Crocodilian-bearing localities and horizons

Transcription

1 24. Crocodilians by Steven W. SALISBURY and DARREN NAISH Having burned my fingers so consumedly with the Wealden, I am fearful for you... Darwin writing to Lyell on 25 November 1860 (Burkhardt and Smith ) Crocodilians are a ubiquitous component of the vertebrate fauna of the Wealden Supergroup of southern England. In the past 160 years, 11 species belonging to as many as nine genera have been recognized. Many of these taxa are represented by well-preserved specimens, often comprising complete skulls and articulated or associated postcranial skeletons. In addition, numerous isolated elements, typically teeth, fragmentary crania and postcranial bones have been collected, and formally or informally referred to various taxa. As is common for reptiles from the Wealden Supergroup, our understanding of these fossils has sometimes become clouded by erroneous referrals and unsubstantiated claims of synonymy. Herein we attempt to alleviate these issues through a detailed review of all the key specimens. In the process we have recognized a number of new taxa, proposed reassignments of others, emended existing diagnoses and flagged several specimens for detailed descriptions and further taxonomic and phylogenetic appraisal. Anatomical abbreviations. alv, alveolus; ang, angular; bo, basioccipital; bs, basisphenoid; can qsqex, quadratosquamosoexoccipital canal; car dermost lat, lateral osteodermal keel; car dermost med, medial osteodermal keel; ch sec, secondary choanae; cond occ, occipital condyle; d, dentary; dermost gast, gastral osteoderm; dermost nuch, nuchal osteoderm; dermost paravert, paravertbral osteoderm; ect, ectopterygoid; ex, exoccipital; fen itp, infratemporal fenestra; f, frontal; fac art ext, external articular surface; fac ext, external surface; fac int, internal surface; fen suborb, suborbital fenestra; for carot post, posterior aperture of the carotid foramen; for eus med, median eustachian foramen; for mag, foramen magnum; for ptp, posttemporal foramen; for sqex; squamosoexoccipital foramen; for stp, supratemporal foramen; fos max, maxillary fossa; h, humerus; j, jugal; l, lachrimal; ls, laterosphenoid; m. (in text), musculus (muscle); max, maxilla; n, nasal; nar, naris; orb, orbit; pal, palatine; palp, palpebral; par, parietal; pars lat, lateral part; pars med, medial part; pf, prefrontal; prezyg, prezygapophysis; pmax, premaxilla; po, postorbital; proc art, articular process; proc caud, posterior process; proc cran, anterior process; proc parocc, paraoccipital process; proc spin, spinal process; proc trans, transverse process; pty, pterygoid; q, quadrate; qj, quadratojugal; rad, radius; rec ot ext, external

2 306 Crocodilians otic recess; sa, surangular; seg latcost, lateral rib segment; seg vertcost, vertebral rib segment; so, supraoccipital; sp, splenial; sq, squamosal; vert cerv, cervical vertebra; vert thor, thoracic rib. Crocodilian-bearing localities and horizons Crocodilian fossils are now known from most of the major units of the Wealden Supergroup. Within the Hastings Group, they are best known from the Grinstead Clay Formation north of Cuckfield, West Sussex (TQ , the original Tilgate Forest localities near Whiteman s Green, and TQ ; Mantell, 1822, 1827; Cuvier 1824; Owen , 1842, 1878; Hulke 1878), and the Ashdown and Wadhurst Clay formations at Hastings, East Sussex (between TQ and TQ ; Owen 1851, 1878; Woodward 1885, 1886; Seeley 1887a; Lydekker 1887a, 1888a; Buffetaut 1975; Buffetaut and Ford 1979; Clark and Norell 1992). Other localities within the Hastings Group that have produced crocodilians include Burwash, East Sussex (TQ ; Ashdown Formation), Brede, East Sussex (TQ ; several sites with exposures of the Brede Bone Bed in the Wadhurst Clay Formation, including Hare Farm Lane, TQ ; Topley 1875; Allen 1949; Benton and Spencer 1995b), and Telham, near Battle, East Sussex (TQ , several sites with exposures of the Telham Bone Bed in the Wadhurst Clay Formation, including Black Horse Quarry, TQ ; Binfield and Binfield 1854; Topley 1875; Woodward and Sherbon 1890; White 1928; Allen 1949; Lake and Shephard-Thorn 1987; Benton and Spencer 1995b). The Weald Clay Group, in contrast, has produced fewer and less substantial remains (mainly osteoderms and isolated teeth). Lower Weald Clay Formation crocodilian-bearing localities include quarries at Keymer Tileworks, Burgess Hill, West Sussex (TQ ; Young and Lake 1988; Cook 1995; Cook and Ross 1996), Longbrook Brickworks, Thakeham, West Sussex (TQ ), Horsham Park, Horsham, West Sussex (the Horsham Stone ; TQ ), Brightling, East Sussex (TQ ), Crowhurst Pit, Rackwell Wood, East Sussex (TQ ; Sweeting 1925; White 1928), and Peasmarsh, Waterfall Wood, East Sussex (TQ ; Allen 1949; Benton and Spencer 1995b). The most productive crocodilian-bearing locality in the Upper Weald Clay Formation is the Smokejacks Brickworks pit, Ockley, Surrey (TQ ; Rivett 1956; Ross and Cook 1995). Within the Wealden Group on the Isle of Wight, important crocodilian fossils have been recovered from the Wessex Formation and the Shepherd s Chine Member of the Vectis Formation, between Compton Grange Chine (SZ ) and Atherfield Point (SZ ) on the south-western coast of the island (Huxley and Etheridge 1865; Owen 1874a; Huxley 1875; Hulke 1878; Lydekker 1887a, 1888a, 1890a; Seeley 1887a; Hooley 1907; White 1921; Buffetaut and Ford 1979; Buffetaut and Hutt 1980; Clark and Norell 1992).

3 Crocodilians THE 'GAVIAL' OF TILGATE FOREST: THE FIRST WEALDEN CROCODILIAN 307 The history of Wealden crocodilians begins around , when Gideon Mantell collected isolated teeth, vertebrae and osteoderms from two quarries near Whiteman's Green, West Sussex, an area he referred to as 'Tilgate Forest', as noted above (see Dean 1995). The fossils from this area derived from a 'Tilgate Stone', one of several bands of calcareous sandstone in the Grinstead Clay Formation, extensively quarried for building and road stone at the time (Gallois 1965). Mantell described his initial collection of Tilgate Forest crocodilian fossils in 1822 (pp ). He identified three types of crocodilian teeth based on size, robustness, curvature and degree of surface striation. 'Variety a' included large, nearly cylindrical teeth with a blunt point; those of 'variety b' were much more delicate, rather compressed and somewhat curved with a sharp apex; while 'variety c' types were more slender and nearly straight. On the advice of William Clift, the then curator of the Hunterian Museum of the Royal College of Surgeons, Mantell (1822) attributed 'variety b' to a 'crocodile or a monitor', and noted the strong similarity of 'variety c' to the Indian gharial, Gaviaiis gal1geticlls Gmelin. Among the postcranial elements, he distinguished thoracic, lumbar and caudal vertebrae, and based on their biconcave nature (i.e. with a slightly concave articular surface at both ends of the vertebral body; the amphicoelous condition) considered them to be most similar to crocodilian fossils from Le Havre and Honfleur in Normandy, north-western France, then in the possession of Georges Cuvier (this material is now regarded as belonging to a thalattosuchian; Buffetaut 2008). Mantell (1822, p. 50) considered the isolated osteoderms from Tilgate Forest to 'perfectly resemble' those of the extant Alligator mississippiensis, even with their irregular rhomboidal outline and sharply pointed lateral process. Mantel! later had some of these crocodilian fossils sent to Georges Cuvier in Paris, who figured them for the first time in 1824 and corroborated Mantell's initial identifications and comparisons (Cuvier 1824, pp ; Buffetaut 2010; Text-fig. 24.1). Of the teeth, Cuvier regarded those of Mantel!'s 'variety a' (Cuvier 1824, pi. 10, figs 25-27; numbers unknown; Text-fig. 24.1A-C) to be similar to some large, obtuse teeth from the Jurassic of Switzerland (now thought to belong to the teleosaurid Machimosallrlls; see Krebs 1967; Buffetaut 2010). With regard to the vertebrae, Cuvier simply confirmed Mantel!'s original identifications (Cuvier 1824, pi. 10, figs 31-34; Text-fig. 24.1E-H) and their resemblance to those of the crocodilians from Ca en and Honfleur (now referred to thalattosuchians; Buffetaut 2008). Mantel! provided further information on his Wealden crocodilian fossils, and illustrated many of them for the first time in 1827, including additional isolated postcranial elements (e.g. NHMUK 3805, a partial paravertebral osteoderm;

4 308 Crocodilians Mantell 1827, pl. 6, fig. 8; Text-fig. 24.1I). His insights into the affinities of the teeth were much the same as they were in 1822, and he seemed pleased that Cuvier had confirmed his earlier conjectures on their identification (Mantell 1827, p. 64). Contrary to his 1822 work, Mantell (1827) only recognized two types of crocodilian teeth in his Wealden collection: the large, obtuse variety a (e.g. NHMUK 2512; Mantell 1827, pl. 5, fig. 10; also pl. 5, figs 1 2, 7, 9, 12; see Lydekker 1888a; Text-fig. 24.1D), and the more slender, gently curved variety b (Mantell 1827, pl. 5, figs 5 6, 8; Buffetaut 2010). His previous variety c seems to have been grouped with variety b. Collectively, he referred to both varieties as the Gavial of the Tilgate Forest (Mantell 1827, p. 64). Further publications by Mantell (1833a, 1839) did not cover his Wealden crocodilian material as thoroughly as in his 1827 book and little in the way of new material from Cuckfield seems to have been added to his collection. In 1837, he was forced to sell his entire collection to the British Museum (see Lyddeker 1888a for a full listing of the specimens in the Mantell Collection of the NHM resulting from this and subsequent sales). It was not until 1841 that fossils of the Wealden gavial were assigned to distinct taxa. In his Odontography ( ), Richard Owen used the name Crocodilus (Suchosaurus) cultridens for Mantell s variety b teeth (vol. 1, p. 287 and vol. 2, p. 16, caption for pl. 62A, fig. 10), and Goniopholis crassidens for Mantell s variety a (vol. 1, p. 289 and vol. 2, p. 16, caption for pl. 62A, fig. 9). According to Woodward and Sherbon (1890, p. xxix), the eight volumes of Odontography were published in 3 parts: part 1, pp , pls 1 50 was published in 1840; part 2, pp. i xl, , pls 51 89A (excluding 62A) in 1841; and part 3, pp. xli lxxiv, pp , pls 1, ii, 62A, in So, although 1841 was the first time that both Crocodilus (Suchosaurus) cultridens and Goniopholis crassidens were used in print, Owen presumably intended parts of Odontography to build on information and descriptions provided in his Report on British Fossil Reptiles, part II (published in April 1842), which he presented on August 2, 1841 at the Plymouth meeting of the British Association for the Advancement of Science. It is in this report that he (Owen 1842) provided what should be regarded as the first official taxonomic assignments of each of the Wealden crocodilians. Ironically, the Wealden Suchosaurus cultridens teeth described by Mantell (1822, 1827, 1833a, 1839), Cuvier (1824) and Owen ( , 1842) have recently been shown to belong to indeterminate baryonychine theropods (see Buffetaut 2007, 2010: Chapter 29). The second tooth type, Mantell s variety a and one that Owen ( , 1842) assigned to G. crassidens, along with isolated vertebrae, osteoderms and other postcranial elements from Tilgate Forest (see Lydekker 1888a), are still regarded as pertaining to crocodilians (but see section below on issues associated with the referral of these teeth to G. crassidens).

5 English Wealden Fossils 309 Text-fig Goniopholididae gen. indet. from the Grinstead Clay Formation near Cuckfield, West Sussex; the gavial of the Tilgate Forest, as figured by Cuvier (1824) and Mantell (1827). A C (numbers unknown). D, isolated teeth, described by Mantell (1827) as variety a ; NHMUK E H, isolated vertebrae. I, a partial paravertebral osteoderm in external aspect; NHMUK A C and E H from Cuvier (1824); D and I from Mantell (1827). Scale bar represents 50 mm. Systematic Palaeontology The following is an account of all the crocodilian taxa that have been recognized in Wealden Supergroup and their revised taxonomic status. Although we have focused primarily on specimens that have been described, figured and assigned to various taxa in the literature, we have also included comments on other specimens that we consider significant. CROCODYLOMORPHA Walker, 1968 CROCODYLIFORMES Hay, 1930 MESOEUCROCODYLIA Whetstone and Whybrow, 1983 NEOSUCHIA Benton and Clark, 1988 GONIOPHOLIDIDAE Cope, 1875 GONIOPHOLIS Owen, 1842 Type species. Goniopholis crassidens Owen, Lectotype. NHMUK 3798, slab and counter-slab preserving the disarticulated postcranial skeleton and most of the left mandibular ramus from a single individual,

6 310 Crocodilians which in life would have been approximately 3 m long. The association of two smaller slabs with the main slab and counterslab cannot be confirmed (see Salisbury 2002, text-figs 3B, 4B). As such, these smaller slabs should not be included in the lectotype until a proper association can be demonstrated. Remarks. Goniopholis is a well-known Mesozoic mesoeucrocodylian, represented by numerous remains attributed to as many as 19 species. These are mainly from Late Jurassic and Early Cretaceous deposits in western Europe, from localities in England, Scotland, France, Germany, Portugal, Belgium, Spain and Denmark (Owen 1842, , 1878, 1879a; Hulke 1878; Dollo 1883; Koken 1883, 1886, 1887, 1896; Hooley 1907; Edinger 1938; Jonet 1981; Huckriede 1982; von Oekentorp 1984; Buffetaut 1986; Norman 1987a; Buscalioni and Sanz 1987a, b; Buffetaut et al. 1989; Cuny et al. 1991; Ortega et al. 1996; Salisbury et al. 1999; Salisbury 2002; Schwarz 2002; Schwarz-Wings et al. 2009). The genus has also been recognized in North America and South-East Asia, but the possibility that the American species are generically distinct has been mentioned on occasion (Clark 1986; Salisbury et al. 1999; Lauprasert et al. 2007; Allen 2007, 2010). Recent revisions of European Goniopholis taxa have shown that most species are either synonymous with G. simus Owen, 1878, or are of uncertain taxonomic validity (Salisbury et al. 1999; Salisbury 2002; De Andrade and Hornung 2011). Salisbury et al. (1999) recognized G. crassidens, G. simus and an as yet undescribed taxon from the Barremian of Bernissart, a small village in the province of Hainaut, Belgium, referred to G. simus by Dollo (1883). Salisbury (2002) raised the possibility that G. simus is a junior synonym of G. crassidens, but comparisons between the two taxa have been limited by the nature of the type specimens. New material from the Berriasian of northwestern Germany reported by Hornung et al. (2009) and Hornung and Reich (2011) supports this idea, but the synonymy is yet to be formalized. Salisbury (2002) also regarded Nannosuchus gracilidens Owen, 1879a from the Purbeck Limestone Group as a valid species of Goniopholis, while Schwarz (2002) named G. baryglyphaeus from Portugal. De Andrade et al. (2008) and De Andrade and Hornung (2011) reported a potentially new species of Goniopholis from the Intermarine Bed (DB 128/9 of Clements 1993) of the Durlston Formation in the Purbeck Limestone Group that is closely allied to G. simus. Owen ( , 1842; see earlier comments for the intended timing of these publications) included several specimens in his initial description and diagnosis of G. crassidens. His account commences with the description of isolated teeth from the Grinstead Clay Formation north of Cuckfield, West Sussex, first described by Mantell (1822): NHMUK 2512 (Mantell 1827, pl. 5, fig. 10; Text-fig. 24.1D); numbers unknown (Mantell 1827, pl. 5, figs 1 2, 7, 9, 12; see Lydekker 1888a, pp for a full listing of the crocodilian teeth in the Mantell Collection at the NHM). He then provided a detailed account of the Swanage specimen (NHMUK 3798), expanding the diagnosis to include postcranial features (Owen 1842, pp ).

7 English Wealden Fossils 311 Owen (1842, p. 70) claimed that two teeth preserved in the mandibular ramus of the Swanage specimen were identical with the obtuse teeth of the Wealden. Owen s (1842) description of the Swanage specimen provides useful characters for the diagnosis of G. crassidens relating to the dermal skeleton, vertebrae and ilium. Considered in combination, the characters that Owen identified in his 1842 description are sufficient to diagnose G. crassidens. However, we agree with Hooley (1907, p. 53) and Salisbury et al. (1999) that dental characters are insufficient to define species of Goniopholis and question the claim of Owen (1878, p. 2) and Koken (1887) that such characters can be used to distinguish species of Goniopholis. Owen (1842) was never explicit in designating a type specimen, such that all the specimens he described should be considered syntypes. The subsequent referral of other specimens to G. crassidens by Hulke (1878; see Goniopholis willetti sp. nov. below) and Hooley (1907; see Anteophthalmosuchus hooleyi gen. et sp. nov. below) led to the inclusion of additional cranial characters in the diagnosis for this species (e.g. Lydekker 1888a; Steel 1973). This has led to confusion surrounding the diagnosis of G. crassidens and, in turn, Goniopholis. Here, we formalize the suggestion of Salisbury et al. (1999) and propose that NHMUK 3798 (from the Purbeck Limestone Group) be designated the lectotype of G. crassidens. Until additional material shows otherwise, the referral of isolated teeth and other cranial elements from Cuckfield and other Wealden and Purbeck localities to G. crassidens is, in our opinion, not possible, and such specimens should be identified only as Goniopholididae gen. indet. The specimens described by Hulke (1878) and Hooley represent distinct taxa (see below), and any characters associated with them should not be linked to G. crassidens. Pending the description of new material from north-western Germany, the formal synonomy of G. crassidens and G. simus would allow cranial remains assignable to the latter to be referred to G. crassidens. Lydekker (1888a, 1890a), Nopcsa (1928a), Buffetaut (1982), Ortega et al. (1996), Salisbury et al. (1999), Schwarz (2002), Tykoshki et al. (2002), Lauprasert et al. (2007, 2009), Hornung et al. (2009) and De Andrade and Hornung (2011) have proposed additional postcranial and cranial characters that are diagnostic of Goniopholis in Europe. Most of the British Goniopholis specimens (including the lectotype of G. crassidens Owen, 1842) are from the Purbeck Limestone Group (see Salisbury 2002), but a number of additional remains are known from the Wealden succession. These specimens are discussed below. Goniopholis willetti sp. nov. Text-figures Derivation of name. Named in honour of Edgar W. Willett, collector of the holotype. Holotype. BMB , a nearly complete skull (approximately 55 cm total length; estimated total body length 3.5 m), missing only portions of the left infratemporal

8 312 Crocodilians Text-fig Goniopholis willetti sp. nov., BMB , holotype skull in A B, dorsal aspect (photograph and schematic interpretation) and C D, occipital aspect (photograph and schematic interpretation). Scale bar represents 100 mm. and preorbital region (jugal, quadratojugal, ectopterygoid, posterior part of the maxilla and the articular surface of the quadrate), part of the left lacrimal, the posterior part of the right maxilla, the right ectopterygoid, and the lateral parts of the pterygoids. Type locality and horizon. Cuckfield, West Sussex (presumably near TQ , but the exact location is unknown), in the Cuckfield Stone of the Grinstead Clay Formation.

