outwards between the anterior and
lateral surfaces of the nasal canals and the skin.

C. and L.—280 (d).

The transverse muscles are in two sets—one short, between
the nasal canals; the other longer and intermixed with a few
oblique radiating fibres, behind the canals, and between them and
the posterior longitudinal muscles.

C. and L.—280 (e).

MUSCLES OF THE EXTERNAL EAR.

Zygomatico-auricularis (portion of attrahens superior, Wat-
son).—A rather strong muscle. Arises from the outer surface of
the root of the zygoma, and is inserted into the posterior surface
of the cartilage of the ear, immediately behind and below the
meatus.

C. and L.—274-5 (XII), “ parotido-aurien.”

Attrahens swperior (upper portion of attrahens superior,
Watson).—A small quadrilateral muscle. Arises from temporal
fascia and bone anterior to the aperture of the ear, where the
cartilaginous and bony meatus are continuous. Inserted into
the aural cartilage, on the lower and anterior surface of the
tube.

Not in C. and L.

Attrahens inferior arises from the root of the zygoma, close to
the meatus, and beneath attrahens superior. Inserted into a
small projection of the auricular cartilage which overhangs the
meatus.

Not in C. and L.

Tragicus arises from the cartilaginous anterior border of the

THE ANATOMY OF THE INDIAN ELEPHANT. 35

meatus. Inserted into the same process of the aural cartilage
as attrahens inferior.

Not in C. and L.

Cervico-auricularis superficialis (retrahens inferior, Watson).—
Arises from the cervical fascia, near the occipital ridge. In-
serted into the back of the pinna, near the middle of its upper
border.

C. and L.—272-3 (IL, III. 2).

Cervico-auricularis medius (retrahens anterior, Watson).—
Arises from beneath the last-named muscle. Inserted into the
back of the pinna, below the insertion of occipito-auricularis.

C. and L—274-5 (X.?).

Cervico-auricularis profundus (retrahens internus, Watson).—
Arises from the fascia beneath the two last-named muscles.
Inserted into the back of the pinna, opposite the external audi-
tory meatus.

C. and L.—274-5 (XI. ?).

Occipito-auricularis (retrahens swperior, Watson).—Arises from
the cranial aponeurosis, along the occipital ridge. Inserted
into the back of the pinna, beneath the insertion of cervico-
auriularis superficialis.

C. and L—274-5 (IX. ?).

Attollens swperior arises from the cranial aponeurosis above
the ear. Inserted into the back of the pinna, in advance of
cervico-auricularis super ficialis.

C. and L.—272-3 (upper part of I. 2).

Attollens inferior consists of two portions, the posterior of
which crosses the anterior superficially and obliquely. Arises
beneath attollens superior from the cranial aponeurosis, also from
the cervical fascia by the mediation of two slight muscular slips.
Inserted into the pinna, above the cartilaginous auditory
meatus.

C. and L.—272-3 (IV., V.?). There is no close correspondence
between Cuvier’s dissection of the muscles of the external ear
and our own.

MASTICATORY MUSCLES.

Temporalis arises from the temporal fossa, and is inserted into
the inner surface of the anterior half of the ascending ramus

36 STUDIES IN COMPARATIVE ANATOMY.

of the mandible. The muscle is very thick where it passes
beneath the zygoma; above this level are numerous tendinous
septa. The anterior fibres are united to a powerful tendon of
insertion, while the posterior fibres are muscular throughout.

C. and L.—274-5 (6) (temporal fascia); 276-7 (b); 287-8 (0).

Masseter—The principal portion of this muscle arises from
the lower border of the zygoma, extending from below the eye
to the insertion of sterno-maxillaris. The fibres pass down-
wards and backwards, to be inserted into the lower two-thirds
of the ascending ramus of the mandible as far as the angle.
Some of the deeper fibres arise from the posterior third of
the internal surface of the zygoma, pass forwards, and are
inserted into the mandible immediately below the sigmoid
notch.

C. and L.—274-5 (7); 276-7 (7,72).

Pierygoideus externus arises from the upper part of the external
surface of the pterygoid bone. Inserted into the neck of the
mandible and into the inter-articular fibro-cartilage.

C. and L.—287-8 (p).

Pterygoideus internus arises from the outer surface of the
pterygoid bone, below and slightly behind pterygoideus externus.
Inserted into the internal surface of the mandible below the
dental foramen.

C. and L.—287-8 (p1); 290-1 (not lettered).

Mayer speaks of the internal pterygoid as “narrow and
short,” while the external pterygoid is “thick and broad.” The
relative strength was exactly reversed in our specimen. Super-
ficial to temporalis are the temporal fascia, a venous plexus, the
temporal artery, and the temporal gland. Beneath the muscle
is an important venous plexus. The masseter is crossed immedi-
ately below the zygoma by a small artery, a vein (which joins
the facial vein), the trunk of the facial nerve, and the parotid
duct.

Buccinator.—Our notes of this muscle are incomplete. Dr
Watson’s description, which we have verified in its principal
features, 1s as follows :—“ Buccinator muscle arises from the
alveolar margins of both jaws, as well as from a powerful elastic
ligament which extends from the styloid process down to the
lower jaw. The fibres pass forward, and blend with the other

9°

THE ANATOMY OF THE INDIAN ELEPHANT. on

muscles surrounding the opening of the mouth. The muscle is
further strengthened by the addition of two muscular bundles,
which arise from the alveolar margin of the upper jaw, between
the tusk and molar tooth. Of these, the anterior passes down-
wards, and blends with the fibres of the zygomatico-labialis,
whilst the other, passing obliquely backwards, is inserted into
the ligament which gives attachment to the fibres of the bucci-
nator muscle in the interval between the upper and lower jaws.
The muscle is pierced by the duct of the parotid gland.”

ELEVATORS OF THE Hyoip BoNE—DEPRESSORS OF THE
MANDIBLE.

Digastricus arises from the skull behind the articulation of the
stylo-hyal. The superficial portion is inserted directly into the
anele of the mandible; the deeper fibres into the posterior
border of the posterior process of the stylo-hyal, and continued
forwards by an equally strong muscular mass which arises from
the anterior border of the same process, and blends with the
superficial portion.

C. and L.—274-5 (q); 276-7 (q); 287-8 (q); 292-3 (q).

No tendinous intersection of digastricus, such as Mayer and
Watson describe, was found.

Stylo-hyoideus may be represented by the deep fibres of
digastricus.

Mylo-hyoideus arises from the inner border of the mandibular
ramus, between the angle and the symphysis. It is inserted
into the lower border of the greater cornu of the hyoid bone,
and into the tendinous arch which crosses the thyro-hyoid mem-
brane; the anterior fibres blend with the muscle of the other
side.

C. and L—287-8 (7); 292-3 (7).

Genio-hyoideus arises from the posterior surface of the mandiI-
bular symphysis, and is inserted into the upper border of the
body of the hyoid bone.

C. and L.—281, fig. 2 (#).

The posterior part of mylo-hyoideus is covered by sterno-maz-
illaris, Gento-hyoideus is seen when mylo-hyoideus is reflected.
Still deeper lie hyo-glossus minor and genio-glossus. The internal

38 STUDIES IN COMPARATIVE ANATOMY.

jugular vein emerges close to the inner side of digastricus. The
parotid gland lies superficial to this muscle, and the external
carotid artery beneath it.

DEPRESSORS OF THE HyorD BONE.

Sterno-hyoideus wanting.

Sterno-thyroideus arises from the first rib, external to its
sternal articulation, and is inserted into the oblique line of the
thyroid cartilage.

C. and L.—276-7 (a1); 281, figs. 1,2 (21) ; 287-8 (a); 292-3
(@).

Thyro-hyoideus arises from the oblique line of the thyroid
cartilage, and is inserted into the basi-hyoid.

C. and L.—276-7 (y); 281, fig. 2 (y); 287-8 (y); 292-3 (y).

These muscles occupy very much their usual position. Both
are overlaid by sterno-maxillaris, which extends nearly to the
middle line. The two thyro-hyoidei are almost continuous, but
the sterno-thyroidet divide so as to expose the larynx between
their internal edges.

MUSCLES OF THE TONGUE.

Genio-glossus arises from the mandibular symphysis in its
whole depth, and is inserted into the under surface of the
tongue. The muscle has its usual fan-like arrangement, the
lower fibres being horizontal, the upper nearly vertical.

C. and L.—281, fig. 2 (@.

Hyo-glossus major (hyo-glossus lateralis, Watson) arises from
the outer surface of the greater cornu of the hyoid bone. It
passes upwards and forwards to the side of the tongue, where
it blends with the other lingual muscles.

C. and L.—276-7 (v) ; 281, fig. 2 (v?) ; 287-8 (v) ; 292-3 (v),
“ cerato-glossus.”

Hyo-glossus minor (hyo-glossus anterior, Watson) arises from
the anterior edge of the body of the hyoid bone, immediately
adjacent to its fellow muscle, and is inserted into the lateral
margin of the tongue, blending with stylo-glossus.

C. and L.—281, fig. 2 (v), “ hyo-glossus.”

Stylo-glossus arises from the apex of the anterior pointed pro-

THE ANATOMY OF THE INDIAN ELEPHANT. 39

cess of the stylo-hyal bone and stylo-maxillary ligament. Inserted
into the lateral margin of the tongue, partly in common with
hyo-glossus minor.

C. and L.—281, fig. 2 (s?).

Dr Watson has described the peculiar relation of the last
three muscles to the pharyngeal pouch. This cavity, lying in
the arch of the hyoid bone, is embraced around its narrow
outlet by the muscles passing from the hyoid to the tongue.
The stylo-giossus and some of the fibres of the superior constrictor
act as a sphincter; the hyo-glossus minor and genio-hyoid reduce
its vertical dimension; while the depressors of the hyoid bone
enlarge the cavity of the pouch.

When the genio-hyoid is divided, the genio-glossus is seen to
form a very narrow ellipse, extending from the mandibular
symphysis to the body of the hyoid. In front, the sub-lingual
glands lie between and beneath the two muscles. The posterior
extremity of this pointed ellipse is embraced by the diverging
hyo-glossi_ minores, which pass forwards on each side of the
tongue, and are joined on their outer sides by the stylo-glossv.
The small interval between the straight portions of the genio-
glossus and hyo-glossus minor is occupied by a large branch of
the hypoglossal nerve and the lingual artery. The nerve 1s
innermost, and runs superficial to (i.¢., beneath) hyo-glossus minor,
while the artery is partly concealed by it. Branches of the
gustatory nerve enter the same space, and form loops with the
hypoglossal. External to the stylo-glossus lies a large vein.
The stylo-glossus is, during part of its course, in contact with the
mucous membrane:of the mouth.

ABDOMINAL MUSCLES. °

Panniculus (abdominal portion).—The muscle forms an ex-
tensive sheet upon the sides of the body, increasing in thickness
below, where it is continued into the fascize of the fore and hind
limbs. A tolerably distinct scapular portion may be traced.

C. and L.—272-3 (6d, 5c).

Rectus abdominis arises by a broad thin tendon from the pos-
terior border of the inferior surface of the pubic symphysis. An
expansion of the tendon is continued forwards, and closely

40 STUDIES IN COMPARATIVE ANATOMY.

attached to the symphysis in its lower three-fourths, as well
as to the adjacent rami of the pubes. The two tendinous layers
form thus a kind of flattened pouch, communicating with the
abdominal cavity, and lined with the peritoneal membrane. The
free part of the tendon adheres to the inner or superior surface
of the thick abdominal aponeurosis. The muscle expands as it
passes forward, and again contracts towards its insertion into
the sternum. Eight tendinous intersections occur.

C. and L.—287-8 (15).

Obliquus externus takes origin from all the ribs, at least as far
forward as the third. The fibres pass downwards, and are inserted
into the abdominal aponeurosis. About six interdigitations with
serratus magnus may be made out.

C. and L.—274-5 (13); 276-7 (13); 282, upper figure (13).

Obliquus internus is less extensive than externus. The fibres
converge to a thin aponeurosis continuous with the sheath of
vectus, and closely attached to the outer surfaces of the cartilages
of the ribs.

