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Morphology of the palate and braincase of Dimetrodon milleri

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DOI: 10.1080/08912963.2012.704918

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Historical Biology
Vol. 24, No. 5, October 2012, 453–459

PUBLISHER’S NOTE

This paper is part of the 12th International Symposium on Early Vertebrates/Lower Vertebrates special
issue (Historical Biology, Vol. 24, No. 4, August 2012). We extend sincere apologies to the authors for
this mistake.

12th International Symposium on Early Vertebrates/Lower Vertebrates

Morphology of the palate and braincase of Dimetrodon milleri

Kirstin S. Brinka* and Robert R. Reiszb1

a
Department of Ecology and Evolutionary Biology, University of Toronto Mississauga, 3359 Mississauga Road, Mississauga, Ontario,
Canada L5L 1C6; bDepartment of Biology, University of Toronto Mississauga, 3359 Mississauga Road, Mississauga, Ontario, Canada
L5L 1C6
(Received 8 March 2012; final version received 18 June 2012)

The palate and partial braincase of the holotype of Dimetrodon milleri (MCZ 1365) are preserved in three dimensions, but have
yet to be described in detail. Here, we describe these structures for the first time for this species, and compare them with the
better-known specimens of D. limbatus. Interesting characteristics of the morphology include the patterns of articulation of the
palatal elements, including the palatine and vomer, and anatomy of the pterygoid in the posterior region of the palatal vacuities.
Dimetrodon milleri is found to differ from D. limbatus in the lack of teeth on the ectopterygoid, the shape of the basal process of
the epipterygoid, and the anterior extent of the palatine and pterygoid. The two species are similar in the relative position of the
basicranial articulation, but differ significantly from that in other sphenacodontids, including Secodontosaurus and
Sphenacodon. The evolution of these cranial features will be the subject of future phylogenetic analyses of sphenacodontids.
Keywords: non-mammalian synapsid; Permian; Sphenacodontidae; Therapsida

Introduction Previous phylogenetic analyses that have considered

Sphenacodontids are derived non-therapsid predatory
synapsids known from Laurasia, with a fossil record that
extends from the Late Carboniferous to the Middle Permian.
The sphenacodontid Dimetrodon Cope, 1878 is an Early
Permian (295–270 Ma) non-therapsid synapsid known from
southern USA and Germany (Reisz 1986; Berman et al.
2001). Dimetrodon is of exceptional importance as it is the
most abundant and speciose sphenacodontid, providing
valuable information on issues related to the origin of
therapsids. Twelve species of Dimetrodon are currently
recognised from hundreds of specimens, but these taxa are
primarily identified on the basis of size and stratigraphic and
geographic locations (Romer and Price 1940; Reisz 1986;
Berman et al. 2001). However, little is known about the
relationships among species of Dimetrodon, as few
characters have been identified that are useful for species-
level phylogenetic analyses.
Dimetrodon at the genus-level have proposed different
hypotheses regarding relationships among sphenacodon-
tids (Reisz et al. 1992; Fröbisch et al. 2011; Benson 2012).
Reisz et al. (1992) hypothesised that Dimetrodon is more
closely related to Secodontosaurus than to Sphenacodon,
whereas two recent analyses suggest a closer relationship
between Dimetrodon and Sphenacodon (Fröbisch et al.
2011; Benson 2012). The relationships within Sphenaco-
dontidae and specifically those among the 12 recognised
species of Dimetrodon are important as this clade is
generally considered to be the sister group to Therapsida
(which includes mammals). Thus, a greater understanding
of the evolutionary relationships within the group will
provide more detailed information about the acquisition of
mammalian characters within Synapsida (Reisz 1986; Liu
et al. 2009; Amson and Laurin 2011).
The stratigraphically oldest-known species of Dime-
trodon, D. milleri Romer, 1937, is known from only two

