Young Et Al., 2014)

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Biological Journal of the Linnean Society, 2014, 113, 854–871. With 11 ﬁgures

Marine tethysuchian crocodyliform from the ?Aptian-Albian (Lower Cretaceous) of the Isle of Wight, UK

MARK T. YOUNG1,2*, LORNA STEEL3, DAVIDE FOFFA4, TREVOR PRICE5, DARREN NAISH2 and JONATHAN P. TENNANT6

1Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, The King’s Buildings, Edinburgh EH9 3JW, UK 2School of Ocean and Earth Science, National Oceanography Centre, University of Southampton, Southampton SO14 3ZH, UK 3Department of Earth Sciences, Natural History Museum, London SW7 5BD, UK 4School of Earth Sciences, University of Bristol, Wills Memorial Building, Bristol BS8 1RJ, UK 5Dinosaur Isle Museum, Sandown, Isle of Wight PO36 8QA, UK 6Department of Earth Science and Engineering, Imperial College London, London SW6 2AZ, UK

Received 1 February 2014; revised 5 May 2014; accepted for publication 7 July 2014

A marine tethysuchian crocodyliform from the Isle of Wight, most likely from the Upper Greensand Formation (upper Albian, Lower Cretaceous), is described. However, we cannot preclude it being from the Ferruginous Sands Formation (upper Aptian), or more remotely, the Sandrock Formation (upper Aptian-upper Albian). The specimen consists of the anterior region of the right dentary, from the tip of the dentary to the incomplete fourth alveolus. This specimen increases the known geological range of marine tethysuchians back into the late Lower Cretaceous. Although we refer it to Tethysuchia incertae sedis, there are seven anterior dentary characteristics that suggest a possible relationship with the Maastrichtian-Eocene clade Dyrosauridae. We also review ‘middle’ Cretaceous marine tethysuchians, including putative Cenomanian dyrosaurids. We conclude that there is insufficient evidence to be certain that any known Cenomanian specimen can be safely referred to Dyrosauridae, as there are some cranial similarities between basal dyrosaurids and Cenomanian–Turonian marine ‘pholidosaurids’. Future study of middle Cretaceous tethysuchians could help unlock the origins of Dyrosauridae and improve our understanding of tethysuchian macroevolutionary trends. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 854–871.

ADDITIONAL KEYWORDS: Dyrosauridae – Ferruginous Sands Formation – Pholidosauridae – Sandrock Formation – Tethysuchia – Upper Greensand Formation.

INTRODUCTION tooth count (e.g. Koken, 1887; Mook, 1933, 1934; Wu, Russell & Cumbaa, 2001; Jouve et al., 2005a, 2006a; Tethysuchian crocodyliforms were a highly successful Barbosa, Kellner & Viana, 2008). Tethysuchia was group, some of which returned to a marine lifestyle one of several crocodyliform clades that survived the during the latter part of the Mesozoic and early end-Cretaceous mass-extinction event, with the Cenozoic. Many species superﬁcially resembled subclade Dyrosauridae continuing to radiate and extant gharials in having enlarged supratemporal diversify during the Palaeocene and Eocene (e.g. fenestrae, an elongate, tubular snout, and a high Buffetaut, 1976, 1978, 1982; Jouve, 2005, 2007; Jouve et al., 2005a, 2006a; Jouve, Bouya & Amaghzaz, 2005b, 2008; Barbosa et al., 2008; Hill et al., 2008; Hastings et al., 2010; Hastings, Bloch & Jaramillo, *Corresponding author. E-mail: [email protected] 2011, in press). However, the origins of this clade are

854 © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 854–871 CRETACEOUS TETHYSUCHIAN FROM UK 855 poorly understood owing to a paucity of fossils from This has important implications for tethysuchian the ‘middle’ Cretaceous. evolution, in particular for our understanding A phylogenetic definition of Tethysuchia was of dyrosaurid origins, and whether dyrosaurids recently proposed by Andrade et al. (2011: S102) as: and the marine ‘pholidosaurids’ Terminonaris and ‘the clade composed of Pholidosaurus purbeckensis Oceanosuchus constitute a single marine radiation or (Mansel-Pleydell, 1888) and Dyrosaurus phospha- several independent ones. ticus (Thomas, 1893), their common ancestor and all One of the major issues hampering resolution in its descendants’. This definition encompasses these analyses is the paucity of ‘middle’ Cretaceous Dyrosauridae and Pholidosauridae, and possibly (Barremian–Turonian) marine tethysuchians, thus Elosuchidae. Although curiously, this is not the affecting our understanding of character polarity. In case in the phylogenetic analysis of Andrade those phylogenetic analyses in which ‘pholidosaurids’ et al. (2011), as Elosuchidae was recovered are paraphyletic, all potential dyrosaurid sister taxa outside the Pholidosaurus + Dyrosaurus clade. (Elosuchus, Oceanosuchus, and Terminonaris) are Moreover, Andrade et al. (2011) proposed that from this time span (see Mook, 1933, 1934; Wu et al., Elosuchidae be used for the clade consisting of 2001; de Lapparent de Broin, 2002; Hua et al., 2007). Elosuchus, Sarcosuchus, and Vectisuchus. Also, a Furthermore, the earliest potential dyrosaurids are phylogenetic definition of Pholidosauridae was Cenomanian in age (Buffetaut & Lauverjat, 1978; recently proposed by Fortier, Perea & Schultz (2011: Buffetaut, Bussert & Brinkmann, 1990). Therefore, S259) as: ‘a stem-based group name including investigating specimens from the Barremian– Pholidosaurus schaumburgensis (Meyer, 1841) and all Turonian stages will be key to elucidating the early taxa closer to it than to D. phosphaticus (Thomas, evolution of Tethysuchia. 1893) or Pelagosaurus typus Bronn, 1841′. No explicit Here we describe a long-known, but previously phylogenetic definition has been proposed for unstudied, tethysuchian crocodyliform. This speci- Dyrosauridae or Elosuchidae. men, the anterior-most part of a right dentary Phylogenetic analyses consistently find (NHMUK PV OR36173), is most probably from the Dyrosauridae to be holophyletic (e.g. Wu et al., 2001; Upper Greensand Formation of England (upper Jouve, 2005; Jouve et al., 2005a, 2006a, 2008; Albian, Lower Cretaceous). Although this specimen Barbosa et al., 2008; Young & Andrade, 2009; was discovered over 150 years ago, it has only been Hastings et al., 2010, 2011, in press; Andrade et al., briefly mentioned once in the literature. Furthermore, 2011; Fortier et al., 2011). The holophyly of it is of importance as a result of its unusual morphol- Pholidosauridae, however, is not always recovered. ogy and geological age. The presence of a marine Pholidosauridae has either been found to be a tethysuchian in the late Lower Cretaceous of the UK paraphyletic grade of taxa closely related to would indicate that tethysuchians moved into the Dyrosauridae (Wu et al., 2001; Jouve et al., 2005a, marine realm earlier than previously realized. 2008; Barbosa et al., 2008; Young & Andrade, 2009; Hastings et al., 2010, 2011, in press; Andrade et al., 2011), holophyletic (Fortier et al., 2011; and in one of ABBREVIATIONS the analyses of the Jouve et al., 2006a data set in INSTITUTIONAL Hastings et al., in press), or holophyletic with MIWG, the Museum of Isle of Wight Geology (now Elosuchus being outside the clade comprising IWCMS – the Isle of Wight County Museum Service, Dyrosauridae and Pholidosauridae (Jouve et al., incorporating Dinosaur Isle museum and visitor 2006a; and in one of the analyses of the Jouve et al., attraction); MNHN, Muséum national d’histoire 2006a data set in Hastings et al., in press). Martin & naturelle, Paris, France; NHMUK, Natural History Buffetaut (2012) and Martin et al. (2014b) also found Museum, London, UK. Pholidosauridae to be holophyletic, but as no dyrosaurids were included in those analyses they never tested whether pholidosaurids constitute a ANATOMICAL natural group. When Pholidosauridae is found to be D1, first dentary alveolus; D2, second dentary alveo- paraphyletic, the sister taxon of Dyrosauridae varies lus; D3, third dentary alveolus; D4, fourth dentary between Terminonaris (Wu et al., 2001; Jouve et al., alveolus; for, foramen; rug, rugose patch; sym, 2005a, 2008; Barbosa et al., 2008; Hastings et al., in symphysis. press), Oceanosuchus (Young & Andrade, 2009), and Elosuchus (Hastings et al., 2010, 2011), although HISTORICAL INFORMATION Oceanosuchus was only included in the analysis of Young & Andrade (2009). As such, the internal evo- The anterior right dentary (NHMUK PV OR36173) lutionary relationships of Tethysuchia are in flux. was purchased by the British Museum (Natural

