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I:

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INDEX. |

Material for the study of the ruminants

=)

MATERIAL FOR THE STUDY

OF

RUMINANTS

EINAR LÖNNBERG.

(PRESENTED TO THE ROYAL SOCIETY OF SCIENCES or Ursara Mas 6 1902.)

UPSALA 1903
PRINTED AT THE UNIVERSITY PRESS
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Material for the Study of Ruminants.

» There is scarcely any natural and
well-defined group in the whole clasf
which presents greater difficulties os
subdivision than this.» (Cavicornia).
Frower and LYDEKKER.

The soft anatomy of the wild ruminants is still imperfectly known.
The occurrence of these animals in far off countries and their often
considerable size have caused this lack in our knowledge and make
it also difficult to obtain, at one time, a large and comprising material
without which the anatomy of the whole group and the mutual affini-
ties between the genera and species cannot be made out in a satis-
factory manner. In such a case one must content oneself if now and
then some smaller contributions can be offered and this may serve as
an excuse for the following paper. In the same the soft anatomy of
some antelopes etc. will be described and compared with that of other
known species. In some instances an adaptation of the intestinal canal
to the diet is stated and certain features of the ontogenetic develop-
ment of the organs of some species is also brought to the light.

Nova Acta Reg. Soc. Sc. Ups. Ser. III. Impr. *?"/o 1902. 1

On the soft Anatomy of the Black-buck of India (Antilope cervi-
capra (LIN.)) compared with that of other Ruminants.

Professor TycHo TULLBERG has kindly placed at my disposition
a young male specimen of Antilope cervicapra from which the following
notes have been taken. The total length of this young buck was.
102 cm.

The palate showed about 16 or 17 pairs of transversal plice
or elevations. The anterior ones of these are most developed and
very feebly crenulated or almost smooth, the posterior ones are quite
smooth. They are thus more similar to those of the sheep than the
same of the Saiga which latter, according to Murin'), are »fringed».
The number of palatal elevations in the Black-buck exceeds, however,
that of both the others. The buccal papillæ of this animal are rather
short and blunt. Almost all of them are single-pointed. In a male
Madoqua saltiana 1 found 14 pairs of palatal elevations in a female
only 12.

The papilla incisiva is roundish, thus different from the triangu--
lar one in the sheep. In Madoqua it is also a round cushion but from
the surrounding groove goes not only, as in the Black-buck, a median
groove forward to the upper lip, but also one towards each side. The
caruncula sublingualis of the Black-buck is formed by a larger triangu-
lar flap anteriorly, close behind which is another smaller one, and
about 3 mm. behind this one is a third of nearly the same shape and
size as the second. In this respect the Black-buck agrees pretty well
with Ovis and Capra. In Madoqua the anterior flap is digitiform and
the posterior ones missing. The salivary glands of this young Black-
buck were destroyed to such an extent or transformed by disease
that it is best not to say anything about them.

The tonsillary glands are flattened, elliptic in outline with the di--
mensions 20 x 15 x 7 mm. There are on one side three, on the
other four tonsillary openings partly closed by wart-like prominences.
In Saiga there is only one such opening on each side according to MURIE..

1) Proc. Zool. Soc, London 1870.

MATERIAL FOR THE STUDY OF RUMINANTS. 9

As is also the case in the Saiga according to MuriE (l. c.) the
anterior portion of the tongue of the Black-buck is strongly pigmented.
The dark colouration is, however, in this animal still more restricted,
being confined to the anterior third or only a little more. The poste-
rior part is perfectly pale. The pepille filiformes are flattened almost
scale-like but pointed, thus similar to those of the sheep. Anteriorly
they extend about 7 mm. on the lower side of the tongue, that is just
as far as the tongue is pigmented. Posteriorly hardly any such pa-
pillae are found on the lateral surfaces of the tongue. The median
region of the posterior half of the tongue is as usual eovered by large
papillæ which are similar to those I have described in Ovibos!). They
are mostly scale-like and blunt at the tip; some of the anterior ones
almost like tubercles. I have had the opportunity to examine two heads
of Salts Dik-dik and may therefore make a comparison between this
species and the Black-buck in some points. The Dik-dik has also
rather scale-like papillæ filiformes which are not even pointed but
rather truneate. But where the papillary covering extends on the sides
of the tongue the papilla are slender and pointed. Posteriorly the
Dik-dik like the Blaek-buck has large wart-like papillæ in the middle
of the back of the tongue but towards the sides they become more
conical or partly scale-like. The papillæ fungiformes are in the Black-
buck by far most numerous and most closely set at the anterior
edge of the tongue, but are in the black region found anteriorly 1!/2
em. and laterally 1 em. from the margin. Underneath they are scat-
tered almost over the whole.black region although, of course, most
numerous near the edge. In the pale region the papille fungiformes
are few and far between, only found near the lateral border of the
tongue leaving a broad central area unoccupied. Some few are also
Iound on the upper part of the lateral surfaces posteriorly. In the
pigmented portion they are black, in the pale region pale. Their di-
stribution is on the whole similar to the same in the Gnu?). Madoqua
saltiana has numerous papille fungiformes scattered all over the ante-
rior portion of the tongue, less densely in the middle, in greater num-
ber at the tip and especially crowded below the same. In the middle
of the posterior portion of the tongue there are no fungiform papilla
in the little Dik-dik, but towards the sides they become fairly nume-

1) Proc. Zool. Soc. London 1900.
?) Conf. LówwbERG : On the soft anatomy of the Gnu. K. Sv. Vet. Akad. Handl. Bd. 35.

4 EINAR LÓNNBERG,

rous and have there a size fully twice as large as that of the ante-
rior ones, or more. There are in the Black-buck basally 7 or 8 pa-
pille circumvallate on each side, but some of these are not quite ty-
pical. They are irregularly distributed in two rows. This number is
low compared with the same in common sheep and goats: but also
in the Saiga these papillæ are »few» according to MuRt (I. c.). In
Madoqua saltiana the papille circumvallate are rather irregularly ar-
ranged and in consequence of the great size of the fungiform papillæ
posteriorly it is often very difficult to decide whether a papilla belongs
to this or that class. In a male specimen I found about a dozen un-
mistakable circumvallate papille arranged in four oblique short series.
In a female the number seems to be about the same but the arrange-
ment is less regular. The Dik-dik differs thus with regard to the ar-
rangement of these papillæ from the Antilope. In the sheep these papilla
sit in four rows but are more numerous as already stated. The Gnu
has a similar number of such papillae as the Black-buck but as the
number of papille circumvallate is rather variable even in the same
species it cannot be considered to be of much classificatory importance.

When seen from the right side the distance from the cardia to
the blind end of the dorsisinistral sack of the paunch is in its present,
moderately distended state 240 mm.!) and the distance from the car-
dia to the blind end of the ventridextral sack is 275 mm.') The
breadth of both sacks is about the same or 12,5 mm.!) from sulcus
longitudinalis dexter to either side. Both sacks are rounded at their
blind ends. Sulcus coronarius of the ventridextral sack is well de-
veloped and the blind sack limited by the same measures about 11
cm. in length by 10,5 in breadth.

When seen from the left side the ventral sack measures 250
mm. and the dorsal 240 mm. in length.

The diameters of the reticulum are about 10,5: 9,5: 7.5 em. It
is almost heartshaped with the rounded point forming its anterior fun-
dus, the communication with the paunch is at one auricle and the
other protrudes like a blind sack beyond the opening to the psalterium.

This latter is very small, beanshaped with its diameters measur-
ing resp. 45; 30; 25 mm.

') These figures agree perfectly with the renewed measurements obtained from the
with air distended and dried stomach. The absolute correspondence of the measurements.
in both cases prove their correctness.

MATERIAL FOR THE STUDY OF RUMINANTS. 5

The abomasus measures along the middle of the curvature about
23 em. its greatest transversal diameter is about 7 cm,

With regard to the general shape the stomach of the Black-
buck seems to differ from most ruminants of which this organ is known
to me. Both sacks of the paunch are almost equal in size as in the
common sheep, but their terminal blind sacks are more separated from
each other and more equally rounded than in the latter. The rounded
shape is also different from that of the Cervicornia as well. The
paunch of the Saiga is termed »bifid» by Muri (l. c.), but he does
not give any detailed description. As in the Gnu sulcus longitudinalis
dexter accessorius is better developed in the Black-buck than sulcus
longitudinalis itself. On the dorsal division of the paunch there is no
sulcus coronarius visible. This depends upon the fact that the entire
portion of this division which forms the rounded posterior blind end,
as described above, is not homologous with the saccus ccecus dorsalis
of the sheep. But if the ventricle of the Black-buck is seen from the
left side there may be detected in the fissure beetween both sacks
a small, somewhat pointed, pocket from the dorsal division which cur-
ves ventrally so that it lies close to the ventral saccus cecus. This is
the homologon to the saccus cecus dorsalis of the sheep and other ru-
minants. From this can be concluded that the dorsal division of the
pauneh of the Black-buck has been subjected to a reduction when
compared with other forms.

The shape of the reticulum as described above seems also to
be characteristic of the Black-buck. The cells of this division are as
in the Saiga, divided by secondary stellate septa.

In the Saiga Murie states that the psalterium is »comparatively
small». In this animal as can be seen from the measurements re-
corded above it is very small and might nearly be termed rudimentary
when its size is compared with that of the other divisions of the sto-
mach. In this respect therefore, the Black-buek differs in a consider-
able degree from Ovis and Capra!).

The abomasus of the Black-buck is provided with about 19
longitudinal folds. "These are best developed on both sides of the lesser
curvature, but short and situated at a greater distance from each
other at the greater curvature. They are, as it seems, a little more

!) I take this opportunity to state that the psalterium of Connochetes gnu is quadri-
plicate which for certain reasons could not be ascertained when I wrote my paper on the soft
anatomy of this animal. K. Sv. Vet. Akad. Handl. Bd. 35.

6 EINAR LÖNNBERG,

numerous than in the sheep. The pyloric tract is to an extent of 7
em, entirely free from folds, but the shorter folds of the greater cur-
vature disappear already at twice that distance from the pylorus.
From the condition of the dorsal division of the paunch and
that of the psalterium as described above it is evident that the ven-
tricle of the Black-buck represents a stage of reduction when compa-
red with that of the sheep. The cause why such a reduction has
taken place must lie in an altered diet. If a ruminant begins to feed
on more nourishing food than before the reservoir formed by the
paunch may be lessened without detrimental effect, and if the food is
softer and more easily crushed by the teeth the grinding action of the
psalterium is not so important and this organ may therefore be re-
duced in size. Unfortunately I do not know anything about the diet of
the Black-buck, but it lives in a climate which allows the animal to
obtain green herbs all the year round and it may therefore have the
opportunity to choose tender leaves and sprouts. BREHM says that
it feeds on »Gräsern und saftigen Kräutern», but it is uncertain how
much such a statement may be relied upon. There is, however, a
different condition of life for the Black-buck than for the wild sheep
and other ruminants which live in a temperate climate so that they
at least during the winter must feed on dry plants, bark of twigs and
similar matter, harder to digest, more difficult to grind and less nou-
rishing. In the latter case a large paunch and a powerful psalterium
is well needed. The variability of the psalterium has already been
observed by previous authors and in the following will be offered
further proofs of this. Garrop') remarks that the organ in question
is minute in »Nannotragus» (Ourebia) and Cephalophus. Boas”) found
the psalterium of Cephalophus maxwelli to be only 2 em. in length, but
in the same animal the abomasus measured 11—12 cm. The same
species was also among those examined previously by Garrop (I. c.).
Boas terms the psalterium of the said animal rudimentary. In the
Tragulids the psalterium is perfectly rudimentary, as is well known.
Boas puts this in connection with the small size of the animals. He
says that it is probable that such small ruminants as those mentioned
above do not choose the coarsest food and that therefore the food
»schon beim Wiederkiiuen so fein gekaut worden, dass eine wieder-
!) Proc. Zool. Soc. London 1877.

*) Morph. Jahrb. XVI. Leipzig 1890. Conf. also the statements below in another
shapter of this paper.

MATERIAL FOR THE STUDY OF RUMINANTS. 7

holte mechanische Bearbeitung eher entbehrlich ist». In this point
I agree perfectly with Dr. Boas. But when be says that the intestinal
canal of the small animals have smaller quantities of food »zu bewäl-
tisen» and that they therefore have less need for a well developed
psalterium I cannot agree witb his reasoning any longer. Of course
the quantities of food needed by a small animal is absolutely smaller
than that required by a larger, but comparatively the former needs
fully as much or more than the latter, and if both feed on the same
kind of food it is as important for the small animal as for the large
to let the food go through an effective treatment from the side of
the digestive organs. It is thus, according to my opinion, not the size
of the animal, but the foodmaterial which in the first instance influen-
ces and causes the different development of the digestive apparatus.
The size of the animals may, however, be of secondary importance
because it is easier for a small animal to find sufficient quantities of
better and more nourishing food while, on the other hand, the large
animal may be obliged to take its refuge in coarser food as well only to
get the quantity needed, and then this influences the development of
the digestive organs. |

The length of the small intestine of the Black-buck is 895 cm.
The duodenal loop is rather short and ductus choledochus opens into
the same about 20 cm. from the pylorus. The general arrangement
of the small intestine is as usual, but the mesentery by which it is
attached to the peripherical coil of the large intestine is very short or
narrow, as a rule only measuring 2'/2—2 cm. In this respect Antilope
cervicapra resembles the ovine group and differs from the common ox
as well as from the Gnu’).

The ileum enters into the cæcum about 12 cm. from the blind
end of the latter. The transversal diameter of the caecum is about
4 em. The colon continues with a similar diameter as the caecum in
a forward direction for about 6 cm. from the ileocæcal opening. Then
it bends back upon itself and forms with a somewhat diminished dia-
meter (27/2 cm.) a loop, ansa proximalis, which extends 10 cm. back-
wards. It then turns again forwards behind the caecum and towards
the median line where the spiral coils begin. If the beginning of the
spiral is counted from this point the colon makes two centripetal coils,

1) Conf. my paper on the soft anatomy of the Gnu K. Vet. Akad. Handl. Bd. 35.
Stockholm 1901.

S EINAR LÓNNBERG,

then bends back upon itself and forms one complete centrifugal coil
and a half turn backwards in close connection with the terminal por-
tion of the ileum. It turns then peripherically and continues rather
narrowly, as is already stated, connected with the jejunum and is in
consequence hereof laid in quite a number of undulations. It forms,
however, no extra loop since it has left the central spiral coils but
follows the jejunum. When it leaves the common mesentery at duode-
num it forms a double loop, ansa distalis 9 em. in length, which is elo-
sely fixed to the ansa proximalis by a very short mesentery, and passes
then into the straight rectum.

If the coils of the central spiral are counted from behind their
number will be 2 centripetal and 1!/» centrifugal.

When the colon leaves the central spiral and is turning peri-
pherically it forms in some ruminants a long loop returning upon it-
self before it makes the last peripherical tour in the common mesen-
tery of the small intestine. This loop which is especially well developed
in the Muskox!) is entirely missing in this animal.

The dimensions of the large intestine are as follows: from the
ileocecal opening to the end of the central spiral 160 em., from this
point to the end of ansa terminalis about 116 cm., rectum 46 cm., thus
the entire length about 322 em. "These measurements, as well as
those recorded above concerning ihe small intestine are made in a
fresh state by means of thread laid along the intestine while still fully
adherent to the mesentery. A comparison between the length of the
small and the large intestine shows that the former is 2,; times the
latter, which is exactly the same figure as is recorded by Murte?) for
this animal and nearly the same as in the Saiga. In Ovis and Capra
the small intestine is fully 3 times as long as the large one although
the latter forms 8 centripetal and 3 centrifugal coils in the spiral.
For comparison with the corresponding measurements of other Rumi-
nants may be referred to papers by GanRop?), Murie“) and the pre-
sent author").

') Conf. LönsBEre: On the Soft Anatomy of the Muskox. Proc. Zool. Soc. London
1900 p. 152, 153:

?) Murie: Proc. Zool. Soc London 1870.

?) Gannop: Proc. Zool. Soc. London 1877.

*) Murie: Proc. Zool. Soc. London 1869 and 1870.

?) LówNbBERG: Proc. Zool. Soc. London 1900; K. Vet. Akad. Handl. Bd. 35. Stock-
holm 1901.