C. and L.—276-7 (14); 287-8 (14).

Transversalis abdominis arises by interdigitations with the
diaphragm, from the ribs between the second or third and the
seventeenth (inclusive); also from the eighteenth, nineteenth,
and twentieth ribs. The ventral portion takes origin as a thick
fleshy mass from the anterior superior spine of the ilium, and
from a small part of Poupart’s ligament adjacent thereto. The
muscle passes thence horizontally inwards towards the middie
line, and is lost upon the inner surface of the abdominal fascia.

C. and L.— 287-8 (17).

The Diaphragm arises by fleshy fibres from all the ribs from
the second or third to the seventeenth. As far back as the
thirteenth rib, the muscle is chiefly attached to the costal
cartilages; behind the thirteenth to the ribs themselves. The
crura arise from the three hindmost dorsal vertebree by tendinous
processes. The tendon of the left crus is somewhat shorter than
the cther. A ligamentum arcuatum, entirely tendinous, and
united to the central tendon, arches over psoas.

Not in C. and L.

Quadratus lumborum arises from the two hindmost ribs, and
passes to the innermost part of the crest of the ilium.

THE ANATOMY OF THE INDIAN ELEPHANT. Al

Not in C. and L.

The Jntercostal muscles present the usual arrangement, but
are continued unusually far downwards between the costal
cartilages.

SUPERFICIAL CERVICAL MUSCLES.

Sterno-maxillaris arises from the first rib, adjacent to the
sternum. Itforms a strap-like band of nearly uniform breadth
(about two inches), but expands greatly, and becomes aponeurotic
at its insertion into the lower jaw, at and in front of the angle.

C. and L—274-5 (x); 276-7 (a); 287-8 (+6); 292-3
(+ 2).

Sterno-mastoideus. —A muscle which may be held to correspond
with this, though it has no attachment either to the sternum or
the mastoid, springs from the first rib, several inches above
sterno-maaillaris (1.e., nearer the vertebral column). It passes
obliquely forwards and outwards, contracting to a thin tendon,
which finally expands upon the root of the zygoma.

©. and L.—274-5 (6); 276-7 (6); 287-8 (6); 290-1 ();
292-3 (0).

MUSCLES OF THE BACK.

Trapezius arises from the ligamentum nuche and vertebral
aponeurosis between the third and fourteenth dorsal vertebre.
It is inserted into the posterior surface of the spine of the
scapula and the superior border of the unciform process.

C. and L.—274-5 (a, a”); 276-7 (a1, a”); 287-8 (a, a}, a’).
An occipital origin is ascribed to this muscle in the Plates.

Latissinvus dorsi arises from the vertebral aponeurosis and the
ribs from the ninth to the fourteenth. The fibres become twisted,
and converge to a tendon which is inserted into the neck of the
humerus. The tendon of insertion is divided; the superficial
portion is inserted above teres major, the deeper and smaller part
behind it.

C. and L—274-5 (2); 276-7 (1); 283, figs. 1, 2 (2) ; 287-8 (2).
Represented as arising from the sixth to the twelfth ribs.

fhomboideus major arises from the ligamentum nuche be-
tween the fourth and six dorsal vertebre, and is inserted into
the postero-superior border of the scapula.

49 STUDIES IN COMPARATIVE ANATOMY.

Rhomboideus minor.—Under this name may be distinguished
a small slip tolerably distinct from the pieceding, which passes
from the ligamentum nuche opposite the seventh cervical ver-
tebra to the posterior angle of the scapula.

C. and L.—(Rhomboidet) 276-7 (c?) ; 283, figs. 1, 2 (c?); 287-8
(2),

Serratus posticus arises from the ligamentum nuche and the
vertebral aponeurosis between the third and tenth dorsal ver-
tebree. It is inserted into the anterior margins of the ribs from
the ninth to the fourteenth inclusive, and also to a strong fascia
with much elastic tissue which underlies the serratc. The pos-
terior fasciculus of the muscle is separable from the rest.

C. and L.—282, upper figure (10); 287-8 (10).

Levator anguli scapule is directed obliquely across the lower
part of the side of the neck. Its most forward attachment 1s to
the front of the transverse process of the atlas by means of a
thin, flat, tendinous slip. This portion of the muscle crosses
superficially the insertion of transversalis colli. The lower por-
tion, which is separated by an interval from the rest, arises by
similar slips from the posterior tubercles of the transverse pro-
cesses of the five lower cervical vertebre. This part of the
origin is overlaid by ¢ransversalis colli. In the dorsal region the
muscle takes origin from the articular processes and angles of
the ribs between the first and fifth dorsal vertebrae. Longissimus
dorsi separates these two origins. The muscle passes down-
wards, and in part backwards, to its insertion into the anterior
border of the scapula.

C. and L—274-5 (d) ; 276-7 (d, d'); 283, fig. 1 (d, should be
d'); 287-8 (d, a).

Splenius—This muscle consists of two superimposed layers.
The superficial layer, which is thinner and narrower than the
other, has a tendinous origin from the ligamentum nuche in the
anterior dorsal region. The tendon narrows as it passes forward,
and reaches the level of the third cervical vertebra. Muscular
fibres are given off from it on both sides, but the internal fibres
much preponderate, especially in the dorsal and posterior cer-
vical region. Towards the occiput the fibres radiate equally on
both sides from the fore end of the tendon. The muscle has an
extensive insertion, first, to a median ligamentous raphe con-

THE ANATOMY OF THE INDIAN ELEPHANT. 43

tinuous with the ligamentum nuche; secondly, to the inner
two-thirds of the superior curved line of the occiput. The deep
layer has a similar arrangement, but its breadth is greater, and
the tendinous centre much larger in proportion to the muscular
part. It is inserted into the whole length of the superior curved
line.

C. and L.—276-7 (K); 282, upper figure (K) ; 287-8 (K).

Sacro-lumbalis and its forward continuations form a thin, flat
sheet, which extends as far as the fourth rib. The tendons arise
primarily from the ilium; subsequently from the ribs between
the sixth and nineteenth (inclusive), a little external to the
angles. The tendons of insertion are somewhat narrower than
the tendons of origin, and pass to the posterior borders of the
ribs between the fourth and sixteenth (inclusive), nearly four
inches external to their angles.

No separate erector spine exists.

C. and L.—282, figs. 1 and 2 (C); 295 (C).

Longissimus dorsi 1s inseparable behind from spinalis. It
diverges from that muscle about the level of the twelfth rib, and
is continued forward to the neck, having the continuation of
sacro-lumbalis on its outer, and semi-spinalis on its inner side.
In the anterior dorsal region longissimus dorsi is enclosed be-
tween the two origins of levator anguli scapulw. It is inserted
along its whole length by a series of tendons into the transverse
processes of the vertebre. The tendons of origin arise similarly
from the transverse processes, but a little to the inner side.

C. and L—282, figs. 1 and 2 (B); 295 (B).

Transversalis colli continues longissimus dorsi forward as far
as the atlas, arising from articular processes, and inserted into
transverse processes.

Not in C. and L.

Trachelo-mastordeus a¥ises by tendinous and fleshy slips from
the articular processes of the hinder cervical vertebre, external
to the slips of origin of complexus. The muscle forms a flat-
tened band, which passes direct to the mastoid process, and is
inserted by tendinous fibres on its superficial aspect, and more
deeply by fleshy fibres. The insertion is overlaid by splenius.
The internal or dorsal border of the muscle is attached to, and
partly sheathed in a layer of yellow elastic tissue, which
separates splenivus from complexus.

44 STUDIES IN COMPARATIVE ANATOMY.

C. and L.—282, upper figure (L’), the origin is incorrect in the
lower figure ; 287-8 (L}).

Spinalis dorsi and multifidus spine form a continuous muscle,
lying internal to longissimus. The fibres take origin in the first
place from the sacrum and adjacent parts of the ilium, from the
under surface of the strong aponeurosis, which extends between
longissimus dorsi and the spinous processes, and also from the
vertebre themselves along the line of the articular processes.
The muscle is inserted by fleshy fibres into the sides of the spines
of the vertebree, as far forward as the fourth dorsal. The slips
corresponding to the fourth, fifth, sixth, and seventh dorsal spines
are tendinous, and form a strong bundle which passes more or
less completely into the ligamentum nuche, and into a loop of
elastic tissue, which extends between the spines of the second
and fourth dorsal vertebrae. The muscle ends ina tendon which
arches over the spines of two vertebree, and is inserted into the
side of the spine of the first dorsal.

C. and L.—282, figs. 1 and 2 (A); 295 (A).

Complexus.—Beneath splenius a thin layer of yellow elastic
tissue is found to enclose a subjacent muscle, and to extend
forwards and outwards from the hgamentum nuche to the
occiput. The muscle thus enclosed is complexus. It arises from
the articular processes of the four hindermost cervical, and about
twelve foremost dorsal vertebree, sightly also from the lLgamen-
tum nuche in the anterior cervical region. It overlies the
funicular portion of the ligamentum nuche, and is inserted into
the inner half of the superior curved line beneath splenius.

C. and L.—282, figs 1 and 2 (L); 287-8 (L); 295 (L).

Ligamentum nuche.—tThis powerful elastic ligament springs
mainly from the spines of the three foremost dorsal vertebre,
which rise successively higher in the order of their numbers.
The ligament is also strengthened by the bundle derived from
spinalis dorsi. Behind the third dorsal vertebree it passes into
the interspinous ligament. At the level of the first dorsal
vertebra the ligamentum nuche consists of two very distinct
masses, one superficial and the other deep, while a less con-
spicuous median partition may be observed. The upper or funi-
cular portion is free from bony attachment between the spine of
the seventh cervical vertebra and the occiput. At its insertion

THE ANATOMY OF THE INDIAN ELEPHANT. Ad

it divides into lateral halves separated by several veins and loose
cellular tissue; each of these again breaks up into many distinct
fasciculi. On the deeper surface of the funicular portion the
yellow elastic is largely replaced by white fibrous tissue. The
deep portion of the ligament forms a flattened mass attached to
the spines of the six hinder cervical vertebre.

Opposite the third dorsal vertebra a short, thick bundle of
fibres is given off, partly from the ligamentum nuche, partly
from the spinous processes, and partly from the interspinous
ligament. This bundle passes to the apex of the scapula, which
is thus strongly connected with the vertebral column.

Semispinalis colli arises in the posterior cervical and anterior
dorsal region by muscular slips internal to the origin of complexus,
and is inserted into the lower border of the spine of the axis.

C. and L.—282, lower figure (L' ?) “ petit complexus.”

Rectus (capitis) posticus major arises by a small round slip from
the spinous process of the axis, and is inserted into the occiput,
external to the hgamentum nuche and beneath complexus.

C. and L—282 (M? ?).

Rectus posticus minor arises from the superior arch of the atlas
close to its fellow muscle. The two diverge as they pass up-
wards, and each is inserted into the occiput beneath rectus posticus
major.

Not identified in C. and L.

Rectus posticus externus, a rather broader muscle than rectus
posticus major, arises in front of itand more deeply. It is inserted
into the occiput, external to rectus posticus minor, and more
superficially. Obliquus superior partly overlies it.

Not identified in C. and L.

Obliquus (capitis) superior, a fan-shaped muscle, arises by two
tendinous slips from the transverse process of the atlas. It
passes upwards and slightly inwards, and is inserted into the
occiput, external to rectus posticus major. A considerable nerve,
an artery, and two or three veins pass between the heads of
origin.

Not identified in C. and L.

Obliquus inferior, a short and stout muscle, arises from the
outer surface of the spinous process of the axis, and passes direct
to the transverse process of the atlas. Towards its inner border

46 STUDIES IN COMPARATIVE ANATOMY.

the fibres are rather closely connected with those of obliquus
superior.

Not identified in C. and L.

fectus capitis lateralis extends from the transverse process of
the atlas to the mastoid process, lying along the anterior border
of trachelo-mastoideus.

Not in C. and L.

Intertransversales are present, stretching as usual between
transverse processes. The longest lie between the atlas and axis.

MUSCLES OF THE TAIL.

Cuvier and Laurillard figure an ischio-caudalis (274-5, Q;
282, lower figure, Q), a sacro-coccygeus superior (274-5, R; 282,
lower figure, R) and a sacro-coccygeus inferior (274-5, R14). The
names indicate the bony attachments.

ANTERIOR VERTEBRAL MUSCLES.