History) in October 1861 from a Mr Simmons. In the tion, and not from one of the Lower Greensand Group NHMUK specimen register, NHMUK PV OR36173 is formations exposed at Shanklin, is important (see listed as an ‘anterior portion of left upper jaw with Fig. 1). In order to determine this, one of the authors four teeth sockets, of a crocodilian reptile. Greensand? (T.P.) examined the fossils in the MIWG (IWCMS) and Shanklin, I. of Wight’. The only mention of the speci- the rocks at Shanklin. Descriptions of the deposits of men we can ﬁnd in the literature is by Lydekker the Lower Greensand and Selborne Groups on the (1889: 179), who identiﬁed NHMUK PV OR36173 as Isle of Wight have been given by White (1921), Wach the pliosaurid, Polyptychodon interruptus, and states & Ruffell (1991), Insole, Daley & Gale (1998), and that it is: ‘apparently the extremity of the left Hopson, Wilkinson & Woods (2008). premaxilla’. Why he referred NHMUK PV OR36173 Numerous marine reptiles are known from the to Pliosauridae, and thought it might be P. Ferruginous Sands Formation (Lower Greensand interruptus, was not stated. Amongst the specimen Group) of the Isle of Wight. The members of the labels is a note left by Dr Leslie Noé, dated 23 August Ferruginous Sands Formation exposed at the lower 1999, in which he states: ‘This jaw fragment is not part of the cliffs at Shanklin (Knock Cliff to Hope pliosaurian [? crocodilian]’. This taxonomic note is Beach) are: the Old Walpen Chine Member (XII), the what brought NHMUK PV OR36173 to our attention. New Walpen Chine Member (XIII), and Member XIV (un-named). If the provenance of NHMUK PV OR36173 is one of these members, we would expect it GEOLOGICAL INFORMATION to be very dark brown or black in colour with a dark The right partial dentary (NHMUK PV OR36173) matrix (this being caused by the iron-rich nature of was discovered at Shanklin, Isle of Wight, UK. The the sediment), as is the case in the exceptionally rare exact formation that yielded it is, however, unclear. It ornithopod bones and a number of ichthyosaur verte- was referred to the Upper Greensand Formation by brae (e.g. MIWG.5376) from Member VI, which are Lydekker (1889), but the NHMUK specimen register dark brown/black. However, we cannot discount that refers to it as from: ‘Greensand?’. Therefore, verifying NHMUK PV OR36173 may have come from an iso- that it is actually from the Upper Greensand Forma- lated calcareous lens in one of these members. As

Figure 1. Map of the Isle of Wight showing Shanklin (A), with the geological column of formations exposed there (B). [A and B are modiﬁed from Hopson (2011).] Photographs of the formations exposed near Shanklin, at Luccombe Chine (C), and Knock cliff (D).

