THE SKULL IN A NEW TYPE O F DINOCEPHALIAN REPTILE.

733

45. On the Structure of the Skull in a new Type of Dinocephalian Reptile.

By R. BROOM,D.Sc., F.R.S., C.M.Z.S., Transvaal Museum.
(Published with the approval of the Director of the Transvaal.
Museum.)
[Received April 20, 1936 : Read June 9, 1936.1

(Text-figures 1-9.)

About a year ago I heard from the Rev. J. D. Vorster, at that time Dutch
Reformed Church minister at Fraserburg Road, Cape Province, that a large
fossil reptile skull had recently been discovered on the farm Stinkfontein,
and he secured for me a considerable portion of the snout. As it was manifest

Text-figure 1.
_----_
--:-:-\
/<--
#
,
-
\

- \\

Side view of the skull of Dinosuchus vorsteri Brown. Slightly under 4 nat. size.
"he thickness of the frontal and the degree in which it overhangs the orbit are
conjectural.

that the skull was that of an unknown Dinocephalian I at once visited the farm.
Mr. C. Botes, the proprietor, very generously allowed me to have the skull
for the Transvaal Museum. When the fragments were cleaned and joined
together it was found that not only had we the most perfect Dinocephalian
734 DR. R. BROOM O N THE STRUCTURE O F THE SKULL IN

skull as yet discovered but also the largest. A short preliminary description
is appearing in the forthcoming part of the ‘Annals of the Transvaal Museum.’
The skull is nearly complete, but much of the nasals, frontals, and parietals
have been weathered off, so that we have no evidence of the thickness of the
bones in these regions. The rest of the skull is practically perfect, though,
owing to the roughness of the surface of the bones, many of the sutures cannot
be satisfactorily traced. Of the lower jaws we only have preserved the back
half of the right mandible. No other parts of the skeleton occurred at the
spot where the skull was found.
Text-figure 2.

Top view of skull of Dinosuchua corsteri Broom. Slightly under & nat. size. The nasal
bones, and much of the frontals and parietals are lost, so that. the restoration of the
parietal and frontal region is largely conjectural.

As will be seen from text-figs. 1, 2, and 3, the skull is of a very unusual

shape. Though the snout is not unlike that of other previously known carni-
vorous Dinocephalians, the great width of the occiput and the very low position
of the temporal arch give the skull an appearance very unlike that of any
Therapsid hitherto discovered.
The greatest length of the skull from the snout to the back of the squamosa.1
is about 740 mm., but as the skull is considerably crushed obliquely some of
 A NEW TYPE O F DINOCEPHAIJAN REPTILE. 735

the measurements must be regarded as only approximate. From the snout

to the occipital condyle the measurement is 636 mm., and the occiput measures
600 mm. across. The skull may be regarded as roughly about 28 inches long
and 24 inches wide.
The premaxillaries are relatively small and considerably overlapped by the
maxillaries. Each carries as preserved the remains of three incisors. In the
right premaxilla these are i l , i2, and i3, while in the left the teeth remains are
i2, is and 14. The first three incisors measure together about 72 mm., though
the measurement must be regarded as only approximate, as the first incisor
Text-figure 3

Under side of skull of Dinosuchus vorsteri Broom.

Slightly under 4 nat. size.

is only represented by part of the root. The whole four incisors probably
measured about 90 mm. There is probably, as in Jonkeria, a long internasal
process of the premaxillaries, but t,his part of the snout has been lost by
weathering.
The maxillary bones are long and well developed. There is a large canine
which measures at its base 43 by 30 mm. Behind it there appear to be five
molars, of which the first, situated 32 mm. behind the canine, has a section
measuring 13 by 11 mm. The second molar is considerably larger and its base
measures 17 by 12 mm. Behind this there are certainly two small molars, and
probably three.
736 DR. R. BROOM ON THE STRUCTURE OF THE SKULL IN