Rectus capitis anticus major arises from a prominent tubercle
upon the sixth cervical vertebra, and is inserted mainly into the
ventral surfaces of the atlas and axis, a small slip passing to the
third vertebra.

C. and L.—287-8 (M) ; 290-1 (+6 2).

Rectus capitis anticus mimor is a small muscle internal to the
last, which arises by thin slips from the third, fourth, and fifth
cervical vertebre. It passes upwards and forwards, converging
towards its fellow, which it meets close to its insertion into the
base of the basilar process.

C. and L.—290-1 (not lettered).

Longus colli is a much stronger muscle, lying symmetrically
upon each side of the middle line and internal to the last. The
muscles of both sides form together a triangular mass in front of
the centre of the cervical vertebrae. They arise from the trans-
verse processes of the third, fourth, and fifth cervical vertebre ;
also from the front of the bodies of the third, fourth, fifth, and
sixth dorsal vertebre, and from the heads of the corresponding
ribs, and are inserted into the tubercle of the atlas.

C. and L—290-1 (0).

THE ANATOMY OF THE INDIAN ELEPHANT. AT

Scalenus anticus arises from the anterior surface of the tubercles
of the second, third, and fourth cervical vertebree by small,
rounded tendons. These unite, and pass towards the thorax,
being inserted into the second, third, and fourth ribs.

C. and L.—287-8 ; 290-1 (6, 6a, 66).

In dealing with the myology of the Elephant, we thought it advis-
able, considering the want of any tolerably complete description, to
note every important detail which came to light in the course of our
dissection. It would be superfluous to treat all parts of the anatomy
with the same fulness. The osteology, for example, has long been
amply made known, nor would any student value minute descriptions
of bones which can be so easily seen and handled. Other parts of
the anatomy are known in various degrees of completeness; some
thoroughly, some superficially, some hardly at all. Vulpian and
Philipeaux have published a lengthy and elaborate description of
the heart; Dr Morrison Watson has minutely described the male
organs of generation, and other important viscera ; the brain has been
figured more than, once; while scattered memoirs contain particulars
of greater or less value respecting other organs. Under these circum-
stances we shall probably employ our space and the reader’s time to the
best advantage by a summary of what is already known, corrected
and supplemented by our own observations. Although such anato-
mists as Cuvier, Camper, and Hunter have preceded us, there is still
much to be gleaned, more than any single exploration of the field is at
all likely to discover.

The osteology and dentition we propose to leave out altogether.
Common text-books already contain descriptions sufficient for the
naturalist or paleontologist.

ALIMENTARY CANAL AND ITS APPENDAGES.

Mours.

The gape of the mouth is small relatively to the bulk of the
animal,—a fact which may receive explanation from the precision
with which food is passed into the mouth by the proboscis, and
the small part which the lips consequently play in the act of
prehension. The upper lip has hardly any separate develop-
ment, but seems a mere lateral expansion of the root of the
trunk; the lower lip is small and pointed, its mucous surface
forming a narrow groove or gutter in which the tongue is lodged.
The cheeks are very capable of distention, but in their ordinary
contracted state they enclose only a very small cavity. -

48 STUDIES IN COMPARATIVE ANATOMY.

The mucous membrane of the mouth is in general smooth ;
that of the palate in particular is quite smooth, and shows none
of the transverse ridges which appear on the hard palate of the
horse and ox. Two shallow triangular depressions placed sym-
metrically immediately in front of the fore edge of the hard
palate lead to Jacobson’s canals. The bony cavity in which the
canals le runs in the suture between the premaxille and
maxille for 7 or 8 inches (Mayer gives 8 inches), taking a
nearly straight direction upwards and backwards ; it is furnished
with a vertical cartilaginous septum and a cartilaginous lining.
Each of the component canals is of about the diameter of a goose-
quill, and is lined by an extension of the mucous membrane of
the mouth. Camper (p. 48) says that these canals when pressed
exude a sticky fluid.

Examination of skulls of different ages seems to show that
Jacobson’s canals are at first nearly horizontal, but that as the
air-cells of the maxilla enlarge, the maxillo-premaxillary suture,
and with it the canals, is tilted more and more, until it
finally gets a steep slope forwards.

TONGUE.

The tongue is thick and rounded towards its base, tapering and
pointed in front. Perrault describes it as 18 inches long,
but in our young example it was much shorter. The tip is
directed downwards, and lies almost invariably in the groove
formed by the lower lip. All observation of the living animal
seems to show that the tongue, like the lips, is of little import-
ance in the act of feeding. The oral surface of the tongue ends
behind in a prominent concave edge, which forms the front and
lower boundary of the pharyngeal pouch. Towards the base are
two, four, or more circumvallate papille of large size, while on
the side, especially behind, are a number of wart-like eminences
and mucous crypts.

SALIVARY GLANDS.

The parotid was small in our example, and measured only
4 inches by 3. Dr Watson gives 8 inches by 5 as the dimen-
sions. The gland is connected by a fibrous band with the

THE ANATOMY OF THE INDIAN ELEPHANT. 49

zygoma; it lies in the space below the zygoma and behind the
ascending ramus of the lower jaw. Steno’s duct passes out from
near the middle of the gland; it is at first of about half an inch
diameter, but gradually contracts to the size of a crow-quill.
Close to its termination it pierces the buccinator, and finally
opens into the mouth near the alveolar margin of the upper jaw
by a simple rounded orifice without papilla.

No sub-maxillary gland was seen either by Dr Watson or our-
selves, though it is described by Mayer. A small lobulated mass,
lying between the genio-hyo-glosst and close to the symphysis of
the mandible, may possibly be a sublingual gland, but we were
unable to discover an efferent duct, or to satisfy ourselves as to
the exact nature of the body.

SoFt PALATE.

The soft palate, which is hardly distinguishable from the base
of the velum palati, extends backwards about 2 inches from
the hind edge of the hard palate. Like the nasal passages
immediately above, it is narrow from side to side. No Jevator
palati was made out. The tensor palate) arises from the front
and outer side of the upper part of the membranous section of
the Eustachian tube. It is a small, spindle-shaped muscle en-
closed in a sheath of fascia. At the groove of the hamular
process 1t becomes tendinous and spreads out in the substance
of the soft palate.

PHARYNX.

The upward extension of the pharynx towards the nasal
passages gradually narrows from side to side as it ascends. The
antero-posterior dimension is much contracted in the neighbour-
hood of the soft palate. Above this level, the pharynx is pro-
longed into the nares in front, and is also continued for some
distance backwards as a gradually diminishing cavity which
extends beneath the basi-sphenoid to near its junction with the
basi-occipital. The ultimate recess (the sinus of Morgagni) just
admits the last joint of the forefinger.

' Dr Watson considers that this muscle is absent in the elephant.
: D

50 STUDIES IN COMPARATIVE ANATOMY.

The pharyngeal openings of the Eustachian tubes lie in the
lateral walls of this part of the pharynx, about an inch above
the hamular pterygoid process, and a little above the level of the
hard palate. The orifices are large enough to admit the little
finger. In its lower part each tube is almost entirely membran-
ous. It runs upwards and a little outwards, and may be
explored by a probe for 7 inches. We have been unwilling
to destroy the surrounding parts for the sake of tracing the tube
further. According to Camper’s figures (pl. xiu. figs. 7, 8), the
bony tube is about 2 inches long and the cartilaginous tube
over an inch. The same author describes and figures an open-
ing from the upper part of the cartilaginous tube into the
nasal passage. We find no corresponding opening in the soft
parts.

The common aperture of the posterior nares occupies a
triangular space 44 inches high and 2 inches wide. The hind
edge of the septum is very thin, and deeply concave above.

Muscles of the Pharyne.

Constrictor pharyngis arises on each side from the thyro-hyal,
from the posterior margin of the thyroid cartilage, and from the
cricoid cartilage below the arytznoid facet. The fibres pass
round the tube of the pharynx, and blend along the middle line
behind. The uppermost (or anterior) fibres form a tolerably
distinct bundle. The lower fibres curve upwards so as to leave
a triangular gap, which is filled by a tapering median bundle of
the longitudinal cesophageal fibres.

C. and L.—261, fig. 1 (s!, a, 2’).

Stylo-~pharyngeus arises from the internal surface of the
anterior branch of the stylo-hyal, close to its origm. The
muscle passes downwards along the side of the pharynx and is
there inserted.

C. and L.—261, fig. 1 (v).

Palato-pharyngeus is largely developed. It arises from the
palate and descends to the pharynx, forming a considerable part
both of the soft palate and the velum palati. It is inserted
laterally on the inner surface of the pharyngeal wall.

A symmetrical venous plexus, which arises by free communi-
cations between the internal jugular, internal maxillary, and

THE ANATOMY OF THE INDIAN ELEPHANT. 51
inferior palatine veins, lies at the back of the mouth, below the
soft palate.

The entrance to the pharynx is bounded above by the antero-
inferior edge of the velum palati (where its descending and
horizontal portions meet) ; in front and below by the sharp, back-
ward directed edge of the dorsum of the tongue; and laterally
by the mucous membrane of the pharynx, with which are con-
nected a number of scattered muscular fibres, and the yellow
elastic pharyngeal wall. There is no palato-glossus. The passage
is very narrow, and cannot in our young example be distended
to admit a cylinder of 2 inches diameter.

The velum palati descends from the soft palate, but its chief
extension, as in many other large quadrupeds, is horizontally
backward. The free posterior edge passes so far back as slightly
to overlap the epiglottis. On each side it is continued into a
thin elastic fold, which is obliquely attached to the pharyngeal
wall, sloping backwards. In the natural position of the parts,
the posterior edge of the velum encloses an oval space, the longi-
tudinal diameter being an inch and a half, the transverse some-
what less. Through this aperture the arytenoid cartilages project.
The lower ends of the lateral bands of the velum are approxi-
mated, but they do not meet; and Dr Watson, therefore, in
speaking of the “central” aperture of the soft palate, is to be
understood as meaning “in the middle line.” The position of
the aperture is anatomically the same as in the human subject
or as in the majority of mammalia, and altogether below the
velum. There is no uvula, but we find a small vertical muscle
in the middle line, passing from the soft palate, to be inserted
into the back of the upper portion of the velum. This may
represent an azygos wvule; its length does not exceed 2
inches.

The muscular layer of the velum palati forms part of the
palato-pharyngeus. Its anterior fibres, arising from the palate,
pass backwards and a little downwards, to be inserted into the
inner surface of the elastic wall of the pharynx, near the free tip
of the thyro-hyal. The posterior fibres gradually take a more
and more transverse direction, and form the thin muscular sheet
which lies in the horizontal valve of the velum. The lateral
bands of attachment contain muscular bundles from the same

5), STUDIES IN COMPARATIVE ANATOMY.

stratum. These bands are separated from the insertion of the
anterior fibres of the palato-pharyngeus by a considerable elastic
pouch, opening backwards into the pharynx, and lined by a con-
tinuation of the mucous membrane of that cavity.

Immediately in front of the epiglottis is a vertical transverse
elastic fold, which projects from the floor, and especially from
the sides, of the pharynx, but not so far as to materially obstruct
the passage. Smaller secondary folds proceed backwards from
this at right angles, passing longitudinally upon the floor of the
pharynx outside the laryngeal aperture. When the pharynx is
seen from above and behind, there are therefore visible three
cavities on each side, divided from each other by mucous folds.
In front of the epiglottis, behind the transverse fold, and inside
the longitudinal secondary fold just described, is a small recess,
- with a small gland in its floor. To the outer side of this is a
larger cavity, in which a larger gland can be seen; it is included
between the transverse fold, the secondary longitudinal fold, and
the lateral aitachment of the velum. Between the lateral attach-
ment of the velum and the proper pharyngeal wall is a third and
much more capacious receptacle, which has been described in
connection with the palato-pharyngeus.

The wall of the pharynx is largely composed of yellow elastic
tissue, overlaid by the constrictor muscles. The palato-pharyn-
geus lies deeper, and is inserted into the inner surface of the
elastic wall.