MATERIAL FOR THE STUDY OF RUMINANTS. 0

The most conspicuous feature in the general arrangement of the
intestine of this animal when compared with the same of the domestic
animals, sheep and ox, is the narrowness of the mesentery, so that
the peripheric coil of the colon lies rather close as well to the central
coils as to the jejunum. The entire length of the intestine of this
young Black-buck is not more than about 12 times that of head and
body. Previous authors (Cuvier) have found the same relation to be
about 15. Even if this latter figure should be correct in some instan-
ces the intestine of this animal is much shorter when compared to
the length of the animal itself than, for instance, in the domesticated
ruminants in which according to ELLENBERGER') the entire length of
the intestine »betrügt das 24—33 fache der Rumpf länge». Both small
and large intestine of the the Black-buck are thus comparatively shor-
ter than in the sheep as can be concluded from this statement com-
pared with the above stated relation between the intestinal traets of
both animals. The shortness of the intestine and the few colic coils
in the Black-buck point to the same as the above stated reduction of
eertain gastric divisions namely that this animal must feed on more
easily digested food than, for instance, the sheep.

The liver of this young Black-buck is rather deeply cleft in a
left and a right lobe. The dimensions of the former is 11,5 by 6 em.,
and of the latter 8 by 10 cm. The greatest thickness is about 21/2
em. On the convex anterior side the umbilical fissure extends half
way through the organ, but on the concave side not quite so far.
The gall-bladder is situated quite close to the fissure so that the well
developed groove or fossa vesicæ fellee which extends from the cervix
of the bladder to the margin of the liver, opens, so to say, in the
fissura umbilicalis at the blind end of the bladder. This situation seems
to be somewhat aberrant from the usual condition in the Cavicornia,
but Murie says about the Saiga that »the slight median notch» is
»close to the fundus of the gall-bladder» so that in that animal the
situation of this organ seems to be similar although the umbilieal fis-
sure is only little developed. In Sus the gall-bladder lies also close to
the umbilical fissure. The shape of the gall-bladder of the Black-buck
is similar to the one which I have figured (I. c.) from the Gnu. It
is not pear-shaped, but abruptly widened near the ductus choledochus
at the cervix and is there broader, about 23 mm. when empty and

') Handb. d. vergl. Anat. d. Hausthiere. Berlin 1900 p. 452.
Nova Acta Reg. Soc. Sc. Ups. Ser. III. Impr. 5/7 1903.

to

10 EINAR LÓNNBERG,

flat. Towards the blind and rounded end it is somewhat narrowed,
IS mm. across. The blind end protrudes very little beyond the free
margin of the liver, unlike the case in the Gnu and the ox. Ductus
choledochus is wide. Its opening in the duodenum has already been
described above. The shape of the gall-bladder is quite different to
that of the Saiga to judge from Murir's description of the same in
which it is termed »short roundish» and the fossa »small and shallow».
Only an undulation of the margin of the right lobe indicates its divi-
sion in a central and a lateral portion. There is a comparatively large
lobus Spigelii with broad base about 4 cm., and rounded free margin
protruding about 1 em. Lobus caudatus must be termed comparatively
large measuring almost 5'/» cm., trihedral in shape and with a large
impressio renalis on its right side. The great breadth of the base of
the Spigelian lobe agrees with the »oviform» type according to GaRROD'S
terminology (I. c.) but as I have remarked before not much stress
can be laid on the shape of such a variable organ. The caudate
lobe also resembles in some degree the same of the sheep. In the
Saiga the Spigelian lobe is, according to Murine (I, c.), »but scantily
developed» and the caudate lobe is »bayonet figured». It is, however,
as it seems from Murin’s description, larger than in the Black-buck
and extending beyond the right border. On the whole, there exists a
certain resemblance in the shape of the liver in these animals.

The pancreas does not present any remarkable feature. Its
duct seems to join the ductus choledochus and I have not been able to
detect any independent duct to the intestine.

The spleen of the Black-buck is triangular in outline, but much
more elongate!) than in sheep and goats. It measures in fresh state
125 mm. in length by 55 in breadth at the broadest place and 10 in
thickness. The longest side of the triangle is convex and somewhat
undulated, the others straight. The triangular shape of the spleen
may be regarded as an ovine characteristic, as it differs so much
from the oval shape of this organ in the Cervicornia, Giraffa and Ovi-

') I take this opportunity of correcting a misprint which has through inadvertence
taken place in my paper »on the soft anatomy of the Muskox» (Proc. Zool. Soc. London
1900). In setting the types a few words have been omitted on page 156. In the middle
of this page it should be read: »The spleen of the Muskox is not elongate as in Bos or
triangular as in Ovis and Capra — — — —», but the words in italics have unfortunately

been dropped.

MATERIAL FOR THE STUDY OF RUMINANTS. 11

bos as well as from the very elongate spleen of the common ox and
of the Gnu.

The glandula thyreoidea of the Black-buck consists of two distinet
portions without any trace of an isthmus. Each body is elongately
beanshaped, about 20 mm. in length, 9 in breadth and 5 in thickness.
They are situated on the dorsilateral side of the trachea rather close
to the oesophagus and a little behind the cricoid cartilage.

The lungs of the Black-buck are divided into the same lobes
as those of the common goat but comparatively more deeply cleft.

The cartilaginous rings of the greater part of the trachea do
not meet behind, which is also the case in the Saiga according to
Murie'). The fibrous interval is, however, in this animal not broad,
as it is stated to be in the latter, but quite narrow. The rings are
thickest in front and become gradually thinner towards the sides and
are finally angularly bent so that the anterior surface is rounded the
posterior flat. A section of the trachea is thus semicircular. This
description holds good for the upper or anterior part of the trachea,
but on a level with the tips of the lungs the condition is altered so
that the rings meet behind and form an edge which becomes sharper
and more pronounced posteriorly.

The larynx (PI. I fig. 1) is rather elongate and compared with
that of a common goat it is large. In its general shape it resembles
the same organ of the Saiga as figured by MurıEe. The length of the
whole organ in this young Black-buck is about 54 mm. and its dorsi-
ventral diameter through pomum adami is about 32 mm. This latter
prominence is more bluntly rounded than in the common goat but not
so much so as in the Saiga. Its apex is situated about 1 cm. from
the hindmargin and this measurment is contained nearly three times in
its distance from incisura thyreoidea superior. This makes a great dif-
ference from the condition found in the common goat in which the
anterior portion of the thyreoid cartilage equals or is at most 11/2
times as large as the posterior one. In the Saiga again, as in this
animal, the anterior portion of the thyreoid cartilage is very long.
Cornua superiora are directed forwards their axis being parallel to the
longitudinal diameter of the larynx. Their length is about 7 mm.
They are rather narrow and in shape more similar to those of the
Saiga than to those of the Capra. Cornua inferiora are somewhat

1) Proc. Zool. Soc. London 1870 p. 495.

12 EINAR LONNBERG,

arcuate and comparatively small, they are shorter both than those of
the Saiga and those of the Capra, only measuring about 8 mm, in
length. The breadth of the cricoid cartilage is ventrally or anteriorly
9 mm. but the posterior or dorsal shield extends to a length of not
less than 33 mm. It has anteriorly a well developed crista. Posteriorly
it is continued in a thin flap. The arytenoid cartilage is stout and
thick as in the Saiga and has also as that of the latter a considerable
nodosity at the external cricoid joint. The larynx of Madoqua has a
quite different shape. It is short, the length of the thyreoid cartilage
measuring in the median line 8 mm. and that of the lateral thyroid
shields at most 12'/2 mm. The pomum adami is situated close to the
posterior margin. Cornua posteriora are short 2 !/» mm. in length by
|!» in breadth. They are directed obliquely forwards and a little
dorsally. Cornua inferiora are slightly arcuate 5 mm. in length. The
cricoid cartilage has no crista but a broadly rounded prominence.
The posterior or dorsal shield is 11 mm. in length and comparatively
broad. In its general shape the larynx of Madoqua is more similar to
that of Capra than to the same of Antilope although the prominentia
laryngea is more squarely truncate than in the goat.

The epiglottis of the Black-buck is triangular with bluntly poin-
ted and recurved apex. Its shape is rather similar to that of the
Saiga and Capra although rather more pointed than that of the latter.
Plica ary-epiglottica is strongly developed. On the whole the larynx
of the Black-buck is most similar to that of the Saiga and the great
leneth of the thyroid cartilage in these animals when compared with
the same organ of other ruminants is remarkable and as far as I
know only superseded by the same in the Muskox. The septum mem-
branaceum of the posterior nasal cavity is well developed in the Black-
buck as in the Gnu, but not to such an extent as in the common sheep.
Madoqua has no such septum membranaceum.

The scrotal sack is small with small testicles. These are elonga-
tely beanshaped or nearly spindle-shaped about 4 cm. in length, but it is
to be remarked that the animal was young, not yet mature. The thickened
terminal portions of the vasa deferentia is about 3 cm., or a little more
than that, in length and their transversal diameter at the thickest place
is between 4 and 5 mm. The lumen is not spongious looking, but
smooth, which, however, may be a sign of immaturity. These termi-
nal portions of the vasa deferentia are not as usual situated in the

MATERIAL FOR THE STUDY oF RUMINANTS. 13
same horizontal plan, but lie one above the other: the left being closest
to the urinary bladder. If this is normal or only an individual varia-
tion is difficult to say. But it may be remarked that on the figure of
the male genital organs of the Saiga reproduced in MuriE's paper
quoted above, a similar condition seems to occur and in that also, the
right tube lies dorsally above the left one. The ductus ejaculatorii open
into the urethra not quite 2 cm. from the urinary bladder. A longi-
tudinal median plica separates the openings of both ducts and another
similar pliea is situated on the lateral side of either opening. These
three plice extend 7 mm. in length but join at both ends and corre-
spond to a colliculus seminalis and it is continued in a distal direction
by a lower plica which at first is double and then simple. This con-
dition is rather similar to that found in the common goat. The »vesi-
cule seminales» are almost compact, glandular bodies consisting of a
number of follicles. When dissected, free from the surrounding con-
nective tissue and straightened, their length will be about 4!/2 em.,
but they are curved and doubled up on themselves so that their length
in this state before dissection is not much more than half of that men-
tioned above. It is undoubtedly these bodies which have been figured
by Murie in his description of the anatomy of the Saiga and termed
»prostate gland», but in the text he is dubious whether he shall re-
gard them as such or as vesiculæ seminales. The muscles, ischiocaver-
nosus, bulbocavernosus, retractor penis ete. are well developed in the
Black-buck. The sigmoid flexure of the penis occurs behind the middle
of the organ as in the Saiga. The præputial cavity is long about 10
em. and of an almost uniform appearance with longitudinal folds. The
distal portion included in the præputial prominence, about 1 cm. in
length, is a little different. It is namely beset with some few, fine
hairs scattered over the surface which does not seem glandular any
longer as is the case with the portion next to it in a proximal direc-
tion. The free end of the penis is short about 4 cm. (PL I fig. 2).
It is pointed but may nevertheless be referred to the ovine type ac-
cording to Garrop’s terminology. The end of the penis is not at all
swollen, but looks as if twisted from the left to the right side and the
end of the urethra is free, and turned up on the left side of the end
of the penis in a groove which then continues on the upper side to
the end. The urethral prolongation extends still half a centimetre
beyond the penis as may be seen on the figure (Pl. I fig. 2). Muries

14 EINAR LÖNNBERG,

description and figure of this organ in the Saiga is not quite clear,
but it seems as if the difference were not important although the free
prolongation of the urethra is shorter.

The kidneys of the Black-buck are short and thick, measuring
55 mm. in length 35 in breadth at the hilus and about 30 in thick-
ness. Exteriorly they are quite smooth and undivided. The right one
is situated a little more forward than the left one and is attached to
the liver by a ligament and capped by lobus caudatus as’ already has
been stated. A longitudinal section reveals the fact that there is only
one large mamilla formed by the fusion of perhaps half a dozen cones
on each side which, however, at their distal bases are well defined.
This is a resemblance with the condition found in the common sheep.
In the Saiga Murte talks of »some half dozen undefined pyramids and
infundibula» so that there seems to be a different arrangement. In
the Gnu as I have shown previously there is no trace of small pyramids.

For comparison may here be appended a few notes on the
male genital organs of Gazella rufina O. THomas and Cobus defassa
(Rürrert). This material has been brought home by the Swedish
zoological Expedition to Sudan and the White Nile. Afterwards pre-
sented to the Zootomical Institute by Dr. JÄGERSKIÖLD it was kindly
placed at my disposition by Professor WIRÉN,

The scrotal sack of Gazella rufina is pendent, constricted proxi-
mally, but short. It is closely covered with short whitish hair. About
21/2 cm. in front of the scrotum a pair of teats are situated 2!/» cm.
apart. The præputial prominence is found about 5 em. in front of the
scrotum. It is sparsely beset with short hair. The testes are elongate,
almost spindleshaped. The general arrangement of the vasa deferentia
is similar to the same of the Black-buck as described above, except
that they do not lie on top of each other but side by side at their
distal ends. They open on a longitudinal colliculus seminalis which,
however, is not divided in longitudinal plicee to such an extent as in
the latter animal. The vesicule seminales are also similar to those of
the Black-buck, but smaller, not measuring more than about 3 em.
when straightened. The glandulæ Cowperi are rather small, too, flat-
tened, roundish about 9 mm. along the greatest diameter. They open,
as is also the case in the Black-buck, in a little pocket, directed caudi-
ventrally, on the dorsal side of the urethra. The muscles are well
developed. The sigmoid flexure of the penis is short and situated at
the middle of the organ. The præputial cavity is about 512 cm. in

MATERIAL FOR THE STUDY OF RUMINANTS. 15

length. It is divided in two regions. The inner which is occupied by
the free end of the penis is about four em. in length, almost smooth
but provided with glands. The remaining distal portion which cor-
responds to the præputial prominence is longitudinally plicated and
towards the opening beset with some few scattered hairs. The end
of the penis is not so pointed as in the Black-buck and not so much
twisted (Pl. I fig. 3). There is a prominent cushion, quite well deve-
loped, which extends on the left side from below and behind, upwards
and forwards over to the right side and to the tip. The urethral pro-
longation lies in a groove mostly on the lower side of the penis, but
turns upwards towards the tip of the same and extends about 6 mm.
beyond it. The shape which may be seen from the figure is thus in-
termediate between that of the Black-buck and common sheep.

The kidneys are similar to those of the Black-buck and may
be termed as Murız calls those of the Saiga »subglobularly oval».
The left one is a little the larger and measures 53 mm. in length 37
in breadth and 36 in thickness. It is perfectly smooth. They were
covered by fat at the hilus as in the common goat.

The male genital organs of Cobus defassa were greatly mutilated
but permit, however, the following notes to be made.

The scrotal sack is pendent about 12 em. in length with vertical
testicles. Its proximal part is well constricted above the testicles. The
portion including the testicles is covered by long whitish hair which
forms a tuft at the apex. The constricted proximal portion is only
scantily beset with long hair, but instead covered by very short hair.
This latter covering extends also over an area in front of the scrotum
in which four small teats are found. The posterior pair of these is
situated 1!/2 em. in front of the scrotum and about 2'/2 em. apart.
The anterior pair is found about 11/2 resp. 2 cm. in front of the others
and sit 3'/2 em. apart. The teats are about '/2 em. in length in pre-
served state. In front of them the hair is again long, single hairs
measuring up to 12 cm. in length. The prepuce forms a prominence
on the ventral surface which in the present contracted state measures
1'/» em. in height 3'/2 em. in transversal and 4 cm. in longitudinal
diameter. This prominence is covered by hair, especially behind the
opening where there is a dense tuft about 4 em. in length, anteriorly
it is less densely hairy. The præputial cavity is differentiated into two
portions. The outer or distal one measures 61/2 em. in length and its
inner surface is strongly plicated longitudinally but is not pigmented and

16 EINAR LONNBERG.

does not appear to be glandular. The proximal or interior portion
about 7 cm. in length is sparsely dark-pigmented and its surface is
laid into partly longitudinal, partly quite irregularly wavy folds which
have a mucous aspect. In this interior portion the end of the penis
with its gland is enclosed when at rest.

The glans penis is black pigmented and has a peculiar shape,
but may be said to be of ovine type according to GARROD'S termino-
logy. It is reproduced in fig. 4 Pl. I and from this figure its shape
may be more easily understood than from a long description. It is
of interest to state that this type of penis also occurs among the an-
telopes of the Cervicaprine section. This type of penis is consequently
widely distributed among the Ruminants. As is stated above it occurs
in the members of the Antilopine section and in sheep and goats,
Garrop!) has deseribed it a little modified from Addax nasomaculatus
and from Cephalophus maxwelli, It thus occurs in the Hippotragine
and Cephalophine sections?) as well. The same author describes also
the same type from Giraffa and Moschus. It seems thus to be an ancient
characteristic not only confined to certain Cavicornia (conf. below where
this question will be more fully discussed).