*Corresponding author. Email: kirstin.brink@mail.utoronto.ca

ISSN 0891-2963 print/ISSN 1029-2381 online
q 2012 Taylor & Francis
http://dx.doi.org/10.1080/08912963.2012.704918
http://www.tandfonline.com
454 K.S. Brink and R.R. Reisz
Figure 1. Dimetrodon milleri, MCZ 1365 palate and braincase, ventral view. ec, ectopterygoid; pal, palatine; pbs, parabasisphenoid;
pm, premaxilla; pt, pterygoid; v, vomer. Diagonal lines indicate plaster reconstruction.
specimens: one nearly complete skull and skeleton (MCZ
1365) and one slightly larger articulated, less complete
skull and skeleton (MCZ 1367). A proportionately tall
skull, a shortened face relative to the temporal region of the
skull, and vertebrae that are anteroposteriorly short
characterise D. milleri, and are also present in the younger
species D. booneorum, D. limbatus and D. grandis. Because
of these similarities, these four species have been suggested
to belong to an evolutionary series (Romer and Price 1940).
One potential autapomorphy of D. milleri is the
morphology of the elongate neural spines, which are
circular in cross section and do not appear to display
pronounced anterior and posterior grooves that are
prevalent in the neural spines of the other species (Romer
and Price 1940; Huttenlocker et al. 2010).
The palate of MCZ 1365 is exceptionally preserved and
nearly complete, and was used to supplement the
description of the palate of D. limbatus (MCZ 1347,
Romer and Price 1940). However, the palatal anatomy of
individual Dimetrodon species has never been examined for
specific taxonomic differences. Therefore, characters that
are potentially useful for species identification may have
been overlooked, and changes in the morphology of the
palate and braincase have never been examined in a
temporal or phylogenetic context. Here, we provide a
detailed description of the newly prepared articulated palate
and braincase of D. milleri (MCZ 1365), and the unique
morphology of these complexes will be used to discern
characters useful for phylogenetic analysis for future
studies of non-mammalian synapsids.
Institutional abbreviations: AMNH, American
Museum of Natural History; FMNH, Field Museum of
Natural History; MCZ, Harvard Museum of Comparative
Zoology.
Description
General structure
The palate and braincase of D. milleri (MCZ 1365) are
preserved in three-dimensional shape with little tapho-

Figure 2. Dimetrodon milleri, MCZ 1365 palate and braincase, dorsal view. bo, basioccipital; cp, cultriform process; ec, ectopterygoid;
ep, epipterygoid; pal, palatine; pbs, parabasisphenoid; pmx, premaxilla; pt, pterygoid; v, vomer. Diagonal lines indicate plaster
reconstruction.
nomic distortion (Figures 1 –4). The braincase and occiput
are positioned on the dorsal margin of the pterygoids,
slightly anterior to the transverse process of the pterygoids,
and may be slightly displaced from the original life
position. When in articulation, the palate is bounded
anteriorly by the premaxilla, laterally by the maxilla,
posterolaterally by the quadratojugal and jugal, and
posteriorly by the braincase. The complex is no longer
articulated with the rest of the cranium, and appears to
have been cut from the articulation with the maxilla and
premaxilla, as small pieces of bone with tooth roots are
attached to the lateral margins of the palatines and to the
anterior margin of the vomers (Figure 3). Nevertheless,
this separation of the palate and braincase from the rest of
the skull has revealed important morphological features
that are described in detail for the first time.
Vomer
The paired vomers are elongate elements that run
anteroposteriorly from their contact with the premaxilla
Historical Biology 455
to their contact posteriorly with the pterygoid and
posterolaterally with the palatine (Figures 1– 3). The length
of the vomer is just under half the total length of the palatal
surface, from the anterior tip of the vomer to the transverse
flange of the pterygoid (43%), which is similar to the
proportions in D. limbatus (45%). The element is slightly
enlarged anterolaterally at its contact with the premaxilla,
narrow for the majority of its length, and expanded again at
its contact with the pterygoid posteromedially and with the
palatine posterolaterally (Figure 1). The paired vomers
come into contact with each other anteriorly at the midline,
and are slightly separated from each other throughout the
remainder of their length. This trough between the vomers
was most likely for passage of air from the external narial
opening into the oral cavity. The vomer has an S-shaped
posteroventral margin with an elongate medial process,
which articulates onto the ventral margins of the pterygoid
and palatine (Figure 1). This morphology is significantly
different from that illustrated for D. limbatus, where the
posterior margin of the vomer is smooth and forms a J-shape
456 K.S. Brink and R.R. Reisz