Dr Watson has described, in connection with the pharynx, a
peculiar structure which throws light on a previously un-
explained faculty of the elephant, viz., the power which the
animal possesses, according to more than one competent observer,
of withdrawing large quantities of water by inserting the tip of
the trunk into the mouth. He observes that

“Tt is evident that were the throat of this animal similar to that of
other mammals, this [withdrawal by the trunk] could not be accom-
plished, as the insertion of a body, such as the trunk, so far into the
pharynx as to enable the constrictor muscles of that organ to grasp it,
would at one give rise to a paroxysm of coughing, or were the trunk
merely inserted into the mouth, it would be requisite that this cavity
be kept constantly filled with water at the same time that the lips
closely encircled the inserted trunk. The formation of the mouth of
the elephant, however, is such as to prevent the trunk ever being

\

THE ANATOMY OF THE INDIAN ELEPHANT. No

erasped by the lips so as effectually to stop the entrance of air into
the cavity, and thus at once, if I may so express it, the pump-action
of the trunk is completely paralysed.
We find, therefore, that it is to some
modification of the throat that we must
look for an explanation of the function
in question, and this we find to be as
follows :—The superior aperture of the
pharynx (fig. 1, A) is extremely nar-
row, so much so as to admit with
difficulty of the passage of the closed
fist. Immediately posterior to this
narrow aperture the pharynx dilates
into a pouch of large size (fig. 2, E),
capable of containing a considerable
quantity of fluid. This pouch is pro-
longed forward beneath the root of
the tongue, and is bounded in the
following manner. The floor extends
from the epiglottis as far forward as
the root of the tongue, being formed
from behind forward by the thyroid
cartilage, thyro-hyoid membrane, and
hyoid bone. Its lateral walls are com-
pleted by the sides of the pharynx
(that is, by the superior constrictor
muscles—fig. 2, F), in addition to the
stylo- (G), and hyo-glossi (H) muscles,
The root of the tongue forms the anterior
boundary, whilst the posterior wall is
completed by depression of the soft
palate, or when the latter is elevated,
the pouch then communicates freely Fic. 1.—A, superior aperture of
with the cesophagus. In connection —Phatynx. _B, root of tongue.
: 3 . ; C, soft palate with larynx pro-
with this pouch is to be observed jecting through the centre. D,
the very peculiar form of the hyoid pharynx.
bone, which, being deeply concave
on its upper surface, forms as it were the greater part of the floor of
this pouch. Between the pouch and the concavity of the hyoid bone,
moreover, there is placed a large quantity of loose and distensible con-
nective tissue, which permits of the expansion of the pouch. The
size of the latter is, moreover, liable to alteration by the actions of
several muscles. These are more especially the hyo-glossi muscles,
and two little additional muscles (fig. 2, K), the homologies of which I
have not yet been able to determine, which, springing from the middle
line of the hyoid bone, in front of the pouch, pass up, one on either
side of the middle line, and blend with the other muscles forming the
root of the tongue. By the action of these muscles the pouch may be
diminished in depth ; but in consequence of the narrow interval exist-
ing between the hyoid cornua, the length of the pouch from before

5A STUDIES IN COMPARATIVE ANATOMY.

backwards cannot be altered, as the thyroid cartilage is thereby pre-
vented from being approximated to the hyoid bone. . . . An elephant
can, as the quotations sufficiently prove, withdraw water from his

Fie. 2.—E, E, pharyngeal pouch. F, superior constrictor. G, stylo-glossus.
H, hyo-glossus. K, small muscle, which diminishes the depth of the
pharyngeal pouch. L, genio-glossus muscle. (For the use of the wood-
cuts of figs. 1 and 2 we are indebted to Professor Turner.)

stomach in two ways: first, it may be regurgitated directly into the
nasal passages! by the action of the diaphragm and abdominal muscles,
the soft palate being at the same time depressed, so as to prevent the
entrance of water into the mouth. Having in this manner filled the
large nasal passages communicating with the trunk, the water contained
in them is then forced through the trunk by means of a powerful
expiration ; or, in the second place, the water may be withdrawn from
the cavity of the mouth by means of the trunk inserted into it. Now,
in this case, it is manifestly impossible that the water can be contained
within the cavity of the mouth itself, as I have already shown that
the lips in the elephant are so formed as effectually to prevent this.
The water regurgitated is, however, by means of the elevation of the
soft palate, forced into the pharyngeal pouch. ‘The superior aperture
of this pouch being much narrower than the diameter of the pouch
itself, and being completely surrounded by the muscular fibres of the
stylo-glossus on each side, and the root of the tongue in front, which
is prolonged backwards so as to form a free sharp margin, we have
thus as it were a narrow aperture surrounded by a sphincter muscle,
into which the trunk being inserted and grasped above its dilated
extremity by the sphincter arrangement just referred to, air is thus
effectually excluded, and the nasal passages being then exhausted by
the act of inspiration, water is lodged within these passages to be used
as the animal thinks fit, either by throwing it over his body, or again
returning it into his mouth.”

1 For remarks on this passage see p. 80.

THE ANATOMY OF THE INDIAN ELEPHANT. 5D

The chief difficulty attending this explanation springs from
the small size of this pharyngeal pouch. In our example, which,
it must be remarked, was by no means of full stature, the pouch
could not be distended so as to hold a pint of water. This
objection is not fatal to the hypothesis advanced by Dr Watson.*
Regurgitation from the stomach may be effected slowly and
continuously until the requisite supply is yielded. It seems to
us that the pharyngeal pouch must be unimportant as a mere
reservoir of fluid, but as a water-tight circular valve it may be
essential to the process of withdrawal. Behind the velum
palati is a somewhat larger cavity, but the entrance to the wind-
pipe lies in its floor, and this is not therefore a very likely re-
ceptacle of fluid. If our examination of an immature elephant
yields data in the least trustworthy, it is hard to suppose that
even in the adult a gallon of water can be retained anywhere
between the stomach and the proboscis.

We do not know enough of the habits of the living animal to
say whether or not the food is moulded and lubricated into a
bolus, but the form, structure, and glandular surface of the
pharyngeal pouch would be well adapted to such a practice.2

‘Dr Watson informs us that he considers the pharyngeal pouch unimportant as
a mere reservoir, ‘‘ though the presence of certain muscles not found in other
animals would appear to indicate that the elephant possesses a certain power of
increasing or diminishing the size of the pouch, and the necessity for this is by
no means evident upon the supposition that the sphincter arrangement is the only
raison d étre of this pouch.”

2 Professor Owen describes the back of the mouth of the Camel in these words:
—‘‘ A broad pendulous flap hangs down from the fore part of the soft palate, and
usually rests upon the dorsum of the tongue. The velum palati extends beyond
this process some way down the pharynx, and terminates by a concave border.
The pharynx behind the velum dilates into a sac. In the rutting male the
palatal flap is greatly enlarged. I have found it extending 10 inches down the
pharynx, passing below the margin of the soft palate and the opening of the
larynx, into the esophagus: in the living animal it is, at this season, occasionally
protruded, with a belching noise, from the mouth. Its surface shows the pores
of innumerable mucous crypts, and in the ordinary state, in both sexes, the flap
may apply its own secretion, and water regurgitated from the storage cells of the
stomach to the extended surface of the pharynx and root of the tongue so as to
allay the feeling of thirst and help the animal to endure the long remissions of
drinking to which it is liable in traversing the desert” (Comp. Anatomy of Ver-
tebrates, vol. iii. p. 395). In transcribing these remarks, we desire to offer no
opinion as to the validity of the explanation offered. The pharyngeal sac
described in the camel may throw light on the similar structure in the ele-
phant.

56 STUDIES IN COMPARATIVE ANATOMY,

A number of pits in the mucous membrane of the floor and
sides of the pharynx are doubtless glandular. They are altogether
absent from the middle line, but become densely aggregated
towards the root of the tongue on the sides of the pharyngeal
floor. Similar glandular crypts line a pouch, in our example
about an inch deep and half an inch in diameter! which les on
each side in the lateral wall of the pharynx in front of the
transverse fold described above. In the lateral spaces behind
the transverse fold are other flattened glands.

The thyro-hyals support the pharyngeal wall laterally, and
their expanded ends can be plainly felt upon its inner surface, in
the recess behind and within the lateral attachment of the velum.
Professor A. H. Garrod has remarked? that the basi- and thyro-
hyals form a lower arch quite distinct from the bifurcate stylo-
hyals, and he adds: “In the elephant, therefore, the deficiency
of the lateral intermediate elements of the hyoid apparatus per-
mits of a much greater movement of the base of the tongue than
in the ungulata, whose nearly rigid stylo-hyals, epi-hyals and
cerato-hyals can allow of little more than an antero-posterior
movement of the base of the tongue in part of the circle of which
the hyo-cranial attachment in the centre.”

(ESOPHAGUS.

Dr Watson states that “the muscular fibres of the cesophagus
are distinctly striated even down to the cesophageal opening in
the diaphragm, and are arranged in two layers—an external, the
fibres of which are distinctly longitudinal in direction ; and an
internal, which consists of two sets of spiral fibres, one of which
passes from right to left, whilst the other passes in the opposite
direction, and thus gives rise to a decussation of the fibres at ail
points.”

Like Dr Watson, we found no trace of a muscle connecting
the trachea with the cesophagus and stomach, such as was
described and figured by Dr Harrison of Dublin.’

* Mayer found this pouch to be 34 inches long and 14 wide.
2 Proc. Zool. Soc. May 1875.
3 Proc. Roy. Irish Acad. vol. iv. p. 133 (1847).

&

THE ANATOMY OF THE INDIAN ELEPHANT.

STOMACH.

An excellent figure of this viscus is given by Camper (pl. ix.
fig. 1).. Mayer’s drawing is less satisfactory.

The stomach is smooth externally, elongate, and nearly
straight. The cardiac end is much prolonged and tapering.
A number of transverse nearly circular folds project inwards
from the cardiac wall; they almost disappear when the stomach
is greatly distended, and are at all times too shallow to serve as
water-cells, though they have been figured and described as
such.!' The gastric follicles are most abundant towards the
cardiac end, as Mayer has observed. In an adult elephant the
stomach is little less than three feet long ; the cesophagus enters
near the middle but rather nearer the cardiac than the pyloric
end. The pyloric valve is well developed.

INTESTINES.

“The duodenum is at first loosely suspended and convolute,
as In some rodents; it is more closely attached at its termina-
tion. The mucous coat of the jejunum is thrown into small
irregular folds, both transverse and longitudinal. There are
oblong patches of agminate follicles. The termination of the
ileum projects as a conical valve” [a very truncate cone] “into
the cecum. The longitudinal layer of muscular fibres is con-
tinued directly from the ileum upon the cecum ; but the circular
layer accompanies the valvular production of the mucous mem-
brane, and is there thicker than on the free gut. The large
cecum is sacculated on three longitudinal bands, which are
continued some way along the colon.”

We find a number of aggregated glands, not unlike Peyer’s
patches, in the rectum. “The occurrence so low down of what
are probably absorbents may be partly explained by the slow
alteration of food passed along the alimentary canal of the
elephant. Even in the large intestine the original form of
many pieces of vegetable food is retained, and grains of maize
were recognisable in the colon of our example, as were hay and
potatoes in the colon of that dissected by Camper.?

1 Emerson Tennent, Natwral History of Ceylon, p. 125; see also Perrault,
as quoted by Buffon, Hist. Nat. vol. xi. p. 109.

2 Owen, Comp. Anat. of Vertebrates, vol. iii. p. 457.

* Further observations are necessary before we can be satisfied that these
appearances are not due to disease.

58 STUDIES IN COMPARATIVE ANATOMY.

Hunter gives 17 feet as the length of the small intestine.
Mayer makes it 37 feet, while he gives the total length of the
intestines as 75 feet. Professor Owen’s measurements, taken
from a young Indian elephant about 7 feet high at the shoulder,
are as follows! :—

Length of the small intestines, ‘ : 38
Circumference of ditto., . , , ; 2
Length of czecum, : 1
Circumference of czecum, : : 5; 5
Circumference of colon, ¢ 6
Total length of colon and saint to-

gether, : f 20 0
Total length of sathosittoel eater deelacive

of the cecum, . : ; : : 58 6

Our own measurements have unfortunately been mislaid,
which is the more to be regretted that the statement of previous
observers are very discordant.

PERITONEUM.

Hunter has remarked that the lymphatics and lacteals are
small. He found no lymphatic glands on the mesentery pro-
per ; there were several on the meso-colon, but these were not
larger than in man.