The anteorbital glands of the young Black-buck consisted of a
6 mm. deep sack and the whole organ was not more than about 11/2
em. in diameter. According to Owen’) this organ can be partially
everted in the adult buck.

The same organs of Madoqua saltiana are, when compared with
the size of the animal itself, exceedingly much larger. They are also
larger in the male than in the female. In the former each anteorbital
gland is a lenticular body 28 mm. in length, 24 mm. in breadth and
Il mm. in thickness. In the female the corresponding measurements
are 20; 19; 8 mm. The interior of the gland as well as its secretion
is intensely black. From a comparatively small central pit wide duets
branch in different directions through the glandular mass. The whole
is surrounded by a fibrous and muscular layer which is not pigmented.
The structure of this organ thus seems to be quite different in these
two animals. In Cephalophus, as wil be mentioned below, there is
still another structure of this gland as it opens through several pores
arranged in a row, composed of many transverse series.

1) Proc. Zool. Soc. 1877.

?) Concerning the latter section farther communications are to be found below.
5) Anat. of Vertebr. III p. 632.

MATERIAL FOR THE STUDY OF RUMINANTS. 17

On the intestinal canal of Cephalophus ogilbyi and C. melanorhoeus.

From my friend, Mr. G. LINNELL I have had the pleasure of
receiving some viscera of the antelopes mentioned above, sent home
from the Cameroons, West-Africa.

The ventricle of Cephalophus melanorhoeus, »isello» of the native
hunters, preserved in formaline shows the four usual portions of a
ruminant stomach but very differently developed. The paunch (fig. 5
PI. I) is divided in the usual dorsi-sinistral and ventri-dextral sacks, but
both are rather shorter and more broadly roundish in a transversal
direction than is usually the case. This may also be seen from the
measurements. The dorsi-sinistral, which as usual is the shorter, has
a longitudinal axis measuring (from the cardia) about 135 mm. in length
and that of the ventridextral is about 170 mm. The breadth measured
from the well developed sulcus longitudinalis dexter is in the former
about 85 mm. in the latter about 97 mm. On the right (median) side
sulci coronarii are not visible, on the left the sulcus coronarius of the
ventri-dextral sack is well developed and forms the limit of a short and
broad saccus cecus measuring about 5!/» em. in depth. In the dorsi-
sinistral sack the saccus cecus is not differentiated by any sulcus coro-
narius. Sulcus longitudinalis sinister is only developed at both ends as
continuations from the right side which do not meet. It may thus be
said that the external sulci of the paunch are only little developed.

The reticulum (fig. 5 I r.) is almost ovate with the diameters
70 X 50 x 45 mm., the narrower end forming a blind sack towards
the abomasus protruding beyond the psalterium.

The latter (fig. 5 Pl. I ps.) is very small and may be termed a
rudiment measuring 25 mm. in length and 15 in thickness.

The abomasus (fig. 5 Pl. I ab.) measures along the middle of
its curvature about 85 mm.

The interior surface of the paunch is densely beset with tongue-
like papille which from a narrow basal stalk by and by broaden to
about 1'/2 mm. or a little more at the rounded free end. Their length
varies, of course, at different places, but seems, as a rule, not to
exceed 3!/» mm. The cells of the retieulum have the usual appearance.
Their diameter is, as a rule, 4—5 mm. and the walls are low, hardly
measuring more than 1 mm. (or less than that) in height. Secondary
septa are seen here and there. The folds of the psalterium seem at

Nova Acta Reg. Soc. Sc. Ups. Ser. III. Impr. !9/z 1903. 3

LS Eryar LONNBERG,

the first look to belong to two cycles. Between the seven large,
»primary», ones are, namely, in some loculi to be found lower »se-
condary», folds, in some these are rudimentary while in others they
can hardly be traced. This makes the condition suspicious and another
interpretation might be tried. It could be possible that all folds, large
and small. amounting to 10 or 11, together represent primary folds
some of which have been reduced more than the others. To judge
from the interspaces between the folds the first opinion seems to be more
readily upheld, but compare below the condition found in the foetus.
The broadest folds measure about 9 to 10 mm. The margins of the
folds especially of those next to the »Schlundrinne» are beset with
comparatively large recurved, and sharply pointed papilla. The lateral
surfaces of the folds have more stout conical papilla. The adjoining
wide portion of the abomasus is provided with about ten longitudinal
folds which, however, in the fundus are partly branching and ana-
stomosing with each other in an irregular manner. The folds cease
4——5 cm. from the pylorus. They are as well few in number as also
low, seldom exceeding 2'/2 mm. in height. The limit between the
psalterium and the abomasus is quite sharp. The ventricle of Cepha-
lophus ogilbyi (the »n'bindi» of the native hunters) differs in several
respects from that of the foregoing species. The transversal expansion
of the paunch is comparatively very considerable as the following
measurements indicate when compared with each other and those of
other ruminants. The distance from the cardia to the blind end of the
dorsi-sinistral sack is about 225 mm. and to the end of the ventri-
dextral ore about 260 mm., but the breadth across the fundus of both
sacks is not less than 290 mm. The ventridextral sack is the broader,
one being about 180 mm. across. Sulcus coronarius of that same sack
is well developed on both sides and defines the rather long and almost
cylindrical saccus cecus which measures about 95 mm. in length and
*5 mm. in basal diameter. On the left side there is also a trace of
a small saccus cecus of the dorsi-sinistral sack and the sulcus longitudi-
nalis is, unlike the condition found in the sheep, better defined in its
ventral branch. From this description may be concluded that the sulci
of the paunch in this animal are, on the whole, well developed and
much more so than in the preceding species.

The shape of the reticulum differs from that of the »isello» therein
that its, towards the abomasus directed, blind sack is the thickest

MATERIAL FOR THE STUDY OF RUMINANTS. 19

portion measuring about 65 mm. in diameter. The length of the organ
is about 110 mm.

The psalterium is much better developed than in C. melanorhoeus.
It is elongate almost cylindrical about 6 cm. in longitudinal and 3 in
transversal diameter.

The abomasus measures about 16 cm. along the middle of its
curvature.

The papille of the paunch have the same general shape as in
the »isello», but are, of course, larger, attaining a length of 5 or 6
mm. in some places, and a breadth of 2'/2 mm. at their apex. The
cells of the reticulum have a diameter of 10—13 mm., but their walls
are low in this antelope, too, hardly measuring 17/2 mm. in height.
The stellate secondary septa are partly well developed. The psalte-
rium is conspicuously thick-walled and muscular, There are 9 primary
folds which seem thick and are strongly armed by stoutly conical but
sharply pointed papillæ. Although the folds are thus in some respects
remarkably well developed, their height is strikingly low, mostly about
10 mm., or less, only one attaining the height of 13 mm. Thanks to
this the free lumen of the psalterium is unusually large in comparison
with the same of other ruminants. Only in three of the lateral loculi
I have seen small rudiments of as it seems secondary folds. The
interspaces between the folds are beset with long, slender, and recurved
papillae. The abomasus which is very sharply defined from the psalte-
rium, is provided with about ten (or twelve) longitudinal folds. These
are much more strongly developed in this species than in C. melanorhoeus
and attain a main height of S—12 mm. The exact number is difficult
to tell, because some folds branch off from others, and other anastomose.
In the fundus-portion there is also a number of smaller transversal
folds. The longitudinal main folds cease in a distance from the py-
lorus of about 8 cm.

A comparison between the ventricles of these two species of
of Cephalophus indicates that they differ in several respects with regard
to some details as has already been mentioned. On the whole, it
seems as if the ventricle of the larger species was a little better
suited to the digestion of coarser food than that of the smaller, but
even the larger of the two differs in several respects from the common
grass-eating ruminants. The capacity of the stomach is not so large
as could be expected when compared with the size of the animal.

20 EINAR LÖNNBERG,

Especially striking is the simplicity of the psalterium which hardly
deserves to be termed duplicate. These features indicate a diet differ-
ing from that of the common grass-feeding ruminants. The paunch
of the Cephalophus ogilbyi which I have had at my disposition was
unfortunately emptied, but I found, however, in the same, several
rather large bean-like (although more or less angular) seeds of pretty
hard consistency, in addition to these were some lumps of a fruit-pulp
which once may have surrounded the seeds and some few pieces of
leaves. Smaller debris could be derived from one or the other of
these three constituents. It is thus evident that fruits and seeds play
important parts in the diet of this antelope. In such a case it is
evident that the paunch does not need to have such a great capacity
as if the food had been composed of grass, leaves and other stuff
chiefly consisting of cellulose. The few but with strong conical pa-
pille beset folds of the psaltertum seem to be very well adapted to
the grinding of the remasticated and by the saliva thoroughly moistened
pieces of seeds.

The paunch of the smaller species C. melanorhoeus was filled
with large chunks of fruit pulp, some of which have the size of half a
walnut or more, large, more or less beanlike, seeds of several kinds,
probably partly belonging to the pieces of fruit pulp. In addition to
these some leaves were found which, however, were so few that
there is no danger in assuming that they have been swallowed with
the fruits mentioned above. From these facts may be concluded that
the »isello» is a more or less exclusively carpophagous animal, perhaps
and probably even more than C. ogilbyi. This may account for the
simple or little differentiated state of the paunch, the small and only
duplicate psalterium and the low folds of the abomasus. I think that it
might be assumed with a high degree of probability that Cephalophus
maxwelt in which Garrop and Boas found the psalterium in a ru-
dimentary state, as is mentioned above, owes this to a similar diet.
Compare also below the description of the foetus of these two species,
C. ogilbyi and melanorhoeus.

In the development of the different parts of the intestine of these
two animals modifications may also be observed which correspond
with those of the ventricle as related above.

The small intestine of Cephalophus ogilbyi has a length of about
1060 cm. its connection with the peripheric colic coil of the large in-
testine is short, as a rule about 2 or 3 em. The ileum enters into

MATERIAL FOR THE STUDY OF RUMINANTS. 21

the cecum about 15!/2 cm. from the blind end of the latter. This is
distinctly narrowed so that its diameter near the end is only about
12 mm. but near the ileocæcal opening about 30 mm. Ansa proximalis
of the colon has about the same width as the adjoining part of the
cecum. The colon forms three centripetal and two and a half cen-
trifugal central spiral coils which greatly cover each other and then
runs out in the common mesentery to form the peripherie coil. The
exact length of the large intestine is not possible to tell because it
has been torn off at the ansa distalis, but assuming the missing portion
(chiefly the rectum) to have been about 30 or 40 cm., in which estimate.
I cannot be much wrong, it might be put to about 315 or 325 cm.

The length of the small intestine of the Cephalophus melanorhoeus
is about 444 cm. The mesentery connecting it with the peripheric
colic coil varies between 1!/2 and 21/2 cm., usually it is about 2 cm.
The ilewn enters into the cecum about 12 cm. from its blind end.
The latter is narrowed at the apex, about 15 mm., but 23 mm. in
diameter near the ileocacal opening. The entire length of the colon
with the rectum is about 170 cm. The colon forms only one and a
half centripetal and one centrifugal central spiral coils, which all
lie in one plane and are comparatively loosely connected with each
other (fig. 7 Pl. I). Ansa proximalis of the colon seems to be com-
paratively long, measuring 7'/» em. which is only 1 cm. less than
the corresponding measurement of the larger species. As I unfortuna-
tely am not in possession of the body-measurements of the specimens
from which these viscera have been taken a comparison in detail
with those is impossible. It appears, however, to be evident that the
small as well as large intestine of C. ogilbyi is, even comparatively, a
good deal longer than the same of C. melanorhoeus, and that the colon
of the former is much more complicated is seen from the description.
There can be no doubt that it is the more exclusively carpophagous
diet which has caused the shortness of the intestine and the fewness
of the colic convolutions of Cephalophus melanorhoeus as well as the
less complicated state of the ventricle.

The liver of Cephalophus melanorhoeus presents the striking feature
of a missing gallbladder. This is in correspondence with Garrop’s')
similar observation on C. maxiwelli and C. pygmeus. It is thus by this
statement made still more probable that the whole genus Cephalophus
has no such organ (conf. also below).

1) Proc. Zool. Soc. 1877.

22 EINAR LÖNNBERG,

In general outline the liver of C. melanorhoeus is less transver-
sally elongate than that of C. maxwelli as figured by Garrop (I. c.).
Its longest diameter from left to right is 91 mm. the greatest width
(right lobe) is about 64 mm. and the greatest thickness (left lobe) is
30 mm. It is thus a rather compact organ. Fissura umbilicalis extends
about 2 cm. from the margin and not quite that on the diaphragmatic
side, and there are no other fissures. The Spigelian lobe is large,
triangular, with the rounded apex extending fully three cm. from the
posterior margin. The caudate lobe is also large, bayonet-shaped and
only coming 1 cm. short of the anterior free margin of the right lobe.
The right kidney is capped by the extreme right portion of the liver
and the caudate lobe.

The question why the gallbladder is missing in some animals
is most puzzling and the reason why it is missing in some but not in
others is certainly not easily explained. Owen put it together with the
shape of the stomach, but got no definite rule. It seems to me more
suitable to put the presence resp. absence in connection with some
certain kind of diet. The presence of a gall-bladder is the primary
condition, that can be concluded from the fact that such an organ is
found as well in the lower vertebrates as in the most primitive mam-
mals for instance Monotremata and Marsupialia. With the exception
of the whales all other mammals!) in which the gall-bladder is missing
feed on vegetable matter of some kind. If the gall-bladder is present
in one, but absent in another of two related genera the latter is, as a
rule more confined to a vegetable diet. An illustration of this is pre-
sented by the genera Sus with and Dicotyles without gall-bladder but
with a complex stomach indicating a vegetable diet. This seems to
indicate that the gallbladder is of less importance for herbivorous ani-
mals so that it may sometimes as in Giraffa or always as in Camelus,
Cervide ete. be absent. It is true that the gail-bladder is present also
in many herbivorous animals, as for instance in most Cavicornia, but
that does not disprove the hypothesis that it could be reduced without
inconvenience, while it seems certain that it would be more incon-

") According to the literature the gall-bladder seems to be absent. in the following
mammals (in addition to the whales) Bradypus tridactylus, Tapirus, Rhinoceros sondiacus,
Equus, Dicotyles, Camelus, Giraffa (sometimes), Cervide, Elephas, Lamnunguia, Mus, Cri-
cetus, Lemmus Echimus, Erethizon, Synetheres, Tamias all of which feed on vegetable sub-

stances, or if exceptions to that rule may take place, as with some members of the genus

Mus, this is secondary.

MATERIAL FOR THE STUDY OF RUMINANTS. Do

venient for earnivorous and omnivorous animals to lose it, as none of
them (except the whales) have reduced it. The lack of the gall-bladder
in Cetacea might be a characteristic inherited from their ancestors, which
may have had a differerent diet (compare the complex stomach).

A consequence of such a theory is that the gall-bladder may
within the same group or even genus become reduced independently
of the condition found in other members of the same group.

The kidneys of Cephalophus melanorhoeus are small, almost pear-
shaped with the narrow end directed backwards. Their axial length
is about 3 em. and the diameters of the anterior thicker end about 2
cm. When they are cut they show only one longitudinal mammilla.

The spleen of Cephalophus melanorhoeus is strongly convex as
in the Cervicornia and some other primitive ruminants but it is trian-
gular in outline wherein its shape somewhat approaches the same of
the ovine tribe; compare the figure (fig. 6. Pl. I) and also the descrip-
tion of the same organ in the foetus.

Notes on the morphological development of some foetal specimens
of Cephalophus melanorhoeus, ogilbyi and silvicultor.

The material to the following study I have had the pleasure of
receiving from my friend Mr. G. LINNELL, who has kindly sent it
home from the Camerons, and I wish therefore to express my grati-
tude to him.

1) A foetus of Cephalophus sp., probably melanorhoeus, 3 a. Mea-
suring from the forehead to the anus 61 mm. and from the snout to
the oeciput 24 mm. It is not in a very good condition, but the viscera
seem, however, to be pretty well preserved (comp. figs 12 & 13).
When the abdominal cavity is opened the liver seems to be the pre-
dominant organ. HFissura umbilicalis is situated in the median line and
the liver reaches perfectly across from right to left extending from
the diaphragma on the left side of the median line more than half
way to the pelvis (11 mm.) and on the right side (14 mm.) almost to
the pelvis. Between the liver and the groin on the right side the
colic spiral is seen quite pressed to the abdominal wall. Laterally from
the colic spiral a portion of the left kidney is seen and in front of it
a portion of the ventricle (the paunch). Behind the kidney the testicle
is situated. On the right side the liver occupies nearly the whole vi-
sible area but quite dorsilaterally the kidney is seen and behind the

24 EINAR LÓNNBERG,

same the testis. A remarkable feature is that the scrotal sack is al-
ready well developed, measuring, 2 mm. in depth and 3 mm. in antero-
posterior direction, Just in front of the scrotal sack is on each side a
single small wart indicating the rudiment of the nipples. The left
lung is fully separated into four lobes, the right into six and an
azygous one.

