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Cranial Osteology and Phylogenetic Relationships of Hamadasuchus Rebouli (Crocodyliformes: Mesoeucrocodylia) from the Cretaceous of Morocco

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Cranial Osteology and Phylogenetic Relationships of Hamadasuchus Rebouli (Crocodyliformes: Mesoeucrocodylia) from the Cretaceous of Morocco Blackwell Publishing LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082© 2007 The Linnean Society of London? 2007 1494 533567 Original Articles HAMADASUCHUS REBOULIH. C. E. LARSSON and H.-D. SUES Zoological Journal of the Linnean Society, 2007, 149, 533–567. With 9 figures Cranial osteology and phylogenetic relationships of Hamadasuchus rebouli (Crocodyliformes: Mesoeucrocodylia) from the Cretaceous of Morocco HANS C. E. LARSSON1* and HANS-DIETER SUES2 FLS 1Redpath Museum, McGill University, 859 Sherbrooke Street W., Montréal, QC H3A 2K6, Canada 2National Museum of Natural History, Smithsonian Institution, NHB MRC 106, PO Box 37012, Washington, DC 20013–7012, USA Received February 2005; accepted for publication June 2006 This paper presents a detailed description of the skull and part of the mandible of the crocodyliform reptile Hama- dasuchus rebouli from the Kem Kem beds (Upper Cretaceous: Albian–Cenomanian) of south-eastern Morocco. This taxon of deep-snouted ziphodont crocodyliform can be diagnosed by a number of autapomorphies. Phylogenetic anal- ysis of a diverse array of crocodylomorph taxa found strong support for a clade comprising H. rebouli, Peirosauridae, and Sebecus. The name Sebecia nom. nov. is proposed for this grouping, which is diagnosed by numerous charac- ters, including the participation of the quadratojugal in the mandibular condyle. The distribution of this diverse and long-lived clade lends further support to the biogeographical hypothesis that faunal connections existed between Africa and South America well into mid-Cretaceous times. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 533–567. ADDITIONAL KEYWORDS: Crocodylomorpha – Hamadasuchus – skull. INTRODUCTION Araripesuchus wegeneri (Buffetaut & Taquet, 1979; Buffetaut, 1981a; Ortega et al., 2000; referred to The evolutionary history of Mesozoic crocodyliform Hamadasuchus by Prasad & Lapparent de Broin, reptiles from Africa is still poorly understood. Most of 2002), the longirostrine Stolokrosuchus lapparenti the relatively few forms known to date are Cretaceous (Larsson, 2000; Larsson & Gado, 2000), and the small in age. Stromer (1914, 1925, 1933, 1936) described a notosuchian Anatosuchus minor (Sereno et al., 2003). series of crocodyliform taxa from the Upper Creta- Furthermore, several crocodyliform taxa have been ceous (Cenomanian) of the Bahariya Oasis in the reported from the Albian–Cenomanian-age Kem Kem Western Desert of Egypt. Most noteworthy among beds of south-eastern Morocco. Lavocat (1955) briefly these are Libycosuchus brevirostris (Stromer, 1914; reported (without illustration) on fragments of a skull Buffetaut, 1976b) and the huge, ‘duck-billed’ Stomato- of a longirostrine form, which he named Thoracosau- suchus inermis (Stromer, 1925, 1936; the holotype was rus cherifiensis; this material, along with more com- destroyed during World War II). Early Cretaceous plete specimens, has now been placed in a new genus strata in Niger have also yielded assemblages of cro- Elosuchus by Lapparent de Broin (2002). Buffetaut codyliform reptiles, including the giant pholidosaurid (1976b) illustrated and referred a partial cranium to Sarcosuchus imperator (de Broin & Taquet, 1966; Libycosuchus sp.. Buffetaut (1994) designated a par- Taquet, 1976; Buffetaut & Taquet, 1977; Buffetaut, tial left dentary with six teeth (Musée des Dinosaures, 1981b; Sereno et al., 2001), the enigmatic Tremato- Espéraza, no. MDE C001) as the holotype of Hamada- champsa taqueti (Buffetaut, 1974, 1976a), the smaller suchus rebouli, which he included in the family Trematochampsidae (see below). Subsequently, Larsson & Sidor (1999) referred isolated teeth *Corresponding author. E-mail: [email protected] from the Kem Kem beds to H. rebouli and other © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 533–567 533 534 H. C. E. LARSSON and H.