“The peritoneum lining the elastic ventral wall of the abdo-
men in the elephant and rhinoceros is of unusual thickness
and strength ; the areolar tissue connecting it to adjacent struc-
tures presents aponeurotic firmness; the free surface of the
serous membrane I found to be white and opaque; it is generally
transparent and opaline or colourless.’? The absence of fat
from all parts of the peritoneum has been remarked by more
than one dissector of the elephant. In our example the great
omentum was quite clear of fat, as indeed were all parts of the
body, except the fibrous pads of the soles of the feet and the
orbits.

1 Comp. Anat. of Vertebrates, vol. iii. p. 458.
2 Owen, Comp. Anat. of Vertebrates, vol. iii. p. 508.

THE ANATOMY OF THE INDIAN ELEPHANT. 59

LIVER.

The liver is divided by the suspensory fissure into two lobes,
of which the right is the larger. There is no gall-bladder, but
the ductus choledocus expands in the wall of the duodenum
into a sacculated pouch, which receives also the secretion of the
pancreas. The termination of the duct projects slightly into the
small intestine, and is surrounded by a sphincter.

PANCREAS.

The pancreas of the elephant differs little from that of most
mammals ; it does not reach the spleen.

SPLEEN.

The spleen is long and flat, broader in the centre than else-
where, and occupies its usual position.

MALE GENERATIVE ORGANS.

Dr Morrison Watson’s account of the male organs of the ele-
phant," evidently founded upon a careful dissection, but neglecting
no advantage which is to be derived from the comparison of
earlier descriptions, is probably the best accessible, and may be
consulted for fuller details than are here given. We extract
from it most of the particulars which follow :—

TESTIS.

The testis is nearly globular, and lies below the posterior ex-
tremity of the kidney. It is entirely invested by peritoneum,
except at the upper and external margin where the vessels enter.
The epididymis lies on its outer side. The veins leaving the
gland are remarkable for their number, large size, and free com-
munications. Ten or twelve vasa efferentia pass into the
epididymis, which is continued without interruption or flexure
into the vas deferens. With the exception of the last five
inches, the vas deferens is convoluted in the whole of its course,
and attached to the superior wall of the abdomen by a double

1 Journ. of Anat. and Phys. vol. iii. p. 65 (1872).

60 STUDIES IN COMPARATIVE ANATOMY.

fold of peritoneum. In the last part of its course it lies upon
the upper surface of the bladder. Here it dilates into an
ampulla two inches long, which is closely connected with its
fellow. It finally unites with the efferent duct of the vesicula
seminalis of the same side to form a common ejaculatory duct,
which opens into the urethra.

VESICULZ SEMINALES.

These have been figured by Camper (pl. v. fig. 1), and described
by Cuvier. Each is six inches long by one anda half in diameter,
and lined internally by a thick membrane thrown into decussat-
ing folds, which become longitudinal towards the urethral end
of the sac. A large transverse fold apparently divides the cavity
into two compartments. Each vesicula is invested by a thick
layer of muscular fibres, continuous with that which surrounds
the membranous part of the urethra and the prostate glands.

PROSTATE GLANDS.

These are four in number, two on each side, and of small size,
the outer ones being the larger. Each gland opens into the
urethra by a single and separate duct.

COWPER’S GLANDS.

These are, as usual, two in number. Each is oval, flattened,
and measures two and a half inches bytwo. The excretory duct
is three inches long, and traverses the spongy body, opening
separately on the floor of the bulbous portion of the urethra.

MALE URETHRA.

The membranous portion is eight inches long. It is invested
by a continuous layer of transverse muscular fibres a quarter of
an inch thick. Towards the neck of the bladder these become
oblique and continuous with thoge which invest the prostate
elands and the vesicuiz seminales. An extension of the erectile
tissue of the bulb surrounds this part of the urethra, thinning
off gradually towards the neck of the bladder. The orifices of

THE ANATOMY OF THE INDIAN ELEPHANT. 61

the various ducts seem to be situated nearly as in the human
subject.

PENIS.

Each corpus cavernosum commences by a slightly dilated ex-
tremity where it is attached to the ischium, and at once unites
with the other crus. An incomplete fibrous septum separates
the two along the middle line, while the interior of each is further
subdivided by conspicuous but irregular fibrous layers, which
are simply enlarged trabecule. The corpus spongiosum begins
by an elongated bulbous enlargement, and gradually diminishes
in diameter towards the free end of the penis, so that its figure
is compared to that of a carrot. An elongated body measuring
three inches by two and a half, situate on the dorsum of the
penis two inches behind its extremity, is believed by Dr Watson
to represent the glans.1 There is a well-marked prepuce. The
orifice of the urethra is Y-shaped, the two limbs being directed
upwards, and the stem downwards.

The levatores penis arise, not from the pubes, but from the
upper and outer side of each corpus cavernosum, as well as, and
principally, from the tuberosity of theischium, The joint tendon
passes in a strong sheath, derived from the tunica albuginea of
the corpora cavernosa, to be inserted, according to Camper, into
the glans. The other muscles are, on each side of the lower
surface of the penis, (1) a bwlbo-cavernosus, next to the middle
line, (2) an ischio-cavernosus, external to the last, and (3) a com-
pressor of Cowper's gland2 The “transverse muscles of the
perineum,” figured by Camper (pl. v. fig. 1, HK, IL), were not
found by Dr Watson.

The dorsal arteries of the penis le on each side of the middle
line, beneath the levator of the same side. The dorsal veins lie
to the inner side of the arteries, and form a plexus, communicat-
ing with the corpora cavernosa, on the dorsal surface of the root
of the penis. The dorsal nerves run on the outer side of the
arteries.

1 Houel, whose authority is not unimpeachable, figures a kind of glans on the
under side of the erect penis.

* Dr Watson describes this as lying between the two last muscles. Camper
figures a ‘‘short accelerator urine,” which may be the same muscle, above the
bulb of the urethra (pl. v. fig. 1, FH, GI).

62 STUDIES IN COMPARATIVE ANATOMY.

It will be of interest to give here some passages from a letter of Dr
William Ogle, in which are quoted the first extant accounts of the
anatomy of the elephant. The description of the generative organs
seems to imply actual dissection by some ancient anatomist.

“ Aristotle most probably, as I think, never saw an elephant him-
self, but the following passages from his Historia Animalium show
that even at that early time some one or other either had or professed
to have examined the inside of one.

“TI. 17. ‘The elephant’s intestine is formed of parts so put together
as to give the appearance of there being four stomachs. Its viscera
resemble those of the hog, but are of course much larger. The
liver, for instance, is four times as large as that of an ox. The
spleen, however, is of small size, considering the large bulk of the
animal.’

“II. 1. ‘The penis of the elephant is like that of the horse, but
small considering the animal’s bulk. The testes are not visible ex-
ternally, but are placed inside, near the kidneys. The pudendum of
the female is placed in the position which in ewes is occupied by the
dugs, but for congress is drawn upwards and directed outwards, so as
to facilitate the action of the male. It has naturally a wide orifice.’ ”

FEMALE GENERATIVE ORGANS.
OVARY.

The ovary in our young example was about one-third larger
than in the adult human female, deeply corrugated on the
surface, and of a flattish oval figure. Many immature Graafian
follicles were distinguished.’

FALLOPIAN TUBES.

The commencement of each Fallopian tube lies within a some-
what capacious pouch, which holds 34 ounces of water when
distended. ‘The fimbriz expand over the membranous walls of
the pouch, and project as ridges from the inner surface. The
outer surface is covered with peritoneum. Scattered muscular
fibres (unstriped), together with vessels and nerves, lie in the
membrane. The ovary les, not im the fimbriated pouch itself,
which directly communicates with the Fallopian tube, but in a

1 Mayer describes the outer surface as smooth “ and only provided with lobes

(Lippchen) at its point of attachment.” He says, further, that there are no
projecting Graafian follicles.

THE ANATOMY OF THE INDIAN ELEPHANT. 63

separate compartment, lined with peritoneum, in whose wall
the fimbriated pouch may be said to be excavated. A valve or
membranous fold separates the two cavities. On the side of
this valve, remote from that part of the pouch which ordinarily
lodges the ovary, the Fallopian tube may be seen, expanding to
its orifice. In the opposite direction it rapidly contracts to a
long, narrow, and tortuous canal, which suddenly expands
again to an outside diameter of about half an inch. From
this point each of the cornua uteri converges towards its fellow,
running parallel therewith for the last 3 inches of its course,
and opening finally into the common uterus. The total length
of each is about 14 inches. A number of uterine glands are
visible towards the lower end of the cornua.

UTERUS.

The cornua unite to form a short tube of about an inch in
length and three quarters of an inch in diameter. This leads
into a somewhat larger chamber about three inches long,
which represents the cervix uteri. The wall of the uterus
is provided with circular muscular fibres. Outwardly the
uterus is with difficulty distinguished from the vagina,
but on opening the tube loose longitudinal folds of the
mucous surface are found to converge towards two well-marked
(anterior and posterior) internal protuberances, which nearly
close the passage from one to the other, forming thus a kind of
os uteri.

Dr Watson finds no constriction corresponding to an os
uteri in the female of Hyena crocuta, and goes on to say :—
“The same remark holds good, so far as I can ascertain, of only
one other placental mammal—that is, of the Indian elephant,
in the female of which, as Mayer pointed out, the vagina is
altogether absent, and the uterus opens directly into the urino-
genital canal.’1 Mayer does not actually use this language, for
he regards the uro-genital canal as vagina (“die mit der Urethra
vereinte Vagina”), but the facts as described by him admit of
Dr Watson’s interpretation. Whether the uterus is really
perfectly continuous with the vagina is another question, and
here we must remark that we find much discrepancy between

1 Proc. Zool. Soc. 1878, p. 424.

64 STUDIES IN COMPARATIVE ANATOMY.

Mayer’s account and the actual part which is now before us.
Shortly below the union of the Fallopian tubes, there is visible
externally a slight constriction. At the same point two con-
siderable and well-marked enlargements project inwards from
the internal wall and almost close the passage. Above this
point the uterine wall is more distinctly and closely plicate
than below. Hence the united Miillerian ducts appear to us
to be plainly divisible (above the uro-genital canal) into
two parts, which are separated by a constriction and differ in
internal structure. The upper part seems to us to represent the
uterus, and the lower the vagina, while the internal thickening
may well represent an os uteri. It would be interesting to
know something of the gravid uterus of the elephant, and in
particular to ascertain by direct observation in what part of the
united sexual ducts the fcetus is lodged, but we have no obser-
vations before us which bear upon this point.

VAGINA.

The next division of the generative canal may be distinguished
as the vagina. It is about 9 inches long, and tapers upwards.
The walls are almost entirely destitute of muscular fibres. On
its inner surface a number of small and irregular ruge are seen,
some of which converge to an indistinct anterior and a posterior
raphe.

Uro-GENITAL CANAL.

Close to the entrance of the urethra into the generative duct
a marked constriction is externally visible, and here, when the
peritoneum is removed, the outer surface, which was smooth
and membranous in the region of the vagina, becomes strongly
marked by circular muscular fibres. On opening the tube a
narrow passage serves as the only communication between the
proper vagina and the vulva, or uro-genital canal. This passage
is divided into lateral halves by a thick rounded cord covered
by mucous membrane, which is apparently a hymen.'

The uro-genital canal is more than twice as long as the pro-
per vagina, and when freed by dissection and extended, it

1 Jt would be well to ascertain, when an opportunity occurs, whether this band
is ruptured by copulation or parturition, as the descriptions of Hunter and Mayer
indirectly lead us to suppose probable.

THE ANATOMY OF THE INDIAN ELEPHANT. 65

projects 14 or 15 inches beyond the anus. In the natural
(unexcited) state the tube is curved forwards and opens on
the under surface of the belly, but by distention of the crura
clitoridis it can be so far straightened that the orifice looks
almost directly backwards, being then beneath the anus, but
separated from it by a considerable interval. Thus the urine is
ordinarily discharged downwards and forwards, but sexual con-
eress takes place in something like the position ordinary among
quadrupeds.’