The paunch measures about 6 mm. as well longitudinally as
dorsi-ventrally. It lies, as is seen in the figure, close to the left body-
wall, ventrally it is bordered by the liver, behind by the colic spiral,
and more dorsally by the left kidney. Both sacks of the paunch lie
practically in the same vertical plane. The dorsal sack seen from
the left side is almost crescent-shaped and the sulcus longitudinalis is
well developed, it is especially deep proximally. Sulci coronari of both
sacks are also well developed and the two sacci ccc? are roundish in
shape and curyed towards each other so that they are in contact.
That of the ventral sack is larger. The reticulum is comparatively
very large, measuring almost 5 mm. in dorsiventral and 4 in antero-
posterior direction. The psalterium measures 31/2 mm. in length by a
little more than 2 in width. The abomasus does not offer any peculia-
rities. The small intestine makes first a quite short loop close to the
hilus of the liver then turns backwards forming a long loop which
reaches on a level with the hind margin of the kidneys then it returns
and enters as jejunum the common mesentery. To follow the intricate
whirls of this organ is not possible, they occupy, however, an almost
central and ventral position in the abdominal cavity. The colic spiral
is not yet fully formed as may be concluded from a comparison with
the following stages. It consists of about 11/2 coil in each direction.
lts situation is already mentioned above. The rectum is almost the
narrowest portion of the whole intestine. It runs quite straight in the
median line in the dorsal part of the abdominal cavity.

The comparative size and the position of the liver has already
been described. It is only divided in two main lobes by the umbilical
fissure. The caudate lobe is large and extends with its tip beyond
the margin of the right lobe. It caps from the ventral side not only
the whole right kidney but also the testicle and the epididymis. The
testicle is smaller than the epididymis and quite globular.

The spleen occupies the normal situation, but it has a different
shape from that in the adult. It is namely elliptic almost equally
rounded at both ends. The greatest convexity is situated near the

MATERIAL FOR THE STUDY OF RUMINANTS. 25

middle of the dorsal edge. It resembles consequently the same organ
of Cervicornia, Giraffa and Ovibos. The ontogenetic development of
this species as well as that of Cephalophus ogilbyi proves therefore,
that the triangular shape of the spleen is a secondarily aquired cha-
racteristic, as I have pointed out before.

2) A foetus of Cephalophus melanorhoeus 3 b., measuring from
the forehead to the anus 114 mm. and from the snout to the occi-
put 47 mm.

The fine bristles above the upper eyelid are beginning to appear
in two series. Some bristles on the sides of the lower jaw are also
developed and on a prominent wart or cushion on the chin, between
the rami of the mandible, four small bristles are developed. Otherwise
the specimen is naked.

The arched row of pores of the anteorbital gland seems to be
divided into 23 transversal groups, but the pores are not quite di-
stinct yet.

When the exterior of this foetus is examined the great size of
the scrotal sack is one of the most striking features. It measures
fully 11 mm. in depth and S mm. in anteroposterior direction. The
prepuce is also large, protruding about 31/2 mm. and with a trans-
versal diameter of equal size. Just in front of the scrotal sack or on
a level with its anterior border is, on each side, a single wart about 2
mm. in diameter. This is the rudiment which later should have de-
veloped into the teats and although there is only one such on either
side, each of them has distally an anterior and a posterior small tip
thus indicating the future division into two teats (see fig. 14 a). This
is important because it indicates that the four teats of Cephalophus
melanorhoeus are developed from only two rudiments, one on
each side, which becomes divided. The warts of this stage are namely
»Anlagen» of teats only, not of udder. This is proved by compa-
rison with a somewhat later stage in which the division is more pro-
nounced and with a still later in which four teats are seen, but no
trace of an udder. If the warts of this stage had represented an
udder and only the small tips had been the rudiments of teats, it
follows that also in a later stage an udder must have been present.
But as this is not the case the above statement must be correct, these
warts are only rudiments of teats and become later divided into two
(compare also next stage). Such a division is of course a rather
remarkable fact as it would seem more probable that a smaller number

Nova Acta Reg. Soc. Sc. Ups. Ser. III. Impr. ?!/7 1903. : 4

26 EINAR LÖNNBERG,

of teats was derived from a greater through reduction than the con-
trary. But on the other hand it is known that at least in the dome-
sticated cow the regular number of four teats may be inereased to six.

Concerning the situation of the abdominal viscera it may be
remarked that it has undergone a considerable change from the one
found in the foetus of the stage described previously under the mark a.

The liver is not by far so large, comparatively speaking. The
right lobe lies longitudinally close to the abdominal wall. The greater
part of the left lobe is situated to the left of the median line, but it
leaves not only the whole paunch uncovered, but also a great deal
of the reticulum and a portion of the abomasus. Behind the ventricle
are seen convolutions of the small intestine, but the colic spiral coils
are now entirely concealed, the intestinal mass having rotated towards
the left so that the colic coils have in this stage been turned inwards
from the former situation at the left wall of the abdominal cavity
which it occupied in stage a., towards the dorsal portion of the same.
The coils thus lie now in a horizontal plane instead of in a vertical
one as before. The left kidney has the same situation as before, but
the testicle has wandered backwards to the inguinal region. On the
right side the liver occupies a greater area and extends more than
half way to the pelvis. Dorsally and behind the liver the kidney is
found partly surrounded by the former organ. The testicle has changed
place as on the left side. Behind the kidney is therefore a coil of
the rectum to be seen between it and the testicle. In a medio-ventral
direction from these organs the cocum is found and then the numerous
convolutions of the small intestine.

If the different organs of this stage are compared with those
of the foregoing the following facts may be discerned. The paunch
has increased considerably in size. Its longest longitudinal diameter
measures about 17!/» mm. thus nearly three times as much as in the
stage a. while the body-measurements of the stage b. are not quite
twice as great as those of the stage a. The sacci cœci are not so
rounded as iu the foregoing stage, but more broadly extended, the
ventral one, however, being narrowed at the posterior end. The
reticulum has increased proportionally to the growth of the body as
its longest diameter is about 10 mm. The psalterium has almost done
the same, its longest diameter being about 6 mm. The reticular
structure of the inner surface of the reticulum is fully developed. The
folds of the psalterium are also developed.. In the abomasus some

bo

MATERIAL FOR THE STUDY OF RuMINANTS. fi
low folds are seen radiating from the opening to the psalterium. Three
such are found on the hepatie surface, two on the opposite and some
smaller ones along the lesser curvature. The beginning portion of
the small intestine forms the same two loops as described in stage a.
The convolutions of the small intestine occupy the central ventral
portion of the abdominal cavity. The caecum is found on the right
side close to the right kidney. It forms a semicircular curve with the
convexity turned forward and towards the kidney, the blind end is
directed backwards and inwards. ‘The beginning colon makes a short
ansa proximalis forwards, then returns and enters from behind, and
on the right side, the colic spiral, all the coils of which lie in one
plane. There are 2'/2 centripetal and 1?/4 centrifugal coils before the
large intestine leaves the central spiral and runs out in the peripheric
mesentery. When the large intestine has left this it makes a rather
long ansa distalis which reaches backwards to the middle of the left
kidney and then returns upon itself, passes from the right side (close
to the short duodenal loop) round and in front of the root of the me-
sentery and then again returns backwards as rectum. It is, however,
not straight but forms a broad transversal coil behind the right kidney
before it continues to the anus in the median line again. It is inte-
resting to note that the foetus in this stage has a more compli-
cated large intestine than the adult as it possesses one centri-
petal and three fourths centrifugal spiral coils more than the latter
and in addition to this a transverse coil of the rectum. This indicates
that Cephalophus melanorhoeus has descended from ancestors which
have had a more complicated arrangement of the large intestine, but
that a secondary simplification!) has taken place later, no doubt
caused by a changed diet, formerly herbi- or graminivorous, now
(at least to great extent) carpophagous.

The situation and extension of the liver of this stage has al-
ready been reported above. The Spigelian lobe is rather large and
trihedral in shape. The caudate lobe is large, but it does not reach
the posterior end of the right kidney, still less the™ testicle. The latter
has grown and is no longer globular but elliptic in outline and some-
what flattened. If the scrotal sack is cut transversally it shows itself
to be fully developed with two apartments separated by the septum
of tunica dartos, In each appartment is an inner bag, the cremaster-

*) Or retrograde adaptation as it also might be called.

28 EINAR LÖNNBERG,

sack, constituted by tunica vaginalis, but this is empty as the testicles
are still situated in the abdominal cavity. The penis has already
nearly assumed its definite shape which will be described more fully
under stage f. The only difference in this stage lies therein that the
filiform urethral prolongation is comparatively less developed. It reaches,
however, a good deal beyond the tip of the glans.

About 11 palatal elevations are developed on either side but
they are all smooth, not crenulated as in later stages.

3) A foetus of Cephalophus melanorhoeus @ c., measuring from
the forehead to the anus 140 mm., and from the snout to the occi-
put 52 mm.

In addition to the hairs and bristles of the stage b. there are
several hairs developed on the lips and on the chin of this foetus,

The arched row of pores of the anteorbital gland shows di-
stinctly about 25 transverse series each consisting of three pores.

There are already two pairs of teats each measuring 2 mm.
in length. The distance between the two pairs is about 4 mm. The
members of the posterior pair sit close together, only 11/2 mm. apart,
the anterior ones 3 mm. apart. A vulva and a small protruding cli-
toris is developed. It is interesting to find that in this female foetus
the teats are four in number at an earlier stage than in the male, as
can be seen by comparing this and the following stages. This is quite
natural as the teats are more important organs for the female than for
the male, in which latter they are only rudimentary organs. This
latter fact could also be taken as an explanation for the condition
found in the male specimens. The development from two to four teats
should then only prove the variability of rudimentary organs.

The general arrangement of the abdominal viscera of this foetus
is nearly the same as in stage b. The kidneys are surrounded at
their posterior ends by considerable quantities of fat. The ovaries are
situated about 1 cm. behind the kidneys. |

The large intestine seems already to have begun its retrograde
development as there are not quite 2!/2 centripetal and 1!/2 centri-
fugal coils. But the transversal coil of the rectum behind the right
kidney still exists. The latter organ is ventrally, to a great extent,
covered by the liver, the caudate lobe of which protrudes beyond the
margin of the right lobe. The spleen is slightly beginning to assume
a triangular shape, although this is as yet only little pronounced.

MATERIAL FOR THE STUDY OF RUMINANTS. 29

The right lobe of the liver does not seem to be so much longi-
tudinally stretched as in the foregoing stage.

4) A foetus of Cephalophus melanorhoeus 3 d., measuring about
212 mm. from the forehead to the anus, and 79 mm. from the snout
to the occiput.

This foetus is hairy all over although the hair is best deve-
loped on the head and the neck. The bristles above the upper eyelid
stand in two series and are about a dozen in number. Behind the
muzzle on the upper lip and on the lower lip and the chin there are
several scattered fine bristles. On the wart under the lower jaw there
are four principal bristles and some small additional ones.

The scrotal sack measures 11 mm. in depth and 9 mm., in
anteroposterior direction. Just in front and on the sides of the scrotum
the teats are seen. They have not yet become fully divided into four.
But the »Anlage» of each side is half way cleft into two (fig. 14 f»).

Concerning the general situation of the abdominal viscera the
most conspicuous feature is that the liver has grown comparatively
less than other organs. The abomasus has increased in size a good
deal. It is very little covered by the left hepatic lobe, but extends
itself somewhat over the paunch.

The fat round the kidneys has increased a good deal, other-
wise the general appearance is about similar to that of stage c.
The testicle of the right side which seems to have grown compara-
tively little occupies the same situation as in stage b, in the inguinal
region of the abdominal cavity, and between it and the kidney the
transversal coil of the rectum is seen.

The longest diameter of the paunch is about 28 mm. The re-
ticulum has almost the shape of a low three-sided pyramid, the longest
basal (transversal) diameter of which is 16 mm. The longest diameter
of the psalterium measures about 81/2 mm.

The colic spiral is formed by 2!/4 centripetal and 11/2 centri-
fugal coils. The reduction from the foregoing stage is consequently
not great, but nevertheless visible.

Concerning the interior of the different divisions of the ventricle
of this stage it may be remarked that the folds of the psalterium
count about ten in number, but some of these are small and low like
secondary folds. The abomasus has about the same number of folds
as in the adult.

30 EINAR LÖNNBERG,

The liver has about the same general shape as in the foregoing
stage. The broad caudate lobe caps the kidney. The Spigelian lobe
is comparatively very large. It is not trihedral as in stage b, but
expanded into a roundish knob which is thicker than its basal junction
so that it seems pedunculated.

The spleen does not seem to have advanced much more towards
its definitive triangular shape than in the foregoing stage.

5) A foetus of Cephalophus melanorhoeus 3 e., measuring from
the forehead to the anus 210 mm., and from the snout to the occiput
77 mm.

Although this foetus is a little smaller than the foregoing, it re-
presents a somewhat later stage which is seen by comparing the two
specimens with each other. The hair of this one is better developed
on the body and legs etc. The teats are fully cleft into two on each
side and descensus testiculorum has taken place. Situs viscerum is about
the same as in stage d. The liver is perhaps a little more moved to
the right side and the kidneys are wholly covered by fat. The trans-
versal coil of the rectum behind the right kidney is fully straightened
and has thus disappeared.

The colic spiral has the same number of coils as in the fore-
going stage. The Spigelian lobe of the liver is in this specimen a
flattened tongue-like body with a triangular outline. A comparison of
the shape of this lobe in the three stages termed b, d, and f. proves
that the shape of this organ is much too variable to be of any syste-
matic value as Garrop') once believed. To use the terminology of
this author the Spigelian lobe of stage e. should be called »oviform»
and that of stage d. »rusiform». The adult specimen I have had at
my disposition has according to the same terminology an »oviform»
Spigelian lobe. Of the two species Garrop of Cephalophus himself
examined one, C. maxwelli, had a »modified rusiform» Spigelian lobe,
the other C. pygmeus, had it »modified or absent».

The spleen is rather flattened but it still retains the elliptic outline.

The penis has assumed its definite shape which will be describ-
ed for the next stage.

6) A foetus of Cephalophus melanorhoeus, 3 f., fully developed
and almost ready to be born: measuring about 25 em. from the fore-
head to the vent, 87 mm. from the snout to the occiput.

1) Proc. Zool. Soc. 1877.

MATERIAL FOR THE STUDY OF RUMINANTS. oil

Along the anteorbital gland there is a shallow groove in the
skin about 15 mm. in length and in this groove there is a series of
23 rather large pores in a nearly straight row and on either side,
upper and lower, of each of these there is also a pore, although much
smaller and less conspicuous. These ducts represent the openings of
the anteorbital gland which is somewhat pigmented and surrounded
by a fibromuscular layer as usual.

The upper lip has a vertical groove which is continued into the
mouth where it separates to form a circular groove round the papilla
incisiva. This latter is ovoid in shape with the narrow end directed
forward. A lateral groove branches of towards each side from the
nasal groove before this has reached the papilla. This is thus a differ-
ent condition from that prevailing in the Madoqua. The palatal ridges
are about ten in number, of which the three or four foremost ones
are much more prominent than the others, but the posterior ones are
more distinctly crenulated. There is no caruncula sublingualis only a
straight fold.

The papille filiformes of the tongue are scale-like, not pointed.
On the posterior back of the tongue they partly form flat warts, partly
broad scales, and there are no conical or eurved ones. In the anterior
half of the tongue the papillary covering extends a good deal over
the lower surface, but is there quite similar to that of the upper surface.
The papille fungiformes are evenly scattered over the tongue with the
exception of the median portion of the posterior half.

The papille circumvallate are arranged in two series with 5—6
in the inner and 3—4 in the outer row of either side. This is a re-
markable difference from » Antilope» (Cephalophus) mergens which ac-
cording to BRÜCHER as quoted by Opps!) is said to have 18—20 small
papillæ circumvallate an either side. But it may have been difficult to
see the difference between such papillae and p. fungiformes.

The glandula thyreoidea is a flattened bean-shaped body situated
on each side of the trachea just behind the larynx near its dorsal sur-
face. There is no isthmus and the dimensions are about 6 x 4 x 2 mm.