9 Crocodilia Diagnosis. Goniopholis willetti sp. novo is distinguished from other species of GOlliopholis based on the following unique combination of characters (autapomorphies, marked with an 'a'): proportionately long, narrow rostrum (maximum width of maxillary rostrum at the fifth maxillary alveoli 25% of maximum rostrallength); five premaxillary (1 </=5<4<3<2) and at least 24 maxillary alveoli; partially fused parietals (shared with G. simlls); prefrontal divides the posteriormost extent of the nasal, such that the rostral portion of the prefrontal is separated from the rostralmost extent of the frontals (a); interorbital ridge restricted to the frontals and does not extend onto the prefrontal (a); contact between the prefrontal and postorbital within the orbit, ventral to the dorsal surface of the cranial table (shared with G. simlls); shallow, triangular groove extends rostrally from the rostrodorsal margin of the orbit onto the jugal and lacrimal (shared with G. simlls, AnteophthalmosuchllS hooleyi and many mesoeucrocodylians); posterolateral part of squamosal on the cranial table forms a distinct, rounded boss (a); well-developed rostral tuberosity on the postorbital bar (shared with many neosuchians and basal eusuchians). Remarks. The specimen was collected, and presumably prepared, by Edgar W. Willett, who showed it to the Geological Society of London around Along with the rest of his collection, it was subsequently accessioned to the BMB, where it has remained ever since (Cooper 2010), contrary to claims that it could not be located or was 'lost' (e.g. Salisbury et al. 1999; Schwarz 2002). John W. Hulke presented a paper on BMB to the Geological Society in February His relatively detailed description and a pencil drawing of the skull in dorsal aspect were published later the same year. Hulke (1878) joined Willett in considering BMB to represent the first known skull of G. crassidens. Hulke's (1878) rationale for referring BMB to G. crassidens seems to stem from Owen's original inclusion of some of Mantell's crocodilian teeth from Tilgate Forest (the obtuse ones) and isolated postcranial elements in his 1842 description (see previous section on the 'Gavial of Tilgate Forest'), along with Willett's belief that it probably belonged to this species. As Hulke (1878, p. 377) correctly pointed out, however, prior to the discovery of this specimen, the skull of G. crassidens was 'quite unknown'. With the possible exception of a partial cranial table, the direct association of which to NHMUK 3798 cannot be confirmed (see Salisbury 2002 and comments below on Anteophthalmosllclllls hooleyi), there is no anatomical overlap between NHMUK 3798 and BMB Although teeth preserved on each specimen (two mandibular teeth on NHMUK 3798 and several partial maxillary and premaxillary teeth on BMB ) are similar and conform to the typical Goniopholis-type, this is not sufficient grounds upon which to refer BMB to G. crassidens. BMB shows a number of characteristics also seen in G. sil11!lsas defined by Salisbury et al. (1999). These include relatively uniform pitting (comprising roughly circular pits) on the cranial table, jugal and lateral half of the quadratojugal, with the remainder of the maxillary rostrum covered in indistinct scarring and smaller,

10 314 Crocodilians shallower pits, which are almost absent from the premaxillae; a transversely elongated squamosoexoccipital foramen; partially fused parietals; and contact between the prefrontal and postorbital within the orbit, ventral to the dorsal surface of the cranial table. The secondary choanae of BMB and G. simus are also of a similar size, shape and position, mid-way between the palatines and pterygoid (see Buffetaut 1982, fig. 7A for the morphology of the secondary choanae in G. simus). BMB also shares partially fused frontals and broad inclusion of the frontoparietal suture within the supratemporal foramen with G. simus and G. gracilidens. Although the rostralmost extent of the maxillae on the palate of BMB is incompletely preserved, the likely M-shaped nature of the premaxillomaxillary suture suggests that a premaxillary fenestra was absent, as is the case in G. simus (Salisbury et al. 1999). Text-fig Goniopholis willetti sp. nov., BMB , holotype skull in ventral aspect. A, photograph. B, schematic interpretation. Scale bar represents 100 mm.

11 Crocodilians 315 TEXT-FIG GOlliopholis willetti sp. nov., BMB , holotype; posterior half of the skull in right lateral aspect. A, photograph. B, schematic interpretation. Scale bar represents 100 mm. In common with G. simus and the Portuguese species G. baryglyphaells, BMB has a laterally exposed quadratosquamosoexocciptial (QSE) canal (the 'cranioquadrate' canal of other authors), continuing posteriorly on the dorsal surface of the quadrate as a shallow sulcus, resulting from a lack of contact between the paraoccipital process, the ventrolateral lamina of the squamosal and the dorsal surface of the quadrate. A similar condition is also seen in Hylaeochampsa vectiana Owen, 1874a (Clark and NoreIl1992; Salisbury et al. 1999; Buscalioni et al. 2001; Delfino et al. 2008) and the Romanian form of Allodaposuchus precedens Nopcsa, 1928a (Delfino et al. 2008). As occurs in all valid species of Goniopholis, the nasals ofbmb are excluded from the naris by the mediolaterally broad and anteroposteriorly short premaxillae, which have a concave posterolateral margin rostral to the suture with the maxillae (see below for the condition in Anteophthalmosuchus hooleyi and 'G. simlls' from Bernissart). It also shares with other goniopholidids a distinct bean-shaped fossa on the posterolateral margin of the maxilla. The maxillary fossa ofbmb is approximately three times as long as it is high in lateral aspect, with a dorsally inflected ventral margin (Text-figs 24.2, 24.4). At least one subspherical concavity is preserved at the rostral end of the fossa; others may have been present posteriorly to this, but this part of the fossa is poorly preserved on both sides. In common with G. gracilidens and most specimens of G. simlls, particularly those from Germany (see Salisbury et al. 1999; De Andrade and Hornung 2011), BMB has a well-developed palpebral that forms the medial dorsal border of the functional orbit.

12 316 English Wealden Fossils Notable (but not autapomorphic) features of BMB include a much more elongated, proportionately narrower maxillary rostrum than other species of Goniopholis (maximum width at the fifth maxillary alveoli 25% of maximum rostral length, compared with c. 40% in G. simlls), somewhat reminiscent of the rostral morphology of PholidosallTllS purbeckensis Salisbury, 2002 and Vectisuchus leptogllatillls Buffetaut and Hutt, There are five premaxillary and at least 24 maxillary alveoli. The sizing of the premaxillary alveoli (1 </=5<4<3<2) differs from that of G. simus (e.g., 1 </=5<2<3=4 for GAB Sch. 3, skull number 3 in the Ballerstedt Collection of the Gymnasium Adolfum, Biickeburg, Germany; Salisbury et al. 1999). The prefrontal of BMB divides the posteriormost extent of the nasal, such that the rostral portion of the prefrontal is separated from the rostralmost extent of the frontals. The interorbital ridge ofbmb is restricted to frontals and does not extend onto the prefrontal, unlike the more extensive crest in G. simus, G. gracilidells and G. baryglyphaeus (an interorbital crest is absent in Al1teophthalmosuchus hooleyi and 'G. simus' material from Bernissart; see below). A shallow, triangularshaped groove extends rostrally from the rostrodorsal margin of the orbit onto the jugal and lacrimal, similar to that on G. simlls and Anteophthalmosllchus hooleyi (see below). The posterolateral part of squamosal on the cranial table of BMB forms a distinct, rounded boss. Finally, there is a well-developed rostral tuberosity on the postorbital bar, similar to, but not as enlarged as, that on Allteophthalmosuchus hooleyi (NHMUK R3876; see below), 'G. simlls' material from Bernissart (IRSNB R1537; see below) and V.leptognathus (SMNS 50984; see below). On the strength of this unique combination of osteological characters, along with other characteristics of the skull discussed above, we propose that BMB represents a distinct species of GOl1iopholis, namely G. willetti. It seems likely that many of the other likely goniopholidid specimens from the Grinstead Clay Formation described and figured by Mantell (1822,1827) and Owen ( ,1842, ,1878), who referred them to G. crassidel1s, also pertain to G. willeti, based on their size and provenance (see Lydekker 1888a, pp for a full list of specimens). The same is also true for most of the Wealden 'G. crassidel1s' specimens in the G. Holmes Collection of the BMB. However, nearly all of these specimens are either associated or isolated postcranial elements, and none is coupled with cranial remains, such that comparisons with BMB are not possible. Of the cranial elements, most are partial mandibles (e.g. NHMUK 37972, a near complete right mandibular ramus containing several broken teeth; NHMUK R974, the rostral portion of a left mandibular ramus of a small individual, showing 13 alveoli with at least two complete teeth in situ). Unfortunately, we have not been able to locate these specimens. However, given that neither is listed as showing anatomical overlap with the holotype of G. willetti, it is unlikely that they can be compared with, and assigned to, the latter taxon. The one exception might be a portion of rostrum figured by Owen ( , vol. 2, pi. 12, figs 1-4). This specimen

13 English Wealden Fossils 317 Text-fig Goniopholis sp. indet., a pair of premaxillae from the Grinstead Clay Formation at Cuckfield, West Sussex; BMB Scale bar represents 100 mm. includes occluding portions of both the maxillary and mandibular rostrum. The mandibular rostrum is proportionately narrow and the mandibular symphysis appears to extend well posterior to the eight or ninth dentary alveolus, suggesting a long, narrow rostrum consistent with that of G. willetti (in more generalized, broad-snouted Goniopholis spp. such as G. simus, the mandibular symphysis extends posteriorly to a point level with the sixth dentary alveolus). Owen ( , vol. 2, pl. 12, fig. 3) only illustrated a small portion of the ventral surface of the maxillary rostrum, limiting direct comparisons with the holotype of G. willetti. Unfortunately, as with many of the other G. crassidens specimens from the Wealden of Cuckfield, we have not been able to locate this specimen. It is hoped that if it is found, comparisons will demonstrate that it can be referred to G. willetti. As stated previously, we do not think it is possible to assign isolated teeth to specific species of Gonipholis; hence, the numerous isolated Gonipholis-type teeth from the Grinstead Clay Formation of Cuckfield and other sites in Sussex cannot be assigned to G. willetti and should be placed in Goniopholididae gen. indet. At least one specimen indicates that more than one species of Goniopholis might be present in the Grinstead Clay Formation. Owen ( , vol 2, pl. 11, figs 1 2; 1878, p. 4, pl. 1, figs 1 2) described and figured a pair of partial premaxillae from the Wealden of Cuckfield as G. crassidens (BMB ; Text-fig. 24.5). Similar to G. simus and possibly G. willetti and IWCMS (see following section on Anteophthalmosuchus hooleyi), this specimen shows no evidence of a premaxillary fenestra. Within the nasal vestibulum there is a small, median, diamondshaped fossa, very similar to that considered diagnostic for G. simus by Salisbury et al. (1999). The sizing of the premaxillary alveoli on this specimen (1<4<2=3) is distinct from both BMB (1</=5<4<3<2) and G. simus (1</=5<2<3=4 for GAB Sch. 3). Unlike G. simus, the dorsal rim of the naris on BMB is not built up such that it is confluent with the rest of the premaxillae, as occurs in BMB However, this part of BMB has been partially reconstructed with putty, and could easily have been worn down prior to fossilization. The most obvious difference between BMB , BMB and other species of Goniopholis

14 318 English Wealden Fossils is the length of the premaxillae and premaxillary suture rostral to the naris in dorsal aspect. In BMB , this suture is approximately equal to the rostroposterior length of the naris, whereas in BMB and G. Sil1111S, it is considerably shorter. Based on these observations we consider BMB possibly to represent a species of Goniopholis distinct from G. willetti and G. simus. However, based on the fragmentary nature of the specimen, we place it in GOl1iopholis sp. indet. ANTEOPHTHALMOSUCHUS gen. novo Derivation of name. Greek ante, forward, ophthall11ol1, eye, and SOUcilOS from cjouxo<;, crocodile, with reference to the forward-facing orbits of the type specimen. Type species. Anteophthalmosllchus hooleyi. Diagnosis. As for type species. Anteophthall11os11chus hooleyi sp. novo Text-figures Derivation of name. Named in honour of Reginald W. Hooley, a collector of dinosaur, pterosaur and crocodilian fossils from the Isle ofwight. Holotype. NHMUK R3876, several large blocks containing portions of a partial, articulated skeleton of a crocodilian that was once m long (total reconstructed skull length c. 600 mm). Elements that are visible include the almost complete cranium and mandible, the axial skeleton (minus the left and right sacral vertebra II, the caudal vertebrae, caudal ribs and haemal arches), coracoids, right humerus, radius, ulna and possibly the right radiale and ulnare, the right ilium and ischium, the ala of the right pubis, the right femur, several gastralia, a portion of the paravertebral shield (including 20 consecutive osteoderms from the right side), and an articulated portion of the gastral shield. Referred material. IWCMS , a partial, disarticulated skeleton from a plantdebris layer in either the basal part of the Vectis Formation or the upper part of the Wessex Formation at Compton Bay (SZ ; S. Hutt, pers. comm. 2011), Isle of Wight, collected by N. Chase; IWCMS , a partial skull, an incomplete right dentary, both lacking teeth, a number of paravertebral osteoderms and several thoracic vertebrae, collected by R. Giles in 2005 from a plant-debris layer in the lower part of the Wessex Formation following a cliff fall just to the west of Brook Chine (SZ ). Type locality and horizon. Atherfield Point, Isle ofwight (SZ ), Shepherd's Chine Member, Vectis Formation (Daley and Insole 1984; Allen and Wimbledon 1991; S. Hutt, pers. comm. 1997; Insole et al. 1998). Referred material (IWCMS ) indicates that the taxon also occurs in the Wessex Formation. Diagnosis. Anteophthalmosllchus hooleyi gen. et sp. novo is distinguished from other goniopholidids based on the following unique combination of characters

15 English Wealden Fossils 319 Text-fig Anteophthalmosuchus hooleyi gen. et sp. nov., NHMUK R3876, holotype skull in A B, dorsal aspect (photograph and schematic interpretation) and C D, occipital aspect (photograph and schematic interpretation). Scale bar represents 200 mm. (autapomorphies marked with an a ): entire cranial table, infratemporal region (excluding the medial surface of the quadratojugal) and the entire maxillary rostrum uniformly covered in similarly sized, regularly spaced, roughly circular pits (shared with G. baryglyphaeus, G. gracilidens and IRSNB R1537); orbital portion of lacrimal, prefrontal and frontal capped by a thin, dorsally pitted, rectangular palpebral that does not project laterally over the orbit (a); rostral process of frontal ends in a sharp point (shared with G. baryglyphaeus, G. gracilidens and IRSNB R1537); interorbital transverse ridge absent (shared with IRSNB R1537); lacrimal

16 320 Crocodilians Text-fig Anteophthalmosuchus hooleyi gen. et sp. nov., NHMUK R3876, holotype skull in ventral aspect. A, photograph. B, schematic interpretation. Scale bar represents 200 mm. and rostral part of jugal with distinct shallow groove extending rostrally from the orbital border (shared with G. simus, G. willetti and numerous mesoeucrocodylians); rostrolateral part of postorbital forms a distinct, laterally bowed process that projects from the cranial table partly to encircle the orbit laterally, extending rostroventrally towards, but not quite contacting the dorsal surface of the jugal arch (shared with IRSNB R1537); dorsal outline of supratemporal foramen subcircular and almost twice the maximum diameter of the orbits (a); pterygoidopalatine suture rises acutely from the posterior end of the suborbital fenestrae (a, but possibly shared with Leiokarinosuchus brookensis gen et sp. nov.; see below); palatines with subparallel lateral borders, bulging laterally slightly mid-way along the suborbital fenestrae (shared with L. brookensis and many eusuchians); internal septum that divides the secondary choanae formed by the pterygoids, extending rostrally to contact the posteriormost extent of the palatines (a, but possibly shared with L. brookensis); occipital condyle c. 75 per cent of the maximum width of the foramen magnum, and c. 10 per cent of the maximum reconstructed width of the dorsal surface of the cranial table (a); external mandibular fenestra absent (shared with G. baryglyphaeus and IRSNB R1537); dorsal crest of the ilium very low anteriorly, and

17 English Wealden Fossils 321 Text-fig Anteophthalmosuchus hooleyi gen. et sp. nov., NHMUK R3876, holotype skull in right lateral aspect. A, photograph. B, schematic interpretation. Scale bar represents 200 mm. close to the supraacetabular crest (shared with G. crassidens and IRSNB R1537 and R1539); ala of coracoid with nearly straight dorsal margin and subparallel ventral margin that flares ventrally slightly only towards its tip (a). Remarks. The holotype of Anteophthalmosuchus hooleyi, NHMUK R3876, was recovered from a large mass of rock that fell from the cliff at Tie Pits, near Atherfield Point on the Isle of Wight in The horizon that the block came from was m below the top of the Vectis Formation. Portions of the skeleton along with the skull were washed up onto the shore during 1905 and passed on to the Sedgwick Museum, Cambridge. The majority of collecting was carried out by Walter White, coxswain of the Atherfield lifeboat (Hooley 1907). At the age of just 21, Reginald W. Hooley presented a detailed description of NHMUK R3876 to the Geological Society of London in November His paper includes labelled photographs of the cranium and mandible (Hooley 1907, pls 2 3) and the assembled skeleton in both right and left lateral aspect (pl. 4). Following comparisons with Goniopholis tenuidens (NHMUK 48300, Owen 1879a; cf. Goniopholis sp. indet, Salisbury 2002), Nannosuchus gracilidens (NHMUK 48217, Owen 1879a; Goniopholis gracilidens, Salisbury 2002), Oweniasuchus (Brachydectes) minor (NHMUK 48304, Owen 1879a; Theriosuchus pusillus, Salisbury 2002), Oweniasuchus (Brachydectes) major (NHMUK 48328, Owen 1879a; Crocodilia

18 322 Crocodilians Text-fig Anteophthalmosuchus hooleyi gen. et sp. nov., NHMUK R3876, holotype, portion of the postcranial skeleton in right lateral aspect, anterior to the right. A, photograph. B, schematic interpretation. Scale bar represents 100 mm. incertae sedis, Salisbury 2002) and the holotype of Goniopholis crassidens (NHMUK 3798), Hooley (1907) assigned the specimen to Goniopholis crassidens. Hooley s (1907) assignment of NHMUK R3876 to G. crassidens was based primarily on similarities he discerned between the cranium of this specimen and some of the bones and impressions preserved on one of the smaller of the four slabs assigned to the holotype of G. crassidens (NHMUK 3798; see Salisbury 2002, textfig. 4). However, the association of the latter smaller slab with the two main slabs of NHMUK 3798 is questionable; although the slab is listed in the catalogue of Mantell s collection as pertaining to the specimen, it is not illustrated, described or even mentioned by Mantell in any of his publications where the specimen is figured (e.g. Mantell 1839, 1851; Owen 1842, , 1878, 1879a, b; Lydekker 1888a;

19 English Wealden Fossils 323 Woodward and Sherborn 1890). The first person to mention it is Hooley (1907), who was made aware of Mantell s catalogue by A. Smith Woodward. As outlined by Hooley (1907, p. 58), the smaller slab associated with NHMUK 3798 preserves some bones, but mainly external moulds of what appears to be the dorsal surface of the cranial table and posterior part of the maxillary rostrum. Two pieces of bone (a portion of bone between these was removed by A. Smith Woodward for Hooley, the location of which is unknown) appear to form the medial sides of the orbits, and include portions of the parietal, frontal, prefrontal and possibly the postorbital. External moulds of the nasals, lacrimals and maxillae are apparent, but the sutures between these bones are difficult to discern. There is a clear suture on the medial margin of the right orbit, most likely the frontoprefrontal suture. Alternatively, as in G. simus, G. willettii and probably G. baryglyphaeus, it could be argued that this suture represents a contact between the prefrontal and the postorbital within the orbit. Central to Hooley s argument that NHMUK R3876 can be assigned to G. crassidens is that the impression of the interorbital region on the smaller slab associated with NHMUK 3798 does not show any indication of a transverse ridge. As discussed previously, a transverse interorbital ridge occurs in G. simus, G. gracilidens and G. baryglyphaeus, extending across the frontal, the prefrontal and lacrimal (a similar ridge occurs in some American goniopholidids and many other mesoeucrocodylians, including extant caimans). In G. willetti, the interorbital ridge is restricted to the frontal. An interorbital ridge is clearly absent from NHMUK R3876. Contrary to Hooley (1907), there is a distinct demarcation in sculpture pitting in this region on the smaller slab associated with NHMUK Posteriorly, on the cranial table, the sculpture comprises relatively large, evenly spaced pits. Between the orbits, however, these pits become more rostroposteriorly elongated, quickly becoming indistinct grooves and striations on the maxillary rostrum (mainly on the nasals and maxillae). Such a change in sculpture pitting is reminiscent of the condition in G. simus (see Salisbury et al. 1999), and, to our mind, is suggestive of a transverse ridge. This pattern of pitting is quite different from that on BMNH R3876 where, in addition to the absence of an interorbital transverse ridge, similarly sized, uniformly spaced, roughly circular pits occur over most of the cranial table and maxillary rostrum. A similar condition also occurs on the skull of one of the G. simus specimens from Bernissart (IRSNB R1537; Salisbury et al Even if it could be demonstrated that the bones and external moulds preserved on the smaller slab associated with NHMUK 3798 do indeed belong to the same individual as that preserved on the two main slabs, we agree with Salisbury et al. (1999) that Hooley s (1907) referral of NHMUK R3876 to G. crassidens based on this feature alone is problematic and difficult to justify. Differences between the interorbital regions of these two specimens make their assignment to a single taxon unlikely.