© 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 854–871 CRETACEOUS TETHYSUCHIAN FROM UK 857 such, the possibility of a Ferruginous Sands Forma- Formation or from an isolated calcareous lens in tion origin cannot currently be completely ruled out. Member XII, Member XIII, or Member XIV of the Overlying the Ferruginous Sands Formation at Ferruginous Sands Formation. Shanklin is the Sandrock Formation (Fig. 1). This Is NHMUK PV OR36173 reworked from an earlier, formation is a mixture of blue silty clay and yellow Jurassic, horizon? Not only is the morphology of sands, and is considered to have been deposited NHMUK PV OR36173 distinct from any known during a time when cyclical sea-level changes pre- Jurassic taxon (see the description below), but a served a number of wide estuaries and near-shore drifted block of Jurassic material would needed to mud flats (see Wach & Ruffel, 1991). Fossil plants have travelled many kilometres from the nearest discovered from this formation are typically pale exposed Jurassic outcrop to have become entombed in (white), fragmentary, and crumble easily. Although we the marine Greensands. We find this to be an unlikely think it unlikely that the Sandrock Formation is the scenario. origin of NHMUK PV OR36173, the lack of any suitable reptilian material for comparison means that we cannot entirely discount the possibility. SYSTEMATIC PALAEONTOLOGY The final, and highest, formation from the Lower CROCODYLIFORMES BENTON &CLARK, 1988 Greensand Group exposed is the Monks Bay Sand- MESOEUCROCODYLIA WHETSTONE &WHYBROW, stone Formation (formerly known as the Carstone 1983 Formation of the Isle of Wight) (Fig. 1). This overlies NEOSUCHIA BENTON &CLARK, 1988 the Sandrock Formation. It is very gritty, with large iron-rich sandstone nodules and has a dark-brown TETHYSUCHIA BUFFETAUT, 1982 coloration. Vertebrate fossils are unreported in this TETHYSUCHIA INCERTAE SEDIS formation, and along with the iron-rich deposits Specimen (which would make fossils darker in colour), we NHMUK PV OR36173, the anterior region of the exclude this as the possible origin of NHMUK PV right dentary. OR36173. The Lower Greensand Group is overlain by the Locality Selborne Group, with two formations exposed (Fig. 1): Shanklin, Isle of Wight, UK. the Gault Formation (Albian) and the Upper Greensand Formation (Albian–Cenomanian). The Selborne Group was deposited in a marine setting, Horizon and age with the Gault Formation forming in a mid- or outer- Most likely from the ‘Malm rock’ or Freestone, Upper shelf environment, whilst the Upper Greensand For- Greensand Formation, Selborne Group (upper Albian, mation was formed in a shallow offshore shelf and in Lower Cretaceous). However, it could be from an lower shoreface zones related to an eastward- isolated calcareous lens from one of three members prograding shoreline (Hopson, 2011). within the Ferruginous Sands Formation, Lower At Shanklin, the clays/mudstones of the Gault For- Greensand Group (upper Aptian, Lower Cretaceous): mation are dark in colour, which results in dark the Old Walpen Chine Member (XII), the New Walpen fossils. The general coloration and matrix of NHMUK Chine Member (XIII), or Member XIV (un-named). PV OR36173 appears similar to the central part of the Moreover, we cannot entirely discount the Sandrock Upper Greensand Formation, the ‘Malm Rock’ or Formation, Lower Greensand Group (upper Aptian– Freestone, a pale grey/green glauconitic sandstone/ lower Albian, Lower Cretaceous), as there are no siltstone that contains iron-rich sandy nodules (divi- known suitable fossils from this formation for sion D of Jukes-Browne & Hill, 1900). This thick comparison. sequence lies above the basal ‘Passage Beds’ (division A of Jukes-Browne & Hill, 1900) and below the Chert DESCRIPTION Beds (division E of Jukes-Browne & Hill, 1900). However, it must be appreciated that a considerable DENTARY amount of time (over 150 years) has passed since Only the anterior region of the right dentary is pre- NHMUK PV OR36173 was collected. Therefore, the served. It is approximately 130 mm in anteroposterior external surface may have altered as it dried out and length. Overall, the preservation is good, other than it oxidized. Based on general specimen coloration and being broken posteriorly. There is no evidence of post- matrix, the Upper Greensand Formation seems like mortem mediolateral compression or shearing. The the most probable origin of NHMUK PV OR36173. As external (= lateral and ventral) surfaces of the stated above, we cannot preclude the possibility that dentary are gently convex, with numerous large, NHMUK PV OR36173 originated from the Sandrock subcircular foramina that are mostly widely spaced

Figure 2. Tethysuchia incertae sedis, NHMUK PV OR36173. Anterior region of the right dentary in lateral view: A, photograph; B, line drawing.

Figure 3. Tethysuchia incertae sedis, NHMUK PV OR36173. Anterior region of the right dentary in ventral view: A, photograph; B, line drawing.

Figure 4. Tethysuchia incertae sedis, NHMUK PV OR36173. Anterior region of the right dentary in dorsal view: A, photograph; B, line drawing.

(Figs 2, 3). The external surface ornamentation com- anteroposterior length of 48 mm and a mediolateral prises numerous, very small, subcircular pits. This width of approximately 38 mm. Between the D1 and ornamentation pattern remains constant along the D2 alveoli, there is a minimum interalveolar space of element. The medial contractions between alveoli 17 mm. The D2 alveolus is notably smaller and also (and the raised alveolar rims, especially the D3 and oval in shape, having an anteroposterior length of D4) suggest that the premaxillary teeth would have 25 mm. Between the D2 and D3 alveoli there is a occluded lateral to the interalveolar spaces on the minimum interalveolar space of 14 mm. The D3 dentary (i.e. in an overbite or interlocking manner) alveolus is smaller than the D2 alveolus and is (Figs 2–4). slightly more circular in shape, with an On the dorsal surface of the dentary, the three anteroposterior length of 21 mm. Between the D3 and anterior-most alveoli are completely preserved, along D4 alveoli there is a minimum interalveolar space of with the anterior section of the fourth (Fig. 4). All of 9 mm. The D4 alveolus is incomplete, but it would the preserved alveoli are very large in proportion to have been considerably larger than the D2 and D3 the overall size of the bone, but vary in size and alveoli. The D4 alveolus has a transverse width of shape. There is also variation in interalveolar approximately 35 mm. Each alveolus has a slight, or spacing. The ﬁrst dentary alveolus (D1) is orientated noticeable, raised rim. In the D1 and D3 alveoli they dorsally and slightly anteriorly (Figs 2, 4–5). The D1 are barely noticeable, in the D2 alveolus the rim is alveolus is oval in shape, orientated along the more strongly developed, whilst the D4 alveolius has anteroposterior axis of the dentary, and has an a very large and well-developed rim.

Figure 5. Tethysuchia incertae sedis, NHMUK PV OR36173. Anterior region of the right dentary in anterior view; A, photograph; B, line drawing.

Figure 6. Tethysuchia incertae sedis, NHMUK PV OR36173. Anterior region of the right dentary in medial view: A, photograph; B, line drawing.