The septomaxillary has only its base preserved, and it is diMcult to follow
the suture between it and the maxilla. The nasal seems to be completely
lost, but i t must have been narrow.
Much of the upper parts of the frontals and parietals are lost, and i t is
impossible to say how thick they may have been or to what degree supraorbital
bosses were present. Fortunately the position of the orbits can be determined
fairly accurately, as we have preserved the back of the left orbit and the front
and lower borders of the right. It is impossible to determine the limits of the
prefrontals, lacrimals, and jugals. The jugal is large and curiously shaped,
but its anterior and upper limits cannot be clearly ascertained. It passes
far back and forms the inner side of most of the temporal arch. How much
of the postorbital arch is formed by it is doubtful. Owing to the great width
of the squamosals the jugal has to pass out as a flattened plate to meet the
squamosal, as shown in the figures.
The whole of the palate and under surface of the skull are beautifully
preserved. As the incisors are directed well forwards and have long roots

Text-figure 4.

Occiput of Dinosuchus vorsteri Broom.

+
Slightly under nat. size.

the premaxillaries have large surfaces showing in the lower view. It is

difficult to be sure of the sutures between the premaxillaries and the prevomers,
but the divisions between the two are probably as indicated in the figure.
The prevomers are well developed and they articulate with the pterygoids
and palatines behind. They form together a well-developed ascending median
plate, as shown in the figure of the median section of the skull (text-fig. 5).
Each bone has a well-marked antero-posterior ridge on its palatal surface.
There are no teeth on the prevomers.
The palatines are essentially similar to those in the better known Dino-
cephalians, but they differ in having each a rounded boss which bears about
half-a-dozen small rounded teeth. A somewhat similar condition was pre-
viously known in Scullya, and in a large skull which I recently discovered
and which appears t o be near to, if not identical with, Anteosaurus magni$cus
of Watson ; there is also a rounded boss on each palatine which bears teeth
 A NEW TYPE OF DINOCEPHALIAN REPTILE. 737
The type of Anteosaurus magnificus was described and figured by Watson as
having three incisors in each premaxilla, but the specimen is in a very bad
state of preservation, and I see in notes I made on the type some years ago
I say “ possibly five incisors.” However it may be with this specimen, there
are certainly five incisors in the new skull recently discovered. If this skull
is Anteosaurus then we have three known genera with dentigerous bosses
on the palatines.
The pterygoids are very large. As in all known Dinocephalians they form
powerful descending pterygoid processes. I can h d no traces of teeth on
any part of the pterygoids. Posteriorly the pterygoids have processes passing
towards the quadrates as in all Therapsids, and between these processes lies
what I consider to be the vomer. In text-fig. 8, C, is shown an oblique section
through the vomer and the posterior part of the pterygoid, and the anterior
part of the base of the epipterygoid. Plates of the pterygoids extend upwards
and together form a very large median structure, as shown in text-fig. 5.
Text-figure 5.

Median section of skull of Dinosuchus vorsteri Broom.

Slightly under 6 nat. size.

The transpalatine is probably large, but it is impossible to trace the sutures

between it and the palatine and pterygoid. There is no suborbital vacuity.
The epipterygoid is well developed and satisfactorily preserved on both
sides. It consists of a long columella, which passes up to the parietal and
a long and wide base which lies on the pterygoid. The shape and the relations
of the bone are shown in text-fig. 6, A. The base is considerably foreshortened
in the figure.
Most of the bones round the brain-region are so thickened and fused that
only a very few of the sutures can be satisfactorily made out. The prootic,
the opisthotic, the supraoccipital, the exoccipital, and the basioccipital are
probably all essentially similar in general arrangements to those in the Thero-
cephalians, but it is only where the bones are broken across that sutures can
in places be seen. The occipital condyle is large and subspherical. It is clearly
composed of exoccipitals and basioccipital, and the jugular foramina are on its
outer sides as in the Anomodonts and Therocephalians. The exoccipitals
are probably as shown in text-fig. 4.
738 DR. R. BROOM ON THE STRUCTURE O F THE SKULL IN