20 324 Crocodilians Text-fig Anteophthalmosuchus hooleyi gen. et sp. nov., NHMUK R3876, holotype, portion of the postcranial skeleton in left lateral aspect, anterior to the right. A, photograph. B, schematic interpretation. Scale bar represents 100 mm. There are other differences between NHMUK R3876 and G. crassidens (NHMUK 3798). One seemingly obvious difference overlooked by Hooley (1907) was the absence of an external mandibular fenestra on NHMUK R3876, but its presence in G. crassidens (Hooley notes the former but not the latter). The endocast of the left mandibular ramus of the left main slab of NHMUK 3798 shows the internal ventral and rostrodorsal outline of an oblong-shaped, external mandibular fenestra, bounded rostrodorsally by the dentary, posterodorsally by the supraangular and ventrally by the angular. The portion of the angular that comprises the ventral border is visible on the right slab (see Salisbury 2002, text-figs 3 4). The shape, size and position of this fenestra on NHMUK 3798 are similar to the condition in

21 English Wealden Fossils 325 G. simus (see Salisbury et al. 1999). However, it is worth noting that the absence of an external mandibular fenestra is not restricted to NHMUK R3876 among European goniopholidids. This feature also occurs in Goniopholis baryglyphaeus and on one of the G. simus specimens from Bernissart (IRSNB R1537; the second specimen, IRSNB R1539 lacks a skull and mandible). Several aspects of the skull and skeleton of NHMUK R3876 confirm that it is a goniopholidid. The biserial paravertebral shield is made of up rectangular osteoderms that are longer in a mediolateral direction than they are wide, with a ventrolaterally inclined lateral part bearing a long, spine-like articular process that articulates with a shallow groove on the internal articular face of the proceeding osteoderm (i.e. a stylofoveal or peg-and-groove articulation; Salisbury and Frey 2001). All the vertebrae are amphicoelous, and it has an extensive, plate-like gastral shield comprising interlocking, polygonal osteoderms. While initially identified by Owen (1842) as being restricted to Goniopholis, this type of postcranial morphology is now also known for Dakatosuchus kingi Mehl, 1941 (Vaughn 1956; Langston 1974), Eutretauranosuchus delfsi Mook, 1967, V. leptognathus (Buffetaut and Hutt 1980; Norell and Clark 1990), Sunosuchus junggarensis Wu et al., 1996b, Woodbinenesuchus byersmauricea Lee, 1997 and Siamosuchus phuphokensis Lauprasert et al., 2007; taxa that are all regarded as members of Goniopholididae. Similarly, the well-developed maxillary fossae, and the obtuse, gently lingually curved and slightly labiolingually compressed teeth with mesial and distal carinae and apicobasally aligned striations on NHMUK R3876 are now also associated with a wide range of goniopholidids and closely related taxa, not just Goniopholis (Hulke 1878; Owen 1878, 1879a; Dollo 1883; Koken 1887; Hooley 1907; Ortega et al. 1996; Krebs and Schwarz 2000; Salisbury 2002; Schwarz 2002; Schwarz-Wings et al. 2009). In common with all goniopholidids for which the mandible is known, the splenial of NHMUK R3876 is well integrated into the mandibular symphysis. Some aspects of the morphology of NHMUK R3876 are shared with some, but not all, Goniopholis species. As noted by Hooley (1907), only a small portion of the frontal of NHMUK R3876 participates in the dorsal margin of the orbit proper. In G. simus and G. baryglyphaeus, the frontal appears to be excluded from the dorsal margin of the orbit proper by a posterolateral extension of the prefrontal that contacts the postorbital: see De Andrade and Hornung (2011) for a detailed discussion of this feature. In G. willetti, this same area is covered by a triangular palpebral, which sits atop the orbital margin proper. In both G. simus and G. willetti, the prefrontal and postorbital contact each other within the orbit, ventral to the dorsal surface of the cranial table. In G. willetti, it is possible to see this contact beneath and medial to the palpebral. In G. gracilidens, the frontal forms a much larger portion of the medial border of the orbital margin proper, but with a large rostral, and much smaller posterior palpebral in place, this margin was probably excluded from the functional orbital border. In addition, the rostral process of the frontal of NHMUK

22 326 Crocodilians R3876 ends in a sharp point as in G. baryglyphaeus, G. gracilidens and IRSNB R1537, rather than in a W-shape as in G. simus and G. willetti. As in G. baryglyphaeus, initially there appears to be no clear evidence for an extensive palpebral in NHMUK R3876. In G. gracilidens and G. willetti, when in place, the palpebral effectively excludes the frontal from the functional margin of the orbit. It also changes the shape of the functional orbital margin, projecting out from the orbital border proper (formed by the prefrontal, frontal and postorbital) over the orbit laterally. The presence of a palpebral in G. simus is more variable (see Salisbury et al and De Andrade and Hornung 2011), but the effect on the orbit is similar to the condition in G. gracilidens and G. willetti. On the left side of the skull in NHMUK R3876, a shallow, slightly bevelled, smooth, rectangular surface on the dorsal surface of the lacrimal, prefrontal and frontal portions of the orbital margin indicates an area that may have formed a point of attachment for a palpebral immediately rostral to the small frontal contribution to the orbital margin proper. On the right side of the skull, this area is level with the rest of the cranial table and similarly covered with evenly spaced circular pits, but separated from the prefrontal by a sediment/putty-filled line that is similar in width and colour to the straight planar sutures on other parts of the skull. This small rectangular piece of bone most likely represents a thin palpebral. Significantly, this palpebral does not appear to have extended out laterally over the orbit; assuming that the right element is complete, its lateral border is continuous with the rest of the orbital margin (i.e. there is no distinction between an orbital border proper and a functional orbital border). Although poorly preserved, this aspect of the skull of NHMUK R3876 is distinct from that of G. simus, G. gracilidens, G. baryglyphaeus and G. willetti. A similar condition with the palpebral to that seen in NHMUK R3876 also occurs on one of the G. simus specimens from Bernissart (IRSNB R1537). Although the right orbit of IRSNB R1537 is damaged, a small palpebral is preserved above the left orbit, partly detached from a shallow, bevelled surface. Similar to the condition in NHMUK R3876, this element extends laterally only a little beyond the orbital margin proper. As noted by Hooley (1907, p. 54), the posterior part of the lacrimal on NHMUK R3876 bears two ridges, converging rostrally and divided by a shallow groove, forming the rostral border of the orbit. The depth of the groove and the height of the ridges accentuate each other. A similar condition occurs in G. willetti, but the groove is broader and clearly extends onto the jugal. The same feature occurs to varying degrees in many extant crocodilians (e.g. many Crocodylus spp.), where the groove forms part of the rostral attachment area for the eyelid. Other aspects of the cranial ornamentation of NHMUK R3876 also set it apart from some, but not all, species of Goniopholis. Unlike G. simus and G. willetti, the entire cranial table, infratemporal region (excluding the medial surface of the quadratojugal) and the entire maxillary rostrum are uniformly covered in similarly sized,

23 English Wealden Fossils 327 regularly spaced, roughly circular pits. A similar condition also occurs on the skull of one of the G. simus specimens from Bernissart (IRSNB R1537; Salisbury et al. 1999), G. baryglyphaeus and G. gracilidens, as well as some American goniopholidids. However, in common with G. simus, and to some degree G. willetti, G. baryglyphaeus and G. gracilidens possess a distinct interorbital transverse ridge; uniform sculpture pitting and the absence of an interorbital transverse ridge is a combination shared exclusively by NHMUK R3876 and IRSNB R1537 among European goniopholidids. As occurs in G. simus, possibly G. willetti, and a pair of premaxillae from the Grinstead Clay Formation of Cuckfield (NHMUK ; Text-fig. 24.5), there is no clear evidence for a premaxillary fenestra on the preserved portion of the premaxillae of NHMUK R3876. A more complete set of premaxillae from a referred specimen, IWCMS , confirms that this is the case. On the latter specimen, as in G. simus (Salisbury et al. 1999) and possibly G. willeti, there is a small, median, diamond-shaped fossa on the floor of the nasal vestibulum. We are not sure what the condition is in IRSNB R1537. Hooley (1907, p. 54) described the nasals of NHMUK R3876 as entering the naris and widening rostrally as they approach the premaxillary-maxillary contact. He additionally stated that the posterior process of the premaxilla on the dorsal surface of the maxillary rostrum of NHMUK R3876 was relatively short and blunt. He used these features to help distinguish NHMUK R3876 from G. simus, where the nasals taper rostrally and are excluded from the naris by a broad contact between the premaxillae, each of which has a long, triangular posterior process on the dorsal surface of the maxillary rostrum. However, our examination of NHMUK R3876 and IWCMS indicates that, similar to other goniopholidids, including G. simus (Salisbury et al. 1999), the nasals of NHMUK R3876 are excluded from the dorsal border of the naris by the premaxillae, and the posterior process of each premaxilla is long and triangular. In NHMUK R3876, however, the rostral extent of the nasals appears to be much closer to the naris than it is in G. simus, G. baryglyphaeus, G. gracilidens, G. willetti and V. leptognathus, similar to the condition that Dollo (1883, p. 318) described for IRSNB R1537. It seems that Hooley (1907) mistook the two posterior premaxillary processes as the rostral extension of the nasals. The suture between each of these two processes and the left and right maxillae is much more distinct than that between the nasals and premaxillae. The maxillary rostrum of NHMUK R3876 appears to have broken off transversely at a point level with the reception notch for the mandibular pseudocanines. This seems to be a common point of fracture for the skulls of European goniopholidids, as evidenced by other specimens of isolated and/or broken premaxillae (e.g. BMB , Text-fig. 24.5) and skulls lacking this portion of the maxillary rostrum (e.g. IRSNB R1537). Hooley (1907) also noted that the external outline of the supratemporal foramen on NHMUK R3876 is subcircular rather than subquadrate, which is the condition in G. simus. Similarly shaped supratemporal foramina are seen in IRSNB R1537 and

24 328 Crocodilians G. gracilidens, but in NHMUK R3876 they are proportionately much larger, being almost twice the size of the orbits. As discussed previously, one of the key diagnostic features of European Goniopholis is a laterally exposed QSE canal. This feature occurs in G. simus, G. baryglyphaeus and G. willetii (this part of the skull is not exposed on the type of G. gracilidens). In NHMUK R3876, as in most other mesoeucrocodylians (see previous discussion on this feature), the squamosal is sutured to the quadrojugal laterally, posterior to the external otic cavity, fully enclosing the QSE canal. In addition to the characteristics discussed above, a suite of apomorphic features of the skull and postcranium of NHMUK R3876 help to distinguish it from other goniopholidids. In his comparative section, Hooley (1907) noted several differences between the palate of NHMUK R3876 and other goniopholidids known at the time. In palatal aspect, the pterygoidopalatine suture on NHMUK R3876 rises acutely from the posterior end of the suborbital fenestrae, rather than horizontally as in G. simus. The palatines of NHMUK R3876 have subparallel lateral borders, bulging slightly mid-way along the suborbital fenestrae, unlike the rostrally flaring palatines of G. simus (see Owen 1878, pl. 5, fig. 2; Buffetaut 1982, fig. 7A). A similar morphology occurs in BGS GSM , a partial skull from the Wessex Formation at Brook Bay, Isle of Wight (Huxley 1875; see below). The secondary choanae of NHMUK R3876 are additionally more elongate and located slightly further posteriorly in the pterygoids relative to the posterior margin of the palatines and the suborbital fenestrae than they are in Goniopholis spp. (e.g. G. simus; see Owen 1878, pl. 5, fig. 2; Buffetaut 1982, fig. 7A). In NHMUK R3876 the secondary choanae are almost fully enclosed by the pterygoids, with the palatines contributing only a small percentage to their rostrolateral margins, as occurs in advanced neosuchians such as Bernissartia fagesii Dollo, 1883 (Buscalioni and Sanz 1990a). The internal septum dividing the secondary choanae, formed by the pterygoids, is broader in NHMUK R3876 than in G. simus, and extends rostrally to contact the posteriormost extent of the palatines. The occipital condyle of NHMUK R3876 is proportionately small relative to the foramen magnum and the overall size of the skull; its maximum width is approximately 75 per cent of the maximum width of the foramen magnum and approximately 10 per cent of the maximum reconstructed width of the dorsal surface of the cranial table. In G. simus (IPB R359, see Salisbury et al. 1999), the same proportions are 118 and 9 per cent, respectively, and in a similarly sized Crocodylus porosus Schneider 140 and 20 per cent. Hooley (1907) noted that the dorsal surface of the occipital condyle of NHMUK R3876 lacked a median groove; a feature that would be unusual for a crocodyliform. However, this part of the occipital condyle of NHMUK R3876 is poorly prepared, such that the expression of this feature is difficult to establish with certainty. An unusual and distinctive feature of NHMUK R3876 relates to the postorbital and the degree to which it encloses the orbit laterally. Typically in crocodyliforms

25 Crocodilians 329 the portion of the postorbital immediately dorsal to the postorbital bar forming the rostrolateral corner of cranial table is gently convex (e.g. eusuchians) or squared off in dorsal aspect, sometimes forming a short rostrolaterally directed process (e.g. Goniopholis spp., Termil10naris robusta Wu et al., 2001). In NHMUK R3876, however, this part of the postorbital forms a distinct, laterally bowed process partially encircling the orbit laterally, extending rostroventrally towards, but not quite contacting, the dorsal surface of the jugal arch. On the left side of the skull it is possible to see that this process is continuous with the dorsal half of the postorbital bar, such that the rostrolateral half of the braincase is obscured from view when the skull is in lateral aspect. Hooley (1907, p. 55) noticed this feature, commenting on the vertical elevation of the orbits, suggesting that anything other 'than a forward vision was precluded: He rightly considered the condition to be in sharp contrast to the everted and laterally directed orbits of most extant eusuchians (GaJlialis gangeticus would be the only possible exception). A similar rostrolateral process of the postorbital to that seen in NHMUK R3876 also occurs on the skull of one of the 'G. simlls' specimens from Bernissart (IRSNB R1537; Salisbury et al. 1999). Vectisuchus leptognathus (SMNS 50984; see below) has similar, rostrally directed orbits, with a broad, rostrodorsally inclined postorbital process (see Text-fig A). Indeed, Buffetaut and Hutt (1980) listed this feature in their diagnosis (see below). However, the rostrolateral corner of the postorbital on the cranial table of V. leptognathus does not form the distinct process seen in NHMUK R3876 and IRSNB R1537. Instead, the lateral enclosure of the orbit seems to result from a rostral expansion of the dorsal part of the postorbital bar. Unfortunately, the full rostral extent of the latter process on SMNS is unclear, because it appears to have broken off. Aspects of the ilium and coracoid of NHMUK R3876 are distinct from other European goniopholidids. The preacetabular portion of the ilium is proportionately very short relative to the ala (Hooley 1907, p. 57). The ala is approximately the same length (c %, measured from its posterior tip to the posterior edge of the acetabular peduncle) as the iliac body (measured from anterior tip of the preacetabular tuberculum to the posterior edge of the acetabular peduncle). In G. crassidens (NHMUK 3798) and an isolated ilium from the Wealden of Hastings (NHMUK R1956), the ala is slightly shorter relative to the iliac body, but still much larger than the condition seen in eusuchians. The supraacetabular crest on the ilium of NHMUK R3876 is anterodorsally positioned, forming a sharp edge anteriorly and a raised, rugose surface posteriorly. Anteriorly, the dorsal crest is very low and close to the supraacetabular crest. Although the latter condition is similar to that seen in G. crassidens (NHMUK 3798), 'G. simus' from Bernissart (IRSNB R1537 and R1539) and NHMUK R1956, the lateral overhang of the dorsal crest and supraacetabular crest is less in NHMUK R3876. A well-developed anterodorsal tuberosity, presumably for the attachment of the m. iliocostalis, is present and also similar to both the

26 330 English Wealden Fossils holotype of G. cmssidens, NHMUK R1956, and to an undescribed Goniopholis specimen from Berriasian of north-western Germany (Hornung et al. 2009). However, in NHMUK R3876, this process is positioned further dorsally from the dorsal edge of the preacetabular tuberculum than it is in G. crassidensor NHMUK R1956. In lateral aspect, the dorsal and ventral margins of the iliac ala of NHMUK R3876 closely mirror each other in terms of curvature, with the dorsal margin being convex and the ventral margin being concave, giving this part of the ilium a symmetrical appearance. The anterior portion of preacetabular tuberculum in NHMUK R3876 is large and bulbous and continuous with the supraacetabular crest, whereas in NHMUK R1956 the two structures are separated by a short, dorsally concave margin. The coracoid of NHMUK R3876 has a proportionately narrow ala relative to the condition seen in extant eusuchians, with a nearly straight dorsal margin and subparallel ventral margin that flares ventrally slightly only towards its tip. A similar morphology is seen in an isolated coracoid from the Wealden of the Isle of Wight (NHMUK R214), referred to Goniopholis minor Koken, 1887 by Lydekker (l890a), presumably because of its small size. Unlike the latter specimen, however, on NHMUK R3876 the groove between the posterior rugosity for the attachement of the m. anoconaeus coracoscapularis and the ventrolateral rugosity for the attachment of the m. coracoantebrachials is not pronounced (Furbringer 1876). The coracoid foramen ofnhmuk R3876 is proportionately smaller than that on NHMUK R214. Having considered Hooley's (1907) referral of NHMUK R3876 to G. cmssidens, along with the numerous cranial and postcranial differences that exist between this specimen and currently recognized species of Goniopholis, we conclude that there are strong grounds for recognizing NHMUK R3876 as a new genus and species of goniopholidid, as argued previously (Salisbury et al. 1999). The new taxon, AnteophthalmoSlIclllls hooleyi gen et. sp. nov., shares several features with material previously referred to 'G. simlls' by Dollo (1883) from the Barremian of Bernissart, Belgium, and two partial skulls from the Wessex Formation of Brook Bay, Isle of Wight (BGS GSM and NHMUK R3876; see below). It is also noteworthy that an orbital morphology comparable to that seen in A. hooleyi occurs in V: leptognathus from Wealden strata of similar age on the Isle of Wig ht, the implication being that it is a result of shared ancestry. It is hoped that future phylogenetic analysis (which is beyond the scope of the present contribution) will shed light on these issues. cf. Anteophthalmosllchlls Text-figure Material. BGS GSM , a relatively complete basicranium and the posterior end of both mandibular rami (Huxley and Etheridge 1865; Huxley 1875, pi. 19, fig. 3, p. 432; Hulke 1878).