The medial border of the D2 and D3 alveoli are extended posterior to the fourth alveolus. Based on the either in the same sagittal plane, or lateral to, the preserved region, the dentary appears to be from a lateral border of the D1 alveolus (Fig. 4). Overall, the mesorostrine or longirostrine taxon (based on the long, dorsal surface of the dentary is mainly flat, with narrow and dorsoventrally shallow dentary). It is numerous large foramina medial to the D2 and D3 similar to other tethysuchians such as ‘Elosuchus’ alveoli. The largest foramen is medial to the D3 felixi (de Lapparent de Broin, 2002) and Dyrosauridae alveolus. (e.g. Hill et al., 2008; Hastings et al., 2010). There is In lateral view (Fig. 2), the dorsal margin of the no trace of the splenial, but we would not expect there dentary is concave between the D1 and D4 alveoli. to be because the splenial does not reach as far This results in the D2 and D3 alveoli being ventral in anteriorly as the fourth alveolus in other tethysuchian the transverse plane to the D1 and D4 alveoli. Moreo- taxa (e.g. de Lapparent de Broin, 2002; Hill et al., ver, the D2 and D3 alveoli are orientated slightly 2008; Hastings et al., 2010, 2011; Adams et al., 2011). dorsolaterally. When seen in lateral view, the ventral In posterior view, the bone is exposed in cross margin is distinctly curved, and is convex. It rises section (Fig. 7). As noted by Lydekker (1889: 179) dorsally anteriorly, and is noticeable ventral to the there are large cancelli in the diploë. The dentary is first two dentary alveoli. highly cancellous, with thicker cortical bone along the In medial view, the preservation is poor but the ventral and lateroventral margins of the bone, symphyseal suture is partially visible (Fig. 6). whereas the dorsal and dorsolateral margins have However, the dentary appears to be ‘hollow’ very thin cortical bone. Adjacent to the cortical bone, posteroventral to the D3 and D4 alveoli. Anteriorly, the cancelli immediately internal to the thickest cor- the dentary is a solid bone, which would have created tical bone are larger than those adjacent to thinner a firm mandibular symphysis. Ventral to the D3 there cortical bone. is a concave surface that forms the boundary from the ‘hollow’ and solid regions of the dentary. Posteriorly, bivalve encrustations and bivalve fossils are pre- DENTITION served within this ‘hollow’ region. Only two, incomplete, in-situ tooth crowns are pre- We cannot determine how much of the symphyseal served (Fig. 4). The in-situ tooth crowns are in the surface is preserved, although it is likely that it first and third alveoli. In the first alveolus there is an

Figure 7. Tethysuchia incertae sedis, NHMUK PV OR36173. Anterior region of the right dentary in posterior view: A, photograph; B, line drawing. erupting replacement tooth crown. The crown is only Russell, 2008), and Polycotylidae (Carpenter, 1996; partially visible, with only the apical half of the Albright, Gillette & Titus, 2007b). lingual surface exposed. In the third alveolus there is 2. Eosauropterygians have a distinctive tooth- a cross section through the base of a fully erupted replacement pattern. Their replacement teeth tooth crown. The crown has a diameter of 11–12 mm. originate in separate temporary alveoli, recogniz- Based on the two in-situ tooth crowns, the dentition able by aligned foramina on the dorsal surface of would have been caniniform in morphology (i.e. single the dentaries. Developing teeth grow within these, cusped and labiolingually compressed). The basal until the temporary and functional alveoli fuse section is oval in cross section, being wider (Owen, 1840; von Huene, 1923; Rieppel, 2001; mesiodistally than labiolingually. The incomplete Shang, 2007). The erupting teeth grow in a preservation of the crowns makes it impossible to shallow groove medial to each functional alveolus. make any further comments on the carinal or overall The separation between primary and secondary crown morphologies. alveoli can be short or large. However, tooth replacement in thecodont groups (including Crocodylomorpha) involves development of the DISCUSSION replacement teeth in shallow pits in the lingual EXCLUDING NHMUK PV OR36173 side of the functional alveolus, and then migration FROM PLESIOSAURIA into the primary tooth pulp cavity through resorp- tion pits in the old base (Edmund, 1960, 1969; We agree with Dr Leslie Noé that the referral of Kieser et al., 1993). Consequently, and in contrast NHMUK PV OR36173 to Pliosauridae and to eosauropterygians, the entire process remains P. interruptus is incorrect. There are a number of hidden inside the dentigerous bone. An exception plesiosaurian characteristics that are lacking in to this pattern is in polycotylids, in which the NHMUK PV OR36173, allowing us to exclude, with secondary alveoli are placed below the functional confidence, NHMUK PV OR36173 from Sauropterygia. teeth and so are hidden within the bone These include the following. (O’Gorman & Gasparini, 2013). NHMUK PV 1. The pattern of large-to-small teeth in NHMUK PV OR36173 is clearly thecodont, with a replacement OR36173 (in which the first and fourth dentary tooth erupting within the first dentary alveolus alveoli are considerably larger than the second and (Fig. 4). third alveoli; Fig. 4) is unknown in Plesiosauria. To 3. No plesiosaurians have a symphyseal region of the our knowledge there is no known plesiosaurian in dentary that is as large and continuously flat as in which the first dentary alveous (D1) is larger than NHMUK PV OR36173 (Fig. 4). In plesiosaurians, the second and third alveoli. In fact, in all the alveoli occupy the largest part of the dorsal plesiosaurian dentaries, the D1 is smaller (or at surface. There is usually a groove medial to the least subequal) than the immediately posterior alveoli, although sometimes the alveoli and groove alveoli. This is seen in Rhomaleosauridae (Smith, are separated by paradental plates (e.g. Benson 2007), Pliosauridae (Andrews, 1913; Tarlo, 1960; et al., 2011; Ketchum & Benson, 2011). In Albright, Gillette & Titus, 2007a; Benson et al., Pliosauridae, replacement alveoli develop within 2011; Ketchum & Benson, 2011; Sassoon, Noé & the symphyseal groove. Only medial to the groove Benton, 2012), Leptocleididae (Druckenmiller & is there a flat dorsal surface. The lack of a