In the upper half of the uro-genital canal, above the root of
the clitoris, its walls are invested by nearly circular muscular
fibres, which are less distinct on the side adjacent to the rectum,
and are also less developed in the lower part of this space. A
strong band of muscular fibres is attached for about 2 inches
to the body of the clitoris, near its root, and extends upon the
sides of the uro-genital canal. These fibres diverge consider-
ably, the lower ones passing nearly transversely, while the upper
ones pass upwards and forwards at an angle of about 45°. They
terminate along a defined line on the intestinal side, so that the
muscles of opposite sides are separated by a clear membranous
space of about 24 inches. Some of the higher fibres are con-
tinued over the side of the rectum, and become continuous with
the anterior fibres of the sphincter ani, while a number of more
scattered fibres proceed from the upper part of the uro-genital
canal, and are lost upon the sides of the rectum. The last nine
inches of the canal have the walls thin and comparatively mem-
branous, but muscular fibres can be still detected, which are
attached to the sides of the clitoris and surround the tube.

When the uro-genital canal is laid open, its inner surface is
seen to be lined by a smooth mucous membrane. An inter-
rupted line of dark spots is visible near the external outlet,
running about an inch apart from the defined line which marks
the junction of the epidermic and epithelial surfaces. Two
sinuses (canals of Malpighi?) open on the superior (intestinal) wall
of the canal about the middle of its course, and can be traced for
about 2 inches upwards. They open towards the outlet of

1 Dr Watson informs us that the evidence of eye-witnesses, though not so
ample or explicit as might be desired, goes to show that the female rests upon
the fore-knees with the hind legs extended in the standing posture.

ay)

66 STUDIES IN COMPARATIVE ANATOMY.

the uro-genital canal. No glandular bodies were discovered in
connection with the sinuses. The orifice of the urethra is a
simple rounded opening on the side remote from the rectum.’

The uro-genital outlet is not situated near the anus, but is
ordinarily drawn forward, so as to reach the symphysis pubis.
Its sides are very loose and do not appear to be provided with
a distinct sphincter. The labia, or loose folds covered with
mucous membrane, form a hood or prepuce to the prominent
glans clitoridis.

CLITORIS.

In our specimen the clitoris measured 14 inches, and
the glans projected nearly 2 inches. The two diverging crura
spring from the rami of the os pubis. Beyond this junction the
clitoris is nearly cylindrical. The vessels and large nerve lie on
either side, while in the middle line, enclosed in a distinct
membranous sheath, is the common tendon of the retractores
clitoridis. These consist of a pair of small tapering muscles,
which unite about 5 inches below their origin in a common
tendon. This passes along the dorsum of the clitoris, and is
inserted into the glans. The cavernous structure of the crura
clitoridis is well marked, and they can be easily inflated. The
glans is terminal and semi-globular.

ACCESSORY REPRODUCTIVE ORGANS.

The mamme are pectoral, and two in number. They le
between the fore-legs. The two glands are contiguous, and the
nipples are nearly intermediate between the middle line of the
thorax and the axilla.

A temporal gland, situated above the zygoma, and _ half-way
between the eye and the ear, may be possibly an accessory
reproductive organ. It is flat, and in our example of squarish
outline. It lies upon the temporal muscle, from which, how-
ever, it is separated by the temporal venous plexus. The excre-
tory duct is short and of very small calibre, opening from the

1 Hunter (Essays and Observations, vol. ii. p. 175), says that ‘‘ at the termin-
ation of the proper vagina its cavity contracts at once, almost into a blind end ;
in the centre of which there are three small openings, neither of them larger than

a crow-quill; the two lateral of these lead to two small sacs which pass a little
way along the sides of the common vagina.”

THE ANATOMY OF THE INDIAN ELEPHANT. 67

lower end of the gland. In the male and during the rutting
season, a copious flow of odorous liquid is said to be discharged
from the orifice. Our example was an immature female, but
the gland was nevertheless well developed and conspicuous. It
measured 34 inches by 3 inches, and weighed about two
ounces.

URINARY ORGANS.

Each kidney showed five lobes in our example, but other
dissectors report various numbers, from two to eight or nine.
The lobes were essentially distinct, each having its own cortical
and medullary substance and a separate calyx, but the external
separation was not very marked. Two or three Malpighian
pyramids, hardly projecting into the calices, were found in each
lobe; on the flattened apex of each the renal tubes were distin-
guished by the eye. The calices of the three anterior lobes
united to form a common canal, which after a course of about
3 inches was joined by a similar tube formed by the union
of the two posterior calices. Here the ureter was much dilated;
it tapered towards the bladder for the first 4 inches of its
course, and then continued with a nearly uniform diameter to
near its termination, where it suddenly narrowed. The whole
length was 18 inches.

The supra-renal capsules were transversely elongate, tapering
to a point on the outer side.

Comparison of different descriptions of the elephant’s kidney
reveals much discrepancy as to the number and distinctness of
the lobes, the number of calices and their mode of entry into
the ureter. The unlikeness of Camper’s figure to our dissection
is extreme.

The bladder presents no remarkable features. The orifices of
the ureters were seven-eighths of an inch apart? (in our young
elephant), and the orifice of the urethra was about two inches dis-
tant from them. The ureters lie for about an inch between the
coats of the bladder. In the young female the urethra is about
an inch long, opening on the front of the uro-genital canal close
to the proper vagina.

1 In Dr Watson’s example the two ureters were separated by a distance of 34
inches, and lay for 23 inches in the wall of the bladder.

68 STUDIES IN COMPARATIVE ANATOMY.

HEART AND VESSELS.!
HEART.

The heart of the elephant has been repeatedly dissected, and
may now be treated briefly. For fuller particulars the reader
may consult Vulpian and Philipeaux,? together with the correc-
tions and additions of Dr Morrison Watson.

The pericardium is pointed forwards, where it is united to the
base of the heart. Two fibrous cords pass from the back of the
pericardium to the tendinous centre of the diaphragm. Dr
Watson finds them specially connected with the anterior of twe
plates of yellow elastic tissue which cover the two surfaces of
the muscular portion of the diaphragm. He observes further
“that the pericardiac band of the right side was entirely com-
posed of that peculiar form of yellow elastic tissue which, so far
as I am aware, has only once before been described, and that in
the ligamentum nuche of the giraffe by Mr Quekett. In the
left band, as well as in the plate into which the bands expanded,
the elastic tissue presented the usual appearance.” Vulpian and
Philipeaux speak of the fibrous band as single.

The base of the heart is depressed forward, and the interven-
tricular septum is nearly vertical and longitudinal. In the fol-
lowing remarks we shall use terms of direction and position with
reference to the natural state of the parts. The base is under-
stood to be anterior, the apex posterior, and the right auricle
dorsal or superior.

With respect to the general form of the heart, the separation
of the apices of the ventricles is the most important feature.®
Vulpian and Philipeaux say: “Il n’y a pas de sillon inférieur
inter-ventriculaire nettement dessiné. Il y a, au contraire, un
sillon supérieur tres profond, qui, du milieu de la base du cceur,
se porte au voisinage dela pointe.” In our example the case was

1 It was our first intention (supra, p. 8) to omit the heart, lungs, and liver,
seeing that Messrs Vulpian and Philipeaux have described them so minutely, but
on comparing their account with our dissection, and with Dr Watson’s notes,
we find not a few discrepancies of greater or less importance. A short notice of
these viscera is therefore included in the present memoir.

2 Ann. Sct. Nat. Zool. 4° sér. tom. v. p. 183 (1856).

3 In some Cetacea the apex of the heart is indented, while in Sirenia, parti-
eularly in the dugong, the separation of the ventricles is even more marked than
in the elephant.

THE ANATOMY OF THE INDIAN ELEPHANT. 69

almost reversed; the superior groove was shallow and unim-
portant, whereas the apical interventricular groove was deep and
conspicuous. The right side of the heart is, so to speak, rotated
upon the centre of the left; the right auricle being thrown to
the dorsal surface of the heart, while the principal axis of the
right side, passing directly through the auriculo-ventricular
orifice, is inclined upwards, instead of lying nearly horizontal,
as does that of the ieft side in the animal as it stands. The
inferior face (anterior of man) gives no indication of this tilting
of the right side, except that the right auricle is displaced; the
two ventricles meet along a straight line and divide the lower
surface nearly equally. Vulpian and Philipeaux found much
fat at the base and on the front of the ventricles—the only
instance we can recollect in which any considerable quantity of
this substance has been met with in any part of the elephant.
The heart dissected by us was perfectly destitute of fat.

fight Auricle.—The walls are thin, but strengthened by pecti-
nated muscles above, as well as in the appendix, which is
hardly separable from the general cavity. Vulpian and
Philipeaux describe the wall as areolated towards the vene
cave, but this was not the case in our example, except behind
the left anterior cava. Two anterior vene cave (right and left)
enter, one towards the base, the other towards the apical end,
and a posterior vena cava on the dorsal and external side, some-
what in front of the left anterior cava. A sigmoid valve passes
from the external side of the right anterior cava, adjacent to
the appendix, curves round the ventral side of the orifice, and
is then continued as a long membranous ridge of slight projec-
tion to the basal or anterior side of the posterior cava; it then
crosses that opening on its ventral margin, becoming somewhat
more prominent, and serving as a proper valve to the posterior
cava; finally, it gradually disappears along the base of attach-
ment of the Eustachian valve. This agrees tolerably well with the
description of Vulpian and Philipeaux; and with Dr Watson’s
figure, though in the text of his description he says that the
valve passed round the wpper margin of the right anterior cava.
A large Eustachian valve separates the posterior from the left
anterior cava. The great coronary vein opens into the left
anterior cava under cover of a pectinated muscle.

Lewd
70) STUDIES IN COMPARATIVE ANATOMY.

The two superior ven cave in the elephant, as in monotremata,
marsupials, many rodents, the hedgehog, and the bat, and by a rare
deviation from the ordinary rule in the human subject also, are
explicable as a retention of an embryonic structure very general in
vertebrates. At first the blood is returned to the heart by an anterior
(jugular) and a posterior (cardinal) pair of venous trunks, as well as
by a median posterior, which persists and constitutes the inferior cava.
The cardinal and jugular veins of each side unite to form a precaval
or canal of Cuvier, and the two precavals are primitively united into a
common trunk which enters the undivided auricle. The common
precaval trunk ultimately forms part of the right auricle into which
the two precavals then open separately, forming the right and left
superior (or anterior) vene cave. A transverse connection is next
established between the jugular veins; the left jugular, below the
transverse branch, and the left precaval contract and become obliter-
ated ; while the enlarged transverse branch becomes the left innomi-
nate vein, and the lower part of the right jugular together with the
right precaval forms the single superior (or anterior) vena cava of the
higher mammalia. The coronary sinus is, according to Mr Marshall,
the only part of the left anterior cava which in these mammals re-
mains pervious. (See Rathke, Meckel’s Archiv. 1830, p. 63; and
Ueber den Bau und die Entwickelung des Venensystems der
Wirbelthiere, Konigsberg, 1838 ; Bardeleben, Miiller’s Archiv. 1848 ;
Marshall, Phil. Trans. 1850, part i.; and Owen, Anatomy of Verte-
brates, vol. iii. p. 551.)

Light Ventricle—The “incomplete septum” of Vulpian and
Philipeaux seems to consist merely of the more or less united
fleshy columns. In our example these differ considerably from
the description of the authors just named. Four or five are
attached, low down, to the external wall; three or four others,
less distinct, from the ventricular wall to the interventricular
septum. All the very numerous tendinous cords spring from
fleshy columns, though these have not on the inner side of the
ventricle any considerable free extent. No corpora Arantii are
present on the semilunar valves of the pulmonary artery. The
wall of the ventricle is cavernous towards the tricuspid valve,
and also along a line leading from the apex to the orifice of the
pulmonary artery, close to the ventral border of the interven-
tricular septum. The tricuspid valve may have one or two (in
our example two) small additional cusps.

Left Auricle——The entry of the pulmonary veins seems to vary.
Vulpian and Philipeaux found two openings into the auricle—
a small internal and a large external, which latter received
three of the pulmonary veins. Dr Watson describes four

THE ANATOMY OF THE INDIAN ELEPHANT. a

Separate openings. In our dissection there was a large central
orifice and a smaller one on each side—one internal and the
other external; but the external vein was not altogether clear
of the central one. A thin ridge upon the internal surface of
the auricle separates the central from the internal orifice. The
veins enter a thin and membranous sinus, which is slightly
separated from the rest of the auricle by a prominent fleshy
ridge. Part of the edge of this ridge forms the “vaivular
structure” noted by Dr Watson. The wall of the left auricle
is strengthened, especially on the anterior and external sides, by
numerous trabecule. The fossa ovalis is distinguishable only
by a slight transparency.