27 English Wealden Fossils 331 Text-fig cf. Anteophthalmosuchus, BGS GSM , partial skull and mandible in A, dorsal, B, ventral, C, occipital and D, left lateral aspects. Scale bar represents 50 mm. Locality and horizon. Brook Bay, Isle of Wight (between SZ and SZ ), presumably from the Wessex Formation. Remarks. In his landmark paper on the evolution of crocodilians, Huxley (1875, p. 432) used a partial crocodilian skull and mandible from the Isle of Wight to illustrate the palatal condition that lies on the boundary line between that characteristic of the Mesosuchia and that which distinguishes the Eusuchia. At the time, the specimen was part of the fossil collection of the Jermyn Street Museum of Practical Geology, where Huxley was based from 1854 to 1872 (Forgan and Gooday 1996). In 1934, BGS GSM (along with the rest of the collection) was transferred to the former Geological Survey Museum, London, before eventually ending up in the BGS headquarters at Keyworth, near Nottingham. The orginal label associated with BGS GSM and its catalogue listing (Huxley and Etheridge 1865, p. 261) states that it was collected from Brook Point, Isle of Wight. No further collection data is available. Brook Point, Brook and Brooke are all terms that were once used for Brook Bay, a stretch of coastline between Hanover and Sudmoor points (between SZ and SZ ) on the south-western coast of the Isle of Wight (White 1921; Buffetaut 1983; Benton and Spencer 1995b). Although the Vectis Formation does crop out in the area, the main Cretaceous fossil-bearing horizon in the cliffs at Brook Bay is the Wessex Formation

28 332 English Wealden Fossils (Insole et nl. 1998). Although it cannot be confirmed, we therefore assume that BGS GSM derives from the Wessex Formation. The specimen is badly water-worn and the external surface of several bones is damaged, such that a number of sutures are obscured or difficult to discern. Although the area around the secondary choanae was prepared prior to Huxley's (1875) paper, there is still a considerable amount of matrix adhering to the specimen. The only part of the maxillary rostrum preserved is that immediately rostral to the orbits. Similarly, only the posterior half of the mandible is present. Most of the dorsal surface of the frontal, prefrontals and lacrimals is missing, as is the dorsal portion of both postorbitals, the left squamosal and the right quadratojugal. The posterior part of each mandibular ramus, including the retroarticular process is also missing, and large parts of the posterolateral part of the pterygoid plate and ectopterygoid wings are obscured by matrix. As described by Huxley (1875), the secondary choanae of BGS GSM are bounded rostrally by the palatines and posteriorly by the pterygoid, indicating that this crocodilian is not a eusuchian. Other features of the skull suggest goniopholidid affinities. The paroccipital process forms a laterally projecting, rounded flange that does not appear to extend beyond the lateral extent of the squamosal in occipital aspect, similar to that of G. simus, G. willetti and A. hooleyi. Although other diagnostic goniopholidid features are not preserved or are obscured by matrix (e.g. maxillary fossae), several aspects of the skull and mandible are reminiscent of A. hooleyi. The posterior margin of each secondary choana is situated close to the posterior margin of the suborbital fenestrae. The palatopterygoid suture extends rostrolaterally from the rostrolateral edge of each opening, such that the palatines contribute only a small amount to each margin. It is not possible to determine where the palatopterygoid suture occurs on the internal septum. Nevertheless, the size, shape and position of the secondary choanae are different from the condition in Goniopholis spp. (e.g. G. simusand G. willetti). The palatines ofbgs GSM are also similar to those of A. hooleyi in that they have subparallellateral borders. But they differ by not flaring slightly rostrally, and there is no obvious bulge along their length. Also in common with A. hooleyi, there is no indication of an external mandibular fenestra, and the occipital condyle is considerably smaller than the foramen magnum. The QSE canal does not appear to have been exposed laterally as it is in GOlliopholis, with the preserved portion of the left squamosal sutured to the dorsal surface of the quadratojugallaterally. Pitting on the preserved portion of the parietal, right squamosal and left mandibular ramus is similar to that seen on the holotype of A. hooleyi. The rostralmost preserved portion of the frontal appears to have been almost smooth, but it is possible that this part of the bone has been abraded and is not the original external surface. Despite BGS GSM being incomplete and poorly preserved, it displays a number of features that compare well with A. hooleyi. Differences such as its smaller

29 English Wealden Fossils 333 size and more gracile, less swollen palatines suggest that it represents a distinct species. Alternatively, such differences could be the result of ontogenetic variation, with this specimen being a younger, less mature individual than the holotype. Presumably deriving from the upper part of the Wessex Formation, it is also slightly younger than NHMUK R3876. Until BGS GSM is more fully prepared or additional material suggests otherwise, we propose that it should be regarded as cf. Anteophthalmosuchus. VECTISUCHUS Buffetaut and Hutt, 1980 Vectisuchus leptognathus Buffetaut and Hutt, 1980 Text-figures Holotype. SMNS 50984, a partial, semi-articulated skeleton from an individual that was once c. 1.2 m long (reconstructed skull length 180 mm), comprising the Text-fig Vectisuchus leptognathus (Buffetaut and Hutt 1980), SMNS 50984, schematic interpretation of holotype skull in A, dorsal and B, ventral aspect. (From Buffetaut and Hutt 1980). Scale bar represents 50 mm.

30 334 Crocodilians cranium and mandible, most of the axial skeleton (minus the sacral and caudal vertebrae), the articular extremities of the right scapular and coracoid, bones of the right forelimb, bones of the right hindlimb, and numerous osteoderms from the paravertebral and gastral shields (Text-fig B). Type locality and horizon. Barnes High, Isle of Wight (SZ ), Red Marls of the Upper Wessex Formation, just below the base of the Vectis Formation (Buffetaut and Hutt 1980; Daley and Insole 1996). Remarks. Steve Hutt discovered the holotype specimen of V. leptognathus in 1977, and it was purchased by the SMNS in The near-complete skeleton was in situ and fully articulated, ventral side up with the neck and skull dorsally flexed, possibly the result of subaerial exposure and desiccation prior to burial (Buffetaut and Hutt 1980). Unfortunately, severe weather conditions at the time of its discovery and subsequent excavation apparently resulted in the loss of the pelvic girdle, hindlimbs and tail when part of the cliff in which the specimen was preserved collapsed (Buffetaut and Hutt 1980). When one of us (SWS) examined the specimen in 1998, however, in addition to the elements described by Buffetaut and Hutt (1980), it included an almost complete right hindlimb. The size of the elements preserved and the nature of the sediment in which they occur are consistent with this specimen belonging to the holotype individual. We assume that it was collected from the locality sometime after the cliff collapse and was part of the sale to the SMNS (only one sale is recorded), but for whatever reason it was not mentioned in the initial description. Buffetaut and Hutt (1980) described V. leptognathus as a new genus and species of goniopholidid, with most of their account focusing on the cranium and mandible, of which they provide a diagrammatic interpretation and schematic reconstruction (Buffetaut and Hutt 1980, figs 1 2, 4 5; Text-fig ). Although relatively complete, the preserved portions of the specimen are crushed and the majority of bones are fractured. Some of the larger breaks have been filled with epoxy putty, which prevents detailed examination of the cross-sectional morphology of many of the bones, particularly in the skull. As a result, many of the sutures on the skull are difficult to identify. Most aspects of the postcranium of V. leptognathus are consistent with those of other goniopholidids. All the vertebrae are gently amphicoelous, with at least ten in the thoracic series and four lumbars. The biserial paravertebral shield comprises rectangular osteoderms that are longer mediolaterally than they are wide, with a ventrolaterally inclined lateral part bearing a long, spine-like articular process. Unlike other goniopholidids, in addition to a low lateral keel between the medial and lateral parts, the paravertebral osteoderms of V. leptognathus bear a second, less-pronounced medial keel, commencing mid-way along the posterior/terminal margin of the medial part of the osteoderm and extending in a craniolaterally convex arc craniomedially to a point just posterior to the external articular surface (Text-fig B). This feature appears to be unique to V. leptognathus. Typical

31 Crocodilial t B car darmost mad '------L----palrs lat car dermost ;e;;;:p,_, lat proc art 4-~,::":ermost D~--p-a-rs-m-ed---~pars lat TEXT-FIG ,rectisuclJlls leptogllllthus (Buffetaut and Hutt, 1980), SMNS 50984, holotype. A, schematic interpretation of the posterior half of the skull in right lateral aspect. B-D, reconstruction of two articulating paravertebral osteoderms from the lumbar region. B, from the right side of the paravertebral shield in external aspect. C, from the right side of the paravertebral shield in lateral aspect. D, from the right side of the paravertebral shield in posterior aspect. Anterior is indicated by the arrow in both Band C. Of note is the presence of a crescent-shaped keel on external surface of the medial part of the osteoderm. Scale bars represent 20 mm in A, 25 mm in B-D. of goniopholidids, the gastral shield of V. leptogllathlls comprises interlocking, polygonal osteoderms. The forelimbs of V. leptognathlls are proportionately elongate relative to the condition seen in eusuchians and most other mesoeucrocodylians of comparable size and body shape. This feature is shared with a number of other goniopholidids. It was noted by Dollo (1883) and Buffetaut (1982) for the two 'G. simus' specimens from Bernissart (IRSNB R1537 and RI539), by Hooley (1907) and Buffetaut (1982) for A. hooleyi (NHMUK R3876), and by Wu et al. (l996b) for Sunosuelms jllllggarellsis (Wu et al. 1996b). In the last of these taxa, the forelimbs are per cent longer than the hindlimbs, whereas in eusuchians they are per cent shorter. In goniopholidids, it is the humerus that produces the difference in the length of the forelimb, being considerably longer proximodistally than the femur. The preserved

32 336 Crocodilians femur associated with SMNS is shorter than the humerus, resulting in limb proportions that are consistent with those in other goniopholidids. Similar to IRSNB R1537 and R1539, the carpal elements of V. leptognathus are also more elongated than in most other mesoeucrocodylians. Several aspects of the skull and mandible of V. leptognathus are consistent with the condition seen in Goniopholis spp. and A. hooleyi. The overall shape of the cranial table and size of the supratemporal foramina are very similar to A. hooleyi. There is broad inclusion of the frontal in the supratemporal foramina, and a prominent boss is present on the posterolateral corner of the squamosal. As in most goniopholidids, the nasals reach the premaxillae, but are excluded from the naris. Although the otic region is poorly reserved, the squamosal of V. leptognathus appears to have been sutured to the quadrojugal laterally, posterior to the otic cavity, thereby fully enclosing the QSE canal: in this respect it is unlike Goniopholis spp. but similar to A. hooleyi and most other mesoeucrocodylians. Buffetaut and Hutt (1980) did not comment on the presence or absence of a maxillary fossa in V. leptognatus, a feature generally considered characteristic of goniopholidids. The portion of each maxilla that would have preserved this feature is poorly preserved, such that it is difficult to say with certainty whether it was present. Buffetaut and Hutt (1980) distinguished V. leptognatus from other goniopholidids based on its small size (reconstructed skull length 180 mm, estimated total body length 1.2 m), and on a suite of characters associated with its slender, moderately elongated rostrum that is sharply demarcated from the posterior part of the skull. The orbits are proportionately large, rounded and directed rostrally and laterally. This condition is similar to that in A. hooleyi. Buffetaut and Hutt (1980) also noted that the postorbital bar was displaced medially and ventroposteriorly. Consequently, the postorbital bar overhangs the posterior part of the jugal arch. Unlike in A. hooleyi, the rostrolateral corner of the postorbital on the cranial table of V. leptognathus does not form a distinct process as it does in NHMUK R3876 and IRSNB R1537. Instead, the lateral enclosure of the orbit seems to result from a rostral expansion of the dorsal part of the postorbital bar, ventral to the dorsal surface of the cranial table (Text-fig A). The full rostral extent of this process is unclear, as it appears to have broken off. The palate of V. leptognathus displays several diagnostic features. As noted by Buffetaut and Hutt (1980), the pterygoid wings are well developed, making the pterygoid almost twice as wide mediolaterally as it is long anteroposteriorly. The secondary choana is proportionately large, and located almost exclusively within the pterygoid, with only a small portion of the palatines contributing to the rostral margin (see Buffetaut and Hutt, figs 4 5). In these respects, the palate is similar to that of A. hooleyi and distinct from that of Goniopholis spp. (e.g. G. simus and G. willetti) where the secondary choanae are narrower and located further rostrally on the palate, with a greater contribution from the palatines.

33 English Wealden Fossils 337 The mandible of V. leptognathus is expanded rostrally, with enlarged and confluent third and fourth alveoli, similar to most species of Goniopholis and other long/slender-snouted mesoeucrocodylians such as Pholidosaurus purbeckensis. All the preserved teeth are similar in size, being long and slender, with a slight lingual curvature. Unlike Goniopholis, the retroarticular process is elongate and more reminiscent of the condition in extant crocodyloids and gavialids than that of other goniopholidids (Buffetaut and Hutt 1980). Numerous isolated crocodilian elements from the Wealden of the Isle of Wight currently housed in the IWCMS have been referred to V. leptognathus. In most instances, these assignments seem to be because the specimens were found in the vicinity of the holotype and are of a comparable size. Given the occurrence of at least one other goniopholidid (A. hooleyi) and three non-eusuchian neosuchians (L. brookensis, Theriosuchus sp. and Bernissartia sp. indet.) in the Wessex Formation, the assignment of isolated vertebrae (e.g. IWCMS 5495) or teeth (e.g. IWCMS 5319) to specific genera is difficult to verify and should be regarded as invalid. Such material should be placed in Mesoeucrocodylia indet. To conclude, V. leptognathus represents a valid goniopholidid neosuchian. It shares a number of unusual characteristics of the orbital region with A. hooleyi, but is otherwise distinct in its possession of a relatively short, narrow rostrum that is sharply demarcated from the rest of the skull. In these respects it approaches G. willetti and the long and slender-snouted pholidosaurids. ATOPOSAURIDAE Gervais, 1871 THERIOSUCHUS Owen, 1879a Type species. Theriosuchus pusillus Owen, 1879a. Lectotype. NHMUK 48330, an almost complete, partly articulated skeleton of an individual that was once 500 mm long, preserved on two slabs set in plaster inside a wooden case (Owen 1879a, pl. 4, , pl. 45; Salisbury 2002, text-fig. 13). Remarks. Owen (1879a) erected Theriosuchus pusillus based on a complete cranium and mandible (NHMUK 48330) and a partially articulated skeleton (NHMUK 48216); but see Salisbury (2002, p. 135) for details concerning the earlier use of the name. Both specimens accompanied material collected by Samuel H. Beckles during the course of his famous Mammal Pit excavations in 1857 at Durlston Bay, Dorset (Owen 1879a, pp. 1, 8, 13). Along with other crocodilian fossils, these specimens come from strata collectively referred to as Beckles residuary marls. These derive from one of three beds in the Berriasian Middle Purbeck Beds of the Purbeck Limestone Group (it is not clear which one): the Mammal Bed (DB 83), the Fern Bed (DB 92/93) or Bed DB 94/96 (Salisbury 2002). The specific epithet pusillus was presumably so-named because of the diminutive size of the species compared with other crocodilians known at the time. Owen

34 338 Crocodilians (1879b) interpreted this as dwarfism, suggesting that it may have had something to do with the small size of the contemporaneous mammals (see Owen 1871) on which he supposed T. pusillus preyed. Owen s idea was later refuted by Joffe (1967). Although Joffe agreed with Owen s size estimates for adult T. pusillus (a total length of c. 500 mm) she opposed the use of the term dwarf in reference to the species, maintaining there was no evidence of size reduction from a larger ancestor (Joffe 1967, p. 639). This supposition has since been supported by Buffetaut (1982), Clark (1986) and Buscalioni and Sanz (1988). At least three other species of Theriosuchus are known from Late Jurassic Early Cretaceous deposits in Europe. Brinkmann (1992) described T. ibericus Brinkmann, 1992 from the Barremian Uña Formation of Uña/Cuenca, Spain, while Schwarz and Salisbury (2005) recognized T. guimarotae in the Kimmeridgian Alcobaça Formation of Guimarota, north-western Portugal. Most recently, Martin et al. (2010) described T. sympiestodon from the Maastrichtian of the Haţeg Basin, Romania, extending the range of the genus by about 58 myr. A fifth species of Theriosuchus, T. grandinaris Lauprasert et al., 2011 has been recognized in the Khorat Group (early Aptian) of Thailand (Lauprasert et al. 2011). Theriosuchus material from other European fossil deposits is rare and of a fragmentary nature, comprising mainly isolated teeth and osteoderms (Buscalioni and Sanz 1984, 1987a, b; Cuny et al. 1991; Evans and Milner 1994; Benton and Spencer 1995b; Thies et al. 1997; Schwarz-Wings et al. 2009). Probable Theriosuchus remains have also been reported from the Lower Cretaceous of the Ordos Basin of Inner Mongolia (Wu et al. 1996b). Theriosuchus can be distinguished from other atoposaurids based on the following unique combination of characters: proportionately short, broad rostrum, with the maxillary rostrum forming between 40 and 45 per cent of the total length of the skull; proportionately small antorbital fenestra; slit-like, horizontally orientated and rostrally positioned external nares, separated from each other by the rostralmost extent of the nasals; shallow sulcus on the dorsal surface of the maxillary rostrum, immediately posterior to the junction between the maxilla, premaxilla and nasal; proportionately long jugal; medial base of the postorbital process formed by the ectopterygoid median crest on the frontal and the parietal in later ontogenetic stages; frontal and parietal partially unfused in early ontogenetic stages; dorsal margin of the supratemporal foramen smaller than the orbit throughout ontogeny; lateral margin of the squamosal bevelled ventrally; proportionately narrow quadrate with a concave mandibular articular surface; secondary choanae bounded by the palatines rostrally and separated by a median septum of the pterygoid; mandibular symphysis that does not extend posteriorly beyond a point level with the sixth dentary tooth; ilium with short preacetabular process and long postacetabular process; biserial dorsal shield comprising parasagittal osteoderms (Owen 1878, 1879a; Joffe 1967; Buffetaut 1981, 1982, 1983; Clark 1986; Buscalioni and Sanz 1990b; Norell and

35 English Wealden Fossils 339 Clark 1990; Brinkmann 1992; Wu et al. 1996b; Salisbury 2001, 2002; Salisbury and Frey 2001; Schwarz and Salisbury 2005). The dentition of Theriosuchus includes three different morphotypes (Owen, 1879a; Joffe 1967; Brinkmann 1992; Salisbury 2002; Schwarz and Salisbury 2005; Schwarz-Wings et al. 2009): (1) slender and conical teeth with basioapically aligned striations that are largely restricted to the lingual face of the crown, found in the premaxilla, rostralmost maxilla, and rostralmost dentary (Owen 1879a; Brinkmann 1992; Schwarz and Salisbury 2005); (2) lanceolate teeth with a fan-shaped distribution of the marginal lingual striations from the middle and posterior portions of the maxilla and dentary (Owen 1879a; Brinkmann 1992; Schwarz and Salisbury 2005); and (3) a third morphology restricted to T. pusillus and T. ibericus in which the teeth are broad and strongly labiolingually compressed, and both the lingual and labial surfaces are covered with striations (although fan-shaped striations are present only on the lingual face) (Owen 1879a; Brinkmann 1989, 1992; Salisbury 2002; Schwarz- Wings et al. 2009) Theriosuchus sp. indet. Text-figure Material. NHMUK R3697 and NHMUK R , isolated teeth from the Ashdown and Wadhurst Clay formations of west Fairlight, and possibly Brede, East Sussex (TQ ; Woodward 1911; Anon. 1912); NHMUK R176, a partial basicranium that includes portions of the parietal, supraoccipital, exoccipital and basioccipital from Brook Bay, Isle of Wight (between SZ and SZ ; see section on BGS GSM ), presumably from the Wessex Formation (Buffetaut 1983). Remarks. Small, mediolaterally compressed, lanceolate teeth with the fan-shaped distribution of the marginal lingual striations that is characteristic of Theriosuchus Text-fig Theriosuchus sp. indet., NHMUK R176, a partial basicranium that includes portions of the parietal, supraoccipital, exoccipital and basioccipital from Brook Bay, Isle of Wight. Schematic reconstruction in A, dorsal and B, occipital aspects. (From Buffetaut 1983). Scale bar represents 20 mm.

