24. crocodilians...24. crocodilians by steven w. salisbury and darren naish“having burned my...

24. CroCodilians

by s t ev e n W. s a l i s BU rY and da r r e n na i s H

“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

1985–1994)

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 proc-

ess 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, basi-

sphenoid; 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 sub-

orb, suborbital fenestra; for carot post, posterior aperture of the carotid foramen;

for eus med, median eustachian foramen; for mag, foramen magnum; for ptp, post-

temporal foramen; for sqex; squamosoexoccipital foramen; for stp, supratemporal

foramen; fos max, maxillary fossa; h, humerus; j, jugal; l, lachrimal; ls, laterosphe-

noid; 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 proc-

ess; pty, pterygoid; q, quadrate; qj, quadratojugal; rad, radius; rec ot ext, external

Crocodilians306

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 300256, the original ‘tilgate

Forest’ localities near Whiteman’s Green, and tQ 273500; Mantell, 1822, 1827;

Cuvier 1824; owen 1840–1845, 1842, 1878; Hulke 1878), and the ashdown and

Wadhurst Clay formations at Hastings, east sussex (between tQ 831095 and tQ

853105; 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 670240; ashdown Formation), Brede, east sussex (tQ

820180; several sites with exposures of the Brede Bone Bed in the Wadhurst Clay

Formation, including Hare Farm lane, tQ 832184; topley 1875; allen 1949; Benton

and spencer 1995b), and telham, near Battle, east sussex (tQ 747160, several sites

with exposures of the telham Bone Bed in the Wadhurst Clay Formation, includ-

ing Black Horse Quarry, tQ 769142; 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 323193; Young and lake 1988; Cook 1995; Cook and ross 1996),

longbrook Brickworks, thakeham, West sussex (tQ 117188), Horsham Park,

Horsham, West sussex (the ‘Horsham stone’; tQ 170300), Brightling, east sussex

(tQ 680210), Crowhurst Pit, rackwell Wood, east sussex (tQ 760124; sweeting

1925; White 1928), and Peasmarsh, Waterfall Wood, east sussex (tQ 862213; allen

1949; Benton and spencer 1995b). the most productive crocodilian-bearing local-

ity in the Upper Weald Clay Formation is the smokejacks Brickworks pit, ockley,

surrey (tQ 113375; rivett 1956; ross and Cook 1995).

Within the Wealden Group on the isle of Wight, important crocodilian fos-

sils have been recovered from the Wessex Formation and the shepherd’s Chine

Member of the vectis Formation, between Compton Grange Chine (sZ 377840)

and atherfield Point (sZ 452790) 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).

Crocodilians

THE 'GAVIAL' OF TILGATE FOREST: THE FIRST WEALDEN CROCODILIAN

307

The history of Wealden crocodilians begins around 1817-1818, 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. 48-54). 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. 161-163; 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;

Crocodilians308

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 col-

lection to the British Museum (see lyddeker 1888a for a full listing of the speci-

mens 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 dis-

tinct taxa. in his Odontography (1840–1845), 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. 1–112, pls 1–50 was published in 1840;

part 2, pp. i–xl, 113–288, pls 51–89a (excluding 62a) in 1841; and part 3, pp.

xli–lxxiv, pp. 289–655, pls 1, ii, 62a, 90–150 in 1845. 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 (1840–1845, 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

(1840–1845, 1842) assigned to G. crassidens, along with isolated vertebrae, osteo-

derms and other postcranial elements from ‘tilgate Forest’ (see lydekker 1888a), are

still regarded as pertaining to crocodilians (but see section below on issues associ-

ated with the referral of these teeth to G. crassidens).

English Wealden Fossils 309

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, 1842.

lectotype. nHMUK 3798, slab and counter-slab preserving the disarticulated post-

cranial skeleton and most of the left mandibular ramus from a single individual,

text-FiG. 24.1. 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 2512. e–H, isolated vertebrae. i, a partial paravertebral osteoderm in external aspect; nHMUK 3805. a–C and e–H from Cuvier (1824); d and i from Mantell (1827). scale bar represents 50 mm.

Crocodilians310

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, 1849–1884, 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 north-

western 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 (1840–1845, 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); num-

bers unknown (Mantell 1827, pl. 5, figs 1–2, 7, 9, 12; see lydekker 1888a, pp. 80–81

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. 70–72).


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 diag-

nosis 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 speci-

mens 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 other-

wise, 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 pos-

sible, 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. crassi

dens. Pending the description of new material from north-western Germany, the

formal synonomy of G. crassidens and G. simus would allow cranial remains assign-

able 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. crassi

dens 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 24.2–24.4

derivation of name. named in honour of edgar W. Willett, collector of the holotype.

holotype. BMB 001876, a nearly complete skull (approximately 55 cm total length;

estimated total body length 3.5 m), missing only portions of the left infratemporal

Crocodilians312

and preorbital region (jugal, quadratojugal, ectopterygoid, posterior part of the

maxilla and the articular surface of the quadrate), part of the left lacrimal, the pos-

terior 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 300256,

but the exact location is unknown), in the ‘Cuckfield stone’ of the Grinstead Clay

Formation.

text-FiG. 24.2. Goniopholis willetti sp. nov., BMB 001876, 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.

Crocodilia115 313

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 1877-1878.