The thymus-gland is formed by two elongate bodies situated
just in front of the first pair of ribs and with their posterior fused
portion entering the thoracic cavity. The length of the whole gland
is about 27 mm. and the separate anterior ends measure about 11

1) Mundhöhle, Bauchspeicheldrüse und Leber, Jena 1900.

32 EINAR LÓNNBERG,

mm. When a comparison is made with the following stage it is evi-
dent that these two organs are not yet fully developed.

The posterior ends of the rings of the trachea are united in
such a way as to form a sharp edge which is directed towards the
right side and backwards. The left lung is divided into three lobes,
the right in five or nearly six and an azygous or intermediate lobe.
The lungs are thus much divided. The lobus apicalis of the right lung
is divided into an anterior and a posterior lobe and each of these has
in its anterior margin a slit which in the anterior lobe extends through
more than three fourths of the lobe. The lobus cardiacus of the right
lung is perfectly divided into a dorsal and a ventral lobe.

The liver has a short umbilical fissure. The right lobe is a
little larger than the left. The caudate lobe is long, trihedral and
reaches the margin of the right main lobe. It caps the kidney. The
Spigelian lobe is only little developed. There is, of course, no trace
of a gall-bladder.

The ventricle and the intestine resemble in their relative size
and arrangement so closely the same of the following stage g. so that
it seems unnecessary to repeat the description of these organs twice.

The penis has a very characteristic structure as may be seen
from the figure (fig. 21). The urethra runs as usual along the lower
side of the organ, but at some distance (about 5 mm.) from the tip
of the glans it makes a sharp transverse twist towards the left, ascends
somewhat on the left side, then bends sharply forwards and runs as
a free filiform and tapering prolongation first in a groove on the left
side of the glans, at the end nearly reaching the median line; then it
continues a good deal (about S mm.) beyond the apex of the glans.
The real glans is formed by a rather large cushion or swelling which
thanks to the general twisting of the whole organ has been turned
over towards the right side. It forms also the whole of the apical
end of the penis and its relation to the urethral prolongation has al-
ready been mentioned and may be seen from the figure. In Cepha-
lophus maxwelli GaRROD has described!) a similar shape of the penis
(termed »ovine» by him) but he says that »it is more symmetrical».
I am inclined to think that this alleged symmetry is due to a less good
condition of his material. The shape of this organ of Cephalophus
melanorhoeus is most similar to that which I have described above

1) Proc. Zool. Soc. London 1877.

MATERIAL FOR THE STUDY OF RUMINANTS. 33

from Cobus defassa. As has been stated above I am inclined to think
that such a development of the penis as this one is the primitive one,
at least among the Cavicornia, and aberrant structures of this organ I
regard as being due to a secondary transformation usually in connec-
tion with a reduction in a more or less high degree, of the urethral
prolongation. In the members of Cervicornia investigated by GARROD
(1. c.) the penis end seems to differ widely from this type, being sym-
metrical, although otherwise varying in shape. But as Moschus and
Giraffa in this respect belong to what GARRoD calls the »ovine» type,
it is probable that the condition found in the Cervicornia also is due
to a secondary reduction. This becomes the more probable when a
comparison is made with the Tragulina which often are regarded as
the most ancient among the Selenodontia. The specimen of Tragulus,
which I have had the opportunity to dissect, was not very well pre-
served, but the following observations could, nevertheless, be made.
There are to be distinguished at the distal end of the penis a cushion-
like organ and a terminal tapering portion. The former which, no
doubt, may he homologised with the cushion found on the penis of the
Ruminants enumerated above has, however, a somewhat different si-
tuation although its main part lies on the upper side of the organ as
usual. It extends along that side of the penis end and tapers in a
forward direction. Posteriorly it is thicker, and its posterior end being
to some extent free, is recurved on itself and fits in into a groove on
the left side between the middle portion of the cushion and the penis
itself as is seen in the figure (fig. 20). The tapering portion includes
the urethra and is homologous with the more or less filiform urethral
prolongation of the above quoted Ruminants. The difference lies only
therein that it is more spirally twisted than the same organ of the
typical Ruminants and less strongly tapering than in the »ovine» type
of GARRoD. In Tragulus this urethral prolongation, to use the same
word as has been used above, at the end of the penis turns sharply
upwards on the left side. When it has come almost to a level with
the next posterior portion of the penis with the above mentioned cushion
it curves forwards and downwards again on the right side. Then it
ascends for the second time on the left side, but much less abruptly
and in a forward-sloping direction. The extreme tip is finally some-
what twisted, although its main direction is straight. The penis-end of
the Tragulus is accordingly shaped like a cork-screw. But in this re-

Nova Acta Reg. Soc. $c. Ups. Ser. III. Impr. 74/7 1903. D

34 EINAR LONNBERG,

spect it highly resembles the likewise twisted organ of Sus!). It is
thus interesting to find that Tragulus also in this respect in its mor-
phological structure forms a connecting link between Suina and the
typical Ruminants. But at the same time it is made more than pro-
bable that this twisted shape of the penis is the original one among
all Artiodactyla. In such a case the »ovine» type (shared by Giraffa,
many antelopes etc.) although it looks more complicated and specia-
lised is proved to be less modified and more primitive than the types
exhibited by different Cervicornia, Bos etc. in which the twisting has
more or less completely disappeared and the urethral prolongation
has been reduced, The »filiform appendage» resp. urethral prolonga-
tion of the Ruminants is consequently homologous with the serew-
shaped part of the penis-end of Sus.

7) A foetus of Cephalophus melanorhoeus © g., of about the same
size as the foregoing, but with somewhat longer legs and with better
developed pellage and most probably quite fully developed.

The anteorbital gland shows about 22 transverse series of pores.
There are 11 fully developed palatal ridges on either side.

The papillary covering of the tongue is as in the foregoing
specimen with the exception that the papille circumvallate are are even
less numerous. To the left there are five in the outer, but only three
in the inner series, but on the right side both rows only contain three
papille circumvallate. 11—12 palatal elevations are well developed

!) The same organ of Hippopotamus has essentially a similar structure. A prepa-
rate of such a one which Professor Wiréx has kindly put to my disposition had the follow-
ing appearance. The free end of the penis consists of two different portions. The proximal
one, measuring about 26!/» cm. in length, is much the thicker with a vertical diameter of
61/2 and a transversal of about 6 cm. This basal half passes rather abruptly into the distal
half, measuring 26 cm. in length. This one tapers gradually towards the blunt tip which
has a diameter of about 1 cm. The proximal end of this distal half is exactly half as thick
as the proximal half in either direction. The distal half corresponds to the »screw» of Sus
and the urethral prolongation of certain Ruminants, but at the first look it does not seem
to be twisted. A closer investigation reveals, however, that it is so, which may be seen
from the fact, that a thin fold which originates at the limit between both portions on the
upper side in the median line, gradually turns over to the right side, and although abruptly
decreasing in size (for the last 12—14 cm. being only a slight longitudinal elevation) termi-
nates on the lower side of the extreme tip. This indicates that the distal end of the penis
is twisted half a turn. The height of the fold mentioned varies, where it is well developed,
from 5 to 7 mm. In spite of the great size and clumsiness as well as certain peculiarities
of this organ it is easy to homologise the different parts with the corresponding ones of Sus
and Ruminants.

MATERIAL FOR THE STUDY OF RUMINANTS. 35

on either side. All of them are crenulated but the eight posterior
more so than the foremost ones.

The glandula thyreoidea is a good deal larger than in the fore-
going and measures 10—11 mm. in length by 6--5 in breadth. There
is no trace of an isthmus.

In a similar way the thymusgland is larger and better developed
than in the foregoing. Its length is about 50 mm. and the paired an-
terior ends reach on either side all the way up to the larynx. This
organ thus continues its development, at least during the whole foe-
tal time.

The lungs are very deeply cleft so that the right lung has six
and the azygous lobe, the left three lobes. The lobes of the right
lung is formed in such a way that lobus apicalis is fully divided into
two anterior and a posterior lobe which latter also has a rather deep
fissure in its anterior margin. Lobus cardiacus forms two lobes, a
dorsal and a ventral, but lobus diaphragmaticus is entire.

The liver is like that of the foregoing specimen. The caudate
lobe is trihedral and very large and massive. The Spigelian lobe is
very short and rounded, hardly developed. The Spigelian lobe seems
thus, as is already stated above, to be subjected to great variation. The
greatest transversal length of the entire organ is about 48 mm. and
its greatest breadth about 32 mm. This is about half the size of the
same organ in the adult. Its situation is somewhat oblique, it is not
quite transversal, but on the other hand not quite longitudinal as in
some antelopes.

The distance from the cardia to the end of the ventridextral
sack of the paunch is about 34 mm. Swlei coronarii may be seen as
well on the left as on the right sack, but much better developed on
the latter, as is also the case in the adult, and its saccus cecus is
broad and well defined while it is small and less defined on the left
sack. Compared with the earlier stages, for instance 6. it seems as
if the ventridextral sack should have grown more than the other.
The whole paunch seems also somewhat broader compared with its
length than in the earlier stages. The longest diameter of the reti-
eulum is about 20 mm. This organ, as well as the paunch, is thus
about twice as large as in stage b. The exact measurement of the
psalterium is a little difficult to ascertain, but it seems to be about 11
mm. and it has then grown rather less than the paunch and the re-
ticulum. The abomasus is, however, the portion of the ventricle that has

36 EINAR LONNBERG,

grown comparatively most, which is quite natural as its functions shall
begin first. If the relative dimensions of the different portions of the
ventricle of this foetus are compared with the corresponding ones of
the adult it is apparent that the psalterium of the latter is compara-
tively much smaller than in the foetus. This is made evident by the
following statements: in stage a. the longest diameter of the psalterium
is 70 °/o of the longest diameter of the reticulum; in stage b. the same
0/0, but in the adult only 35 ?/o. The
psalterium of the adult is by this proved to be a reduced organ, which
fact no doubt stands in connection with the changed diet as has been

relation is 60 °/o; in stage g. 55

said above.

The general arrangement of the intestine is about the same as
in the adult. The centripetal coils are two in number, and the cen-
trifugal ones 17/2 The rectum is quite filled by fecal matter and
this may at least to same extent account for the fact that the trans-
verse coil of the rectum behind the right kidney which seemed redu-
ced in the foregoing stages is well developed in this specimen. This
explanation is, however, not quite satisfactory, for in stage f. the rectum
is expanded as well, without forming a coil but only a very short
curve. It seems consequently to prevail some variability with regard
to this.

The kidneys are quite simple without any trace of exterior
lobulation and have only one longitudinal mamilla.

With regard to the development of the teeth in Cephalophus melano-
rhoeus the following notes may be made. In the earliest stages represen-
ted in the collection there are, of course, no teeth developed as yet. In
stage b. for instance a low and even ridge-like prominence in the incisive
tract of the mandible includes the rudiments of the future teeth. This
edge is at the lateral and posterior end bordered by a rather large termi-
nal papilla (see fig. S) which already may be traced in stage a. In stage
c. the ridgelike prominence is beginning to divide into separate papillæ
corresponding to the different teeth, but the large posterior papilla is
unaltered. In stage d. (see fig. 9) this division into papillæ is complete,
and on either side of the median line five papilla may be counted,
four of these are derived from the continuous edge and the fifth is the
posterior terminal papillæ. All five papillæ of either side sit close to
each other and have quite the same appearance. It could therefore
be supposed in this stage that they were homologous all of them and
that the fifth posterior papilla contained the rudiment of same premolar

MATERIAL FOR THE STUDY OF RUMINANTS. aah

that did not develop further. That is, however, not the case. Sections
through the whole mandible of stage d. reveal that there are only to
be found rudiments (»Anlagen») of the four normally present teeth of
the milk-dentition and the small rudiments of the corresponding four
teeth of the permanent dentition. The fifth posterior papilla has no-
thing to do with those teeth. In stage d. the anterior median papilla
corresponding to the median incisor is the largest, because this incisor
is the largest, but the other four papilla are about equal. In stage e.
the teeth are beginning to cut the gum and can be felt with a pair
of pincers, the fifth posterior papilla is now the smallest. It is, however,
well conspicous even in later stages, for instance in stage f. (see fig. 10),
and also in stage g., but now it is quite evident that it has nothing
with the teeth to do as those are already fully developed as far as
their visible parts are concerned.
The structure of the incisors
of Cephalophus melanorhoeus is quite
interesting with regard to the distri-
bution and development of their
enamel. There is namely on the
anterior surface of the teeth quite
a thick layer of enamel all the way
to the cutting edge of the tooth.
But on the posterior or inner sur-
face of the incisor the enamel-layer
is of such extreme thinness that it
ean be'said to be practically ab-
sent"). This is thus a structure
that in some degree reminds one e. enamel layer of the outside, Longitudinal section
DL TIG ame he ncrsonspofthp- a tee meer 0 pme melon showing sedat.
Rodents and it is much more ad-
vanced in such a direction than for instance the same of the lamb or
common ealf. Concerning the two latter I have shown before, that
the enamel on the inner side of the incisors is strongly reduced if
compared with the condition prevailing in such a primitive Ruminant
as Ovibos, in which the difference in thickness of the enamel-layer of
the anterior and posterior (interior) surface of the incisors is not so

*) The preparates, on which this statement is based, have been made of the median
incisors of a foetus fully developed, but not yet born so that the absence of the enamel
could not be due to wearing!

38 EINAR LONNBERG,

very remarkable!). Cephalophus now proves to be the most advanced
type of those mentioned here as far as the development of the inci-
sors is concerned. An arrangement of the enamel of the incisors in
such a manner as has been described above for Cephalophus melano-
rhoeus serves of course to keep the cutting edge of the teeth sharp
while the teeth are worn. The great value of this sharpness is appa-
rent, because, thanks to this, the animal is able to cut off pieces of
such fruits that constitute its food.

It is evident that the brain of the above described specimens
could not be in very good condition as the specimens have been pre-
served in toto, but some short notes about this organ may never-
theless be of value as nothing is known before about it. In stage a.
the furrows do not seem to be developed as yet. The brain of stage 5.
is figured from the side and the left half from above (fig. 23 a,b). In
this stage the mesencephalon is not yet covered by the prosencepha-
lon. On the side of the prosencephalon a triangular depression is
seen, which corresponds to fissura Sylvii. The upper point of this
triangle is according to Krurg’s?) terminology processus acumimis. The
anterior and posterior lower angles are produced resp. in proc. anterior
and posterior as the author quoted describes from the sheep-foetus.
The processus posterior is confluent with and continued without inter-
ruption with the fissura postrhinalis to the posterior end of prosence-
phalon. In this respect the brain of this Cephalophus seems to differ
as well from that of sheep and ox (Kmvza l. c.) as that of Ovibos?).
In the middle of the triangular depression the uncovered insula Reili
is seen. Higher up on the lateral side of the prosencephalon sulcus
suprasylvius is seen. At its posterior end it is forked. The upper
branch of this fork corresponds to what Krure has termed processus
posterior s. ss. The lower branch of the fork extends somewhat more
downwards and backwards than in the sheep and ox (KRUEG) but less
so than in Ovibos (RETZivs) or Sus. It is however probably an an-
cient characteristic. ROGNER*) found in some instances such a branch
also in the sheep. The anterior continuation of the sulcus suprasylvius

1) Compare descriptions and figures in my paper. »On the Structure and Anatomy
of the Muskox». Proc. Zool. Soc. London 1900.

?*) Ueber die Furchung der Grosshirnrinde der Ungulaten. Zeitschr. f. wiss. Zool.
Bd. XXXI. Leipzig 1878.

?) Comp. Rerzivs: Biol. Unters. N. F. Bd. IX N:o 8. Stockholm 1900.

*) Zeitschr. f. wiss. Zool. XXXIX. Leipzig 1883.

MATERIAL FOR THE STUDY OF RUMINANTS. 39

is seen in the figure (fig. 23 a) to form a rather deep depression cor-
responding to sulcus coronalis from which a short branch turns laterally
downwards but the main sulcus continues forwards and downwards
finally dividing in two short branches. This arrangement does not
seem to agree fully with the corresponding one in foetal specimens
of sheep and ox (Kruzs). On the other hand it offers some likeness
with the brain of Tragulina and some deer in which sulcus coronalis
and suprasylvius are connected. On top of the prosencephalon of the
Cephalophus-foetus is seen an almost straight furrow which according
to KRUEGS terminology should be sulcus lateralis and which has about
the same situation as in foetal specimens of sheep and ox. Just at
the border between the upper and median surface of the prosencepha-
lon extends a furrow beginning at the hind-end of the organ on the
posterior surface and running as has been mentioned. The posterior
portion is rather better visible from above than the anterior fourth but the
foremost end curves up again on the upper surface not far from the
anterior end of the foremost fourth of the prosencephalon. This furrow
must correspond to sulcus splenialis and its situation as described above
is of great interest because is it quite different from that in Ovis and
Bos in which genera the said sulcus is confined to the median sur-
face even in foetal stages. According to Rerzius this latter is the
case in the adult Ovibos and according to KruEG in Ovis, Capra, Rupi-
capra and other Cavicornia as well and also in the Cervicornia which
he has investigated. But on the other hand Gazella dorcas »Tragulus
sp.» and Moshus moschiferus have this furrow visible from above as in
the foetus of Cephalophus melanorhoeus. This is no doubt an important
feature indicating the Cephalophus to be rather primitive in this respect.