36 340 Crocodilians were first reported from the English Wealden in an anonymous news article in Nature during 1912 (Buffetaut 1983). The teeth were collected by Pierre Teilhard de Chardin and Félix Pelletier (see Chapter 32) during their time in Hastings, apparently from the Ashdown and Wadhurst Clay formations of west Fairlight and possibly Brede, East Sussex (cf. Woodward 1911; Buffetaut 1983). Charles Dawson also collected similar teeth from the same area in 1909 (Buffetaut 1983). Some of the teeth collected by Teilhard de Chardin are currently housed in the HASMG. NHMUK R176, the skull fragment from the Wessex Formation of Brook Bay, Isle of Wight, provides more substantial evidence of Theriosuchus in the Wealden Supergroup. The specimen formed part of the Hooley Collection in the NHMUK, and was first noticed by P. Welnhoffer in Buffetaut (1983) provided a detailed description and referred the specimen to Theriosuchus. The shape of the preserved portion of the cranial table, its ornamentation, and in particular the morphology of the parietal are a very close match with that of the lectotype (NHMUK 48330) and paratype (NHMUK 48216) of Theriosuchus pusillus from the Purbeck Limestone Group (Buffetaut 1983). Also consistent with T. pusillus, the medial margin of the supratemporal foramen on NHMUK R176 is slightly elevated above the rest of the cranial table, a feature not apparent in T. guimarotae (Schwarz and Salisbury 2005). However, given the fragmentary nature of the specimen, we agree with Buffetaut (1983) that a specific identification is not warranted, and placement in Theriosuchus sp. indet. is the most appropriate assignment. Family incertae sedis BERNISSARTIA Dollo, 1883 Type species. Bernissartia fagessi Dollo, Lectotype. IRSNB R46, an almost complete, mounted skeleton. In addition to the axial skeleton (which lacks only the posteriormost portion of the tail), the appendicular skeleton and much of the dermal skeleton is present. The latter includes the dorsal shield and articulated portions of the gastral shield. Both the left and right hands are missing, as are the bones of the right foot. Many of the bones, in particular the cranium and mandible, are distorted (Dollo 1883; Buffetaut 1975). Remarks. Bernissartia fagesii is an advanced neosuchian crocodyliform from Western Europe, attaining an adult length of approximately 600 mm. The first two specimens of B. fagesii were found in 1879 in the Early Carboniferous Loronne coalseam at the Sainte-Barbe coal mine in Bernissart, Belgium (Dollo 1883). The fissure filling ( cran ) in which the specimens were discovered is thought to be associated with Early Cretaceous (Barremian) marls (Casier 1978; Norman 1980; Yans et al. 2005). It also contained two skeletons of a larger crocodyliform, referred to G. simus by Dollo (1883; IRSNB R1537 and R1539; see previous sections on Goniopholis and

37 English Wealden Fossils 341 A. hooleyi for a discussion of these specimens). The most complete of the two B. fagesii specimens, IRSNB R46, was given a preliminary description by Dollo (1883). A more detailed description of the Bernissart B. fagesii material was published by Buffetaut (1975), who designated IRSNB R46 as the lectotype and IRSNB R1540 as the paralectotype. Based on an examination of the former specimen, Norell and Clark (1990) presented a revised diagnosis for B. fagesii. In the early 1980s, a third skeleton of B. fagesii was found in a Barremian clay pit in the Galve province of Teruel, central Spain (Buscalioni et al. 1984; Buscalioni 1986, 1991; Buscalioni and Sanz 1990a). Isolated teeth and other fragmentary remains referred to Bernissartia sp., cf. Bernissartia and Bernissartidae indet. have also been recorded from a range of Upper Jurassic and Lower Cretaceous localities across Western Europe. In England, material is known from the Berriasian Purbeck Limestone Group (Owen 1878, 1879a; Ensom et al. 1991, 1994; Salisbury 2002), the lower Valanginian of the Hastings Group and the Hauterivian Barremian Weald Clay Group in East Sussex, and the upper part of the Valanginian Barremian Wessex Formation on the Isle of Wight. Material from the latter three units is described below. In Spain, Bernissartia material is known from the Hauterivian Barremian El Castellar and Camarillas formations of Galve/Teruel (Estes and Sanchiz 1982; Buscalioni et al. 1984; Sanz et al. 1984), the Barremian Uña Formation of Uña/Cuenca (Brinkmann 1989, 1992) and Pio Pajarón (Winkler 1995), the upper Barremian La Huéguina Limestone Formation of Beuenche de la Sierra (Buscalioni et al. 2008), and the Barremian Aptian Artoles Formation of Vallipón and La Cantalera, Teruel (Ruiz-Omeñaca and Canudo 2001). In Portugal, material is known from the Guimarota-Strata of the Alcobaça Formation (Brinkmann 1989). In France, Bernissartia-like material is known from Cenomanian, Turonian and Santonian of Charente-Maritime, Maine-et-Loire, Indre-et-Loire and Vendée (Buffetaut and Pouit 1994), and the Kimmeridgian of Boulonnais, northern France (Cuny et al. 1991). Most recently, Bernissartia sp. has been identified in the Berriasian assemblage of the Rabekke Formation of Bornholm, Denmark (Schwarz-Wings et al. 2009). Ever since its discovery, B. fagesii has been considered to bear strongly on the origin of eusuchians, mainly because of the position of its secondary choanae, the configuration of its dorsal shield and the morphology of its vertebrae (Dollo 1883, 1887, 1909, 1914, 1922; Lydekker 1887a; Seeley 1887a; Nopcsa 1928a; Kälin 1955a, b; Buffetaut 1975; Benton and Clark 1988; Frey 1988; Buscalioni and Sanz 1990a; Norell and Clark 1990; Clark and Norell 1992; Wu and Brinkman 1993; Ortega and Buscalioni 1995; Wu et al. 1996a, b; 1997; Brochu 1997b; Russell and Wu 1998; Salisbury and Frey 2001). Detailed accounts of the numerous systematic positions that have been proposed for B. fagesii can be found in Buffetaut (1975), Norell and Clark (1990), Buscalioni and Sanz (1990a), Salisbury and Frey (2001) and Salisbury et al. (2006).

38 342 Crocodilians Bernissartia sp. indet. Text-figure Material. NHMUK R37712, a partial skeleton from the Hastings Group (most probably the Ashdown Formation) at Hastings, East Sussex (between TQ and TQ ; Owen 1851, , vol. 2; Woodward 1885, 1886; Lydekker 1888a; Buffetaut and Ford 1979; Text-fig A C); BMB , , , , isolated teeth from the Lower Weald Clay Formation at Keymer Tileworks, Burgess Hill, West Sussex (TQ ; Cook 1995; Cook and Ross 1996, fig. 9C); NHMUK R9296 and MNHN , about 40 isolated teeth from the Wessex Formation in an area mid-way between Compton Grange Chine and Hanover Point (SZ ) at beach level, and from the Unio bed on the cliff at Sudmore Point about 1.6 km further east, both on the south-west coast of the Isle of Wight (Buffetaut and Ford 1979, pl. 122, figs 1 26). Remarks. Fossil material from the English Wealden succession now recognized as belonging to Bernissartia sp. was first reported by Owen in He described and figured (Owen 1851, p. 45, pl. 15, figs 1 2) two slabs of sandstone from Hastings, at the time in the Museum of a Mr Saull, preserving the disarticulated remains of a Text-fig Bernissartia sp. indet. A C, NHMUK R37712, from the Hastings Group (probably the Ashdown Formation) at Hastings, East Sussex. A B, partial skeleton. C, close-up of an isolated tooth on the same specimen. Scale bars represent 100 mm in A B, 5 mm in C.

39 English Wealden Fossils 343 small crocodilian. Although he regarded the specimen as coming from a green-sand, Woodward (1886) later confirmed that it was from Wealden strata. As with many other specimens collected from Hastings at the time, it very likely derives from the Ashdown Formation. The larger of the two slabs, now in the collections of the NHM, contains numerous ribs, portions of both of the girdles, the fore- and hind-limbs, osteoderms and a single tooth (NHMUK 37712; Text-fig A C). The smaller slab includes the rostral part of a left mandibular ramus. The current whereabouts of the latter specimen is not known, and its precise relationship to the larger slab is unclear. Owen (1851) assigned NHMUK to Crocodilus?, but noted similarities in the form of the mandible with G. tenuidens (NHMUK 48300; now considered to be cf. Goniopholis by Salisbury 2002). He also noticed that the osteoderms lacked the spine-like articular process characteristic of Goniopholis. Owen ( , vol. 2, index, p. vi) subsequently named the specimen Crocodilus sauli. Woodward (1885) was the first to suggest that NHMUK was similar to Bernissartia; a view supported by Lydekker (1888a, p. 77). Buffetaut and Ford (1979), after identifying a single Bernissartia-type tooth among the preserved elements, assigned NHMUK to Bernissartia. Buffetaut and Ford (1979) also assigned about 40 teeth from the Wessex Formation to Bernissartia sp. indet. Isolated teeth and osteoderms assignable to Bernissartia sp. have since been found in the Lower Weald Clay Formation in the quarry at Keymer Tileworks, West Sussex (Cook 1995; Cook and Ross 1996). The complete dentition of B. fagesii is known from the lectotype (IRSNB R46). The three posteriormost teeth in both the maxilla and the dentary are low, mesiodistally elongate and completely blunt, while the next two teeth further rostrally are more rounded (Buffetaut 1975; Buffetaut and Ford 1979). The remaining teeth become progressively more conical and pointed rostrally. Buffetaut and Ford (1979) coined the term tribodont (which literally means crushing tooth ) in reference to the unusual blunt rounded teeth in the rear of the jaw of B. fagesii. In addition to B. fagesii, tribodont teeth occur in a number of Late Cretaceous and Cenozoic alligatoroids from North America and Europe (e.g. Brachychampsa spp., Albertochampsa langstoni, Stangerochampsa mccabie and Allognathosuchus spp.) now considered part of a clade known as Globidonta in reference to the possession of bulbous teeth in the rear of the jaw (Brochu 1999). Enamel ornamentation on Bernissartia-type teeth is largely restricted to the apical half of the tooth crowns, and ranges from vertical striations on the posteriormost teeth to more or less unordered wrinkles on the button-shaped ones. A smooth band is present around the tooth base. The characteristic striated/wrinkled tribodont tooth morphology of Bernissartia is not known from any other contemporaneous crocodyliform, such that teeth of this type from Late Jurassic Early Cretaceous deposits in Western Europe can be assigned to that genus with relative confidence (Buffetaut and Ford 1979; Salisbury 2002; Schwarz-Wings et al. 2009). As has been pointed out by Brinkman (1992) and

40 344 Crocodilians Schwarz-Wings et al. (2009), the teeth of Bernissartia can be distinguished from those of Theriosuchus by virtue of their lower degree of labiolingual compression, absence of distinct mesial and distal carinae, and possession of vertical or near vertical lingual striations. We therefore agree with the previous assignment of NHMUK R37712 and isolated teeth from the Lower Weald Clay and Wessex formations to Bernissartia sp. indet. Clark and Norell (1992, p. 12) raised the possibility that some of the Wealden teeth referred to Bernissartia sp. by Buffetaut and Ford (1979) could belong to Hylaeochampsa vectiana (see below). Although the posteriormost alveoli of H. vectiana are greatly enlarged relative to the condition in other neosuchians, no teeth are preserved in the holotype. However, the posterior maxillary teeth of the hylaeochampsid Iharkutosuchus makadii (Ősi et al. 2007; Ősi 2008; see section below on H. vectiana) are very unlike those of Bernissartia fagesii, with a crown that is square in occlusal view and bears multiple rows of mesiodistally aligned cusps (Ősi and Weisampel 2009); the more rostrally positioned teeth of I. makadii also appear to deviate from the typical neosuchian shape, being incisiform (Ősi and Weishampel 2009). Assuming that the teeth of H. vectiana were similar to those of I. makadii, then identification of Bernissartia sp. indet. still seems appropriate for the Wessex Formation teeth identified by Buffetaut and Ford (1979). cf. Bernissartia Material. Two isolated paravertebral osteoderms (NHMUK PV R16317 and PV R16318) and a gastral osteoderm (BMB 18423; Cook and Ross 1996, fig. 9B) from the Lower Weald Clay Formation at Keymer Tileworks, Burgess Hill, West Sussex (TQ ). Remarks. The additional occurrence of Bernissartia-like osteoderms at the Keymer Tileworks site adds further support to the likely presence of Bernissartia in the English Wealden. The dorsal shield of B. fagesii extends from the middle of the nucha to the base of the tail. In the middle of the trunk, it comprises two sagittal rows of rectangular, double-keeled osteoderms that are longer mediolaterally than they are wide. Lateral to these osteoderms there is an additional sagittal row of square, singlekeeled osteoderms, one on each side of the trunk (Dollo 1883; Buffetaut 1975; Frey 1988; Ortega and Buscalioni 1995; Salisbury and Frey 2001). The doubled-keeled osteoderms lack the articular process found in goniopholidids, atoposaurids, pholidosaurids and other advanced neosuchians, and the portion of the dorsal shield that they comprise has been interpreted as homologous to the paravertebral shield of eusuchians, and the smaller, lateral osteoderms to the accessory osteoderms of eusuchians (Frey 1988; Salisbury and Frey 2001; Salisbury et al. 2006). Buscalioni (1991) and Ortega and Buscalioni (1995) provided a preliminary account of the dermal skeleton of two crocodilians from the Early Cretaceous (late Barremian; Pérez-Moreno and Sanz 1997) Las Hoyas Konservatlagerstätte in the

41 English Wealden Fossils 345 Serranía de Cuenca, Cuenca, Spain (Museo de Cuenca LH 7287 and LH 13370). The dorsal shield of both Las Hoyas crocodilians is similar to that of B. fagesii, except that there is an additional sagittal row of accessory osteoderms. The double-keeled paravertebral osteoderms differ from those of B. fagesii in being more sharply rectangular, with considerably smaller and fewer pits (Salisbury 2001). The double-keeled osteoderms from Keymer are remarkably similar to the paravetebral osteoderms on the lectotype of B. fagesii, and their referral to Bernissartia sp. indet. seems justifiable. However, based on their isolated nature and the fact that similar osteoderms occur in the broadly coeval Las Hoyas crocodilians from Cuenca, Spain, we conclude that they should be regarded as cf. Bernissartia. The osteoderms preserved on NHMUK R37712, while rectangular in outline and lacking a sharp articular process, are exposed in internal aspect, such that any external pitting or keels are not visible. It is possible that these osteoderms come from the lateral surface of the tail, because such osteoderms are known to occur in some atoposaurids and other mesoeucrocodylians with extensive dermal skeletons. The occurrence of such osteoderms in Bernissartia fagesii is yet to be confirmed. Cook and Ross (1996, fig. 9B) assigned an isolated gastral osteoderm (BMB 18423) from the Keymer Tileworks pit to Bernissartia sp. This osteoderm has a square outline, is flat, with a well-defined articular surface and numerous round pits on the external surface. It is about 10 mm long mediolaterally and 8 mm wide anteroposteriorly. Overall, its morphology and size are consistent with the gastral osteoderms of Bernissartia fagesii. However, similar osteoderms occur in the anteriormost part of the gastral shield of numerous goniopholidids (including Goniopholis spp.), pholidosaurids and most eusuchians (the exact morphology of the gastral osteoderms in two Las Hoyas crocodilians is not known). The presence of Goniopholis in the Keymer assemblage and at other sites within the Lower Weald Clay Formation, along with the likely occurrence of an indeterminate basal eusuchian in the Hastings Group (probably the Ashdown Formation; see below) mean that there are several taxa to which this type of gastral osteoderm could be assigned. As with the paravertebral osteoderms, we therefore propose that this specimen is identified as cf. Bernissartia. Although only represented by fragmentary and isolated material, there is compelling evidence that an indeterminate species of Bernissartia is present in the Wessex and Weald sub-basins of southern England. Family incertae sedis LEIOKARINOSUCHUS gen. nov. Derivation of name. Greek, leioka/rinos or leioka/rhnos from λειο-κάρηνος, smooth-headed/bald-headed; souchos from σοῦχος crocodile; with reference to the smooth cranial table of the holotype specimen.

42 346 Crocodilians Type species. Leiokarinosuchus brookensis. Diagnosis. As for type species. Leiokarinosuchus brookensis sp. nov. Text-figures Derivation of name. Old English, brōc, small stream, and Latin, -ensis, from; with reference to Brook Bay on the Isle of Wight, where the holotype was discovered. Holotype. NHMUK 28966, a partial skull (basicranium and posterior part of the palate), the posterior part of the left and right mandibular rami, four cervical vertebrae and associated ribs, and five paravertebral osteoderms. Locality and horizon. Brook Bay, Isle of Wight (between SZ and SZ ), presumably from the Wessex Formation. Diagnosis. Leiokarinosuchus brookensis gen. et sp. nov. is distinguished from other neosuchians based on the following unique combination of characters (autapomorphies marked with an a ): dorsal surface of cranial table smooth and continuous with medial, anterolateral and posterolateral surfaces of supratemporal foramina (a); dorsal outline of supratemporal foramen square or slightly rectangular, with its long axis aligned mediolaterally (shared with Pholidosaurus spp., Anglosuchus geoffroyi, G. willetti and G. baryglypheaus); frontal and parietal fully fused (shared with many mesoeucrocodylians); diameter of occipital condyle smaller than foramen magnum (shared with A. hooleyi and cf. Anteophthalmosuchus); posteriormost maxillary and mandibular teeth slender, gently curved lingually and uniformly covered in weak basioapically aligned striations (shared with many long/slender-snouted mesoeucrocodylians); lateral surface of posterior part of angular and ventral half of surangular covered with deep, circular, closely spaced pits 7 10 mm wide, but adjacent lateral surface of posterior process of dentary smooth (shared with A. hooleyi and cf. Anteophthalmosuchus); external mandibular fenestra absent (shared with many mesoeucrocodylians); anteriormost part of paravertebral shield bi serial, with each osteoderm being slightly longer in a mediolateral direction than it is wide, lacking a sagittal keel and an articular process, with the lateral part inclined at approximately 40 degrees to the medial part (shared with B. fagesii and some eusuchians); cervical vertebrae amphicoelous. Remarks. NHMUK was originally part of the Hastings Collection, purchased by the British Museum in Other than that it comes from the Wealden of Brook, Isle of Wight (Lydekker 1888a, p. 88), little else is known about its collection history. As with other specimens from this locality, we assume that it was found at Brook Bay (between SZ and SZ ), presumably from the Wessex Formation (see section on cf. Anteophthalmosuchus, BGS GSM ). Although he never figured it, Lydekker (1887a, p. 311) provided a brief description of NHMUK and referred it to Pholidosaurus meyeri Dunker, ,