Figure 3. Dimetrodon milleri, MCZ 1365 palate and braincase, right lateral view. bo, basioccipital; cp, cultriform process; ec,
ectopterygoid; ep, epipterygoid; m, maxilla; pal, palatine; pbs, parabasisphenoid; pm, premaxilla; pt, pterygoid; v, vomer. Diagonal lines
indicate plaster reconstruction.

along the lateral edge of the medial process (Romer and
Price 1940), and from the morphology described for
Sphenacodon ferox, where the articulation between the
vomer and palatine is V-shaped (Spielmann et al. 2010). In
D. milleri, the bone surface of the vomer is highly rugose in
the area of its articulation with the palatine and pterygoid.
The element is edentulous except for one tooth located
within the rugose bone of the left vomer (Figure 1). This
tooth is incomplete, but the root is identical to those on the
palatine and pterygoid.
Dorsally, the vomer articulates with the anterior
process of the palatine, where the lateral edges of the
vomer are expanded dorsally and curve around the lateral
edge of the palatine (Figures 2 and 3). Posteriorly, the
vomer is arched ventrally, following the deep curve of the
palatine (Figure 3). The dorsal surface of the vomer along
the anterior process is grooved for the passage of blood
vessels (Romer and Price 1940). The medial margin of the
vomer is dorsoventrally expanded to form the dorsal
lamina of the vomer, which is continuous with the dorsal
lamina of the palatine and pterygoid and forms a medial
separation of the nasal cavity (Figure 3). The height of the
dorsal lamina is similar to the height of the lamina in S.
ferocior (Eberth 1985).

Figure 4. Dimetrodon milleri, MCZ 1365, palate and braincase, occipital view. bo, basioccipital; ep, epipterygoid; pal, palatine; pbs,
parabasisphenoid; pt, pterygoid. Diagonal lines indicate plaster reconstruction.

