Morphological analysis of Protosuchus haughtoni and Protosuchus richardsoni in relation

to modern crocodilians and identification of a second P. haughtoni specimen

Branden Adams1, Kevin H. Lin1, Chi Zhang1 and Farish Jenkins, Jr.1

Abstract
Protosuchus comprises an extinct genus of dimunitive crocodylian carnivores that
prevailed from the early Jurassic to the early Cretaceous (Wu et al., 1997). Protosuchus
demonstrates several morphological markers observed in modern-day crocodiles and thus
represents the second stage of crocodylian evolution, arising from the sphenosuchians.
Furthermore, this genus serves as the basal root from which modern crocodiles are derived
(Clark et al., 2004). Two species of Protosuchus from the fossil record have formerly been
extensively characterized: P. haughtoni, from the Lower Jurassic Stormberg series in southern
Africa and P. richardsoni, from the Mesozoic Moenave Formation in North America (Gow,
2000). Since the skull morphologies of P. haughtoni and P. richardsoni are very distinct, with
the prior reminiscent of a terrestrial species and the latter an aquatic or amphibious organism,
these crocodilians likely occupied separate niches, thereby reducing interspecies competition as
they cohabitated the Jurassic with the ornithodyrans. Later, terrestrial competition pressures with
these larger ornithodyrans coerced the smaller protosuchians to readapt to an aquatic lifestyle
that is exhibited in today's crocodilian ambush predators. A third Prosuchus individual,
comprised of a skull and partial skeleton, was excavated in 1979 from fine-grained sandstone of
Dinosaur Canyon in the Moenave Formation but has yet to be uncharacterized. As this third
specimen shares key morphological skull features with P. haughtoni from southern Africa, it is
likely another P. haughtoni specimen.

Comparative analysis of P. richardsoni and P. haughtoni morphologies

Although members of the same genus, P. richardsoni and P. haughtoni demonstrate
remarkably distinct skull morphologies indicative of adaptations to different niches. P.
richardsoni exhibits a high degree of dorsoventral compression of the skull—such as that present
in extant crocodiles—which facilitates aquatic locomotion by reducing the area of contact with
water. This compression is lacking in P. haughtoni, which retains the ancestral heightened skull

1: Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA

* Corresponding Author: kevin.lin12@college.harvard.edu
 Adams, Lin, Zhang, and Jenkins 2

form of sphenosuchians. Furthermore, P. richardsoni displays dorsally-directed external nares, a

feature of present day crocodilians, whereas these elements in P. haughtoni are anteriorally
oriented, indicating that would be submerged upon entering an aquatic environment. However,
P. haughtoni does possess some diagnostic crocodilian features, including the palpebral bones
above the orbits to shield the eyes during violent feeding behaviors (Walker, 1968) that are
noticeably lacking in P. richardsoni.
Both species have bilateral mandibular fenestrations, but those in P. richardsoni are
significantly more prominent, thereby capable of housing larger adductor muscles and providing
more room for powerful muscle contractions than their P. haughtoni counterparts. Interestingly,
the presence of antorbital fenestra only in P. haughtoni may have served as a compensatory
feature for the greatly reduced mandibular fenestra in P. haughtoni, enabling adductor muscle
attachments or even serving to heighten pneumaticity and thermoregulation by housing soft-
tissue air sacs (Wedel, 2009). P. haughtoni does not appear to possess temporal fenestra, though
the poor preservation of the posteriad cranial features in both the Gow specimen and MCZ 6727
prevents definitive analyses. Conversely, P. richardsoni presents prominent temporal
fenestrations that are also present in modern crocodilians. These aid in increasing gape by
serving as attachment sites for longer muscle fibers (Frazzetta, 1968).
Thecodont dentition is observed in both species with the notable rostral notch in the
maxilla enabling jaw occlusion by providing space for the enlarged lower caniniform tooth of the
dentary in P. haughtoni, an attribute absent in P. richardsoni. Dental morphologies in both
species are exclusively caniniform and conical and are adapted for carnivorous diets, much like
in extant crocodiles, but the absolute number of teeth are fewer in protosuchians than in their
modern day relatives, likely a result of shorter snouts relative to the total skull length.
The skeletons of both species are pelaged with dorsal scutes, or osteoderms, that provided
protection from larger predators at the expense of mobility (Fig. 2). The cost of mobility would
have been reduced or negated by aquatic lifestyle, in which the buoyancy of water would have
offset the additional mass of denser dermal arrangements. However, the neural and haemal
spines on the caudal vertebrae are not adequately elongated to enable propulsion from the tail,
and since the caudal vertebrae themselves are still highly gracile in morphology, they appear to
be more suitable as counterbalances for cursorial species (Colbert and Mook, 1951). While the
characterization of the fore and hindlimbs in P. haughtoni is lacking on account of fossil
 Adams, Lin, Zhang, and Jenkins 3