43 English Wealden Fossils 347 Text-fig Leiokarinosuchus brookensis gen. et sp. nov., NHMUK 28966, holotype. Partial skull and mandible in A B, dorsal aspect (photograph and schematic interpretation) and C D, ventral aspect (photograph and schematic interpretation). Scale bar represents 100 mm. a taxon originally based on the external and internal mould of a skull from the Obernkirchen Sandstone of the Berriasian Bückeburg Member of north-western Germany (Dunker ; Meyer 1846). Lydekker s assignment of NHMUK to P. meyeri was based soley on comparisons with Dunker s drawing (op. cit., pl. 20, p. 74). Other than this referral, English pholidosaurids are best known from the Berriasian Purbeck Limestone Group of Dorset (Owen 1878; Mansel- Pleydell 1888; Watson 1911; Andrews 1913b; Salisbury et al. 1999; Salisbury 2002) and the Bathonian Great Oolite of Oxfordshire (Owen , vol. 3). Salisbury (2002) placed all the previously recognized Purbeck pholidosaurids (Steneosaurus

44 348 Crocodilians Text-fig Leiokarinosuchus brookensis gen. et sp. nov., NHMUK 28966, holotype. Partial skull and mandible in left lateral aspect. A, photograph. B, schematic interpretation. Scale bar represents 100 mm. purbeckensis Mansel-Pleydell, 1888, Pholidosaurus decipiens Watson, 1911 and P. laevis Andrews, 1913b) in a single taxon, P. purbeckensis; an assignment we follow here. As noted by Salisbury et al. (1999) and Salisbury (2002), both pholidosaurids from the Obernkirchen Sandstone, P. meyeri and P. schaumburgensis Meyer, 1841, are remarkably similar to each other and to P. purbeckensis (P. decipiens and P. laevis in Salisbury et al. 1999). Watson (1911) also commented on the similarity between P. decipiens and P. schaumburgensis. Contrary to Andrews (1913b), the only difference between these three forms relates to the size and density of sculpture pits on the skull table, a trait that often shows a high degree of intraspecific variation among extant crocodilians. All other characteristics of the crania and postcrania associated with these taxa are virtually identical, and P. meyeri and P. schaumburgensis should be considered synonymous with P. purbeckensis. More detailed descriptive work on all three taxa is still required to confirm this, however. If this synonymy is formalized, then P. schaumburgensis Meyer, 1841 is the name that would have priority. NHMUK shows none of the characters typical of P. purbeckensis, P. schaumburgensis and P. meyeri. As noted by Clark (1986), Pholidosaurus is essentially a long/slender-snouted version of Goniopholis with reduced pitting on the cranial table. In P. purbeckensis and P. schaumburgensis, the cranial table is approximately times broader than it is long, and in dorsal aspect the lateral margins are gently convex lateral to the supratemporal foramina, but become concave posteriorly. The medial margin of the orbit is roughly level with a line that bisects the supratemporal foramen, such that the interorbital plate is relatively broad, but not as broad as in Goniopholis. The supratemporal foramina of P. purbeckensis and P. schaumburgensis are squarish in outline, and proportionately larger compared with those of Goniopholis. As in the latter, however, the dorsal surface of the cranial table is clearly demarcated from the internal surface of the supratemporal foramen (the supratemporal fossa of other authors). Ornamentation on the cranial table of P. purbeckensis and P. schaumburgensis specimens usually consists of indistinct, randomly spaced, pits and grooves 2 3 mm wide, best developed on the parietal

45 English Wealden Fossils 349 and squamosals, fading to depressions on the rostral part of the cranial table and maxillary rostrum. Unlike Goniopholis spp., there is no evidence of an interorbital ridge, and no palpebrals have ever been found with any species of Pholidosaurus, although a well-defined indentation along the medial edge of the orbit suggests that the palpebral may have been present. The type of Anglosuchus geoffroyi Owen, 1884 appears to possess a single palpebral that was accommodated in this notch (Owen , vol. 4, pl. 18, fig. 1). The preserved portion of the skull of NHMUK includes the posterior part of the basicranium and palate. Most of the surfaces are water worn and the majority of sutures are difficult to discern. Despite its poor preservation, the specimen can nonetheless be compared with other Wealden and Purbeck crocodilians. One of the most distinctive aspects of NHMUK is the morphology of the cranial table, the entire preserved portion of which is smooth and devoid of any ornamentation. While this could be the result of weathering and abrasion, there are no inconsistencies in the bone surfaces to suggest that this is the case, and well-defined pits are present on the preserved portions of the mandible, which are similarly worn (see below). This condition is distinct from all the other Wealden crocodilians, along with other mesoeucrocodylians from Late Jurassic Early Cretaceous deposits in Western Europe. Although some specimens of P. purbeckensis have reduced pitting on the cranial table (e.g. NHMUK R3414; see Salisbury 2002, text-fig. 11), this characteristic seems to be variable within the species and may relate to localized differences in diet or the ontogenetic stage of individual specimens. In addition, the medial, rostrolateral and posterior margins of the supratemporal foramina on NHMUK are essentially continuous with the dorsal surface of the cranial table. This condition is distinct from that seen in P. purbeckensis, P. schaumburgensis, P. meyeri (as best as can be determined from an external and internal mould), Goniopholis spp., A. hooleyi, cf. Anteophthalmosuchus and V. leptognathus and Anglosuchus spp., where these margins are sharply demarcated. Also distinct from G. simus, G. gracilidens, V. leptognathus, A. hooleyi and cf. Anteophthalmosuchus, the dorsal outline of each supratemporal foramen on NHMUK was most likely square or slightly rectangular, with its long axis aligned mediolaterally. The latter feature is shared with P. purbeckensis, P. schaumburgensis, P. meyeri, Anglosuchus geoffroyi, G. willetti and G. baryglyphaeus. It is unclear to us how Lydekker (1887a) was able to compare the size of the supratemporal foramen with the orbit, as neither has a complete margin. The posterior portion of the parietal and all of the supraoccipital are missing, exposing the tympanic cavity and surrounding foramina dorsally. The full extent of the cranial table relative to the supratemporal foramen cannot, therefore, be determined. The preserved portions of the parietal and squamosal indicate that the serrated suture between these two bones was oblique to the sagittal plane, rather than subparallel as in Goniopholis spp., A. hooleyi, Anteophthalmosuchus sp., Pholidosaurus spp. and V. leptognathus.

46 350 Crocodilians In common with most neosuchians, the frontoparietal suture on NHMUK is situated deep within the supratemporal foramina. Both the frontal and parietal are fully fused, unlike the condition in G. simus, G. gracilidens, G. willetti, and some specimens of P. purbeckensis and P. shaumburgensis (this feature cannot be discerned on the holotype of P. meyeri). The preserved portion of the occipital condyle on the basioccipital of NHMUK is narrower than the foramen magnum; this is also the case in A. hooleyi and cf. Anteophthalmosuchus, but not in Goniopholis spp., Pholidosaurus spp. and Anglosuchus spp. Only small portions of the right postorbital and jugal are preserved. It is not possible to determine if the postorbital bar was inset from the lateral edge of the jugal arch as it is in P. purpeckensis and P. schaumburgensis. The secondary choanae of NHMUK are similar in size, shape and position to those of A. hooleyi (NHMUK R3876) and cf. Anteophthalmosuchus (BGS GSM ). The rostral margin of each secondary choana is located just rostral to the posterior margin of the suborbital fenestra, and the posterior margins slightly posterior to it. The palatines appear to contribute only a small percentage to the rostrolateral margin of each choana, as is also the case in A. hooleyi. The pterygoidopalatine suture appears to rise acutely from the posterior end of the suborbital fenestrae, and the median septum that divides the two oval-shaped openings seems to be formed entirely by the pterygoids. Rostral to the choanae, the palatines have subparallel lateral borders with a slight bulge midway along their length. This palatal morphology is very different from that seen in P. schaumburgensis and P. purbeckensis: in these taxa, the choana forms a broad, single opening, equivalent in width to the maximum width of the palatines, and the palatines form the entire rostral and rostrolateral margins (see Owen 1878, pl. 6, fig. 2; Koken 1887, pl. 4 [33]; the nature of the secondary choanae of P. meyeri is currently not known). Although their full posterior extent is difficult to determine because of overlying matrix and glue, the pterygoid plate and ectopterygoid wings of NHMUK appear to have been well developed and rostroposteriorly elongated. Only a small portion of each maxilla is preserved, just ventral to each orbit and sitting in partial occlusion with the mandibular rami. There are four teeth in situ in the right portion of the maxilla and one in the left. All of the teeth are of similar size and shape, being slender, gently curved lingually and uniformly covered in weak, basioapically aligned striations. No clear carinae can be discerned. In these respects the teeth appear similar to those of V. leptognathus and Pholidosaurus spp., and are distinct from the more obtuse teeth that occur in the posterior part of the tooth row in Goniopholis spp. and A. hooleyi. The alveoli are closely spaced, indicating that the teeth did not interlock to any great degree. The only preserved portion of the mandible is the posterior part of each ramus, rostral to the mandibular fossa and retroarticular process, comprising the posterior part of the dentary, the rostral half of the surangular and angular, and a small portion

47 Crocodilians 351 portion of the posterior part of the splenial. The preserved portion of the splenial, dentary and the dorsal half of the surangular are smooth, while the angular and ventral half of the surangular are covered with deep, circular, closely spaced pits that are 7-10 mm wide. This pattern is similar to that on the posterior part of the mandible of A. hooleyi (NHMUK R3876; Text-fig. 24.8) and cf. Anteophthalmosuchus (BGS GSM ; Text-fig D). There is no external mandibular fenestra: this structure is also absent in G. baryglyphaeus, A. hooleyi and Theriosuchus pusillus (it is not known whether Pholidosaurus purbeckensis, P. meyeri and Anglosuchus spp. had an external mandibular fenestra). The three posteriormost alveoli are preserved on the left dentary. These are of a similar size and shape to the maxillary alveoli and contain partially erupted teeth, the apices of which appear similar to the preserved maxillary teeth. At least five, possibly six, paravertebral osteoderms are associated with NHMUK 28966, preserved just posterior to the occiput. Two are complete and preserved in articulation with each other in what was presumably a single transverse row. All the osteoderms are slightly longer mediolaterally than they are anteroposteriorly, with a trapezium-shaped outline in dorsal aspect. The external surface is uniformly covered with deep, circular, closely spaced pits, slightly smaller in size (5-7 mm) than those on the dentary and surangular. There is an angulated lateral part that is inclined at about 40 degrees to the medial part. A lateral sagittally aligned keel does not appear to be present, nor is a well-defined articular process. Based on their position and shape, these osteoderms probably derive from the anteriormost part of a biserial paravertebral shield, and in life would have been situated on the nape. The absence of an articular process distinguishes them from those of described goniopholidids, where this feature occurs in all but the anteriormost pair (best seen in IRSNB R1537, R1539 and A. hooleyi; Salisbury and Frey 2001). Also distinct from described goniopholidids is the lack of a well-defined lateral keel. Although the presumed anteriormost paravertebral osteoderms of P. purbeckensis also lack a sharp articular process and well-defined lateral sagittal keel, they are unlike those of NHMUK in that they are almost flat, with no angulation between the medial and lateral part (Salisbury and Frey 2001). The anteriormost paravertebral osteoderms of Theriosuchus pllsillus are similar to those of NHMUK R3876 in also lacking a well-defined articular process and in being nearly flat, but differ in their possession of a lateral sagittal keel. In these respects, the anteriormost paravertebral osteoderms ofnhmuk R3876 therefore appear more similar to those of advanced neosuchians such as B. fagesii and basal eusuchians, and distinct from those of other non-eusuchian mesoeucrocodylians described from Late Jurassic and Early Cretaceous deposits in Western Europe. The four cervical vertebrae associated with NHMUK R3876 are preserved in articulation at the posterior end of the palate. Some additional vertebrae may also be present adjacent to the occiput; however, the poor preservation of the specimen in this region makes their identification difficult. All of these vertebrae are

48 352 Crocodilians amphicoelous with neural arches that are proportionately higher than those described for goniopholidids and pholidosaurids. The anteriormost vertebra of the articulated series has a well-developed diapophysis and parapophysis (this portion of the body is obscured on the other vertebrae). It is not possible to determine if a hypapophysis was present. A number of cervical ribs also appear to be present, but the only one that can be commented on in any detail is that interpreted as the right cervical rib I, which is preserved at the rear right of the skull, posterior to the pterygoid. This element is flat, c. 90 mm long and mm wide, and lacks a distinct tuberculum. The overall shape is similar to that of cervical rib I of the G. simus specimens from Bernissart (IRSNB R1537 and R1539). Despite its poor preservation, NHMUK R3876 displays an interesting mélange of osteological characters that, in combination, appear to indicate that it is distinct from other Wealden crocodilians. The proportionately large supratemporal foramina and the slender, uniformly sized teeth are suggestive of a long/slender rostrum, and may have been part of the reason that Lydekker (1887a) identified the specimen as a pholidosaurid. Other than these features, any resemblance to P. meyeri, P. schaumburgensis or P. purbeckensis seems superficial. The size of the occipital condyle, pattern of pitting on the preserved portion of the mandible, absence of an external mandibular fenestra, and the position and shape of the secondary choanae are features it shares with A. hooleyi and cf. Anteophthalmosuchus. However, the unusually smooth cranial table and apparent lack of an articular process and sagitally aligned keel on the anteriormost paravertebral osteoderms indicate that it is distinct from these taxa. We acknowledge that the smooth cranial table may be the result of weathering, but in the absence of additional material suggesting otherwise interpret it as a legitimate osteological characteristic of the specimen. On the strength of this combination of features we propose that NHMUK R3876 represents a new genus and species, Leiokarinosuchus brookensis. Based on the shared apomorphic features discussed above and provenance in the Wessex Formation, we suspect that L. brookensis is closely related to Anteophthalmosuchus and, therefore, most likely to be a goniopholidid, but we refrain from assigning it to this clade pending the collection of more substantial material. EUSUCHIA Huxley, 1875 HYLAEOCHAMPSIDAE Andrews, 1913b HYLAEOCHAMPSA Owen, 1874a Hylaeochampsa vectiana Owen, 1874a Text-figure Holotype. NHMUK R177, an imperfectly preserved posterior portion of skull from a crocodilian that was once m long, lacking most of the maxillary rostrum,

49 English Wealden Fossils 353 Text-fig Hylaeochampsa vectiana (Owen, 1874a), NHMUK R177, holotype, an imperfectly preserved posterior portion of skull in A, dorsal, B, ventral, C, occipital and D, left lateral aspects. Scale bar represents 50 mm. portions of the left and right infratemporal regions, and the posterolateral parts of the palate (Owen 1874, pl. 2, figs 23 25; Lydekker 1887b; Andrews 1913b, fig. 2; Clark and Norell 1992, figs 2 9). Type locality and horizon. Brook Bay, Isle of Wight (between SZ and SZ ), presumably from the Wessex Formation. Remarks. The holotype of Hylaeochampsa vectiana was collected by William Fox some time prior to Details concerning the exact timing and location of Fox s discovery are unclear. Clark and Norell (1992, p. 2) stated that the specimen derives from the Vectis Formation, but gave no further information. The original label attached to it states that it was collected at Brook, an area now known as Brook Bay (see section on BGS GSM ). As with other crocodilian fossils collected from this part of the Isle of Wight (e.g. BGS GSM and NHMUK R176), we therefore assume that NHMUK R177 is probably from the Wessex Formation. The specimen is well preserved but most of the bones are very fragmented, making the identification of many of the sutures almost impossible. The maxillary rostrum has been distorted slightly by dorsoventral compression, and the right side

50 354 Crocodilians of the skull has been dislocated dorsally (Clark and Norell 1992). Prior to 1991, portions of the palate, otic regions, occipital area and lateral braincase were covered in matrix. As a result of these issues, there has been considerable controversy surrounding the interpretation of H. vectiana and its evolutionary significance. Owen (1874a) interpreted the large, medially positioned openings in the palate as the secondary choanae, the two rostroposteriorly oval-shaped openings lateral to these as the suborbital fenestrae, and the circular, median opening at the rear of the palate as the median Eustachian opening. Unsurprisingly, he thus considered the remarkable modifications of the palate of H. vectiana to be unique among crocodilians, comparing it to that of some squamates. Based on other characteristics of the skull (such as the size and shape of the supratemporal foramina, and the general form of the cranial table) he considered H. vectiana to compare best to long/slendersnouted thalattosuchians such as Teleosaurus, Metriorhynchus and Pelagosaurus. Andrews (1913b) was the first to recognize that the oval-shaped openings Owen (1874a) had taken to be the secondary choanae were in fact the suborbital fenestrae, such that the true secondary choana was what Owen had interpreted as the median Eustachian opening, the latter being much smaller and located further posteriorly. With the secondary choana situated well within the pterygoid, the palate of H. vectiana conformed to the pattern considered characteristic of Eusuchia, the suborder that included all extant crocodilians (Huxley 1875). But with the lateral openings still partially filled with matrix, Andrews (1913b) followed Owen (1874a) in considering them to have opened internally, thereby referring to each as a foramen. He considered each foramen to be bound almost entirely by a bifurcating transpalatine bone (ectopterygoid). Williston (1925), Nopcsa (1928a), Kälin (1955a), Romer (1956), Steel (1973) and others followed Andrew s (1913b) interpretation of the palate of H. vectiana, with the unusual form of the ectopterygoid and the associated foramina as the main defining features of the taxon. Further preparation of the skull in the early 1990s allowed Clark and Norell (1992) to provide a much needed redescription of H. vectiana, including a detailed reassessment of its palate. Removal of matrix from the ectopterygoidal foramina revealed them to be not as unusual as had long been assumed. Rather than opening beneath the orbit, a thin wall of bone roofs each opening, both of which are almost contiguous with what had previously been assumed to be the posteriormost alveoli of the maxilla. Based on these observations, Clark and Norell (1992) interpreted these openings to be confluent alveoli, presumably once housing greatly enlarged posterior maxillary teeth. The description of Iharkutosuchus makadii Ősi et al., 2007, a closely related hylaeochampsid from the Late Cretaceous Csehbánya Formation of Iharkút, western Hungary, by Ősi et al. (2007) and Ősi (2008) confirmed Clark and Norell s (1992) reinterpretation of palatal foramina of H. vectiana as enlarged alveoli. The posteriormost maxillary alveoli of I. makadii are also greatly enlarged, each housing

51 English Wealden Fossils 355 distinctive multi-cusped teeth. Each tooth has a primary mesiodistally-aligned row of cusps surrounded by many radially orientated rows of smaller cusps (see Ősi et al. 2007, fig. 3; Ősi 2008, figs 6C, 7; Ősi and Weishampel 2009). Similar teeth are also known in species of the putative basal alligatoroid Acynodon from Late Cretaceous deposits in Spain, southern France, northern Italy and Romania (Buscalioni et al. 1997; López-Martínez et al. 2001; Martin and Buffetaut 2005; Martin et al. 2006; Martin 2007; Delfino et al. 2008), which Turner and Brochu (2010) recently suggested may also be a hylaeochampsid. If this idea is correct, it would seem that enlarged, multi-cusped teeth are characteristic of hylaeochampsids, and that such teeth probably also occurred in H. vectiana. Owen (1874a) and Clark and Norell (1992) thought that the rostromedial curvature of the preserved maxillary alveoli of Hylaeochampsa vectiana indicated that the rostrum was sharply demarcated from the rear of the skull, and that it was either very short (similar to extant blunt-snouted taxa such as Osteolaemus tetrapsis Cope) or slender (similar to V. leptognathus). Although the latter type of rostrum does not occur in I. makadii or Acynodon spp., both taxa have a proportionately short rostrum. In I. makadii, the rear of the skull is considerably broader than the rostrum, with the broadening commencing just rostral to the level of the robust ectopterygoids, and dorsal to the enlarged maxillary teeth. In light of its similarly enlarged posterior maxillary alveoli, it is likely that the rostrum of H. vectiana was probably similar to that of I. makadii. In H. vectiana, however, the posterior end of the maxillary tooth row curves medially, bringing the large posterior teeth into a position directly ventral to the robust prefrontal pillar (Clark and Norell 1992). In contrast, in I. makadii the posterior end of the tooth row curves laterally, where it abuts the enlarged lateral process of the maxilla (Ősi 2008). These differences may indicate different types of food processing in both taxa, with dorsal (H. vectiana) or posterolateral (I. makadii) bracing of the enlarged posterior teeth during occlusion: crushing of harder food items in H. vectiana and grinding and smashing of softer food items using lateral jaw movements in I. makadii (Ősi 2008). Other characteristics of the rostrum of H. vectiana, originally thought to be autapomorphic by Clark and Norell (1992), also occur in I. makadii and now diagnose Hylaeochampsidae. This family was first proposed by Andrews (1913b), and following Williston (1925) was thought to encompass both H. vectiana and Heterosuchus valdensis Seeley, 1887a. However, with Clark and Norell s (1992) conclusion that Heterosuchus valdensis was either a junior synonym of H. vectiana or undiagnostic (Clark and Norell 1992; see section below on Heterosuchus), the clade became taxonomically redundant. In addition to the dental characters discussed above, H. vectiana and I. makadii have a prominent boss on the occipital surface of the paraoccipital process of the exoccipital and a large protuberance on the ventral surface of the quadrate. The latter is thought to be equivalent to muscle scar A of Iordansky (1964), and indicates a well-developed m. adductor mandibulae caudalis