symphyseal dorsal groove in NHMUK PV Leiokarinosuchus, and Hylaeochampsa lack the ante- OR36173, and its large flat dorsal surface, does not rior dentary, and are thus not comparable with support referral to Pliosauridae. NHMUK PV OR36173. Based on the size and shape of the supratemporal fenestrae of L. brookensis,itwas likely to have been a longirostrine form (Salisbury & NHMUK PV OR36173 WITHIN CROCODYLOMORPHA Naish, 2011). The most commonly discovered marine crocodylo- Although no dentaries are known for H. vectiana, morph clade from the Mesozoic is Thalattosuchia. the anterior dentary is known for two Albian However, the Aptian–Albian age of NHMUK PV hylaeochampsid species: Pachycheilosuchus trinquei OR36173 makes a thalattosuchian identification from Texas, USA, and Pietraroiasuchus ormezzanoi unlikely. Teleosaurids are currently only known from the southern Apennines, Italy. Both species are from the Jurassic, as the only definitive Lower Cre- interpreted as living in shallow, near-shore, brackish taceous specimen (from the Valanginian of France) environments (Rogers, 2003; Buscalioni et al., 2011). was recently redescribed and reidentified as a However, their dentaries do not resemble the metriorhynchid (Young et al., 2014). Moreover, the Shanklin specimen (NHMUK PV OR36173). In anterior dentary morphologies of teleosaurids do not hylaeochampsids, the anterior tip of the dentary is match NHMUK PV OR36173 (Fig. 8). In teleosaurids laterally convex in dorsal view, resulting in a broad the anterior region is as follows: spatulate, with the anterior dentary (D1–D6 region), with each succes- maximal width being present at the level of the D3 sive alveolus being lateral to the preceding one alveolus; the D3 and D4 alveoli are closely set; there (Fig. 8O). Also, their symphyses are short (terminat- are no enlarged foramina medial to the alveoli; and ing level to the fourth or sixth dentary alveolus) and the alveolar margin is convex at the D3–D4 region, the dentaries lack concavities (i.e. festooning) along resulting in those alveoli being positioned dorsal to the alveolar margins. Furthermore, the D1 and D4 the D1 and D2 alveoli (e.g. Fig. 10C; Andrews, 1913; alveoli are not larger than the D2 or D3 alveoli. Hua, 1999; Lepage et al., 2008; Martin & Vincent, Rogers (2003: 132) described P. trinquei as having: 2013). Accordingly, we can safely disregard a ‘Rough pitting and grooves sculpture the ventral teleosaurid origin for this specimen. surface and a row of nutrient foramina parallels the However, at least four lineages of the other labial margin’; this differs from the low-relief orna- thalattosuchian clade, Metriorhynchidae, survived mentation pattern and the large foramina that are into the Early Cretaceous (Young et al., 2014), and arranged across the lateral and ventral surfaces of there is an indeterminate specimen from the the dentaries in NHMUK PV OR36173. As such, we Barremian of Spain (Parrilla-Bel et al., 2012). We can can confidently exclude NHMUK PV OR36173 from exclude NHMUK PV OR36173 from pertaining to Hylaeochampsidae. any of these lineages (Cricosaurus, Dakosaurus, The only longirostrine clade of eusuchians known Geosaurus, and Plesiosuchina) as they all have the from the Cretaceous are Gavialoidea. Gavialoids are following dentary characteristics: the D1 and D4 known from the Maastrichtian (Late Cretaceous) to alveoli are not enlarged relative to the other anterior the present day, with numerous ‘thoracosaurine’ alveoli; festooning along the alveolar margin is either species known from Maastrichtian–Paleocene marine absent or only subtle; no enlarged foramina are deposits (e.g. Brochu, 2004, 2006; Hua & Jouve, 2004; present medial to the alveoli; and the dentary alveoli Delfino, Piras & Smith, 2005; Jouve et al., 2006b). lack raised rims (e.g. Fig. 8B; Fraas, 1902; Gasparini The Shanklin specimen, however, does not resemble & Dellapé, 1976; Pol & Gasparini, 2009; Young & basal gavialoids. The anterior region of the dentary Andrade, 2009; Young et al., 2012; Herrera, Gasparini of the Maastrichtian species Eothoracosaurus & Fernández, 2013). Therefore, there is no reason to mississippiensis has circular D1 alveoli; the D2 assume a thalattosuchian origin for NHMUK PV alveoli are larger than the D1, D3, and D4 alveoli; a OR36173. diastema is present between the D2 and D3 alveoli; From non-marine Barremian–Aptian deposits of the the D3 and D4 alveoli are closely set, although they Isle of Wight, numerous crocodylomorph clades are do not form a ‘couplet’; and large foramina on the known. There is a new bernissartiid (genus and species dorsal surface of the dentary adjacent to the alveoli currently in press), the atoposaurid Theriosuchus sp., are absent (Fig. 8P; Brochu, 2004). The anterior the goniopholidid Anteophthalmosuschus hooleyi, the region of the dentary of the Palaeocene species, ?goniopholidid/elosuchid/pholidosaurid Vectisuchus Eosuchus lerichi, has: circular D1 alveoli; D1–D4 leptognathus, the enigmatic neosuchian Leiokarino- alveoli of similar diameters, with the D3 alveoli being suchus brookensis, and the basal eusuchian slightly smaller; a diastema between the D2 and D3 Hylaeochampsa vectiana (see Salisbury & Naish, alveoli; and large, widely separated foramina on the 2011; Sweetman et al., in press). Vectisuchus, dorsal surface of the dentary, immediately medial to

Figure 8. The anterior ends of right dentaries of select crocodylomorphs compared with NHMUK PV OR36173 (some are reversed relative to the original figures); all are shown in isolation from the left dentary. A, NHMUK PV OR36173; B, metriorhynchid Plesiosuchus manselii (after Young et al., 2012); C, teleosaurid Machimosaurus hugii (after Martin & Vincent, 2013); D, ‘pholidosaurid’ Sarcosuchus imperator (after Buffetaut & Taquet, 1977); E, dyrosaurid Phosphatosaurus gavialoides (after Hill et al., 2008); F, dyrosaurid Hyposaurus ‘morphotype 1’ (after Jouve, 2007); G, dyrosaurid Hyposaurus ‘morphotype 2’ (after Jouve, 2007); H, dyrosaurid Rhabdognathus sp. (after Jouve, 2007); I, ‘pholidosaurid’ Terminonaris robusta (after Mook, 1934); J–K, elosuchid Elosuchus cherifiensis (after de Lapparent de Broin, 2002); L, ‘pholidosaurid’ ‘Sunosuchus’ thailandicus (after Buffetaut & Ingavat, 1980); M, ‘Goniopholis’ phuwiangensis (after Buffetaut & Ingavat, 1983); N, goniopholidid Goniopholis baryglyphaeus (after Schwarz, 2002); O, hylaeochampsid Pietraroiasuchus ormezzanoi (after Buscalioni et al., 2011); P, gavialoid Eothoracosaurus mississippiensis (after Brochu, 2004). The reconstructions are not to scale. the alveoli (Delfino et al., 2005). The anterior region of phologies seen in basal gavialoids differ from the dentary of the Paleocene species, Argochampsa NHMUK PV OR36173, combined with the long krebsi, is poorly preserved. However, the alveolar inferred ghost range, we cannot refer the specimen to diameters are similar, with the D2 alveoli being that clade. slightly smaller than the D1, D3, and D4 alveoli The Shanklin specimen (NHMUK PV OR36173) (Jouve et al., 2006b). As the anterior dentary mor- does not resemble any goniopholidid. In

Figure 9. Goniopholis sp., NHMUK PV R1807. Anterior region of the dentary in dorsal view: A, photograph; B, line drawing.