The brain of a more advanced stage (stage f.) is still more
badly preserved, but the following can be observed. The general
shape of the whole organ is very elongate. The length of each he-
misphere is about 44 mm. measured in a straight line and in the same
way the greatest width of the dorsilateral surface is only about 23
mm. Each hemisphere has thus the shape of a section of a pear.
This elongate shape may also be regarded as a primitive characteristic.
Rhinencephalon is very large. Each of the bulbi projects about 8 mm.
in front of the prosencephalon. The furrows of this stage have pro-
bably attained almost the same development as in the adult.

The fissura Sylvii is represented by a deep vertical cleft cor-
responding to the processus acuminis of KruEG. Below, it divides as

40 EINAR LÖNNBERG,

usual in an anterior and a posterior branch. The former of these can
be followed just above fissura rhinencephali almost to the anterior end
of the prosencephalon just as in Ovibos according to Rerzius’ figures
(l. c. Pl. XXIII figs 3 and 5). The sulcus suprasylvius is very strongly
curved around the proc. acuminis fiss. Sylv. In this respect Cephalophus
differs widely from Moschus (Krurg’s Taf. XXI 1. c.), but agrees better
with Gazella. It is uncertain whether the strongly curved shape of
this furrow is a primitive characteristic among the Ungulates or not,
when one considers that it is comparatively straight in Moschus and
Cervicornia but strongly curved in Sus, Ovibos and in some Antelopes
ete. On the whole it seems to vary a good deal in Cavicornia. From
the anterior portion of the sulcus suprasylvius extends in this Cephalophus
a branch which continues vertically to the median line. This is the
processus (KRUEG) or ramus (RETZIUS) superior of this furrow. How
the posterior ramus of this furrow ends cannot be ascertained. The
sulcus coronalis Stands in open connection with the upper ramus of
sulcus suprasylvius. In this respect it seems to agree with the condi-
tion found in Gazella according to Krure (1. c. Taf. XXII), but differs
from the ovine and bovine tribes. The condition prevailing in Cepha-
lophus seems to be the primitive one as it is also to be observed in
several Cervicornia as well as in Moschus and Tragulus according to the
drawings of the author quoted.

As the homologon of the sulcus ansatus I take a short trans-
verse furrow which in front of the ramus superior sulc. suprasylv. from
the median line extends transversally towards sulcus coronalis without
quite reaching the same.

On the posterior half a well developed sulcus lateralis is seen.
Parallel to the same and between it and the sulcus suprasylvius another
well developed furrow is seen, although shorter. Such a furrow is in
a similar situation represented in Krura’s figures of the brain of Ga-
zella and of that of his Lophotragus (Elaphodus). But in the others the
arrangement seems to be different. On the median side of the sulcus
lateralis there is a faint longitudinal impression indicating that in the
adult a furrow might be developed there as in Gazella, Rupicapra,
Capra ete. according to KruEG. The sulcus splenialis is also in this
stage somewhat visible from above as in Gazella.

Although the short notes about the brain of these specimens
are incomplete they tend to show that there is a somewhat greater
likeness between Cephalophus and Gazella with regard to this organ

——— —

MATERIAL FOR THE STUDY OF RUMINANTS. 41

than between the former and other Cavicornia. At the same time the
brain of Cephalophus shows several primitive characteristics and is less
specialised than the same of the ovine and bovine sections.

8) A foetus of Cephalophus ogilbyi © h., Measuring from the fore-
head to the anus 180 mm. and from the snout to the occiput 75 mm.
It is quite naked, the rhinarium and the posterior third of the ear is
pigmented. The bristles above the upper eyelid begin to appear and
the cushion between the rami of the mandible is quite plainly promi-
nent, although the bristles are not developed.

The pores of the anteorbital gland form a straight series of 13
groups of pores. Each group seems to be constituted by a trans-
versal (vertical) row of three or four pores, which sometimes are not
regularly arranged.

The situs viscerum is similar to that of the specimen described
below and termed 7., but the liver is comparatively larger. A com-
parison of the different organs of this foetus and the following termed
i. will be made further below. |

9) A foetus of Cephalophus ogilbyi © i., Measuring from the fore-
head to the anus about 250 mm., and from the snout to the occiput
about 98 mm. The eyelids are closed and the body is naked, but on
the head there is some hair, especially on the top of the same, on
the lips and the chin. From the region between the eyes short, partly
blackish, but also partly rufous, hair covers the forehead up to the
horn-centres or the places where the horns should appear later. It
is directed backwards but on the sides obliquely so that, so to say, a
fanshaped whorl is formed. In the median line the black, on the sides
the rufous colour is dominating. Posteriorly the hair increases in
length especially towards the horn-centres. A similar »fan» of alto-
gether rufous hair begins between the ears and is directed forwards.
This posterior and the anterior system of hairs meet along a straight
transversal line drawn from one horn-centre to another and along
that line all hairs on either side point towards the horn-centre. The
area between the upper eyelelid and the ear is covered by short
rufous (and some few blackish) hairs with an oblique downward and
backward direction. At the anterior ear-margin these are a little denser
than elsewhere. The ears are pigmented on the back and posterior
third but quite naked. On the eyelids some scattered hairs are found
becoming more numerous towards the margins. About a dozen fine
bristles are found situated in two series above the upper eyelid. In

Nova Acta Reg. Soc. Sc. Ups. Ser. III. Impr. 27/7 1903. 6

49 EINAR LÖNNBERG,

front of these a little above the anterior corner of the eye is a single
bristle situated. Below and a little behind the posterior corner of the
eye two bristles are placed together. These correspond with the white
spot with a couple of bristles below the eye in Tragelaphus and the
foetus of Boselaphus. Below, and in front of these, there is along the
lower jaw a series of 5 or 6 single bristles. The most posterior of
these corresponds to a white spot with a single bristle in the foetus
of Boselaphus (conf. below). In addition to these there are some scattered
bristles and long hairs on the chin and lower lip. Quite a conspicuous
organ is found between the rami of the mandible a little in front of a line
drawn through the corners of the mouth. It forms a prominent cushion
about four mm. in diameter with fine bristles arranged in a horse-shoe
along the frontmargin. The chin and lower lip are hairy. The upper
lip is also hairy along the margin, and behind the rhinarium a number
of bristles are seen arranged in four vertical series. The body is
naked, as is already mentioned, but on the dorsal surface of the neck
and on the foreleg in the region between the carpal and the elbow-
joint some few scattered hairs are beginning to show their tips, and in
less degree on the hindleg in the neighbourhood of the knee-joint.

The row of pores along the anteorbital gland of the face is
quite straight not curved as in Cephalophus melanorhoeus. The pores
are not well conspicuous, but they seem to be fewer in number than
in the smaller species, or about 15. At least in some instances there
seems to be upper and lower small pores in addition to the main pores.

The vertical groove of the upper lip extends inwards to papilla
incisiva round which it forms a circular: groove. There are about
16—17 palatal ridges, but the posterior ones are only little prominent
although as in C. melanorhoeus they are more distinctly crenulated
than the anterior ones. All of them are more or less distinctly inter-
rupted in the middle. The buccal papilla have a very peculiar shape
and may be said to be compound as they have three to five roundish
tips, the middle one of which is larger than the others and may be
termed digitiform.

The caruncula sublingualis is represented by a narrow fold, broadly
and shortly triangular in shape. Behind the same a row of flattened
papillae extend backwards along each side of the tongue.

The papille filiformes of the tongue are as yet roundish warts,
posteriorly larger, as usual. The papille fungiformes are evenly scat-
tered over the tongue with exception of the posterior median portion.

MATERIAL FOR THE STUDY OF RUMINANTS. 43

The papille circumvallate are 12—14 in number and somewhat irre-
gularly arranged in two (or three) series, thus more numerous than
in the foregoing species.

The glandula thyreoidea has the usual situation just behind the
larynx and near the dorsal surface of the trachea. It is formed by
an ovate body on each side, measuring about 11 mm. in length and
with the narrower end pointing backwards. There is no isthmus be-
tween these two bodies.

The thymus-gland extends from the heart nearly to the larynx.
It is a rather thick body 11—14 mm. in the thoracic cavity, but ex-
tends in front of the first pair of ribs as a 2—3 mm. thick cord an
each side of the trachea.

The left lung is divided in three and the right in four lobes
and an azygous lobe. The lobulated structure of the lung is very
conspicuous,

Concerning the position of the abdominal viscera may be re-
marked that about two thirds of the left lobe of the liver lies on the
left side of the median line, but half of the reticulum, the whole paunch
and a portion of the abomasus are seen to the left of the same.
Behind the paunch the left kidney is visible and behind the abomasus
and the left lobe of the liver the coils of the jejunum are seen. On
the right side the liver occupies about half of the visible area and
behind the same are seen the coils of the small intestine, as well as
the cæcum and some parts of the large intestine. In the inguinal
tract the ovary and part of the right cornu uteri is conspicuous.

‘As in the adult the sulci coronarti are not quite so well deve-
loped on the right side, but on the left side there is a well developed
sulcus coronarius on the ventri-dextral sack. On the right side it is
best visible from the inside. The saccus cecus thus determined is of
about the same comparative size as in the adult and measures more
than a third of the length of the whole ventri-dextral sack. On the
dorsi-sinistral sack there is also a short sulcus coronarivs on the left
side but none on the right. The inside of the paunch is smooth, the
papillæ being not yet developed. The reticulum is comparatively larger
than in the adult; its diameter being about two thirds that of the
paunch. The reticular structure of its inner surface is already well
developed. A very remarkable feature is that the psalterium is com-
paratively so large viz. measuring 14 mm. in length by 8 mm. in
thickness. Its length is thus about two fifths of the length of the

4

14 EINAR LONNBERG,

paunch. In the adult the length of the psalterium is hardly a fourth
of the length of the paunch. This proves that the psalterium of Ce-
phalophus ogilbyi has become secondarily reduced in correspondence
with its carpophagous diet as stated above. Eight large primary folds
are developed here, and in addition to them a few (two on one, one
on the other) rudiments of, as it seems, secondary folds in the lateral
interspaces. To judge from the large interspaces between these rudi-
mentary folds and the adjoining large folds, which fully equal the
interspaces between two large folds at other places, I am inclined to
think that also the small folds belong to the primary system, but have
become suppressed for the purpose of leaving better room for the ad-
joining folds as they are situated at the lateral curving of the wall.
The height of the middie folds is about 4 mm. and they leave a com-
paratively smaller free lumen than in the adult. The abomasus is, as
is normal in a foetus, large, its length measured along the curvature
being about twice the length of the paunch. The wide fundus-portion
of the abomasus shows on its inner surface ten broad or high (8—4
mm.) folds (the same number as in the adult), Between these large
folds a number of lower transverse folds run crosswise from one large
longitudinal fold to another. In the narrowed pyloric portion the folds
disappear or are only continued as quite low ridges towards the py-
lorus. The inner structure of the abomasus is thus similar to the
same in the adult.

The general arrangement of the intestine is already the same
as in the adult. The blind end of the cæcum is narrowed and the
large intestine forms three centripetal and two and a half centrifugal
coils before it runs out in the peripheric mesentery. When the large
intestine has left the connection with the small intestine it does not
run as a straight rectum to the anus, but makes from the region of
the left kidney a transversal coil dorsally and to the right side and
then back ventrally and to the median line where it proceeds in the
pelvic region to the anus,

The situation of the liver has already been mentioned. The
right lobe which is much the largest extends longitudinally along the
body wall. Except the umbilical fissure there is no other fissure di-
viding the two main lobes. The caudate lobe is very large, thickly
trihedral and even extending beyond the margin of the right lobe.
Together with the margin of the latter the caudate lobe not only caps,

MATERIAL FOR THE STUDY OF RUMINANTS. 45

but almost wholly covers the right kidney on the ventral side. There
is no trace of a gallbladder.

The spleen is strongly convex on its anterior (diaphragmatic)
surface; the dorsal and median portion is much thicker than the other
parts which become quite thin towards the margin. It is triangular
in outline.

The kidneys are quite simple and show when cut only a single
longitudinal mamilla.

A comparison of the organs of foetus h. and foetus i. reveals
the following facts. The glandula thyreoidea of the former is even
comparatively smaller than that in foetus i. measuring only 6 mm.
But, on the other hand, the thymus gland of the younger foetus is
comparatively better developed as it reaches quite up to the larynx
where it is broadened again. The lungs are similar in both. The
reticular structure of the inside of the reticulum is feebly developed
in the foetus 7. The psalterium of foetus h. is remarkable for its com-
paratively great size which is not far from equal to the same in the
foetus i. It measures namely, not less than 12 mm. in length by 6
in width. Eight folds are visible although one of the lateral ones is
less developed. The size of the psalterium in this young foetus (its
length being not far from half the length of the paunch) adds further
strength to the theory expressed above that the psalterium of the
adult Cephalophus ogilbyi is a, in consequence of the changed diet,
reduced organ. The abomasus has in the younger foetus the same
interior structure as in the older. With regard to the arrangement of
the colic coils there is an interesting difference between the two foe-
tus. In the older one (i.) the coils are more drawn out longitudinally
and partly cover each other. In the younger (h). all coils lie nearly
in the same plane so that hardly any are covered. The coils form,
therefore, together a more broadly rounded figure, which, however,
is not quite regular because the spiral is more flattened just opposite
the tip of the cæcum. In the older foetus and in the adult, as is
mentioned above, there are three centripetal and two and a half cen-
trifugal central coils, in the younger one (h.) again there are three and a
half centripetal and almost three centrifugal central coils. This indicates
that by assuming the definite arrangement half a colic coil in either
direction is reduced. It may therefore be supposed that the ancestors
of Cephalophus ogilbyi has had the large intestine more spirally coiled

46 EINAR LÖNNBERG,

than it has now and I do not think I err in placing this reduction in
connection with the choice of a different food or with other words
the change from a herbivorous or graminivorous diet to a carpopha-
gous one.

Concerning the liver of the younger foetus (h.) it may be re-
marked that the caudate lobe is less developed than in the older stage
(4). It does not reach the margin of the right lobe and in consequence
of this the capping of the right kidney is almost as much effected by
the lateral or posterior portion of the right lobe as by the eaudate lobe.

The spleen of the foetus /. displays also quite an interesting
difference in shape when compared with that of the foetus 7. In the
former it is not triangular as in the latter, but elongately elliptie,
rounded at both ends although somewhat broader at the outer poste-
rior end. It resembles thus more in its general shape the spleen of
Giraffa and Cervicornia although the same organ of these animals is
comparatively shorter. It seems from this, however, as I have ex-
pressed before!) that the elliptic shape of the spleen is in the Rumi-
nantia an ancient characteristic and that the triangular ovine shape is
secondarily aquired.

10) A young foetus of Cephalophus silvicultor 3 measuring from
the forehead to the anus about 180 mm. and from the snout to the

occiput about 75 mm. It is perfectly naked as yet — not even a
bristle is seen — and may be regarded to represent almost the same

stage as that named b. of C. melanorhoeus (conf. above).

15—16 palatal elevations are developed and the four foremost
of those reach continually across the palate the others are arranged
more or less regularly in pairs. On the latter the crenulation of the
posterior edge is beginning to show. The continuity of the foremost
palatal ridges is a difference from the smaller species described above.

The thymus-gland extends all the way from the heart up to
the larynx. In the thoracic cavity it is a rather thick body measuring
about 6 mm. across, but then it is continued as a narrow cord, 1!/2
mm. in diameter on either side of the trachea. On a level with the
thyreoidea and dorsally of the same it is again expanded to a thickness
of 3 mm., at the same time reaching a little beyond the thyreoidea.

The latter gland forms a bean-shaped body on either side of
the trachea just behind the larynx. Its length is about 61/2, its breadth

1) Proc. Zool. Soc. 1900.