52 356 Crocodilians (Ősi and Weisampel 2009). Both taxa also have an unusual lacrimal that is considerably shorter than the prefrontal. Among the suite of features that distinguish H. vectiana from I. makadii is the former s possession of a laterally exposed QSE canal (Clark and Norell 1992). A similar condition also occurs in European Goniopholis spp. (Salisbury et al. 1999; see above section on Goniopholis), Allodaposuchus precedens (Delfino et al. 2008) and Acynodon iberoccitanus (Martin 2007). While slightly different expressions of this feature now appear to have arisen multiple times among neosuchians, the presence of a laterally exposed QSE canal in Acynodon iberoccitanus adds further weight to Turner and Brochu s (2010) proposition that this genus may be a hylaeochampsid. On the basis of its pterygoid-bounded secondary choana, H. vectiana has long been thought to represent the world s oldest eusuchian (e.g. Andrews 1913b; Nopcsa 1928a; von Huene 1933; Mook 1934; Steel 1973; Buffetaut 1975). Its phylogenetic position as a basal eusuchian has been confirmed in numerous cladistic analyses (e.g. Benton and Clark 1988; Clark and Norell 1992; Wu and Brinkman 1993; Clark 1994; Salisbury and Willis 1996; Wu et al. 1996a; Brochu 1997a, b, 1999; Salisbury et al. 2006; Ősi et al. 2007; Turner and Buckley 2008), and its possession of other characters support this placement (e.g. the basisphenoid is expanded rostroposteriorly ventral to the basioccipital and is also broadly exposed ventral to the basioccipital in occipital aspect; this results in the pterygoid being dorsoventrally tall ventral to the median eustachian opening; Brochu 1999; Salisbury et al. 2006). Earlier propositions that H. vectiana evolved a eusuchian-type palate independently (e.g. Buffetaut 1975) seem unlikely. While a number of recently discovered, probable non-crocodylian eusuchians may be further away phylogenetically than H. vectiana (e.g. Isisfordia duncani Salisbury et al., 2006; but see Pol et al for an alternative interpretation of Isisfordia s phylogenetic position), its occurrence in the Wessex Formation on the Isle of Wight and likely early Barremian age (Allen and Wimbledon 1991) means that it is still the stratigraphically oldest-known eusuchian, and that the clade must have originated sometime prior to this. However, as to exactly when and where eusuchians first appeared remains unclear, with a Laurasian (Western Europe or North America) or eastern Gondwanan (Australia) point of origin being equally likely (Salisbury et al. 2006). cf. EUSUCHIA Huxley, 1875 Text-figure Material. NHMUK 36555, an articulated series of 12 procoelous cervical, thoracic, lumbar and sacral vertebrae from the Hastings Group (probably the Ashdown Formation) at Hastings, East Sussex (between TQ and TQ ; Seeley 1887a, pl. 12, fig. 7; Lydekker 1888a; Clark and Norell 1992); NHMUK 36524, an isolated procoelous caudal thoracic vertebra from Brook Bay, Isle of Wight (between

53 English Wealden Fossils 357 SZ and SZ ), presumably from the Wessex Formation (Seeley 1887a; Lydekker 1888a); NHMUK 36525, an isolated procoelous thoracic vertebra, also from Brook Bay and presumably the Wessex Formation (Seeley 1887a, Lydekker 1888a); NHMUK 36526, three articulated thoracic vertebral bodies, at least one of which is procoelous, from the Grinstead Clay Formation north of Cuckfield, West Sussex (TQ ; Seeley 1887a); NHMUK 36527, an isolated vertebra, possibly procoelous, also from this locality (Seeley 1887a; Lydekker 1888a); NHMUK 36528, an isolated thoracic vertebra from a Cuckfield Stone, again within the Grinstead Clay Formation north of Cuckfield, but the exact location is unknown (Lydekker 1888a); NHMUK R188, a disarticulated agglomeration of postcranial bones, including procoelous thoracic vertebrae, the left ilium, vertebral segments of thoracic ribs, long bones and an isolated paravertebral osteoderm from Brook Bay, Isle of Wight, and presumably the Wessex Formation (Lydekker 1888a); NHMUK R4418, an isolated procoelous thoracic vertebra from the Wadhurst Clay Formation at Peasmarsh, north of Rye, East Sussex (Lydekker 1888a); NHMUK R10080, an isolated procoelous vertebra from the Upper Weald Clay Formation at Smokejacks Brickworks pit, Ockley, Surrey (TQ ). Remarks. Procoelous crocodilian vertebrae have been collected from several horizons in the Wealden Supergroup. The best-known example is NHMUK 36555, a series of 12 articulated vertebrae from the Hastings Group (Text-fig ). This specimen was collected by Gideon Mantell and described as Heterosuchus valdensis by Seeley (1887a). Although badly water worn and obviously fragmentary, Seeley (1887a) considered that the procoelous vertebrae of NHMUK were sufficient Text-fig cf. Eusuchia, NHMUK 36555, an articulated series of 12 procoelous cervical, thoracic, lumbar and sacral vertebrae from the Hastings Group (probably the Ashdown Formation) at Hastings, East Sussex, previously assigned to Heterosuchus valdensis by Seeley (1887a). Scale bar represents 50 mm.

54 358 Crocodilians grounds upon which to name it, as no crocodilian with this type of vertebral morphology was known at the time from the British Purbeck or Wealden successions. Seeley (1887a) also proposed that a radially constricted vertebral condyle and a vertebral body that is slightly concave laterally were features that could be used to distinguish He. valdensis from other eusuchians. Other procoelous crocodilian vertebrae collected by Mantell that Seeley (1887a) assigned to He. valdensis included NHMUK and (from the Tilgate Forest ) and NHMUK (from Brook Bay on the Isle of Wight). The following year, Lydekker (1888a) referred three other specimens from the Mantell Collection in the NHMUK to He. valdensis: NHMUK 36526, R4418 and 36528, even though the latter is clearly amphicoelous. Lydekker (1888a), Kälin (1955b), von Huene (1956), Romer (1956, 1966), Steel (1973) and Carroll (1988) have all proposed that He. valdensis is a junior synonym of Hylaeochampsa vectiana. This synonymy has rested upon the occurrence of the two taxa in roughly contemporaneous beds, coupled with the likelihood that a crocodilian with the eusuchian-type palate (H. vectiana) should also have procoelous vertebrae (Buffetaut 1983; Clark and Norell 1992). However, we agree with Buffetaut (1983) and Clark and Norell (1992) in considering such a synonymy invalid, on the grounds that the respective specimens do not share any overlapping characters (one being represented by a series of vertebrae and the other by a partial cranium). We also concur with Clark and Norell (1992) in regarding NHMUK 36555, the specimen that Seeley (1887a) designated as the holotype of He. valdensis, as undiagnostic; both of the features listed by Seeley (1887a) as being diagnostic of He. valdensis are variably expressed throughout the vertebral column of most eusuchians through ontogeny. As such, He. valdensis should be regarded as a nomen dubium. Furthermore, although they both derive from the Wealden Supergroup, the Hastings Group and the Wessex Formation are not wholly contemporaneous. Each unit occurs in a different sub-basin, with the upper part of the Wessex Formation being approximately 10 myr younger than the youngest unit in the Hastings Group (i.e. the Upper Turnbridge Wells Sands Formation; Allen and Wimbledon 1991; Daley and Insole 1996; Rasnitsyn et al. 1998). Other isolated procoelous vertebrae from DB83 of Beckles Residuary Marls in the upper Lulworth Formation of the Purbeck Limestone Group, such as NHMUK (Seeley 1887a, p. 215; Clark 1986, p. 345), are at least 10 myr older than the holotype of H. vectiana (see Salisbury 2002 for the likely origin and age of the Beckles Residuary Marls ). Buffetaut (1983) also suggested that some of the isolated procoelous vertebrae referred to He. valdensis (namely NHMUK and 36525) from the Wessex Formation at Brook Bay may belong to Theriosuchus. As discussed above, isolated teeth (NHMUK R and 3697) and a partial cranial table (NHMUK R176) from Brook Bay can be referred to an indeterminate species of Theriosuchus. Given that the type species of Theriosuchus, T. pusillus, is known to possess some procoelous vertebrae, it seems reasonable to assume that the isolated procoelous vertebrae

55 English Wealden Fossils 359 from the Wessex Formation could pertain to this taxon. Nevertheless, only the anteriormost cervical vertebrae of T. pusillus are unequivocally procoelous; the anteriormost thoracic vertebrae have gently concave posterior ends, the remaining thoracic vertebrae are amphicoelous, the second sacral vertebra has a flat anterior end and a concave posterior end, and the first caudal vertebra is opisthocoelous (Salisbury and Frey 2001, pp ). Considering that NHMUK and are either thoracic vertebrae or lumbar vertebrae, Buffetaut s (1983) suggested referral of these specimens to Theriosuchus cannot be confirmed. A second possibility for the taxonomic identity of the isolated procoelous vertebrae from Brook Bay is NHMUK R188, a disarticulated agglomeration of postcranial bones, presumably belonging to a single individual, which was purchased by the British Museum in 1882, and likened to He. valdensis by Lydekker (1888a). Also from the Wessex Formation of Brook Bay, the specimen was originally identified by the British Museum as Pholidosaurus meyeri, but subsequently had its label changed to that of an Eocene dyrosaurid from Nigeria, Rhabdognathus rarus Swinton, Some of the thoracic vertebrae associated with this specimen are procoelous, and the isolated paravertebral osteoderm is square in outline, with a low keel that is slightly oblique to the sagittal plane. Both of these features are suggestive of a eusuchian, but pending additional preparation, further details concerning the morphology of this specimen and its precise identification cannot be established. Although it seems likely that, based on multiple occurrences of isolated procoelous vertebrae, eusuchians are present in the Wealden Supergroup, it cannot be established whether these specimens belong to H. vectiana or represent a second, as yet unnamed taxon. Until the collection of additional, more complete material suggests otherwise, we propose that these specimens be identified as cf. Eusuchia. NHMUK 36528, the isolated amphocoelous thoracic vertebra from the Grinstead Clay Formation at Cuckfield, is assigned to Mesoeucrocodylia indet. Other crocodilian taxa from the Wealden Supergroup Lydekker (1890a) assigned a small collection of crocodilian fossils from the Isle of Wight and Ecclesbourne, East Sussex, to Goniopholis minor. The Isle of Wight material was originally part of the Fox Collection, purchased by the British Museum in Unfortunately, no collection data were associated with any of this material, which includes about 15 elements. Identifiable bones include osteoderms from the paravertebral shield, vertebrae, a right coracoid, a right humerus and an articular, all sharing the number NHMUK R214. The Ecclesbourne material includes an isolated vertebra and a femur, both bearing the number NHMUK R608. Based on its provinance, these specimens probably derive from either the Ashdown Formation or the Wadhurst Clay Formation. Except for the osteoderms and the cervical vertebra, all the Wealden G. minor material belongs to individuals that, in life, were probably less than 1 m in length, and

56 360 Crocodilians this seems to have been the main reason for Lydekker s (1890a) referral. Salisbury et al. (1999) synonymized G. minor with G. simus. None of the specimens listed above shares characteristics that would allow it to be assigned to G. simus, and we follow Salisbury et al. (1999) in regarding it as of indeterminate origin (see section on A. hooleyi for comments on the morphology of the humerus, NHMUK R214). A possible exception might be the more complete of the two paravertebral osteoderms from the Isle of Wight (NHMUK R214). Based on the fact that it is almost square in dorsal view, and has a very deep, vertically inclined lateral part and a well-developed sagittally aligned keel between the medial and lateral parts, it probably derives from the thoracolumbar part of the shield. Pitting on this osteoderm consists of shallow, circular pits 2 3 mm wide. The keel between the medial and lateral parts extends caudally beyond the rest of the osteoderm, such that it would have overlapped the corresponding articular process to a considerable degree. There is also a second, much smaller, crescent-shaped keel on the medial part. This osteoderm is very similar to those of V. leptognathus. In light of its isolated nature, we assign this specimen to Goniopholididae cf. Vectisuchus. DISCUSSION Taxonomic composition of the Wealden crocodilian fauna The Wealden Supergroup of southern England contains one of the best-known and well-represented Mesozoic crocodilian faunas in Western Europe. Yet the full taxonomic composition of the fauna, its palaeobiogeographic relationships with other crocodilian faunas of similar age and the temporal ranges of various taxa that comprise it have been difficult to gauge. Some of the main issues that have plagued our understanding of this fauna are the antiquated nature of much of the literature in which the fossils are described, and the fact that many of the key specimens are housed in small, regional collections, which, for whatever reason, are rarely visited by researchers. Despite recent advances in our understanding of crocodilian phylogeny and evolution, many Wealden crocodilian taxa have languished in the poorly understood wastebasket as new discoveries steal the limelight. Prior to this study, the Wealden Supergroup was known to contain nine genera and eleven species of crocodilians (Table 24.1). The fauna was dominated by neosuchian mesoeucrocodylians, including four goniopholidid species (Goniopholis crassidens, G. minor, Vectisuchus leptognathus and?oweniasuchus), an atoposaurid (Theriosuchus sp. indet.), a pholidosaurid (Pholidosaurus meyeri) and an indeterminate species of the advanced neosuchian Bernissartia. At least one (Hylaeochampsa vectiana) and possibly two (Heterosuchus valdensis) basal eusuchians were also thought to be present. Some of these taxa, such as G. crassidens, G. minor,?oweniasuchus, Theriosuchus, P. meyeri and Bernissartia, are shared with other Late

57 Crocodilians 361 Jurassic-Early Cretaceous faunas in Western Europe, whereas V. leptognathus, H. vectiana and He. valdellsis are unique to the Wessex-Weald Basin. In light of this reappraisal, we conclude that the overall taxonomic fabric of the Wealden crocodilian fauna is largely unchanged (Table 24.1). Goniopholidids still dominate, both in terms of taxonomic diversity and numerical abundance, while Theriosllchus, Bernissartia and H. vectiana remain valid components of the fauna. However, many of the key specimens (e.g. BMB , NHMUK R3876 and 28966) have been reassigned to new taxa, such that G. crassidens and P. meyeri are no longer considered part of the fauna. Among goniopholidids, the only Goniopholis species that occurs in the English Wealden is G. willetti, and only the holotype (BMB ) from the Grinstead Clay Formation can confidently be assigned to this taxon. The possibility that a second Goniopholis species occurs in this formation seems likely based on BMB , but this specimen is too fragmentary to be assigned to an existing species or to form the basis of a new one. Until additional material suggests otherwise, all other specimens from the Hastings Group previously assigned to either Goniopholis or G. crassidens should be placed in Goniopholididae gen. indet. It is likely that much of this material pertains to G. willetti. GOl1iopholis willetti is distinctive among valid species of Goniopholis in its possession of a considerably more elongated and more slender rostrum, and a greater number of teeth. Although not as tubular as in other long-snouted crocodilian taxa (e.g. G. gangeticus), the elongated rostrum of G. willetti nevertheless suggests a shift within Goniopholis to a more specialized feeding behaviour, perhaps focused on smaller, faster-moving prey. The size of BMB is comparable to that of the largest G. simus specimens (e.g. NHMUK R5814), and indicates that, despite its slightly different head shape, G. willetti probably filled a similar niche (that of a 3 m+ semi-aquatic ambush predator) during deposition of the Hastings Group in the Weald Sub-basin to that of A. hooleyi in the Wessex Sub-basin. The previous recognition of G. crassidens in the Wessex Sub-basin by Lydekker (1888a) and Hooley (1907) is not supported by this study. The main specimen upon which the occurrence of G. crassidens in the Wessex Formation is based, NHMUK R3876, is here referred to a new genus and species of goniopholidid, Al1teophthalmosuchus hooleyi. IWCMS , a recently collected partial disarticulated skeleton, indicates that A. hooleyi may also be present in the Wessex Formation. Anteophthalmosuchus hooleyi differs from Goniopholis in numerous aspects of its cranial and postcranial morphology, and appears to have been the largest crocodilian in the Wessex Sub-basin. BGS GSM , a partial skull from the Wessex Formation of Brook Bay, figured by Huxley (1875), may represent a second species of Anteophthalmosuchus or a closely related taxon. Anteophthalmosllchus /woleyi shares several distinctive features with two skeletons previously assigned to 'G. simus' from contemporaneous Barremian strata in the Mons Basin at Bernissart.