Figure 10. Theriosuchus pusillus, NHMUK PV OR48262, referred specimen. Anterior region of the left dentary in dorsal view: A, photograph; B, line drawing. goniopholidids, the anterior tip of the dentary is erupted tooth crowns, owing to the characteristic laterally convex in dorsal view, resulting in it shift in alveolar shape/morphology (Brinkmann, being wider than the region posterior to the D4 alveo- 1992; Schwarz & Salisbury, 2005; NHMUK PV lus (Fig. 8M, N; Salisbury et al., 1999; Salisbury, OR48262). NHMUK OV OR36173 lacks this shift in 2002; Schwarz, 2002). The D1 alveolus is alveolar shape from subcircular (pseudocaniniform mediolaterally wider than it is anteroposteriorly long; morphotype) to oval (labiolingually compressed teeth) the D3 and D4 alveoli are both larger than the D1 and alveoli. The D1 alveoli border the symphysis at its D2 alveoli; and the ventral and lateral surfaces of the most anterior point; the gap between the D1 and D2 anterior part of the dentary are covered with small, alveoli is approximately equal to that between the D2 closely spaced pits (Salisbury et al., 1999; Salisbury, and D3 alveoli; and the D3 to D7 alveoli are conflu- 2002; Schwarz, 2002). Additionally, the anterior-most ent. The lateral margin of the anterior snout is part of the dentary is often out-turned in heavily ornamented, and has numerous heterogene- goniopholidids (e.g. Goniopholis sp. (Fig. 9) and ously spaced foramina, all approximately 0.5–1 mm Goniopholis tenuidens; Owen, 1879, Pl. 1, fig. 1; in diameter, as is also the case in the other western Salisbury 2002 as cf. Goniopholis sp.). In view of European species Theriosuchus guimarotae (Schwarz these marked differences, there is no reason to regard & Salisbury, 2005) and Theriosuchus ibericus NHMUK PV OR36173 as a member of the (Brinkmann, 1992). Theriosuchus differs from Goniopholididae. NHMUK PV OR36173 in the relative sizes of the The Shanklin specimen can additionally be first four alveoli, with the D1–3 alveoli all being excluded from the semi-aquatic basal neosuchian equal in size and smaller than the D4 alveoli, and group, Atoposauridae (Gervais, 1871), which per- the aforementioned heterogeneous spacing. Addition- sisted from the Middle Jurassic to the latest Creta- ally, the symphyseal surface has a slight dorsal cur- ceous (Evans & Milner, 1994; Martin, Rabi & Csiki, vature anteriorly, unlike the linear surface of 2010; Martin et al., 2014b). Multiple individuals of NHMUK PV OR36173. Based on these marked dif- Theriosuchus pusillus (Owen, 1879) are known from ferences and the overall size differences, we can the Lower Cretaceous of the UK. The mandibular exclude NHMUK PV OR36173 from being regarded rostrum of one well-preserved specimen (NHMUK as a large marine atoposaurid. PV OR48262) is 10 mm long and has seven The anterior end of the dentary is well preserved in alveoli (Fig. 10). Theriosuchus has a heterodont den- several taxa conventionally or putatively considered tition, which is visible even without preserved or as members of the Pholidosauridae. In P.

© 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 854–871 864 M. T. YOUNG ET AL. schaumburgensis from the Berriasian of Germany, lateral view, which in both ‘E.’ felixi and NHMUK PV the anterior region is spatulate and widest at the OR36173 is dorsal to the D2 and D3 alveoli. Note that level of the D2 alveoli (Koken, 1887). There is a de Lapparent de Broin (2002: fig. 2N) incorrectly sizeable gap between the D1 and D2 alveoli and the showed in a figure the D4 alveolus as being ventral to D2 and D3 alveoli, whereas the D3 and D4 alveoli are the D2 and D3 alveoli, and orientated somewhat closely set (Koken, 1887). Moreover, the D3 and D4 posteriorly. This is probably a result of the post- alveoli are only slightly larger than the other dentary mortem deformation of the holotype (MNHN.F INA alveoli, all of which are similar in size (Koken, 1887). 25); however, the two other lower jaws are from much In the large-bodied ‘pholidosaurids’, Sarcosuchus larger individuals (MNHN.F INA 21 and MNHN.F hartii, Sarcosuchus imperator, and ‘Sunosuchus’ INA 22) and they share the same D4 position as thailandicus, the anterior dentaries are spatulate NHMUK PV OR36173. There are, however, some with the maximal width at the level of the D4 alveoli noticeable differences between ‘E.’ felixi and NHMUK (Fig. 8D; Buffetaut & Taquet, 1977; Buffetaut & PV OR36173: (1) ‘E.’ felixi lacks the elongate Ingavat, 1984; Martin et al., 2014a). In these taxa, anteroposterior axis of the D1 alveolus that is found the D1 and D2 alveoli are small, and substantially in NHMUK PV OR36173; (2) the anterior dentary is smaller than the D3 alveoli. In ‘S.’ thailandicus the gladius-shaped in ‘E.’ felixi – the anterior dentary of D4 alveoli are greatly enlarged relative to the D1–D3 the holotype has a slight concavity along the lateral alveoli (Buffetaut & Taquet, 1977; Martin et al., margin in dorsal/ventral views, but in the larger 2014a), whereas in S. hartii and S. imperator the D3 specimen, MNHN.F INA 21, this concavity is much and D4 alveoli are both enlarged (Buffetaut & Taquet, more pronounced, and in both specimens the anterior 1977). dentary is widest at the level of the D4 alveoli; and (3) In the marine ‘pholidosaurids’ Terminonaris browni the gladius-shaped anterior dentary results in the and Terminonaris robusta, the anterior region of the bone tapering anterior to the D4 alveoli, whereas in dentary is slightly spatulate in shape, its maximal NHMUK PV OR36173 the D2–D3 region is almost width being at the level of the D3 alveoli (Fig. 8I; straight and is only slightly lateral to the D1. Mook, 1933, 1934). The D1 and D2 alveoli are notably There are several anterior dentary characteristics smaller than the D3 and D4 alveoli and the rest of the which suggest a close relationship between NHMUK alveoli adjacent to the symphysis. The anterior region PV OR36173 and Dyrosauridae: of the dentary is unknown in the ‘pholidosaurids’ Oceanosuchus boecensis (Hua et al., 2007; Lepage 1. The D1 alveolus is enlarged relative to the D2 and et al., 2008) and V. leptognathus (Buffetaut & Hutt, D3 alveoli (Fig. 8). This is also seen in: 1980; Salisbury & Naish, 2011). However, the Arambourgisuchus khouribgaensis (Jouve et al., spatulate premaxilla of O. boecensis suggests that 2005a), Dyrosaurus maghribensis (Jouve et al., the anterior region of the dentary was also spatu- 2006a), Hyposaurus ‘morphotypes 1 and 2’ late, being similar to Terminonaris and other and Rhabdognathus sp. (Jouve, 2007), and ‘pholidosaurids’. Phosphatosaurus gavialoides (Hill et al., 2008). The elosuchid/‘pholidosaurid’ Elosuchus cherifiensis 2. The D1 alveolus is mainly dorsally orientated, but also has a spatulate anterior dentary region, the there is also a slightly/moderate anterior orienta- maximal width being at the level of the D2 and D3 tion. This is also seen in: Cerrejonisuchus alveoli (Fig. 8J, K; de Lapparent de Broin, 2002). The improcerus (Hastings et al., 2010), D. maghribensis D1 alveoli are enlarged compared with the D2–D4 (Jouve et al., 2006a), D. phosphaticus (Jouve, 2005), alveoli, with the D3 alveoli being the smallest of the Hyposaurus ‘morphotypes 1 and 2’, and anterior alveoli (de Lapparent de Broin, 2002). The Rhabdognathus sp. (Jouve, 2007). This morphology line drawings of de Lapparent de Broin (2002) make is also seen in basal gavialoids such as A. krebsi the shape of the D1 alveoli between the different (Jouve et al., 2006b) and E. mississippiensis E. cherifiensis specimens somewhat hard to judge, but (Brochu, 2004). they appear to be subcircular to suboval in shape. 3. Numerous, large foramina on the lateral and The holotype of ‘E.’ felixi and the two lower jaws ventral surfaces of the dentary that are widely referred to this species, share some similarities with spaced. This is also seen in: A. khouribgaensis NHMUK PV OR36173. The ‘E.’ felixi specimens and (Jouve et al., 2005a) and D. phosphaticus (Jouve, NHMUK PV OR36173 share: (1) a D1 alveolus that is 2005). larger than the D2 and D3 alveoli; and (2) noticeably 4. A concave dorsal margin of the dentary between enlarged D4 alveoli relative to the other alveoli (de the D1 and D4 alveoli, resulting in the D2 and D3 Lapparent de Broin, 2002; MNHN.F INA 21, alveoli being slightly dorsolaterally orientated (i.e. MNHN.F INA 22, MNHN.F INA 25). Another shared a festooned anterior dentary). This is also seen in: characteristic is the position of the D4 alveoli in D. phosphaticus (Jouve, 2005), Hyposaurus