Palatine
The palatine forms part of the dentigerous roof of the oral
cavity. It articulates with the posterodorsal margin of the
vomer anteriorly. An anterior wing-shaped process,
ventral to the contact with the vomer, articulates laterally
with the maxilla, and remains slightly dorsoventrally
expanded for its length while abutting the alveolar ridge of
the maxilla (Figures 2 and 3). The palatine extends
posteriorly to contact the ectopterygoids posterolaterally
and the pterygoids medially and posteriorly. The
denticulate portion of the ventral surface of the palatine
is convex. The anterior portion of the palatine is slightly
elongated, extending one quarter the length of the vomer,
as in D. limbatus (Figure 3). A ridge runs diagonally along
the dorsal margin of the element towards the lateral contact
with the maxilla, and continues anteriorly to abut the
pterygoid and vomer medially and form the dorsal lamina.
The palatine is dorsoventrally expanded in this region,
and is about twice the height of the vomer (Figure 3).
This contribution is more than the contribution to the
dorsal lamina in D. limbatus, where the palatine is about
the same height as the vomer (Romer and Price 1940).
Ectopterygoid
The left and right ectopterygoids are partially preserved in
MCZ 1365 (Figures 2 and 3). This element is bounded
anteromedially by the palatine and posteromedially by the
pterygoid, forming a small portion of the roof of the
mouth. Laterally, the ectopterygoid thickens at its
midpoint where it contacts the maxilla, and is thinner
when separated from the maxilla anteriorly and posteriorly
by a small portion of the pterygoid.
The ectopterygoid is illustrated by Romer and Price
(1940) as possessing irregularly arranged teeth in
D. limbatus based on the presence of two non-articulated
bones preserved in AMNH 4363 (Case 1910). This
element has not been described for any other species of
Dimetrodon. Here, the ectopterygoid of D. milleri does not
bear teeth, and although incomplete, the distribution of the
teeth in D. limbatus suggests that the portions of the bone
that are preserved in MCZ 1365 should bear teeth, but do
not. Ectopterygoid teeth are not preserved in S. ferocior
(Eberth 1985), Haptodus garnettensis and H. baylei
(Currie 1979; Laurin 1993), and the ectopterygoid appears
to be edentulous in S. ferox (Spielmann et al. 2010) and
Secodontosaurus (Reisz et al. 1992).
Pterygoid
The pterygoid is the largest bone of palatal complex, and has
a triradiate shape typical of basal synapsids (Romer and
Price 1940; Reisz 1986). In the anterior region, the dorsal
margin of the pterygoid is dorsoventrally expanded to form
Historical Biology 457
the tall dorsal lamina, which, together with the dorsal
lamina of the palatine, creates a tall median separation into
the orbital region of the skull, as in the nasal cavity. The
anterior extension of the dorsal lamina is longer than
illustrated for D. limbatus (Romer and Price 1940),
extending about half the length of the vomer, twice as
long as the anterior extension of the palatine (Figure 3).
Toothed triangular-shaped pterygoid flanges demarcate the
posterior margin of the pterygoid. Each transverse flange
has six – seven teeth, which is less than the number
preserved in D. limbatus and Ctenospondylus ninevehensis
(Berman 1978), ranging from 10 to 11 (AMNH 4363, MCZ
1347). However, the presence of six – seven teeth is the same
as the pterygoid tooth counts reported for Sphenacodon
(Eberth 1985; Spielmann et al. 2010) and Haptodus (Laurin
1993), possibly representing the primitive condition.
Although the number of teeth on the transverse flange
may be species-specific (Case 1907), it is also possible that
it may be a factor of size (Reisz et al. 1992). However, these
two alternatives have not been examined in detail.
The pterygoid flanges of D. milleri are not angled sub-
horizontally towards the midline as in D. limbatus (Case
1910; Romer and Price 1940), but rather are more
horizontal, as in Haptodus (Laurin 1993). The orientation
of the transverse process is difficult to interpret in
Sphenacodon. Additionally, the posterior margins of the
transverse flanges are closely apposed, which differs
significantly from the large interpterygoid vacuities
described for D. limbatus (Romer and Price 1940).
However, both the angle and positioning of the flanges
could be due to taphonomic distortion. The quadrate
processes of the pterygoid are only partially preserved
posterodorsally, but the bases of the processes suggest that
the pterygoid flares laterally posterior to the braincase
(Figure 4). The pterygoid does not contribute to the
basicranial articulation, as in Haptodus (Laurin 1993).
Epipterygoid
The epipterygoids are preserved in place as large sheets of
bone anterior to the braincase, dorsal to the pterygoid. The
slender, rod-like dorsal portion of the epipterygoid is not
preserved in MCZ 1365. As in other species of Dimetrodon,
the basicranial articulation is largely restricted to the mid-
point of the epipterygoid, and is located far dorsally from
the ventral surface of the palate. This dorsal position of the
basicranial joint is significantly different from other
sphenacodontids in which this area of the skull is preserved,
and may be an autapomorphy of the genus Dimetrodon.
The relative position of the basicranial articulation is
anterior to the transverse flange of the pterygoid, which
differs from the reconstructions of D. limbatus (Case
1907, 1910; Romer and Price 1940) and S. ferocior
(Eberth 1985) where the basicranial articulation is farther
back in the skull relative to the transverse flange. This
458 K.S. Brink and R.R. Reisz
positioning could be due to taphonomic distortion. In
posterior view, the basicranial articulation is visible where
the parabasisphenoid articulates laterally with the
epipterygoid via the basipterygoid process, lateral to the
basal tubera (Figure 4).
The flange of the epipterygoid ventral to the
basicranial articulation is wide and significantly different