Left Ventricle :—The mitral valve forms a continuous mem-
branous ring, but indications of a separation into internal and
external cusps are visible. Vulpian and Philipeaux found only
two fleshy columns in the left ventricle—one above and the other
below. We can distinguish four or more, all nearly on the same
level, but divisible into an internal and external set. The
aortic valves are continuous with the inner mitral.

Coronary Vessels—Camper was probably mistaken in saying
that there is only one coronary artery. Like several previous
observers, we find two. The great coronary vein opens into the
left anterior cava.

Pulmonary Artery—Three dilatations, corresponding to the
sinuses of Valsalva, are distinctly visible externally. The artery
passes forwards, upwards, and to the left, curving round the
aorta and dividing in the concavity of the aortic arch into two
branches, one to each lung. The nearly obliterated ductus
arteriosus passes obliquely from right to left, from the pulmo-
nary artery shortly before it divides to the aorta, which it enters
immediately to the left of the left subclavian artery.

Aorta—tThe arch of the aorta gives off a very short innomi-
nate trunk, which subdivides into the right subclavian and the
two carotids, and secondly, a left subclavian! An arteria
thyroidea wma proceeds from the point of separation of the two
carotids.

1 Cuvier and Mayer seem to have found three trunks, viz., right subclavian,
carotid, and left subclavian. Hunter, Owen, Vulpian and Philipeaux, and
Watson agree with the statement in the text.

72 STUDIES IN COMPARATIVE ANATOMY.

We have traced the arteries throughout the body, but the
details of distribution offer few significant features.

Anterior Vene Cave—Dr Watson observes that “each was
formed by the junction of three large trunks a short distance in
front of the arch of the subclavian artery. Of-these, one came
_ from the outside, a second came from the direction of the middle
line, whilst an intermediate one passed directly backwards.
The vena cava of each side, thus formed, passed directly back-
wards, receiving in its course several smaller veins, one of which
was the trunk formed by the union of the companion veins of
the mammary artery, and finally opened into the right auricle.
In addition to these, the right anterior cava received the azygos
vein immediately before piercing the pericardium. There was
no trace of a small or left azygos vein: the posterior cava,
immediately after piercing the diaphragm, opened into the
auricle.”

VEINS AND VENOUS PLEXUSES.

The most striking peculiarity of the veins lies in the plexuses
and free anastomoses which occur in nearly all sheltered parts
of the body. We find extensive plexuses in the superficial and
deep temporal, pharyngeal, pectoral, anterior and internal femo-
ral, popliteal, axillary, and brachial regions, besides less impor-
tant communications elsewhere. The veins are in general large
and capacious. In some cases, at least, valves are wanting in
the plexuses, but we were unable to test the freedom of com-
munication in different directions by a general venous injection.

Superficial Temporal Plecus—This lies above the zygoma and
behind the eye, beneath the temporal gland and superficial to
the temporal muscle. Its communications are with the tem-
poral vein by means of a superficial branch which crosses the
posterior end of the zygoma, with the internal maxillary and
facial veins, and with the deep temporal plexus. ‘The temporal
artery crosses the zygoma immediately behind the vein noticed
above, and gives branches, which do not inosculate, to the area
of the venous plexus.!

1 Neugebaur has described a temporal venous plexus in the goose (Nova Acta,
vol. xxi. p. 521. 1845). In Froriep’s Wotizen, Oct. 1832, p. 39, Otto has ae-

THE ANATOMY OF THE INDIAN ELEPHANT. 04

Deep Temporal Plecus.—On reflecting the temporalis muscle, a
large and intricate venous plexus is seen. The largest of the ©
contributory veins cross the temporal fossa longitudinally under
cover of the zygoma, and communicate in front with the inter-
orbital veins. Two considerable branches pass in front of the
neck of the lower jaw to communicate with the superficial tem-
poral plexus, as also do some smaller branches at the anterior
border of the temporalis.

Pterygoid Pleauses.—There is a complicated network of veins
overlying the pterygoideus externus, and communicating (1) with
the longitudinal veins of the deep temporal plexus ; (2) with the
internal maxillary vein; and (3) with the superficial temporal
plexus, by means of the veins passing in front of the neck of the
lower jaw.

When the pterygoideus externus is reflected, a free anastomosis
of large veins is found, which receives branches from (1) super-
ficial temporal; (2) deep temporal; (3) superficial pterygoid
plexuses; (4) inferior dental veins; (5) venze comites of menin-
geal artery. The plexus discharges into the internal jugular
vein by four large trunks.

Pharyngeal Plecus.—This lies at the back of the mouth, and
has been noticed in the description of the pharynx. On each
side it receives muscular and vertebral veins; the internal
jugular, with lingual, muscular, and facial branches ; the internal
maxillary ; and a descending palatine vein. A single transverse
branch connects the plexuses of the two sides.

Pectoral Pleaus.—Beneath the pectoralis is a plexus of great
extent and intricate arrangement, which effects communications
with the intercostal, internal jugular, axillary, and internal
mammary veins.

These particulars will probably suffice respecting vascular
arrangements whose physiological interpretation is as yet so
obscure. The plexuses of the extremities lie usually in the hol- ©
lows of joints, and are both numerous, intricate, and capacious.
Our notes and drawings preserve many details which it is not
thought needful to describe in print.

scribed the superficial temporal plexus of the elephant as arterial, an error cor-
rected by Dr Watson. The same mistake is repeated in Owen’s Anatomy of Ver-
tebrates, vol. i. p. 548.

74 STUDIES IN COMPARATIVE ANATOMY.

Van der Kolk and Vrolk,! Hyrtl,? and others * have collected
many examples of vascular plexuses in vertebrate animals, and
have suggested several possible theories of their physiological
effect. It seems necessary to distinguish and classify before any
explanation is attempted, for no single explanation can possibly
apply to all the recorded cases. Arterial and venous plexuses
must be carefully separated, though this has not always been
attended to. With respect to the arrangement of their com-
ponent vessels, some plexuses arise by the breaking up of a
trunk into many small branches, which subsequently reunite,
and may be compared to a rope untwisted in the middle (funt-
form plexuses). Others form a network of communication be-
tween different vessels (vetiform plexuses). Others, again, are
due to the sudden breaking-up of an artery into many small
branches, which may anastomose, but do not reunite; or to the
entry of many small veins into a common trunk at or near the
same place (distributive plexuses).

Such plexuses as we have seen in the elephant are all venous
and retiform. We have no confident opinion respecting their
physiological meaning. It may be worth while to remark that
they are so extensive as to constitute in the aggregate a not
inconsiderable reservoir of blood, which may remain nearly
motionless, or move sluggishly when the animal is at rest, but
must be impelled towards the heart by energetic contractions of
the adjacent muscles. The small development of the lymphatic
system is also an important, and not impossibly a related fact.
The lymphatic glands are few and small, and the thoracic duct
was not, in our example, materially larger than in an adult man.
That the venous blood lodged in these many plexuses may have
an absorbent office is a possible view, but one upon which we do
not venture to lay emphasis.

1 ‘Recherches sur les plexus vasculaires chez différents animaux,” Zool. Soc. of
Amsterdam, transl. in Ann. Sci. Nat. Zool. 4° ser. tom. v. p. 111 (1856).

2 ‘¢ Neue Wundernetze und Geflechte bei Vogeln und Saugethieren,” Kais. Akad.
d. Wiss. Math. naturw. Bd. xxii. (1863). Also reprinted separately, 1864.

3 Miiller, Von Baer, Carlisle, Breschet, Milne Edwards, Turner, Murie, &c.

~I
on

THE ANATOMY OF THE. INDIAN ELEPHANT.

RESPIRATORY SYSTEM.

LARYNX.

The thyroid consists of two rhomboidal ale, whose superior
and inferior borders respectively converge forwards to the
middle line at an obtuse angle. They are continuous in front
for the upper half of their depth, and strongly connected by
yellow elastic tissue in the lower half. The cornua project very
slightly. To the postero-superior angle the cartilaginous tip of
the thyro-hyal is articulated by a capsule lined with synovial
membrane ; the postero-inferior angle is similarly connected with
the prominent lateral facet of the cricoid cartilage. The cricoid
and arytenoid cartilages differ in no material respect from the
same parts in man. Professor Owen observes that “ the cricoid
extends posteriorly over the first three tracheal rings.”’ This is
not the case ; the three upper rings of the trachea are obliquely
cut off behind, and their edges are connected with the lower
border of the cricoid. The epiglottis is very thin and flexible.

The crico-thyroidet are closely united in the middle line (C. and
L.—261, fig. 2, 2!). Crico-arytenoideus posticus passes vertically
to the base of the arytenoid cartilage, and the muscles of oppo-
site sides blend. There is no cerato-cricoideus. Arytenoideus
consists of a single set of fibres, which converge from each side
to a transverse median tendon. The upper fibres are continuous
with thyro-arytenoideus, so that the two muscles form a com-
plete ring round the larynx. Crico-arytenoideus springs from
the upper half and edge of the side of the cricoid cartilage ; it is
almost continuous with thyro-arytenoideus towards its insertion.
Thyro-arytenoideus arises from the whole depth of the thyroid
cartilage in the middle line, including the elastic tissue which
closes the inferior vertical notch. The fibres pass downwards
and backwards, to be inserted into the outer and posterior sur-
faces of the arytenoid cartilages. The muscle is continuous
behind with the upper fibres of arytwnoideus. No depressor of
the epiglottis was made out. Hyo-epiglottideus (elevator of the
epiglottis) is a strong muscle arising close to its fellow, from
which it is separated by a septum of yellow elastic tissue. It

1 Anat. of Vertebrates, vol. iii. p. 591.

76 STUDIES IN COMPARATIVE ANATOMY.

springs from the basi-hyoid close to the middle line, and passes

directly backwards to the anterior surface of the epiglottis,
having a length of about three inches. The two muscles under-

lie the thyro-hyoid membrane and support this part of the floor

of the pharyngeal pouch. Camper’s drawing (pl. xii. fig. 4) is

fairly correct, except that he does not indicate any separation of

the muscles of opposite sides.

The upper or false vocal cord is rounded and hardly apparent
except behind. The lower is straight and well-defined, with a
sharp vibratile edge. A small laryngeal pouch leads backwards
from the ventricle, and extends for a short distance under cover
of the antero-external border of the arytenoid cartilage.

The superior laryngeal nerve pierces the anterior border of
the thyroid cartilage near its centre.

Below the arytenoid cartilages and vocal cords the internal
surface of the larynx is covered with longitudinally wrinkled
yellow elastic tissue similar to that of the trachea. The mucous
membrane above the vocal cords resembles that of the pharynx.

TRACHEA.

The trachea consists of about thirty rings. These are incom-
plete behind, and occasionally subdivided, the joints being
enclosed in synovial capsules. The tube is lined by yellow
elastic tissue thrown into firm and very narrow longitudinal
folds. The space between the ends of the rings behind is occu-
pied by two distinct layers of unstriped muscle, the outer layer
being transverse and the inner vertical in the central part, but
radiating outwards laterally.

BRONCHI.

“The right bronchus consisted of eight, the left of six rings.
In addition to these, however, there were several cartilaginous
nodules of small size representing rudimentary rings. No
accessory bronchus, such as is so common among the ruminants,
was met with, the azygos lobe of the right lung receiving its air-
tube from the right bronchus after it had entered the substance
of the lung.’—( Watson.)

THE ANATOMY OF THE INDIAN ELEPHANT. Wi

LUNGS.

The shape of the lungs is adapted to the deep and narrow
thorax ; the whole of the ventral surface is in contact with the
diaphragm, which extends forwards to the second or third rib.
The visceral and parietal layers of the pleura are closely con-
nected together by matted elastic tissue, while the pleural sac is
at the same time pretty generally adherent to the thoracic wall
on the one hand and to the surface of the lungs on the other.
The right lung, according to Vulpian and Philipeaux, has a
small upper lobe, a triangular internal lobe, which rises towards
the trachea, and a very considerable third lobe. Dr Watson
found only two lobes, of which the smaller lay on the left side
and projected transversely from the lung immediately below
the hilum. In our example the arrangement was nearly the
same. The left lung is undivided. ‘The lobules are very easily
separable. Dr Watson observes that the bronchi branch irre-
gularly, and that upon entering the lung they immediately lose
the cartilaginous lamine. This was the case in our subject also.