-D. SUES multicuspid teeth to indeterminate crocodyliforms, mentary braincase of a smaller specimen; ROM 52045, and Prasad & Lapparent de Broin (2002) discussed nearly complete right dentary of a large individual; and the microstructure of teeth of H. rebouli. ROM 52047, left dentary of a small specimen. In this paper, we describe a series of exquisitely pre- Revised diagnosis: differs from other known cro- served specimens that are referable to H. rebouli and codyliforms in the following combination of characters are housed in the vertebrate palaeontological collec- in adult specimens. Contribution of nasals to interna- tions of the Royal Ontario Museum (ROM) in Toronto. rial bar exceeding 50%; dorsomedial edges of They comprise the complete skull of a large individual supratemporal fenestrae level with skull table; (ROM 52620), the interorbital region of the skull roof tapered distal squamosal prong; large posteroventral of another larger specimen (ROM 54585), an associ- process on postorbital that contacts quadrate and ated snout and partial left mandibular ramus of a quadratojugal; external auditory meatus fossa extend- smaller individual (ROM 49282), posterior portions of ing anteriorly over entire length of postorbital; two well-preserved crania (ROM 52059, with associ- supratemporal fossa covering most of bony bar ated left jugal and quadratojugal, and ROM 54511), a between supratemporal fenestra and orbit; thickened fragmentary braincase (ROM 54113) of smaller speci- premaxillary extension over posterodorsal corner of mens, a nearly complete right dentary of a large indi- external naris forming notch; small incisive foramen; vidual (ROM 52045), and a left dentary of a small palatine–pterygoid suture extending to posterior specimen (ROM 52047). This wealth of superb new angle of suborbital fenestra; ectopterygoid–maxilla material permits, for the first time, a detailed account suture approaching posteromedial margin of maxil- of the cranial structure of H. rebouli and an assess- lary tooth row; absence of posterior ectopterygoid pro- ment of the phylogenetic position of this distinctive cess along ventral surface of jugal; absence of taxon within Crocodyliformes. prominent crest on dorsal surface of distal end of The fossils reported here were recovered by local col- quadrate; and prominent bilateral posterior projec- lectors in south-eastern Morocco from predominantly tions on posterodorsal surface of supraoccipital. An red continental sandstones, which are known as the autapomorphy not previously reported in other cro- Kem Kem beds (Sereno et al., 1996). These strata are codyliforms is the presence of shallow dorsal and ven- generally considered to be between Albian and Cen- tral grooves extending anteriorly from the antorbital omanian in age because they are conformably overlain fossa (the fossa is only present on one side in the by limestones of late Cenomanian age (Neolobites single complete cranium currently known for vibrayeanus Zone; Courville et al., 1991). The exact Hamadasuchus). provenance for the fossils cannot be established. The Kem Kem beds have yielded abundant often exquis- Distribution: Kem Kem beds, south-eastern Morocco. itely preserved, if typically disassociated, skeletal Age: Cretaceous (Albian–Cenomanian). remains representing a diverse assemblage of fishes and reptiles (Russell, 1996; Sereno et al., 1996). DESCRIPTION SYSTEMATIC PALAEONTOLOGY SKULL CROCODYLIFORMES CLARK IN BENTON & CLARK The following anatomical description is based prima- (1988) MESOEUCROCODYLIA WHETSTONE & rily on the superbly preserved skull ROM 52620 WHYBROW, 1983 SENSU CLARK IN BENTON & CLARK (Figs 1–5). The only preservational deficiencies of this (1988) METASUCHIA BUFFETAUT, 1981 SENSU CLARK specimen are either damage to or loss of several teeth, IN BENTON & CLARK (1988) the loss of the palpebral bones (the former presence of which is indicated by articular facets on adjoining cra- SEBECIA NOM. NOV. HAMADASUCHUS REBOULI nial elements), an oblique (repaired) fracture through BUFFETAUT, 1994 the snout, some breakage in the palatal region, and Referred specimens: ROM 52620, complete skull of a minor damage to the braincase in the proximal region large individual (Figs 1–5); ROM 54585, interorbital of the right paroccipital process. As a result of distor- region of the skull roof of another larger specimen; tion during fossilization, the sides of the snout are no ROM 49282, associated snout and partial left man- longer symmetrically aligned so that the rostrum dibular ramus (Fig. 6) of a smaller individual; appears skewed towards the right when viewed from ROM 54512, fragmentary left maxilla of a smaller indi- the front. vidual; ROM 52059, posterior portion a well-preserved In dorsal view, the outline of the cranium is that braincase with the left jugal and quadratojugal of an elongate triangle. (See Table 1 for selected (Fig. 7A–C); ROM 54511, posterior portion a well- measurements of ROM 52620.) Snout length in preserved braincase (Fig. 7B–D); ROM 54513, frag- ROM 52620 is about 70% of the basal skull length – the © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 533–567 HAMADASUCHUS REBOULI 535 Figure 1. Cranium of Hamadasuchus rebouli (ROM 52620). A, dorsal and C, ventral view. B and D, outline drawings cor- responding to each view. Scale bar = 10 cm. Anatomical abbreviations are defined in Appendix 1. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 533–567 536 H. C. E. LARSSON and H.-D. SUES Figure 2. Cranium of Hamadasuchus rebouli (ROM 52620).
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