Figure 11. Stratigraphic ranges of relevant Lower Cretaceous crocodylomorph lineages, plotted against time and compared with the possible age range of the Shanklin specimen (NHMUK PV OR36173). Dotted lines represent parts of lineages for which fossils are unreported; arrows show that lineage persisted beyond the end of the Turonian.

‘morphotypes 1 and 2’, and Rhabdognathus sp. tive collection of extant reptiles), and are possibly (Jouve, 2007). related to an interlocking dentition and vertically 5. Large foramina on the dorsal surface of the festooned tooth row at the anterior-most region of the dentary, medial to the D2 and D3 alveoli. This is rostrum. also seen in: P. gavialoides (Hill et al., 2008). In summation, there were numerous clades of 6. The external ornamentation on the dentary crocodylomorphs living during the ‘middle’ Cretaceous is of low relief and not conspicuous. This is also (Fig. 11). However, NHMUK PV OR36173 differs seen in: A. khouribgaensis (Jouve et al., 2005a), considerably from the basal eusuchian clade Hyposaurus ‘morphotypes 1 and 2’, and Hylaeochampsidae and the neosuchian clades Rhabdognathus sp. (Jouve, 2007). Goniopholididae and Atoposauridae in alveolar con- 7. The anterior end of the dentary is not spatulate (as figuration and dentary shape, and thus cannot be it is in the ‘pholidosaurids’ and teleosaurids dis- considered a member of these clades (Figs 8–10). cussed above) or gladius-shaped (as in ‘E.’ felixi), Additionally, the anterior dentary morphology of and the dentary is not laterally convex in basal gavialoids and metriorhynchid thalattosuchians dorsal view (which results in a distinctly wide differs from NHMUK PV OR36173 (Fig. 8). Further- anterior region, such as in goniopholidids and more, NHMUK PV OR36173 also lacks the hylaeochampsids, discussed above). NHMUK PV mediolaterally expanded anterior dentary seen in OR36173 has a narrow anterior dentary, like that ‘pholidosaurids’ and teleosaurids, and also differs in of longirostrine dyrosaurids (e.g. Jouve, 2007; Hill alveolar configuration (Fig. 8). The alveolar configu- et al., 2008) and basal ‘thoracosaurine’ gavialoids ration and dentary shape of NHMUK PV OR36173 (e.g. Brochu, 2004; Delfino et al., 2005; Jouve et al., are similar to those of Dyrosauridae (Fig. 8), and 2006b) (see Fig. 8). numerous other characteristics (listed above) suggest a close relationship. There is one anterior dentary Note that characteristics 1–5 are also seen in characteristic shared by dyrosaurids that NHMUK Crocodylus spp. (NHMUK Earth Sciences compara- PV OR36173 lacks: a diastema/gap between the D2