from the slender base described for D. limbatus (Case
1910; Romer and Price 1940). The shape of the base of the
epipterygoid of D. milleri is similar to that of
Secodontosaurus and Haptodus, and is slightly triangular
in lateral view (Reisz et al. 1992; Laurin 1993). The paired
epipterygoids are separated medially by the cultriform
process of the parabasisphenoid (Figure 2).
Parabasisphenoid
The parabasisphenoid has been described previously as a
disarticulated element for Dimetrodon limbatus, Sphena-
codon and Ctenospondylus casei, and is not significantly
different in MCZ 1365 (Baur and Case 1899; Case 1904;
Vaughn 1964; Eberth 1985). Here, it is preserved in
articulation laterally with the epipterygoids at the
basicranial articulation. A small incomplete portion of
the basioccipital is articulated on the dorsal margin of the
body of the parabasisphenoid. The cultriform process is
broken anteriorly and the preserved height of the process is
similar to the height of the parabasisphenoid process. The
true height of the element is not known in this specimen.
At the connection of the cultriform process to the body of
the parabasisphenoid, there is a small dorsal fossa, the
retractor pit, which is relatively deep in S. ferox (Eberth
1985) and C. casei (Vaughn 1964), but the depth of this
fossa is not well understood in MCZ 1365 as the cultriform
process is broken (Figure 3). Posterior to the retractor pit,
the symmetrical dorsal pillae are preserved, and would
articulate dorsally with the prootic to form the base of the
braincase (Reisz 1986).
Posteriorly, the basal tubera extend ventrolaterally and
are thickened and marked with muscle scars ventrally,
likely for the attachment of prevertebral musculature
(Romer and Price 1940). These elements are convex
ventrally (Figure 4). The basal tubera are shorter than the
elongate processes considered to be the primitive
condition, as seen in Haptodus (Laurin 1993). Anterior
to the basal tubera, two small foramina are visible on the
ventral surface of the parabasisphenoid, marking the
passage of the internal carotid artery. These foramina lie
medial to the basipterygoid processes that articulate
laterally with the epipterygoids (Figure 4).
Discussion
Several features of the palate and partial braincase of
Dimetrodon milleri described here can be considered as
autapomorphies for the species. These include the
enlarged ventral process of the epipterygoid, and the
anterior and dorsal extension of the palatine and pterygoid
and their contribution to medial separation of the nasal
cavity. As these characters have only been described in
some detail for D. limbatus, their presence in other species
of Dimetrodon is ambiguous, and have the potential to be
considered as synapomorphies with the examination of
new specimens in future analyses.
The morphology of the other elements of the braincase
and palate show many similarities to those described for S.
ferocior (Eberth 1985) and D. limbatus (Romer and Price
1940), and we tentatively interpret these similarities as
sphenacodontid features. The position of the basicranial
articulation relative to the ventral surface of the palate
appears to be slightly higher in D. milleri than in
Sphenacodon. This is likely due to the derived shape of the
skull, a feature that may characterise the genus
Dimetrodon. Interestingly, this relatively high position of
the braincase is derived in Dimetrodon, as the dorsal
location of the basicranial articulation relative to the
transverse process of the pterygoid is more similar to
Tetraceratops than to Haptodus (Amson and Laurin 2011).
The lack of ectopterygoid teeth has been used in
previous phylogenetic analyses as an apomorphy for
Therapsida (Laurin and Reisz 1990; Liu et al. 2009;
Amson and Laurin 2011). This study suggests that the
presence of ectopterygoid teeth is variable among
sphenacodontids, and may be a useful character in
understanding the relationships among species of Dime-
trodon, as well as the relationship of sphenacodontids with
therapsids, but this remains to be tested within a
phylogenetic context. Currently, the lack of ectopterygoid
teeth is only known in D. milleri, but this anatomy remains
unknown in other species, such as D. natalis, D. grandis
and D. giganhomogenes. The teeth are present in
specimens of D. limbatus (Romer and Price 1940) and
D. loomisi (FMNH UC 40). However, whether the
presence of teeth on the ectopterygoid is related to skull
size and width has not been examined in detail.
The articulation of the vomer, palatine and pterygoid at
the posterior margin of the nasal cavity suggests a very
strongly buttressed skull in the region of the marginal
dentition. Such buttressing may have served to reinforce
this region of the skull, resisting deformation forces during
prey capture. A somewhat unexpected feature of this
palate is the close apposition of the posterior region of the
transverse processes of the pterygoids, resulting in a small
interpterygoid vacuity. It is very difficult to determine at
this stage if this is a feature that characterises this taxon, or
whether this is a more widely distributed character among
Dimetrodon species, or among sphenacodontids in
general. This feature is found to be present in the stem
therapsid Tetraceratops (Amson and Laurin 2011), and its
presence in this sphenacodontid may have a significant

impact on our understanding of the acquisition of therapsid
features in the Early Permian.
Acknowledgements
The authors would like to thank Farish Jenkins, Jessica Cundiff
and Larry Flynn of the MCZ for facilitating the loan of the
specimen, Diane Scott for assistance with specimen preparation
and illustration, and Jessica Hawthorn for manuscript edits. Two
anonymous reviewers provided comments that greatly improved
the manuscript. Funding provided by the Natural Sciences and
Engineering Research Council of Canada to KSB and RRR, and
the MCZ for an Ernst Mayer Travel Grant to KSB.
Note
1. Email: robert.reisz@utoronto.ca
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