MATERIAL FOR THE STUDY OF RUMINANTS. 47

about 4!/2 and its thickness about 3 mm. In this species of Cepha-
lophus as well as in the others there is not the slightest trace of an
isthmus. In this respect Cephalophus shows a remarkable difference
from the condition found in bos the calf of which has a well developed
isthmus. In Ovis »der Isthmus ist meist nicht mit Sicherheit nach-
zuweisen» !).

The left lung is divided in three lobes, and the right in five
lobes and the azygous, in such a way that the apical lobe is divided
in an anterior and a posterior and the lobus cardiacus in a dorsal
(smaller) and a ventral (larger) lobe, but /obus diaphragmaticus is entire.

Concerning the general arrangement of the abdominal viscera
may be remarked that the liver is very large and its right lobe has
an almost longitudinal position along the abdominal wall. It extends
so far back that it almost covers the whole right kidney from a ven-
tral view. Behind the right kidney is seen a transversal loop of the
rectum directed towards the right side. The middle and posterior
portion of the left half of the abdominal cavity is occupied by the
coils of the intestine behind which a part of the left kidney protrudes.
To the left of the liver the greater part of the reticulum and the
paunch are visible and along the posterior margin of its left lobe pro-
jects a strip of the abomasus.

Sulcus coronarius of the ventridextral sack is well developed,
less so on the dorsi-sinistral. The distance between cardia and the
posterior end of sacci ceci 1s. about 32 mm. The reticulum seems
large, almost semiglobular with a diameter of 16—17 mm. The psal-
terium is also quite large with a longitudinal diameter amounting to
about 13 mm. and a transversal of 9 mm. When compared with the
reticulum it appears to be considerably larger than in the correspon-
ding stages of the smaller species, which indicates that the psalterium
is for Cephalophus silvicultor a more important organ than for the smaller
species, no doubt depending upon differences in the diet. The ar-
rangement of the folds is, however, quite the same as in Cephalophus
ogilbyi, thus eight well developed primary folds beside some rudimen-
tary ones. Secondary folds are not developed; at least not in this
stage. The abomasus is large, measured along the curvature it is
about a fourth longer than the longest diameter of the paunch. Nine
broad folds are developed in its fundus region and in addition to those

!) ELLENBERGER. und Baum: Handbuch d. vergl. Anat. d. Hausthiere. Berlin 1900.

48 EINAR LÖNNBERG,

some smaller ones, partly situated at the lesser curvature. This organ
is consequently similar to that of the other species.

With regard to the arrangement of the intestine the colon and
its coils absorb the most interest. There are thus, counted from the ansa
proximalis not less than four centripetal and three centrifugal central
coils, or, counted from behind three coils and a half in either direction.
This is thus a good deal more than what is found in the smaller car-
pophagous species and may in the same way as the larger psalterium
be put in connection with another diet. The most peripherie coil of
the colon runs, since it has left the central spiral, in quite close con-
nection with the small intestine (jejunum) and must therefore be very
long. From «ansa distalis the large intestine runs straight backwards
in the median line, but forms as already is mentioned a transversal
coil behind the right kidney.

The liver is in consequence of its situation, already alluded to
above, in a still higher degree than in the other species bent in a
right angle. The longitudinally placed right lobe constitutes also in
this species still more than in the others the main mass of the liver.
Its bulk may be understood from the following measurements length
38 mm., breadth 33 mm. and thickness 18 mm., while the same of
the left transversal lobe are resp. 26; 30; 10 mm. The two lobes
are quite undivided as in the other species. The caudate lobe is
large, and thickly trihedral, but not extending to the margin of the
main right lobe. Together with the posterior end of the latter it caps
the right kidney. The Spigelian lobe is comparatively large and broad,
as broad at the ends as at the base.

There is not the slightest trace of any gallbladder to be seen.
This is of interest to note, because it could otherwise have been
suspected that the missing gallbladder was a feature only characteri-
stic for the smaller species of this genus. But now it is proved as
well for the largest as for the smallest species of Cephalophus and
is thus no doubt a common characteristic for all members of this
genus.

The spleen is strongly convex on its diaphragmatic surface, |
much thicker at its dorsal side. It has already assumed a triangular
outline although the foetus represents such an early stage.

The scrotal sack is very large measuring 16 mm. in length and
13 in breadth. That this organ is very early developed has already
been stated in describing the anatomy of the young foetal stages of C.

MATERIAL FOR THE STUDY OF RUMINANTS. 49

melanorhoeus, but it is interesting to find that, in striking contrast to
what was the case in C. melanorhoeus, descensus testiculorum already
has taken place, while this in the small species did not happen before
the foetus is quite hairy all over and almost ready to be born. This
difference is the more peculiar, as, of course, even at the end of the
foetal life, the large species, C. silvicultor, is more remote from sexual
maturity than the small one, ©. melanorhoeus, which latter most pro-
bably becomes fully developed within a year, which cannot be assumed
concerning the large species.

Another difference of a similar kind is shown therein that the
two teats on either side in the young foetus of C. silvicultor, although
sitting quite close together and almost touching each other, are quite
separate to the base. As is described above the teats of C. melano-
rhoeus are developed from a common rudiment on either side and are
not in the male foetus fully divided into two before the foetus is hairy
all over and almost fully developed.

The kidneys are not quite so simple as in the other species,
as, at least in this stage, the different cones may be distinguished al-
though they seem to have the tendency to fuse together and form
only one mamilla. This is not completed in this stage, but it might
follow in a later, although this cannot be ascertained.

It is an important fact that in all the foetal specimens of these
three species of Cephalophus I have found a pit between the toes, as
well of fore as of hind legs, leading into a small bag corresponding to
the sinus interdigitalis of the sheep. This pit and bag are better de-
veloped in the advanced foetal stages and they are thus no doubt
persisting structures in the adult of this genus. This fact tends to
indicate that foot-glands are, and probably have been, still more gene-
rally distributed among the Ruminants than they usually are believed
to be. It is even probable that where they are missing this is due
to reduction, rather than that they should have been acquired inde-
pendently in different genera.

Nova Acta Reg. Soc. Sc. Ups. Ser. III. Impr. ?9/s 1903. i

50 EINAR LÖNNBERG,

Some observations on a foetus of Boselaphus tragocamelus PALLAS.

The dimensions of this foetus are: Length of head about 14 em.

Distance from the neck to the root of the tail: 38 cm.

At the first look it seems almost naked with the exception of
the head, the tail and the hindportion of the tighs from the tail and
downwards to about midway between knee and heel-joints. A close
examination reveals, however, that short hairs are just beginning to
grow out everywhere. The hair that covers the face is dark brown.
The hair of the nose is reverted or directed upwards up to a point
between the orbits where it diverges like a fan. In that same point
the reverted hair from the nose meets the hair from the frontal whirl,
in the middle of the forehead. From this latter the hair radiates in all
directions. The hair on a strip above the orbits is directed backwards
in continuity with the hair of the nose. The hair of the occipital re-
gion is directed forwards, diverging somewhat in the parietal region.
Just above and behind the orbits where the horns are in the adult is
a bare spot towards which the hair radiates from three directions viz.
the mid-frontal whirl, the supraorbital strip and the parietal region.
On the sides of the head the hair is directed downwards from the
temporal region to the snout. The upper eyelid is provided with a
number of bristles distributed in three or four irregular longitudinal
series, below these it is lighter in colour. Just above the corner of
the eye there is also a light spot. Right below the eye on the middle
of the cheek is a light spot with two or three bristles. A little below
and in front of this is another light spot with one bristle. There are
also some scattered bristles on the sides below the lower lip. The
chin and a spot on each side of the black naked rhinarium is white
haired. In the middle of the surface of the lower jaw there are three
white bristles. The distal half of the ears is mostly blackish on the
outer surface but within it is white-haired with two black spots at the
posterior margin. .

From this description compared with that given above concerning
the foetus of Cephalophus ogilbyi is apparent that the arrangement of
the hair of the head is different in these two genera. The bristles of
the eyelid are also in the Boselaphus-foetus arranged in more rows
than in Cephalophus. But, on the other hand, the sensitive spot above
the anterior corner of the eye, that below the eye, that underneath

.

the lower jaw etc. correspond to similar spots with bristles in the

MATERIAL FOR THE STUDY OF RUMINANTS. 51

Cephalophus-foetus as has been mentioned above, and indicate that
these organs are of ancient origin.

The papillae of the tongue have perhaps not yet attained their de-
finite shape. Without comparison with the tongue of an adult animal
this can, however, not be decided. In the present stage the papille
filiformes are rather short, scale-like with truncated or rounded tips.
They are thus similar to the same of the Madoqua, although a little
larger than those. They increase in size towards the posterior half
of the tongue and become also stouter than anteriorly. In the me-
dian portion of the posterior half of the tongue the papille assume a
flattened wartlike shape which also agrees with the same in the
Madoqua although they are more rounded in the Nilgai-foetus. Another
resemblance to the Dik-dik lies therein that the lateral surfaces of the
tongue are, to a great extent, covered with more slender and pointed
papille filiformes. On the hindmost part of the tongue of the Nilgai-
foetus the papille are conical or almost cylindrical and soft. The
papille fungiformes are numerous and scattered all over the anterior
half of the tongue but sit more densely anteriorly and are as usual
especially crowded at the tip and below the same. On the posterior
half of the tongue the papillæ fungiformes are few in number and only
found towards the lateral borders. They are here, however, a good
deal larger in size. The papille circumvallate are numerous, 17 on
one side, 19 on the other. 'They form one long series on each side,
but, in addition to this there are also some scattered ones. Some of
these have a rather peculiar appearance, because the ringlike sur-
rounding wall is not smooth as usual but itself carries a smaller or
greater number of papille. In other cases the surrounding wall is
not ringlike but consists of an oblique longitudinal fold on each side.
It looks therefore as if these organs had originated through a combi-
nation of papille fungiformes and foliate.

There are about 18 pairs of palatal elevation in the Nilgai-
foetus, some incomplete posterior ones not counted. They are con-
spicuously fringed, even the posterior ones. The buccal papillæ are
partly conical, partly flattened with small secondary papilla at their
margin. The papilla incisiva has a rounded posterior and a bluntly
angular anterior outline.

There is no septum membranaceum developed in the posterior
nasal cavity.

H2 EINAR LÖNNBERG.

The thyroid cartilage of the larynx is anteriorly emarginate and
measures in the median line 14, at the side 16 mm. The greatest
width of each lateral shield is about 14 mm. Prominentia laryngea is
comparatively large but rounded. It is situated at the posterior end
of the cartilage. The cricoid cartilage forms anteriorly or ventrally a
rounded but rather stout arch. Its posterior or dorsal shield mea-
sures about 9 mm. in length. It has a short crista anteriorly and is
comparatively thick even at its posterior end. The epiglottis is trian-
gular with pointed apex.

The lungs are divided by very deep fissures. The left lung
consists of three lobes. The large posterior lobe is perfectly free from
the others and only by a little connective tissue connected with the
middle lobe. This is dorsiventrally very long, tonguelike and rather
narrow. Anteriorly it is, by a deep fissure, divided from the upper or
anterior lobe. The right lung shows four well defined, almost entirely
separated lobes and an azygous lobe. The spleen of the Nilgai-foetus
(fig. 15) has an elongated triangular shape which reminds one of
that in the Black-buck, but is somewhat different from the shortly
triangular ovine type and still more from the oval (primitive) cervine
(including Giraffa and Ovibos) and the elongate taurine (also shared
by Connochetes and Bubalis sp. foetus) type. It measures 55 mm. in
length by 27 mm. in breadth.

All. four divisions of the ventricle are well developed, although
of course the abomasus is dominating in size. A dorsal and ventral
sack of the paunch is interiorly already differentiated by the develop-
ment of the large longitudinal fold (»Lüngspfeiler»). Both sacks are
subequal in size. The interior folds corresponding to the sulci coronarti
of the two main divisions are also developed and indicate that each
is provided with a saccus cecus. The reticular structure of the reti-
culum is also developed. In the present stage it is a good deal larger
than the psalterium which latter has developed four systems of folds.
It is thus quadriplicate, but according to Garrop') the same organ of
the adult Nilgai is quinqueplicate. The abomasus is provided with a
large number of folds. The exact number cannot, however, be indi-
cated because they are irregularly arranged. The largest folds are
more or less distinctly longitudinal, but there are also a good many
oblique or transversal folds. From the pylorus the duodenum runs to

Ty PZ. SMS TT:

MATERIAL FOR THE STUDY OF RUMINANTS. 53
the liver at which it forms a loop as short as possible. Then it turns
backwards as far as to the level of the hind-end of the right kidney,
bends then over to the other side and runs again forward in close
connection with the proximal part of the rectum to the level of the
anterior end of the left kidney where it enters as jejunum the common
mesentery and becomes highly convoluted. The ileum makes, just
before entering the cæcum, a short loop bending back upon itself. The
length of the small intestine is about 334 cm. The length of the caecum
is about 5 em. and its diameter about 12 mm. The colon forms first
a rather large ansa proximalis which descends toward the middle of
the cæcum and then enters into the spiral labyrinth. In this it forms
three centripetal and 2'/» centrifugal coils and then goes directly, not
making any loop on itself, out in the mesentery as the peripheric coil
in connection with the jejunum. This peripherical coil is situated at
a similar distance as well from the jejunum as from the colic spiral.
When the colon leaves the common mesentery is goes backwards and
forms the ansa distalis which is rather long and connected as well
with the ansa proximalis as also with the descending part of the duo-
denum. The anterior end of the ansa distalis curves over to the left
side and passes at the anterior end of the left kidney into the rectum
which is connected with the ascending portion of the duodenum. On
a level of the posterior end of the left kidney the rectum forms a
short loop and there becomes widened, whereupon it runs straight to
the anus.

The length of the colon is about 129 em. The small intestine
is thus about 2,5 times longer than the large one. The different parts
of the intestine have not of course yet attained their definitive pro-
portions. According to Garrop') the small intestine of the Nilgai is
not fully twice as long as the large one. That the large intestine is
strongly developed in this animal may already be stated in this foetus
because the number of spiral coils is so great, nearly equalling the
same in the sheep and exceeding that of Bos, Connochetes and Antilope.

The liver of this Nilgai-foetus is not situated transversally”), but
its greatest expansion lies in a vertical longitudinal plan, the same which
also passes through the right kidney. There can thus be distinguished

1) Proc. Zool. Soc. 1877.

?) Nevvitte (Bull. Mus. hist. nat. T. 3. 1897) found in Adenota kob and Tragela-
phus gratus that the liver did not occupy a transversal but a longitudinal position. Compare
also the description of the foetal specimens of Cephalophus above.

51 EINAR LONNBERG,

a lateral convex and a median concave surface corresponding resp.
to the anterior and posterior surfaces of a liver in normal position.
Although the situation is altered it seems nevertheless suitable to
retain the attributes »right» and »left» for the different lobes in
correspondence to the same parts of a normally situated liver. The
umbilical fissure extends through half the organ on the concave but
only through one third of the convex side. The right lobe is much
larger than the left one and shows on its concave surface a fissure
in which the gallbladder is situated. This fissure lies much nearer
to the free (renal) border of the right lobe than to the umbilical
fissure. The /obus caudatus has the shape of an elongate trihedral
pyramid. The right kidney is wedged in between the caudate and
the right lobe. GaArrop has already stated!) that the Spigelian lobe
of this animal is absent.

The kidneys are perfectly compact and show when cut only
one longitudinal mamilla.

Concerning the genital organs there is not much to be said.
The testes have not yet descended into the scrotum, but are situated
near the base of the scrotal sack at the ventral wall of the abdominal
cavity. 'The glans penis (Fig. 11) shows a well developed cushion on
the upper (dorsal) side of the end of the organ, which extends down-
wards and forwards on the right side and forms the tip of the penis.
The urethral tube is somewhat free at the end and is situated on the
left side, but it hardly extends beyond the tip of the penis. It may
thus be said that this organ to a certain degree resembles the ovine
type as this is modified in the Gazella, but it has no filiform appendage.

A little in front of the scrotal sack four teats are seen; the
posterior pair sit much nearer the median line than the anterior
pair.

From this description is apparent that the Nilgai in its visceral
morphology does not show any distinct close affinity to any special
generic group among the Cavicornia, but at the same time it does not
exhibit any striking aberrant characteristic.

“) Proc. Zool. Soc. 1877.

MATERIAL FOR THE STUDY OF RUMINANTS. 55

Notes on the morphological anatomy of the Anoa.

The viscera of a young bull of Anoa have given me the oppor-
tunity of making the following notes.

The tongue must be regarded as rather short, only measuring
15 em. in length. Its breadth is almost the same posteriorly, and in
front about 31/2 em.