The opisthotics form powerful paroccipital processes which support the

squamosals. Above each process is a relatively small temporal foramen,
and above the foramen is another powerful lateral process. This upper process
is probably entirely supraoccipital, though no trace of a suture between this
supposed supraoccipital and the opisthotic can be seen. This lateral supra-
occipital process is closely clasped by the squamosal and the tabular, as seen
in the section figured in text-fig. 6, C .
Above the supraoccipital is a relatively small median interparietal, but
the suture dividing it from the supraoccipital cannot be made out.
The tabular is a large bone which is curiously shaped. I n the upper part
of the occiput i t can be seen between the outer squamosal and the median

Text-figure 6.

Details of skull of Dinoauchus worsteri Broom. All slightly under f nat. size.
A. Front view of the two epipterygoids whose bases rest on the pterygoids and whose
upper ends articulate with the parietals. The vertical groove in the upper part
of the view is the pineal canal seen in section.
B. Front view of right quadrate and quadratojugal.
C. Transverse section through the left squamosal tabular and part of the supraoccipital.

interparietal, and the dividing sutures can be clearly traced. Inferiorly the
tabular sends down a narrow process which rests on the outer process of the
supraoccipital. It just fails to reach the opisthotic.
The squamosal is a huge bone which forms almost the whole of the outer
half of each side of the occiput. A descending ridge formed above by the tabular
and below by the squamosal divides the occiput proper from the squamosal,
and probably as in some other Therapsids the external auditory canal lay on
the outer aide of this ridge. Probably the muscles supporting the head were
all attached to the occiput proper and the squamosals stood much out from the
sides of the back of the head. The squamosal forms most of the back wall
 A NEW TYPE OF DINOCEPHALIAN REPTILE. 739

of the temporal fossa. The temporal arch is situated low down, and the
process of the squamosal which forms its outer side passes far forward on the
jugal, probably as far as indicated in the figure.
The quadrate is large, measuring 185 mm. in height and 143 mm. wide
a t its base. It is a nearly flat bone of moderate thickness. On its outer side

Text-figure 7.

Views of posterior half of right mandible of

Dinosuchus vorsteri Broom. Slightly under & nat. size.
A 1. The outer and under side. 2 R. Upper view. 3 C. Inner view. 4 D. Under side.

there is articulated to it a small quadratojugal, as shown in the figure. The two

bones articulate rather loosely with the squamosal and lie in a cavity in the
front of the lower half of the bone.
~ O C ZOOL.
. SOC.-1930. 50
740 DR. R. BROOM ON THE STRUCTURE O F THE SKULL IN

A median section of the skull is shown in text-fig. 5 . The prevomer and

the pterygoid are situated in somewhat the same positions as in the Thero-
cephalians, but the back part of the base of the skull is remarkably specialized.
I have figured the bones as they occur, and give the determinations that appear
to me to be most probable.
The basioccipital is large and the suture between i t and the basisphenoid
in front can be clearly made out. Above the occipital the exoccipitals meet
in the middle line as shown in the section.
The basioccipital appears to be small, but the sutures between it and the
prootics cannot be made out, and part of the suture between it and the vomer
(parasphenoid) is also obscure. There is a clearly marked suture between the
ba.sioccipita1 and the vomer below, as indicated in the figure, and the vomer
passes a little way under t.he basioccipital.

Text-figure 8.

Some details of the skull of Dinoauclm~vorsteri Broom.

A. Posterior end of the right mandible seen from behind. Slightly under 4 nat. size.
B. Section through right mandible at the line marked 5 in C of text-fig. 7. Slightly under
4 nat. size.
C. Oblique section through the posterior part of the pterygoid and the vomer. Slightly
under 4 nat. size.