© 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 854–871 866 M. T. YOUNG ET AL. and D3 alveoli. In view of this combination of dentary (Koken, 1887), O. boecensis (Hua et al., 2007; Lepage characters, we conservatively identify NHMUK PV et al., 2008), and T. robusta (Mook, 1934), and the OR36173 as Tethysuchia incertae sedis. More com- dyrosaurid C. lateroculi (Jouve et al., 2005b; Hill plete material and inclusion within a comprehensive et al., 2008). Therefore, in some aspects, cranial vari- phylogenetic analysis would enable testing of whether ation between ‘pholidosaurids’ and basal dyrosaurids or not NHMUK PV OR36173 is the sister taxon to is less marked than originally thought. Dyrosauridae, or if it is a ‘pholidosaurid’-grade However, one region of the skeleton is profoundly marine taxon. different between the marine ‘pholidosaurids’ and dyrosaurids: the dorsal osteoderms. Like S. hartii, the marine taxa T. robusta and O. boecensis: (1) have ‘MIDDLE’CRETACEOUS MARINE TETHYSUCHIANS subrectangular paravertebral dorsal osteoderms that The best known ‘middle’ Cretaceous marine are wider than they are anteroposteriorly long; (2) tethysuchians are T. browni and T. robusta from retain the stylofoveal joint (has a anterolateral North America (middle Cenomanian–middle process); (3) have a longitudinal keel; and (4) do not Turonian) and O. boecensis (lower Cenomanian) from have accessory osteoderms (i.e. a biserial series, e.g. France (Mook, 1933, 1934; Wu et al., 2001; Hua et al., Buffetaut & Taquet, 1977; Wu et al., 2001; Hua et al., 2007; Lepage et al., 2008; Adams et al., 2011). From 2007). Dyrosaurids, however, lack the stylofoveal joint the upper Cenomanian of Bavaria, Germany, a large, and longitudinal keels, they have accessory incomplete upper jaw from a longirostrine taxon has osteoderms lateral to the paravertebral series, and been referred to Terminonaris cf. browni (Buffetaut & have a lateral process which creates a joint between Wellnhofer, 1980). Terminonaris and Oceanosuchus the paravertebral osteoderm and the accessory differ from dyrosaurids in having: (1) more strongly osteoderm (see Schwarz, Frey & Martin, 2006). As ornamented skull and lower jaw bones; (2) a such, while cranial variation between ‘pholidosaurids’ premaxilla without an anterodorsally projecting and basal dyrosaurids is less marked than is initially process anterior to the external nares; (3) a dorsally presumed, their difference in dorsal osteoderm mor- orientated external nares; (4) five premaxillary alveoli phologies are significant. These differences have the instead of four; (5) a subvertical anterior margin of potential to be very useful in identifying dyrosaurids the premaxilla that extends ventrally relative to the solely, or primarily, based on osteoderms. rest of the element (i.e. the ‘pholidosaurid beak’); (6) Although dyrosaurids are well known from the absence of the D2–D3 diastema/gap seen in Maastrichtian onwards, their pre-Maastrichtian fossil dyrosaurids; and (7) absence also of the small D7 record is relatively poor. Two maxillary fragments alveoli seen in dyrosaurids. The anterior region of the from the Campanian of Egypt pertain to a dentary is not preserved in Oceanosuchus, but the longirostrine crocodyliform tentatively referred to anterior region of the dentary of Terminonaris is Dyrosaurus (Churcher & Russell, 1992; Churcher, slightly spatulate (Mook, 1933, 1934; Wu et al., 2001; 1995). The earliest potential dyrosaurids are Adams et al., 2011). As such, these taxa still had Cenomanian in age. A partial dentary (mid- classic ‘pholidosaurid’ characteristics (Hua et al., symphyseal region) from the middle Cenomanian 2007; Fortier et al., 2011). marine deposits of Portugal was noted to be reminis- It should be noted that some putative pholidosaurid cent of dyrosaurids (Buffetaut & Lauverjat, 1978). (sensu Fortier et al., 2011) and elosuchid (sensu However, its incomplete nature means that no firm Andrade et al., 2011) apomorphies are also found in conclusions can be drawn as to its affinities. Skull the basal-most known dyrosaurids Chenanisuchus fragments (frontal, parietal, and left postorbital) and lateroculi (Jouve et al., 2005b) and a new genus and incomplete vertebrae from Cenomanian lacustrine species (Hastings et al., in press). Fortier et al. (2011) deposits of Sudan were referred to Dyrosauridae stated that the medial margin of the orbit (in dorsal indet. by Buffetaut et al. (1990). The dorsal vertebrae view) is mostly formed by the prefrontal, with the have very large hypapophyses, as is the case in frontal participating only slightly in the medial dyrosaurids (i.e. blade-like, flattened laterally, margin, and is a pholidosaurid apomorphy. However, rounded ventrally, and anteroposteriorly long) (e.g. this also occurs in C. lateroculi (Jouve et al., 2005b), see the figures in Storrs, 1986; Jouve & Schwarz, gen. et sp. nov. (Hastings et al., in press), and 2004; Jouve et al., 2005a; Schwarz et al., 2006). goniopholidids (see the figures in Andrade & Terminonaris robusta also has noticeable Hornung, 2011). Andrade et al. (2011) stated that the hypapophyses on some vertebrae, but they are much frontal being concave, with the medial borders of the lower and are a different shape (see fig. 4 in Wu et al., orbit being upturned, is an elosuchid apomorphy. 2001). The skull fragments were considered to be However, this also occurs in the pholidosaurids P. from a primitive dyrosaurid by Buffetaut et al. (1990), purbeckensis (Salisbury, 2002), P. schaumburgensis as they have: (1) more strongly ornamented cranial

© 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 854–871 CRETACEOUS TETHYSUCHIAN FROM UK 867 bones; (2) a smaller anterolateral process on the left dyrosaurid or an especially close relative of that postorbital than that of other dyrosaurids; (3) a clade, especially as some putative apomorphies for broader intertemporal bar than dyrosaurids (i.e. ‘Pholidosauridae’/Elosuchidae are also found in the between the supratemporal fenestrae); and (4) a basal-most known dyrosaurids. Only future studies of dorsal region of the postorbital pillar that is only new discoveries, and/or unrecognized museum speci- slightly medially inclined. Unfortunately, the mens from this key time span, will elucidate the anterolateral process of the left postorbital is broken, tethysuchian radiation into the marine realm and the so its size cannot be properly judged. However, large origin of Dyrosauridae. anterolateral processes of the postorbitals are also seen in T. browni (Mook, 1933), T. robusta (Wu et al., ACKNOWLEDGEMENTS 2001), and O. boecensis (Hua et al., 2007; Lepage et al., 2008). These examples further highlight how We would like to thank Phil Hurst (Image Resources, certain marine ‘pholidosaurid’ and dyrosaurid cranial NHMUK) for photography of NHMUK PV OR36173, morphologies overlap and may be homologous. As and Ronan Allain (MNHN) for collection access. We such, only the vertebrae (which Buffetaut et al., 1990 also thank Alexander Hastings (Halle, Germany) and could not be certain were associated with the skull Stéphane Jouve (Marseille, France) for their com- fragments) reliably suggest a dyrosaurid, not indeter- ments on NHMUK PV OR36173, and Leslie Noè minate tethysuchian, origin for this material. Note (Bogotá, Colombia) for making his taxonomic note that the presence of the Sudanese material in non- which highlighted this specimen. We also thank an marine deposits does not imply that during the anonymous reviewer and Steve Salisbury (University Cenomanian some putative dyrosaurids were of Queensland, Australia) whose comments improved restricted to freshwater environments, as there is the quality of this paper. MTY received support from growing evidence that dyrosaurids successfully the SYNTHESYS Project http://www.synthesys.info/ exploited both freshwater and saltwater environ- which is financed by European Community Research ments (see Khosla et al., 2009 and the references Infrastructure Action under the FP7 ‘Capacities’ therein). Program. REFERENCES CONCLUSIONS Adams TL, Polcyn MJ, Mateus O, Winkler DA, Jacobs Herein we describe a tethysuchian crocodyliform from LL. 2011. 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