As in Antilope and some other antelopes the anterior portion,
not quite half the tongue, is pigmented!). The papille filiformes are
flattened, seale-like with blunt tips, thus differing from those of Bos
which are more narrow and pointed. On the posterior thick portion
of the tongue the enlarged gapillæ filiformes are in the centre more
or less bluntly conical, partly curved. Towards the sides they become
more flattened and truncate, but also here recurved. Their general
shape differs here as well from those of bos as those of Ovis and
Antilope. The papille fungiformes are rather evenly scattered all over
the upper surface of the anterior portion of the tongue, being, how-
ever, as usual more closely set at the margin. Near the transverse
groove dividing the anterior thin and posterior thick portion these pa-
pille are scarce in the middle, but appear again further posteriorly.
On the whole it can be said that these papilla are comparatively nu-
merous and more evenly distributed than in other Cavicornia (except
Bos and Ovibos) instead of leaving a central space free as is usually
the case. HELLER (l. c.) says, however, that the papille fungiformes
are comparatively better conspicuous in the young Anoa than in the
old. The papille circumvallate are arranged in two rows?) the outer
consisting of nine or ten and the interior of five papillæ. In this re-
spect the Anoa agrees pretty well with Bos and its number of such
papilla exceeds that found in Antilope, Connoch«tes ete.

The thymus-gland had been cut to pieces on the preparate so
that only the anterior ends were left. These were situated on either
side of the larynx and resembled in appearance the same organ of
the common calf.

The thyreoidea has the shape of a flattened reddish body situated
at the sides of the trachea, just behind the larynx and near the dorsal

1) This may be variable to judge from the statements by Herter in »Der Urbüffel
von Celébes».

?) Heiter talks about a third »row» consisting of 1 papilla, this is not found in
this specimen.

56 EINAR LÓNNBERG.

surface. Its length is about 16, its breadth about 11 and its thick-
ness about 6 mm. It must thus be termed, comparatively, very small.
The most interesting feature with regard to the thyreoidea of this
animal is, however, that it is provided with an isthmus. This has a
width of almost 3'/2 mm. when it leaves the main lateral body of the
thyreoidea, but proceeding transversally along the surface of the tra-
chea it is gradually narrowed so that it hardly can be traced macro-
scopically across the median line. This is a feature which reminds
one about the condition prevailing in Bos in which the isthmus is well
developed while in no species of antelope investigated by me such an
isthmus could be traced. It may thus to a certain extent be regarded
as a bovine characteristic.

The larynx of the Anoa is very small. Taken as a whole it
measures only 47 mm. in length and its dorsiventral diameter through
pomum adami is only 88 mm. The length of cartilago thyreoidea is
laterally 36 mm. Pomum adami forms a rather broad and flattened
area, triangular in outline. Its apex is situated on a distance of 11
mm. from the hindmargin, 20 mm. from incisura oralis. This triangular
area is hard and ossified while the other parts have the usual cartilagi-
nous consisteney. The most remarkable fact concerning this cartilage
is that the cornua anteriora are coalesced with the main cartilage in
such a way that (as in the horse) a foramen thyreoideum is formed.
Such an arrangement I have not seen in other Ruminants. The cornua
posteriora are long (15 mm.) and arcuate. The dorsal plate of the
cricoid cartilage has a rather strong keel and measures about, 30 mm.
in length. The epiglottis is broad with rounded outline thus resembling
the same organ of the Gnu!), but differing from the triangular one of
Bos etc. From this description and the figure (fig. 17) may be seen
that the larynx of the Anoa does not closely resemble the same organ
of other Ruminants as far as is known.

The tracheal rings are thinner behind, and meet dorsally in such
a way that an edge is formed, as is also the case in the common ox
and the Gnu.

The number of tracheal rings is quite small, only 48, which is
the smallest number indicated by ELLENBERGER & Baum’) for any
domesticated ruminant. The small number of tracheal rings of the

') Lüxxserc: Studies on Ruminants. K. Vet. Akad. Handl. Bd. 35.
”) Handbuch der vergl. Anat. d. Hausthiere. Berlin 1900.

MATERIAL FOR THE STUDY:OF RUMINANTS. 57
Anoa stands, no doubt, in connection with the shortness of the neck
of this animal. Some of the rings had even become confluent with
each other, which may indicate that a secondary reduction in number
has taken place.

The left lung has three lobes, the right four and an azygous
lobe. The division is thus similar to that of the common ox. The
Anoa-lungs are also distinctly lobulated as in cattle.

When the Anoa-ventricle was moderately expanded the following
dimensions were recorded: total length of dorsi-sinistral sack counted
from the cardia 43 cm., distance from the cardia to the posterior end
of the ventri-dextral sack 46 cm.'). The diameters of the reticulum
were resp. 15, 10, 10 and those of the psalterium 9, 5, 7. The dorsi-
sinistral sack does not project so far backwards as the other but its
saccus caecus is more pointed while that of the ventri-dextral sack is
more broadly rounded. The latter saccus cecus is also larger, mea-
suring 12 cm. as well in length as in breadth and is well defined all
round by a sulcus coronarius. The saccus cecus of the left sack is only
10 em. in length and 8 em. across at its middle. Its swleus coronarius
is not so sharply defined. With regard to the backward extension of
the sacci ceci the Anoa thus differs from bos taurus, but although the
left saccus cecus is a good deal smaller than the right it seems to be
better developed than the same in sheep and in antelopes (as far as
the latter are known).

The small size of the psalterium when compared with that of
the reticulum is as HELLER”) already has noted a difference from the
prevailing condition in Bos taurus. In this respect the Anoa resembles
the less specialised Ruminants as antelopes, goats, deer.

According to HELLER (l. c.) the psalterium of the Anoa is tri-
plicate.

The abomasus is about 22 cm. long and 9 cm. broad.

The small intestine of this specimen measured only about 890
em. while adherent to the mesentery and in an empty state. This
is remarkably short which may be deduced from the fact that I have

!) When Hetter (in: »Der Urbüffel von Celebes») from a blown up ventricle has
recorded that the left sack is longer than the right this is quite correct when one does not
count from the cardia but only from the anterior end of the right sack. If so is done on
this specimen the length of the ventridextral sack is found to be about 37 cm.

*) Urbiiffel von Celebes. Dresden 1889.

Nova Acta Reg. Soc. Sc. Ups. Ser. II. Impr. ?"/s 1903. 9

5S EINAR LÖNNBERG.

found the same measurement of a, by no means, large sheep to
be 1515 cm.

The length of the cæcum is 11 cm. It is broadly rounded at
the blind end and is widest near that end. In a flat and empty state
it measures 6 em. across, a little above the blind end. HELLER found
that the by him measured caecum had a length of 16 em. but as it
was quite isolated it might have been distended, or it might perhaps
also have belonged to a larger animal. In either case, however, the
cecum must be termed small as the same organ of a sheep mea-
sured about 36 and that of a young buck 22 cm.

The length of the large intestine measured from valvula ileo-
cecalis to the anus is about 250 cm. This measurement as well, is
very short as is seen from the comparison with the same organ of a
sheep or a buck which, measured in the same way, attain a length
of 490 cm. Already before I had measured the large intestine of this
Anod-specimen it was apparent to me, that it must be very short
when I considered its general arrangement, which was as follows.
After having formed a wide but short ansa proximalis, in which it has
a width of 3!/z—4 cm. when flat and empty, the colon enters the
central spiral and forms 1/2 centripetal and 11/2 centrifugal coil in
which it has a width of about 18 mm. But when this central spiral
has been formed the colon does not run out in the common mesen-
tery to constitute a peripheric coil near the jejunum as in sheep,
antelopes and deer but forms only a rather short ansa distalis and
runs then straight backwards as rectum to the anus. This arrange-
ment reminds one of that found in Bos taurus, but is still simpler.

On the whole it may thus be said that the Anoa with regard to
the arrangement of its intestine to some extent resembles that of Bos
taurus, but the intestine of the former is shorter which, no doubt, is
due to a diet consisting of less coarse fibre than that of the common ox.

The absence of the peripheric colic coil seems to be a feature
of much importance as this coil has been found in all members as
well of Cavicornia (except Bos, as is already mentioned, and the Bu-
balidine section) as of Cervicornia which I hitherto have had the oppor-
tunity to dissect. I have also stated the presence of this peripherie
coil in members of Tragulina. The absence of this coil in Anoa, Bos
and Bubalidine antelopes is therefore due to a secondary reduction.

The liver of the Anoa is divided in a left and a right lobe and
the fissure between them is best developed on the diaphragmatie side.

MATERIAL FOR THE STUDY OF RUMINANTS. 59

The leit lobe is rather small next to the fissure it is about 12 cm.
and the greatest distance from the fissure to the left margin is about
10 cm., but hardly half of that at the dorsal border. The right lobe
measures from the fissure to the right free border about 1612 cm.
There is a small fissure indicating the limit of a right central lobe
which measures 13 cm. from the dorsal to the ventral free border.
The pathologically enlarged gallbladder is connected with the right
lateral lobe which is prolonged at this junction so that its longest ex-
tension from the dorsal border to the free margin is about 1612 cm.
The caudate lobe is trihedral, but short (51/2 em.), and caps together
with the posterior dorsal portion of the right lobe the kidney in a
similar way as in the Gnu’). The Spigelian lobe is only slightly de-
veloped with a broad base and rounded tip.

The spleen is elongate but at the same time broad and with
both ends equally rounded. Its length is about 25 em. and its width
about 9 cm. at the thicker fixed end, and 8 cm. at the other. It
thus, to a certain degree, resembles the same organ of the Gnu and
at the same time its shape is intermediate between the very elongate
one of Bos taurus and the primitive elliptic one as exhibited by Ovibos,
Cervicornia, Giraffa etc. being, however, somewhat more taurine than
that of the Gnu.

Both kidneys of the Anoa are beanshaped and of almost the
same size not quite attaining a length of 9 em. and with a breadth
of 4'/ cm. or a little less (the left). Their greatest thickness is about
3'/2 cm. Exteriorly they show a surface which is not quite even, but
provided with impressions and furrows, which although less strongly
developed than in the kidneys of Bos, remind one about the condition
of those and give a hint that the kidneys of the Anoa are not so
simple as those of the antelopes, sheep etc. A longitudinal section
reveals also the remarkable fact that the Anoa-kidney interiorly is just
as complicated as that organ of the common ox, that is; every cone
is distinct and opens each with a separate mamilla in a tubular pro-
longation of the renal pelvis. HELLER (l. c. p. 29) who has shortly
described some of the viscera of a quite young calf of Anoa says that
the kidney of that was 4,5 cm. long and without lobulation. He has,
however, as far as can be seen from his words, not made any section
of the kidney, but only inspected its exterior surface. It is, however,

1) Studies on Ruminants. K. Vet.Akad. Handl. Bd. 35.

60 EINAR LÓNNBERG,

in consequence of HELLERS statement possible that the exterior lobu-
lation of the Anoa-kidney makes itself more conspicuous in a some-
what later stage. The structure of the Anoa-kidney, as it appeared
on my preparate described above, is a characteristic of considerable
importance from a classificatory point of view. As this characteristic
of having at least interiorly perfectly lobulated kidneys among the
Cavicornia is shared only by the common ox and its nearest relatives
it may be regarded as a true bovine characteristic indicating affinity
and common origin.

As the animal dissected was not fully adult the genital organs
were not fully developed and in addition to this they were, before they
fell into my hands, damaged as they had been roughly cut out from the
body of the dead animal. Some notes about them may nevertheless be
appended. The testicles are small. Together with the epididymis they
measure about 4'/2 cm. in length and 2 cm. in thickness. The thick-
ened terminal portions of vasa deferentia are comparatively long,
about 5 em. in length by 5 mm. in thickness, and the interior of the
same within the muscular coat has a plicated and spongious-looking
appearance. The colliculus seminalis is well developed and the openings
of the ductus ejaculatorii are protected by broad plicæ and in addition
to this by a double papilla on either side at the distal end. The vesi-
cule seminales are small, but have the elongate shape of those of the
common ox. Their length in this young Anoa is about 24 mm. and
their thickness about 8 mm. They lie straight and are not doubled
up as, for instance, is the case in Antilope cervicapra.

The glandulæ cowperi have been cut away in this preparate and
cannot therefore be described.

The penis has a short flexura sigmoidea. The end of the organ
(fig. 16) is bluntly pointed and has no urethral prolongation as may
be seen on the figure (fig. 16) wherein we find a certain likeness to
the same organ of Bos taurus. The length of the in the prepuce in-
cluded glans is about 35 mm. The almost slit-like opening is situated
on the left side about 4 mm. from the blunt tip. Dorsally towards the
right side there is a prominent cushion although not so strongly deve-
loped as in the Nilgai with the penis of which that of the Anoa also has a
certain likeness. The asymmetry of the organ with the urethral opening
on the left and the dorsilateral cushion on the right side is an ancient
characteristic although the urethral prolongation has been reduced.
The penis of the Anoa is, however, of a more primitive ruminant type

MATERIAL FOR THE STUDY OF RUMINANTS. 61

than that of Bos taurus. All characteristics taken together it seems
evident that the Anoa has an unmistakable affinity with the Bovine
group or section. Among characteristics of importance for the passing
of such a judgement the structure of the kidneys and the general
arrangement of the large intestine seem especially valuable, because
Bos (s. str.) and Anoa agree in these respects with each others at
the same time as they widely differ from all other Cavicornia, the Bu-
balidine section excepted with regard to the colic morphology. But at
the same time it must be admitted that the Anoa, with regard to the
morphology of its soft parts as well as with regard to its skeleton and
the structure of its horns, is less specialised than the typical members
of the genus Bos and has retained more ancient characters and thus
remained on a more primitive stage. In this respect its seclusion on
an island may have played an important part. But in spite of the fact
that the Anoa is a generalised type it cannot as yet, with the present
state of our knowledge, be said with which, if with any, of the antelo-
pine groups the Anoa might be more closely related than the typical
members of the genus bos are. The Anoa is thus a true member of
the Bovine section. The characteristics with regard to which it re-
sembles Connochetes and Bubalis (shape of spleen, simplicity of colic
spiral ete.) are shared by the typical Bos as well, and these might
indicate that the ancestors of the Bovine and Bubalidine sections ori-
ginally branched off from the common Cavicorn stem in one branch!).
In a later time, however, this branch was divided into two, differing
through several distinguishing features, as well internal as external.
Among the former the Bovine section (incl. Anoa) acquired the lobu-
lated kidney, the isthmus of the thyreoidea etc.

!) In a similar manner the Antilopine and Ovine sections might once have together
formed another branch.

Explanation of figures.

EX
Larynx of a young Black-buck (Antilope cervicapra). Nat. size.
Distal portion of the penis of a young Black-buck (Antilope cervicapra). Nat. size.
The same organ of Gazella rufina.
The same organ of Cobus defassa.

”

The stomach of Cephalophus melanorhoeus. 1/2 nat. size oe. oesophagus, p. paunch

r. reticulum, ps. psalterium, ab. abomasus.

The spleen of the same. Nat. size.

The colic spiral coils of the same. 1/2 nat. size.

Distal end of the lower jaw of a young foetus of Cephalophus melanorhoeus stage
b.; upper view.

The same organ of a somewhat older foetus, stage d.

The same organ of an older foetus, stage f.; side view.

Distal end of the penis of a young Nilgai-foetus. Nat. size.

PI:

Situs viscerum of a young foetus of Cephalophus melanorhoeus, stage a., with the
right side exposed to the view. Nat. size.

The same with the left side exposed. Nat. size.

Different stages of development of the teats of Cephalophus melanorhoeus, & from
stage b.; B from stage d.; y from stage f.

The spleen of a Nilgai-foetus. Nat. size.

Distal end of the penis of Anoa. Nat. size.

Thyreoid cartilage of Anoa. Lateral view. Nat. size.

Glandula thyreoidea of Anca. Nat. size.

Longitudinal section through the kidney of Anoa.

Distal end of the penis of Tragulus sp. Nat. size.

Distal end of the penis of a foetus of Cephalophus melanorhoeus, stage f. ?/1.

The spleen of a foetal specimen of Cephalophus melanorhoeus, stage a. 2/35

The brain of a foetal specimen of Cephalophus melanorhoeus, stage b. Nat. size. a
from the left side 5. from above.

E. Lönnberg, Study of Ruminanis Taf.l.

Nova Acta Reg. Soc. Sc. Ups. Ser. Ill.

M

N
N

Fig8

L. Ljunggren, Upsala.

| Nova Acta Reg Soc. Sc. Ups. Ser. III.

E. Lönnberg, Study of Ruminants Taf. I

722

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