preservation, such structures have, in fact, been documented by Colbert and Mook in P.
richardsoni. This species presents limbs that are more slender and elongate than those in modern
crocodilians, but shows an elongated radius and ulna in the forefoot and enlarged calcaneus and
astragalus in the hindfoot, all indicative of an advanced crocodilian (Colbert and Mook 1951).
These fossil limb patterns reveal a transition from a long and slender cursorial form to a shorter
and stouter aquatic form capable of tucking in limbs closer to the body to reduce drag while
swimming while also supporting the larger mass of the sculling tail.
Based on these protosuchian features, the modern day crocodilian is an admixture of both
species, deriving skull architecturual elements from both P. richardsoni and P. haughtoni.

Identification of MCZ 6727

Several significant cranial morphologies indicate that specimen MCZ 6727 (Fig. 1) is of
the species P. haughtoni, as originally described by Gow (2000). In particular, the maxilla of
MCZ 6727 exhibits a rostral notch bordered by premaxillary and nasal bones (Fig. 1C, 1F). This
notch receives an enlarged caniform tooth rooted in the dentary that, although absent from the
right side of the Gow specimen, has been preserved in MCZ 6727.
Small mandibular fenestra are present at the intersections of the angular and squamosal
bones (Fig. 1C, 1F), corresponding to the preserved right lateral mandibular fenestrum on the
original P. haughtoni. Additionally, MCZ 6727 retains antorbital fenestra (Fig. 1C, 1F) as in the
primitive diapsid condition (Benton, 1985) and displays a pair of parebrals (Fig. 1A, 1C, 1F) that
are similar in shape and placement to those of the Gow specimen. The skull of MCZ 6727 is
approximately half the size (40 mm) of that described by Gow, indicative of a juvenile condition.
This also provides a potential explanation for the lack of an ossified articulation between the
nasal and premaxillary bones at the anterior-most position of the rostrum (Fig. 1C, 1F).
The overall cranial architecture of MCZ 6727 is similar to that of P. haughtoni. The ratio
of the width at the widest point (the lateral-most margins of the jugal bones) to length (parietal to
the tip of the rostrum) is 0.70 while the corresponding ratio for P. haughtoni is 0.78, well within
an appropriate range for specimens of different ages. However, these calculations may be
inaccurate on account of geologic distortion of the cranium during fossilization, as evident in
Figure 1B. Therefore, while the lateral view of MCZ 6727's cranium reveals a more dorso-
ventrally compressed skull than that described by Gow, this discrepancy may result from
 Adams, Lin, Zhang, and Jenkins 4

compression and shearing during fossilization rather than an actual morphological differences
between the two specimens. This set of diagnostic morphological features and the general cranial
architecture indicate that MCZ 6727 is indeed of the species P. haughtoni.

Discussion
While the morphologies of the P. haughtoni and P. richardsoni skulls reveal two
physiologically distinct species, the ultimate contributions from keystone adaptations in these
early crocodilians to their present-day brethren are shared. Some traits that appear in the family
Crocodylidae are manifested in P. haughtoni but not in P. richardsoni, and vice versa. This
implies that crocodilian evolution from protosuchians may not be a monospecific process but
rather a conglomerative process among multiple distinct species, which may account for the
morphological diversity of modern crocodilian species.
But what impetus would spur such a divergence in the morphology of the two
protosuchians? One hypothesis suggests that niche specialization reduced competition among
species. This phenomenon is most famously known from the variation in the beak shapes of
Darwin's finches but can be applied to all species as a means of maximizing available resources
when sharing a common geographic environment. Hence, the more depressed skull of P.
richardsoni was likely more suited for aquatic habitats than P. haughtoni, which had cranial
features adapted for terrestrial hunting. While P. richardsoni may not have been fully aquatic
given its highly cursorial skeletal structure, these cranial adaptations may have enabled it to hunt
piscine prey from the shore while P. haughtoni competed with contemporaneous ornithyodyrans
for terrestrial prey. As a result, P. richardsoni and P. haughtoni may have developed into
separate species on account of niche specialization—even if they evolved from a common
sphenosuchian ancestor—and independently acquired aquatic adaptations on account of
competition pressures from the larger terrestrial ornithodyrans during the Jurassic, forcing an
exodus to a more spacious and resource-laden aquatic environment.
Protosuchians represent an ancestral form from which modern crocodiles arose and serve
as particularly convincing models for the transitions of some organisms from land back to water,
as evidenced in skull and limb morphologies. The two characterized species each possess distinct
crocodilian features that may have developed as a result of selective pressures from
 Adams, Lin, Zhang, and Jenkins 5

ornithodyrans and show independently-acquired traits suitable for aquatic lifestyles as a result of
niche specialization.

References
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