The lower part of the vomer is essentially similar to that in the Thero-
cephalians, but above there is a most remarkable development. From a
narrow base the bone passes far up and forms a median fan-like plate. This
appears to be all part, of the vomer (parasphenoid). I n front of it is a long pit,
which is formed between lateral plates of the vomer and the prootics, in which
doubtless lay the pituitary ; but whether the pituitary extended right down
to the base of the narrow pointed funnel must remain at present uncertain.
The section shows the long canal for the pineal organ.
There is preserved the nearly perfect posterior half of the right mandible.
Of this I give five views and five sections. It will be seen that the jaw agrees
in essentials with that already described (1923) in Titunosuchw cloetei (Jonkeria
cloetri), but that it differs in being curiously twisted and in having a huge boss
on the upper part of the angular,
 A NEW TYPE OF DINOCEPHALIAN REPTILE. 741

Text-fig. 7 , 1 , shows the jaw as viewed from the outer side and slightly from
below. The angular has the outer plate well developed as in all typical Therap-
sids, and the large inner part has a wide articulation with the articular and the
surangular. Above, the l u g e elongated boss is seen.
Text-fig. 7, 2, is an upper view of the jaw. It shows the upper part of the
dentary, which behind meets the surangular to form the coronoid proqess.
The short broad articular is shown covered in part on its outer side by the sur-
angular and on its inner side by the prearticular. Nearly the whole length
of the prearticular is seen.
Text-figure 9.

Sections of the right mandible of Dinosuchwr corsteri Broom,

at the points indicated in C, text-fig. 7. Slightly under 4 nnt. size.

Text-fig. 7, 3, is an inner view of the jaw. It shows the relations of the

dentary, coronoid, prearticular, and splenial, the lower part of the angular
and near the whole of the inner side of the large surangular. Nos. 1, 2, 3, 4 ,
and 5 indicate the positions of the cross-sections shown in text-figs. 8 and 9.
Text-fig. 7, 4, is an under view of the jaw.
In text-fig. 8, A, we have a posterior view of the jaw. It shows the broad
articulation, the coronoid process, and the large angular boss.
The sections in text-fig. 9 show very clearly the relations of the bones near
the middle of the jaw. Thc little bone situated near the lower end of the
prearticular in text-fig. 9, 3, is evidently a portion of the angular. These
60+
742 THE SKULL I N A NEW TYPE OF DINOCEPHALIAN REPTILE.

sections and that seen in text-fig. 8, B, show that the Dinocephalian jaw is
essentially similar in structure to that of the Therocephalians.
The skull and mandible appear to indicate that the Dinocephalians have
been evolved from a proto-Therocephalian ancestor.

REFERENCES
TO LITERATURE.
WATSON, D. M. S. The Bases of Classification of the Theriodontia. P. Z. S. 1921,
pp. 35-98.
BROOM,
R. On the Structure of the Skull in the Carnivorous Dinocephalian Reptiles.
P. Z. S. 1928, pp. 661-684.
BROOM,R. On the Carnivorous Mammal-like Reptiles of the Family Titanosuchidze.
Ann. Transvaal Museum, xiii. i . 1929, pp. 9-36.
Other references will be found in
BROOM,
R. The Mammal-like Reptiles of South Africa. London, H. F. & G . Witherby,
1932.

.References to Lettering.
Ang., angular ; Art., articular ; B.O., basioccipital ; B.S., basisphenoid ; Co.,
coronoid ; Cor.F., carotid foramen ; D., dentary ; E.O., exoccipital ; E.Pt., epi-
pt,erygoid ; Fr., frontal ; I.P., interparietal ; Ju., jugal ; Mx., maxilla ; O.O., opisthotic
(paroccipital) ; O.Ang., outer plate of angular ; Pa., parietal ; Pal., palatine ; €'.Art.,
prearticular ; Pmx., premaxilla ; P.O., prootic ; Pt., pterygoid ; Q., quadrate ; Q j . , quad-
ratojugal ; S. Ang., surangular ; S.O.,supraoccipital ; Sp., splenial ; Sq., squamosal ;
St., stapes ; Tb., tabular ; T.p., transpalatine ; Vo., vomer (parasphenoid).