58 362 Crocodilians TABLE Summary of taxonomic assignments for Wealden Supergroup crocodilians Specimen(s) Previous taxonomic assignments Taxonomic status in this study NHMUK Crocodilus (Suchosaurus) cultridens Owen, 1842 [syntypes] Theropoda, Baryonychinae (Buffetaut 2007, 2010) Goniopholis crassidens Owen, 1842 NHMUK other isolated teeth (see Lydekker 1888a for a complete list) Goniopholis crassidens Owen, 1842 (Owen , , , 1878, 1879a; Lydekker 1888a) Theropoda, Baryonychinae gen. indet. Goniopholididae gen. indet. BMB Goniopholis crassidens Owen, 1842 (Owen ) Goniopholis sp. indet. BMB Goniopholis crassidens Owen, 1842 (Hulke 1878) Goniopholis willetti sp. nov. [holotype] NHMUK R3876 Goniopholis crassidens Owen, 1842 (Hooley 1907) Anteophthalmosuchus hooleyi gen. et sp. nov. [holotype] IWCMS No previous assignment Anteophthalmosuchus hooleyi gen. et sp. nov. IWCMS No previous assignment Anteophthalmosuchus hooleyi gen. et sp. nov. Wealden Crocodile (Huxley and Etheridge 1865; Huxley 1875) SMNS Vectisuchus leptognathus Buffetaut and Hutt, 1980 [holotype] Goniopholididae cf. Anteophthalmosuchus Vectisuchus leptognathus Buffetaut and Hutt, 1980 [holotype] NHMUK R214 (paravertebral osteoderm) Goniopholis minor Koken, 1887 (Lydekker 1890a) Goniopholididae cf. Vectisuchus NHMUK R214, R608 Goniopholis minor Koken, 1887 (Lydekker 1890a) Mesoeucrocodylia indet. NHMUK R898?Oweniasuchus sp. (Lydekker 1888a) Mesoeucrocodylia indet. NHMUK R3697, R Theriosuchus (Anon. 1912) Theriosuchus sp. indet. (Buffetaut 1983) Theriosuchus sp. indet. NHMUK R176 Theriosuchus sp. indet. (Buffetaut 1983) Theriosuchus sp. indet. Bernissartia sp. indet. NHMUK R37712 Crocodilus? (Owen 1851) Crocodilus sauli Owen, [holotype] Bernissartia sp. indet. (Buffetaut and Ford 1979) NHMUK R10080 No previous assignment in a publication, but labelled as Heterosuchus valdensis Seeley, 1887a cf. Eusuchia

59 English Wealden Fossils 363 Specimen(s) Previous taxonomic assignments Taxonomic status in this study NHMUK R9296, MNHN Bernissartia sp. indet. (Buffetaut and Ford 1979) Bernissartia sp. indet. BMB , , , Bernissartia sp. (Cook 1995; Cook and Ross 1996) Bernissartia sp. indet. BMB Bernissartia sp. (Cook and Ross 1996) Neosuchia cf. Bernissartia NHMUK PV R16317, PV R16318 No previous assignment Neosuchia cf. Bernissartia NHMUK Pholidosaurus meyeri Dunker, (Lydekker 1887a) Leiokarinosuchus brookensis gen. et sp. nov. [holotype] NHMUK R177 Hylaeochampsa vectiana Owen, 1874a [holotype] Hylaeochampsa vectiana Owen, 1874a [holotype] NHMUK Heterosuchus valdensis Seeley, 1887a [holotype] nomen dubium (cf. Eusuchia) NHMUK Heterosuchus valdensis Seeley, 1887a (Seeley 1887a) cf. Eusuchia NHMUK 36525, Heterosuchus valdensis Seeley, 1887a (Seeley 1887a) cf. Eusuchia NHMUK 36526, R4418 Heterosuchus valdensis Seeley, 1887a (Seeley 1888a) cf. Eusuchia NHMUK Heterosuchus valdensis Seeley, 1887a (Lydekker 1888a) Mesoeucrocodylia indet. NHMUK R4418 Heterosuchus valdensis Seeley, 1887a (Lydekker 1888a) cf. Eusuchia NHMUK R188 cf. Heterosuchus valdensis Seeley, 1887a (Lydekker 1888a) Labelled initially as Pholidosaurus meyeri Dunker, , but subsequently changed to Rhabdognathus rarus Swinton, 1930 NHMUK R10080 No previous assignment in a publication, but labelled as Heterosuchus valdensis Seeley, 1887a cf. Eusuchia cf. Eusuchia

60 364 English Wealden Fossils It is possible that future work may demonstrate that the Bernissart specimens are congeneric with Anteophthalmosltchus, or even belong to A. hooleyi. At present, there is no evidence to indicate that Al1teophthalmosuchus occurred in the Weald Sub-basin. We agree with Buffetaut and Hutt (1980) that Vectisuchlls leptognathus is a valid goniopholidid taxon, apparently restricted to the Wessex Formation. Vectisllchus leptogl1athus recalls G. willetti in representing a departure from the typical broad-snouted goniopholidid body plan, with a slender, moderately elongated rostrum that is sharply demarcated from the posterior part of the skull. The holotype indicates a total length of c. 1.2 m, such that it was considerably smaller than adults of A. hooleyi (total length m). Its smaller size and moderately elongated, slender rostrum would have resulted in ecological separation from A. hooleyi. Material from the Isle of Wight and the Hastings region previously referred to G. minor and?oweniasllchus cannot be identified with any certainty and is placed in Mesoeucrocodylia indet. However, a single paravertebral osteoderm from the Wessex Formation is assigned to Goniopholididae cf. Vectisllchus. Our study confirms earlier reports that Theriosuchus and Bernissartia are present in the Wessex Sub-basin (Theriosuchus sp. indet and Bernissartia sp. indet. in the Wessex Formation) and the Weald Sub-basin (Bernissartia sp. indet. and Neosuchia cf. Bernissartia in the Lower Weald Clay Formation; Theriosuchus sp. indet and Bernissartia sp. indet. in the Wadhurst Clay and Ashdown formations), although both taxa are represented only by isolated teeth and postcranial elements. Theriosuchus and Bernissartia were both less than 1 m in length with highly distinctive dentitions, and probably exploited habitats differently relative to taxa such as A. hooleyi, G. willetti, V. leptognathus and L. brookensis. Compared with the larger goniopholidids, Theriosuchus and Bernissartia fossils are extremely rare, which may be a further indication that they occurred in different parts of the drainage basins, possibly further away from the areas where fossils mostly accumulated and which were presumably inhabited by the larger crocodilians, the remains of which are more abundant and typically better preserved. There is presently no evidence that pholidosaurids were present in the Wessex Weald Basin. NHMUK 28966, a partial basicranium from the Wessex Formation, shows none of the features characteristic of Pholidosaurus meyeri, the taxon to which it was assigned by Lydekker (1887 a). Despite its poor preservation, this specimen shows a suite of features not seen in any other Wealden crocodilian, so we feel justified in assigning it to a new genus and species, Leiokarinosuchus brookensis. Rather than pholidosaurids, L. brookensis seems to share several characteristics with Al1teophthalmosuchus and, thus, may be a goniophoiidid. More complete material is required to confirm this assignment. The slender, uniformly sized teeth and enlarged supratemporal foramina of L. brookensis suggest that it may have had an elongated rostrum. Regardless of whether it was a goniopholidid or not, L. brookel1- sis could (along with V. leptognathus and G. willetti) probably represent the third

61 Crocodilians 365 long-snouted crocodilian taxon in the Wealden Supergroup. The predominance of non-pholidosaurid long- and slender-snouted neosuchians in the succession may explain the absence of pholidosaurids in this fauna. Hylaeochampsa vectiana is still the only named eusuchian in the Wealden Supergroup. Although its palatal morphology and phylogenetic relationships to other basal eusuchians from the European Cretaceous are now much clearer than when it was first described by Owen (l874a), no further evidence of this crocodilian other than the holotype has surfaced in any part of the Wealden succession; it remains restricted to the Wessex Formation of the Isle of Wight. Isolated procoelous vertebrae from the Wessex, Ashdown, Wadhurst Clay, Grinstead Clay and Upper Weald Clay formations provide further evidence suggestive of eusuchians in other parts of the Wealden Supergroup, but none can be referred to H. vectiana, and the taxon to which many were previously assigned, Heterosuchus valdensis, is considered to be a nomen dubium. Assuming that its dentition was similar to closely related taxa from Eastern Europe, H. vectiana was very likely a dietary specialist, with an enhanced capacity to crush hard food items such as molluscs, crayfish and small turtles. As with other specialist crocodilians in the Wealden fauna such as Theriosuchus and Bemissartia, the rarity of H. vectial1a may reflect a more specific habitat usage within the Wessex Sub-basin than that of generalist species such as A. hooleyi. The diversity and abundance of crocodilian taxa within the Wealden Supergroup is not uniform across all formations (Table 24.2). The highest diversity (a minimum of sb:: and a maximum of 11 taxa) occurs in the Wessex Formation of the Isle of Wight. Four taxa are known from the Ashdown and Wadhurst Clay formations, three from the Lower Weald Clay Formation, and two each from the Upper Weald Clay and Vectis formations. The Wessex and Vectis formations have also produced TABLE Stratigraphic distribution of crocodilian taxa in the Wealden Supergroup Horizon Wessex Sub-basin VectisFm WessexFm Weald Sub-basin Upper Weald Clay Fm Lower Weald Clay Fm Grinstead Clay Fm Wadhurst Clay Fm AshdownFm Taxa AllteaphthalmaSllclllIs haaleyi, Goniopholididae gen. inde!. AllteaphthalmasllclllIs haaleyi, Goniopholididae cf. AnteaphthallllOsllchllS, Vectisllchus leptagllatlllls, Goniopholididae cf. VectisllclllIs, Goniopholididae gen. indet., Leiokarinosllchus brookensis, Theriosuclllls sp. indet., Bernissartia sp. indet., Hylaeochampsa vectiana, cf. Eusuchia, Mesoeucrocodylia indet. cf. Eusuchia, Goniopholididae gen. indet. Bemissartia sp. inde!., Neosuchia cf. Bernissartia, Goniopholididae gen. indet. Galliaphalis willetti, Galliaphalis sp. inde!., Goniopholididae gen. indet., cf. Eusuchia, Mesoeucrocodylia inde!. Theriasuchus sp. inde!., Bernissartia sp. inde!., Goniopholididae gen. inde!., cf. Eusuchia, Mesoeucrocodylia inde!. Theriasuclllls sp. inde!., Bemissartia sp. inde!., Goniopholididae gen. inde!., cf. Eusuchia, Mesoeucrocodylia indet.

62 366 English Wealden Fossils the best-preserved fossils, with A. hooleyi and V. leptognathus being represented by nearly complete, articulated skeletons. The only other formations to produce partial crocodilian skeletons are the Grinstead Clay Formation (material assigned to Goniopholididae gen. indet.) and possibly the Ashdown Formation (NHMUK R37712). All other formations in the Wealden Supergroup have mostly yielded isolated crocodilian elements in 'bone bed' assemblages. Differences in the diversity and abundance of taxa between the two sub-basins and within each formation undoubtedly reflect habitat differences associated with various depositional settings (Chapters 2-5) and localized taphonomic processes. All of the depositional settings associated with Wealden rocks would have provided suitable habitat and food for crocodilians. Although rare, smaller, potentially more specialized taxa such as Theriosuchus, Bernissartia and indeterminate eusuchians occur in the faunas of both sub-basins. The distribution oflarger, more generalized taxa seems to have been more basin-specific. Goniopholis willetti is thus far the only large (adults >3 m in total length) crocodilian in the Weald Sub-basin. A second, similarly sized Goniopholis species may also have occurred, but the exact head shape and likely feeding behaviour of this taxon cannot be discerned from existing material. In the Wessex Sub-basin, A. hooleyi appears to have been the dominant large crocodilian (adults >3.5 m in total length). Although the geographic ranges of many extant crocodilian taxa overlap, distinct syntopic zones, where two species share the same habitat within a zone of sympatry, are rare (Medem 1971; Webb et al. 1983; Meyer 1984; Gorzula 1987; Hollands 1987; Magnusson et al. 1987; Magnusson and Lima 1991; Kofron 1992; Ouboter 1996; Meshaka et al. 2000; Cedeno-Vazquez et al. 2008). Some sympatric species use the same or adjacent habitats at different times of the year (e.g. asynchronous breeding of Crocodylus novaegllineae Schmidt and Cr. porosus in the Sepik River in Papua New Guinea; Hollands 1987). Others tend to separate themselves ecologically and, as a rule, are likely to have slightly different feeding behaviours. Morphologically, differences in feeding behaviour among crocodilians are generally manifested in terms of overall size, head-shape or both (Meyer 1984; Brochu 2001). Most extant crocodilian faunas usually include one, possibly two, large dietary generalists, and a second or third smaller-bodied, broad-snouted form and perhaps a long-snouted dietary specialist (Meyer 1984; Brochu 2001). For instance, while the large, broadsnouted generalist Cr. porosus and the smaller, slender-snouted Cr. johnsolli Kreft are sympatric over much of their range in northern Australia (Messel et al. 1981; Webb eta!' 1983, 1987; Messel and Vorlicek 1987; Read eta!' 2004), syntopy is essentiallya wet -season phenomenon when Cr. johnsoni disperse from dry-season refugia into tidal, saline waters that are normally dominated by the highly territorial Cr. POroSllS. As water levels recede, the majority of Cr. johnsoni retreat to their preferred habitat in the upper reaches of rivers, freshwater swamps, floodplain lakes and lagoons (Webb et al. 1983). Competitive exclusion of Cr. johnstoni by Cl: porosus

63 English Wealden Fossils 367 from coastal floodplains and the lower reaches of tidal rivers is usually mediated through aggressive behaviour, as occurs in other reptiles (Schoener 1977). Even in the vast Amazon Basin, aggression and predation among crocodilian species with overlapping geographic distributions keeps sympatry to a minimum (Meyer 1984). The largest Amazonian crocodile, the large, broad-snouted generalist Cr. intermedius Graves, frequently eats the slightly smaller, broad-snouted Caiman crocodilus crocodilus, which in turn preys on blunt-snouted dwarf caimans (Paleosuchus spp. Gray) in different habitats (Medem 1971). In Central America, the geographic ranges of the morphologically similar Cr. acutus Cuvier and Cr. moreleti Duméril and Duméril overlap, but the slightly larger Cr. acutus typically occurs in coastal mainland habitats and offshore islands, while the smaller Cr. moreleti is displaced into more secluded freshwater habitats (Cedeño-Vázquez et al. 2006). Where both species occur in brackish/saline mangrove swamps in the Yucatan Peninsula of Mexico, hybridization occurs (Ceneño-Vázquez et al. 2008). In the Ganges Basin, Ross (1974) observed the long, slender-snouted, dietary specialist G. gangeticus and the broad-snouted, dietary generalist Cr. palustris Lesson in the same river, and noted aggression and avoidance but not biting. This is one of the few documented instances of aggression without predation between extant crocodilian species utilizing the same habitat. Given the thickness of each formation within the Wessex-Weald Basin and the lack of specific stratigraphic context and taphonomic data associated with many specimens (a problem that afflicts other fossil assemblages containing multiple crocodilian species: e.g. the Middle Eocene Messel and Gieseltal formations in Germany and the Miocene La Venta Formation of Columbia; see Salisbury and Willis 1996 and references therein), it is difficult to determine if any Wealden crocodilians were syntopic, or even sympatric. Assuming that the behaviour and ecology of fossil crocodilians was similar to that of their living counterparts, the presence of one large habitat generalist in each sub-basin (G. willetti in the Weald Sub-basin and A. hooleyi in the Wessex Sub-basin) may have precluded the occurrence, or at the very least, restricted the abundance of others. Similar to small-bodied taxa (Theriosuchus sp. indet. and Bernissartia sp. indet.), the occurrence of species of intermediate size such as V. leptognathus, H. vectiana and L. brookensis alongside A. hooleyi in the Wessex Sub-basin is best explained in terms of their modified cranial morphology and probable more specialized feeding behaviours, which may have separated them ecologically from the larger-bodied generalists. Palaeobiogeography The general taxonomic composition of the crocodilian fauna of the Wealden Supergroup is similar to that of other faunas which inhabited Western Europe during the Late Jurassic Early Cretaceous. Three of the genera that occur in the Wealden succession of southern England, namely Goniopholis, Bernissartia and

64 368 Crocodilians Theriosuchus (usually all three, but sometimes just two), also occur in the following localities: (1) the Berriasian Purbeck Limestone Group, Swanage, Dorset (Owen 1878, 1879a; Ensom et al. 1991, 1994; Salisbury 2002); (2) the Kimmeridgian Alcobaça Formation of Guimarota and Andrès, Portugal (Brinkmann 1989; Schwarz 2002; Schwarz and Salisbury 2005; Malafaia et al. 2006); (3) the Barremian Uña Formation of Uña/Cuenca and Pio Pajarón, Spain (Brinkmann 1989, 1992; Winkler 1995); (4) the late Barremian La Huéguina Limestone Formation of Buenache de la Sierra, Spain (Buscalioni et al. 2008); (5) the Barremian Aptian Artoles Formation, Vallipón and La Cantalera, Teruel, Spain (Ruiz-Omeñaca and Canudo 2001); (6) the Hauterivian Barremian El Castellar and Camarillas formations of Galve/Teruel, Spain (Estes and Sanchiz 1982; Buscalioni et al. 1984; Sanz et al. 1984; Buscalioni and Sanz 1987a, b; Sánchez-Hernández et al. 2007); (7) the Tithonian/Portlandian Formations gréseuses Montrougue and La Rochette II of Boulogne-sur-Mer and Wimillelle, Boulonnais, France (Sauvage 1874, 1882; Buffetaut 1986; Salisbury et al. 1999; Cuny et al. 1991); (8) marls equivalent to the Berriasian Purbeck Limestone Group at Cherves-de-Cognac, France (Pouech et al. 2006; Mazin et al. 2006, 2008; Mazin and Pouech 2008); (9) the Kimmeridgian Langenberg Formation of Langenberg/Oker, north-western Germany (Thies et al. 1997; Thies and Broschinski 2001; Karl et al. 2006); (10) the Barremian marls of Bernissart, Belgium (Dollo 1883; Buffetaut 1975; Salisbury et al. 1999); and (11) the Berriasian Rabekke Formation of Bornholm, Denmark (Schwarz-Wings et al. 2009). The occurrence of shared genera between most of these sites is based on the presence of isolated teeth and osteoderms. However, where relatively complete material occurs and distinct species have been recognized, differences emerge between the various assemblages. Goniopholis baryglyphaeus and T. guimarotae are unique to the Alcobaça Formation, G. gracilidens is unique to the Purbeck Limestone Group, and T. ibericus is unique to the Uña Formation. The only exceptions are the shared occurrence of G. crassidens, G. simus, T. pusillus and P. purbeckensis in Dorset and Cherves-de-Cognac, France. Goniopholis simus and potentially P. purbeckensis also occur in the Obernkirchen Sandstone of the Niedersachsen Basin in north-west Germany (Salisbury et al. 1999). The shared faunal content of these three basins is not surprising given their Berriasian age and relatively close proximity. The Wessex-Weald basin does not appear to share any crocodilian species with other Early Cretaceous basins in Western Europe. Goniopholis willetti (Grinstead Clay Formation), A. hooleyi, V. leptognathus, L. brookensis and H. vectiana (Wessex Formation) are endemic to the basins in which they occur. The potential referral of G. simus material from Bernissart to Anteophthalmosuchus may indicate the latter taxon also occurs in the Mons Basin. The shared occurrence of Bernissartia and the ornithopod Iguanodon bernissartensis at Bernissart and in the Wessex Formation may be a further indication of some degree of faunal continuity between the Mons Basin and Wessex Sub-basin.

65 English Wealden Fossils 369 Acknowledgements. Thanks to the following people who helped with the examination of specimens in their care: Martin Bernhards (Schaumberg-Lippischer Heimartverein E.V., Marburg, Germany); Pierre Bultynck (IRSNB); Angela Buscalioni (Departamento de Biologia, Universidad Autonoma de Madrid, Spain); Sandra Chapman (NHM); John Cooper (BMB); Mr Gicseße (Gymnasium Adolfinum, Bückeburg, Germany); Günter Kauffman (Phillips-Universität Marburg Institut für Geologie und Paläontologie, Germany); Kate Hebditch (The Dorset County Museum, Dorchester); Wolf-Dieter Heinrich (Museum für Naturkunde der Humboldt-Universität, Berlin, Germany); Hans Janke (Institut und Museum für Geologie und Paläontologie der Georg-August-Universität, Göttingen, Germany); Steve Hutt (MIWG/Dinosaur Isle); Gilbert Nee (Le musée de l Iguanodon, Bernissart, Belgium); Steve Tunnicliffe (BGS); Wann Langston Jr (Texas Memorial Museum, University of Texas at Austin, USA); Angela Milner (NHM); Trevor Price (MIWG/Dinosaur Isle); Detlev Thies (Dinosaurier-Freilichtmuseum Münchehagen, Germany); and Steve Tunnicliffe (BGS). Harry Taylor photographed all the NHM and BMB specimens for SWS. For comments on earlier versions of this chapter and discussions on Wealden crocodilians, localities and stratigraphy, we thank Eric Buffetaut (Centre National de la Recherche Scientifique, Paris, France), Alex Burton (Cambridge University), Nick Chase, Steve Hutt (MIWG/Dinosaur Isle) Dave Martill (University of Portsmouth), Jay Nair (University of Queensland, Australia) and Trevor Price (MIWG/Dinosaur Isle). Steve Salisbury gratefully acknowledges financial support from The Royal Society (Banks Alecto Fellowship Scheme), the Deutscher Akademischer Austauschdienst, the University of New South Wales (Australian Postgraduate Award) and The Linnean Society of New South Wales (Joyce W. Vickery Scientific Research Fund).

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European Arachnology 2003, Arthropoda Selecta, Special Issue, 1, * Wealden News can be found at:

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