NERVOUS SYSTEM.

We are not able to give any useful information respecting the
nervous system of the elephant. The brain was not removed
until eight months after death, when its internal structure was
much broken down. We have traced the chief nerves, but find
no peculiarities sufficiently remarkable to justify a minute
description."

ORGANS OF SPECIAL SENSE.

EYE.

Mayer describes a special depressor of the lower eyelid, and
Dr Watson confirms the statement. It arises with the recti
and obliqui from the bony canal posterior to the orbit, and
passes forwards beneath the eyeball to be inserted into the
cartilage of the lower lid. Dr Watson notices also “a very

1 For the relative sizes of the cranial nerves, Dr Harrison’s description may be

consulted. Quoted (with figure) in Sir J. Emerson Tennent’s Natural History of
Ceylon, p. 95.

78 STUDIES IN COMPARATIVE ANATOMY.

extensive and well-developed periosteal muscle,” which “ cor-
responds exactly in position to the similar muscle in the sheep
and deer.”! The muscles which pass from the upper and lower
eyelids to be inserted into the third eyelid are not, according to
Dr Watson, distinct muscles at all, but fibres of the orbicwlaris
palpebrarum.

Camper, Harrison, and Watson agree in stating that no true
lachrymal apparatus is present. The Harderian gland lies
between the inner wall of the orbit and the rectus internus ; its
excretory duct opens on the surface of the third eyelid.

EAR.

The membrana tympani is of oval form, and looks downwards,
outwards, and a little forwards. The malleus is attached above
the centre to the apex of the shallow funnel, which, as in other
mammalia, projects inwards from the tympanic membrane.
The apex of the funnel is directed upwards and forwards. In
our young example the long diameter of the membrane was
Z inch, and the short diameter } inch less. Sir Everard Home
gives 37 inch as one dimension, and rather more than an inch
as the other.2. This was probably from an adult animal. The
middle layer contains the usual radiating and circular fibres.
No peculiarity of importance was observed with respect to the
ossicles.

NOsE.

The cartilaginous nasal septum continues forwards the plane
of the vomer and of the perpendicular plate of the ethmoid.
It ends in front by a tapering projection which extends in
advance of the bony nares, and supports the alinasal cartilages.
The cartilage of the septum is not prolonged into the fibrous
and muscular septum of the proboscis. Its upper edge gives
origin to a set of longitudinal muscular fibres, which increase in
number below, and ultimately blend with the under surface of
levator proboscidis. The same edge gives off on each side a
narrow lateral process, continuous along the whole length of the
septum, and to these processes the alinasal cartilages are attached

1 Described by Professor Turner in the Proc. Royal Phys. Soc. Edinburgh, Dec.

19, 1861, and Nat. Hist. Review, Jan. 1862.
2 Phil. Trans. for 1800, p. 4.

THE ANATOMY OF THE INDIAN ELEPHANT. 79

in front of the bony nares by a simple hinge-joint, without
synovial capsule. In our young example each alinasal cartilage
measures four inches in length and less than one inch in width ;
it has a slight curvature forwards and outwards. Its external
margin is connected with the bony nares beneath levator pro-
boscidis by a thin, tough, and extensive sheet of fibrous mem-
brane, which can be traced downwards into the lining of the
air-tubes of the proboscis. The membrane is capable of disten-
sion, and forms a kind of pouch, which greatly enlarges the
capacity of the nares at this point. Transverse muscular fibres,
continuing the series of the radiating fibres of the proboscis,
connect the sides and under surface of this pouch with the
margin of the bony nares. Another set of transverse fibres
(dilator naris), extending across the alinasal cartilage and tak-
ing origin from it, passes to the adjacent border of the pouch,
leaving a clear space about an inch in width between them and
the lower transverse fibres. This upper set (dilator naris)
would, when in action, tend to raise the free or external edge of
alinasal cartilage, and thus increase the capacity of the pouch,
while the lower transverse fibres would assist them to enlarge
the pouch by drawing its sides outwards. The alinasal pouch
is the only considerable chamber in the nose or trunk of the
elephant ; it may serve an important purpose in the storage of
liquid. Behind the anterior bony nares we find a second lateral
(aliseptal) cartilage. Itis not perfectly continuous either with
the nasal septum or the alinasal cartilage. It is connected with
the former by a fibrous hinge-joint ; with the latter by a strong
elastic sheet, in which are a number of muscular fibres, which
take origin from the upper border of the bony nares, and con-
verge towards the fore end of the aliseptal cartilage. The posi-
tion and form of this cartilage are those of a turbinated bone,
but it is wholly unossified ; its size is small in comparison with
the relative development of the turbinated bones in most mam-
mals. Its chief extension is in the direction of the nasal passage,
and this dimension is about four inches in our specimen; the
greatest vertical depth is two inches. The anterior half of the
cartilage falls away outwards from the plane of the posterior
half, so as to give rise to a decided step in the middle of the
length. The general direction of the cartilage is vertical, but

80 STUDIES IN COMPARATIVE ANATOMY.

somewhat irregular, and the lower edge is rather nearer to
the median plane (especially in its posterior half) than the
upper. This aliseptal cartilage forms part of the outer wall of
the nasal passage, as well as the inner wall of a transversely
contracted channel which leads from the nasal meatus to a
large frontal sinus. The muscular fibres attached to the fore
edge of the aliseptal cartilage, which are continuous with the
lower set of the transverse fibres of the alinasal pouch, when
they contract, draw the cartilage towards the bony or outermost
wall of the nasal chamber, and very effectually close the
entrance into the air-sinus. The lower border of the cartilage,
for somewhat more than the anterior half of its extent, is con-
nected with the wall of the nasal chamber by a strong band of
yellow elastic tissue, which forms part of the dividing septum
between the proper respiratory passage and the entrance to the
frontal air-sinus; it serves to antagonise the muscular fibres
which close the valve. The posterior extremity of the cartilage
is connected to the lateral wall by a mass of fibrous and mucous
tissue, which forms the anterior boundary of the special respira-
tory chamber, and separates this from the channel leading to
the frontal sinus. The aliethmoidal volutes are disposed in
about seven laminz, which gradually diminish in size backwards,
and closely resemble the bony framework of the olfactory organ
in the horse. Behind this point the nasal passage lies between
the hard palate and the spheno-palatine surface. Its vertical
extent in our young example is nearly three inches, but the
transverse width is extremely small, and hence the nasal pas-
sages seen from behind present the appearance of greatly elon-
gated vertical slits. (See p. 50.)

Dr Watson believes that the valve-like aliseptal cartilage is
adapted to obstruct the flow of water into the air-sinuses of the
skull during its passage along the narial tubes. We are disposed
to question the normal passage of water along this highly
sensitive tract. Examination of the parts discovers no valve or
other provision for preventing water flowing from behind for-
wards! from gaining free entrance into the olfactory recesses.
Moreover, the only important receptacle above the stomach is
constituted by the pouch adjacent to the alinasal cartilage. We

* See the quotation from Dr Watson’s memoir on p. 54.

THE ANATOMY OF THE INDIAN ELEPHANT. 81

are inclined to suppose that the water lodged here (if, as we see
no reason to doubt, the pouch in question actually serves as a
receptacle for liquid) is pumped up through the proboscis, and
never passes backwards beyond the anterior bony nares. The
water which, according to the testimony of many observers, is
regurgitated from the stomach, would, we imagine, be withdrawn
exclusively by the tip of the proboscis inserted through the
mouth into the pharyngeal pouch.

CONCLUDING REMARKS.

It may be worth while to direct the attention of future dis-
sectors to certain points, which have been passed over or
inadequately treated by us. Most of these are superficial or
easily made out, and it is to be hoped that the first opportunity
may be seized of correcting our notes.

1. The cutaneous muscles should be redissected and more
fully described.

2. The sterno-humeral and superficial cervical muscles (pector-
alis, masto-humeralis, sterno-maailaris, sterno-nastoideus) were
insufficiently noted by us and in the first hasty examination of
the thorax, made before the possibility of a complete dissection
was entertained, these muscles were mutilated beyond recovery.
We have attempted to piece together and interpret the remain-
ing shreds of muscle, but the results are not altogether trust-
worthy.

3. The large intestine should be studied microscopically while
quite fresh.

4, The brain should be removed immediately after death, and
minutely investigated. The parts liable to be injured in the
process of extraction are now well known.

The difficulties attending the anatomical examination of a
single example of a very large animal must be our excuse for
many shortcomings. If we shall be found to have aided the next
observer as much as Camper, Cuvier and Watson have aided us,
we shall at least have repaid our debt. We do not doubt that
much, very much, remains to be done before the structure of this
single species is adequately made known. Minute and accurate
knowledge of the anatomy cf the elephant will be found of
special value, and competent anatomists are happily not so few : _

82 STUDIES IN COMPARATIVE ANATOMY.

that we need grudge the time and labour which minute and

accurate knowledge is sure to cost.

The foregoing memoir was originally contributed to the
Journal of Anatomy and Physiology, where it appeared in
January, April, and October 1878. A short notice of the male
generative organs has since been added.
word “inaccurate” (p. 28 line 34) has been accidentally changed

to “accurate.”

Postscript.

EXPLANATION OF PLATES.

Prats I,

Fig. 1. Inner side of right fore-limb.

h.
CO.
ol.
pet.
i

sm.
SC.
Sp.
tm.
bi.
cb.
tr.
dé.
ba.
ecr.
ep.
Jer.
pi.
jid.
pr.
br.

Humerus.

Coracoid process.
Olecranon.
Pectoralis.

Radius.

Serratus magnus.
Sub-scapularis.
Supra-spinatus.
Teres major.
Biceps.
Coraco-brachialis.
Triceps.
Dorso-epitrochlearis.
Brachialis anticus.
Extensor carpi radialis longior.
Extenser proprius pollicis.
flexor carpi radialis.
Palmaris.

Flexor longus digitorum.
Pronator radii teres.
Brachial artery.

Fig. 2. Flexor side of left fore-arm.
to)

U.

el.

Ulna.

Elastic ligament.

ecu. Hatensor carpi ulnaris.
feu. Llecor carpi ulnaris

1 This should pass in front of flexor longus digitorwm, to be inserted into the

pisiform bone at x.

In reprinting, the

THE ANATOMY OF THE INDIAN ELEPHANT. 83

an. Anconceus.

m. Seep. 15, line 5.

Jp. lexor longus pollicis.
fod. Flexor brevis digitorum.
amd. Abductor minimé digit.
emd. Hatensor minimae digitt.
ra. Fadial artery.

un. Ulnar nerve.

mn. Median nerve.

Other references as in fig. 1.
* * Fig, 2 is drawn on a larger scale than fig, 1.

Puate II.
Fig. 1. Dissection of the root of the proboscis. Side view.

Ip. Levator proboscidis (reflected).
dn. Dilator narium.
an. Alinasal cartilage.
tf, ‘Transverse muscular fibres of alinasal pouch.

Fig. 2. Front view.

Puate IIL
Section through nasal passages, pharynx, and larynx.

js. Frontal sinus.
ce. Cranial cavity.
Kustachian tube.

nf .p. Pharyngeal pouch.

jc. Jacobson’s canal.

bs. Basi-sphenoid.

mz. Maxilla.

pma. Premaxilla.

mn. Mandible.
hy. Hyoid.

an. Alinasal cartilage.

as. Aliseptal cartilage.

ae. Aliethmoidal volutes.
th, Thyroid cartilage.

cr. Cricoid cartilage.

cp. Constrictor pharyneis.
S. Hind edge of nasal septum.

, The proportions, especially of the bony parts, will be found
to alitter from those of the adult elephant.
Puate IV.
Fig. 1. Uro-genital organs (female). General view.

sp. Supra-renal capsule.

i Tio haresy,

84

STUDIES IN COMPARATIVE ANATOMY,

b. Bladder.
o. Ovary.
u. Uterus.
v. Vagina.

ug. Uro-genital canal.
cl. Clitoris.
7. Rectum.

Fig. 2. Ovary and commencement of Fallopian tube.

o. Ovary.

. 3. Commencement of uro-genital canal (laid open).

h, Hymen? —

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