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 001876 to the Geological Society in February 1878. 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 001876 to represent the first known skull of G. crassidens.

Hulke's (1878) rationale for referring BMB 001876 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 001876. Although teeth preserved on each specimen (two mandibular teeth on NHMUK 3798 and several partial maxillary and premaxillary teeth on BMB 001876) are similar and conform to the typical Goniopholis-type, this is not sufficient grounds upon which to refer BMB 001876 to G. crassidens.

BMB 001876 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,

Crocodilians314

shallower pits, which are almost absent from the premaxillae; a transversely elon-

gated 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 001876 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 001876 also shares partially fused frontals and broad inclusion of the fronto-

parietal suture within the supratemporal foramen with G. simus and G. gracilidens.

although the rostralmost extent of the maxillae on the palate of BMB 001876 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. 24.3. Goniopholis willetti sp. nov., BMB 001876, holotype skull in ventral aspect. a, photograph. B, schematic interpretation. scale bar represents 100 mm.

Crocodilians 315

TEXT-FIG. 24.4. GOlliopholis willetti sp. nov., BMB 001876, 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 001876 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 001876 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 001876 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 001876 has a well-developed palpebral that forms the medial dorsal border of the functional orbit.

316 English Wealden Fossils

Notable (but not autapomorphic) features of BMB 001876 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, 1980. 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 001876 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 001876 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 001876

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 001876 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 (1840-1845,1842,1849-1884,1878), who referred them to G. crassidel1s, also pertain to G. willeti, based on their size and provenance (see Lydekker 1888a, pp. 80-83

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 001876 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 (1849-1884, vol. 2, pI. 12, figs 1-4). This specimen


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, suggest-

ing a long, narrow rostrum consistent with that of G. willetti (in more general-

ized, broad-snouted Goniopholis spp. such as G. simus, the mandibular symphysis

extends posteriorly to a point level with the sixth dentary alveolus). owen (1849–

1884, 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 (1849–1884, vol 2, pl. 11, figs

1–2; 1878, p. 4, pl. 1, figs 1–2) described and figured a pair of partial premaxil-

lae from the Wealden of Cuckfield as G. crassidens (BMB 004430; text-fig. 24.5).

similar to G. simus and possibly G. willetti and iWCMs 2001.446 (see following

section on Anteophthalmosuchus hooleyi), this specimen shows no evidence of a pre-

maxillary fenestra. Within the nasal vestibulum there is a small, median, diamond-

shaped 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 001876 (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 004430 is not

built up such that it is confluent with the rest of the premaxillae, as occurs in BMB

001876. However, this part of BMB 004430 has been partially reconstructed with

putty, and could easily have been worn down prior to fossilization. the most obvi-

ous difference between BMB 004430, BMB 001876 and other species of Goniopholis

text-FiG. 24.5. Goniopholis sp. indet., a pair of premaxillae from the Grinstead Clay Formation at Cuckfield, West sussex; BMB 004430. scale bar represents 100 mm.


is the length of the premaxillae and premaxillary suture rostral to the naris in dorsal aspect. In BMB 004430, this suture is approximately equal to the rostroposterior length of the naris, whereas in BMB 001876 and G. Sil1111S, it is considerably shorter. Based on these observations we consider BMB 004430 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 24.6-24.10

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 3.5-4 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 2001.446, 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 350855; S. Hutt, pers. comm. 2011), Isle of Wight, collected by N. Chase; IWCMS 2005.127, 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 384836). Type locality and horizon. Atherfield Point, Isle ofWight (SZ 452790), 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 2005.127) 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


(autapomorphies marked with an ‘a’): entire cranial table, infratemporal region

(excluding the medial surface of the quadratojugal) and the entire maxillary ros-

trum 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 fron-

tal ends in a sharp point (shared with G. baryglyphaeus, G. gracilidens and irsnB

r1537); interorbital transverse ridge absent (shared with irsnB r1537); lacrimal

text-FiG. 24.6. 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.

Crocodilians320

and rostral part of jugal with distinct shallow groove extending rostrally from

the orbital border (shared with G. simus, G. willetti and numerous mesoeucroco-

dylians); 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 subcircu-

lar and almost twice the maximum diameter of the orbits (a); pterygoidopalatine

suture rises acutely from the posterior end of the suborbital fenestrae (a, but pos-

sibly shared with Leiokarinosuchus brookensis gen et sp. nov.; see below); palatines

with subparallel lateral borders, bulging laterally slightly mid-way along the subor-

bital 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

text-FiG. 24.7. Anteophthalmosuchus hooleyi gen. et sp. nov., nHMUK r3876, holotype skull in ventral 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 1904. the horizon that the block came from was

24–27 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 1906. 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

text-FiG. 24.8. Anteophthalmosuchus hooleyi gen. et sp. nov., nHMUK r3876, holotype skull in right lateral aspect. a, photograph. B, schematic interpretation. scale bar represents 200 mm.

Crocodilians322

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 pri-

marily 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, text-

fig. 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, 1849–1884, 1878, 1879a, b; lydekker 1888a;

text-FiG. 24.9. Anteophthalmosuchus hooleyi gen. et sp. nov., nHMUK r3876, holotype, por-tion of the postcranial skeleton in right lateral aspect, anterior to the right. a, photograph. B, schematic interpretation. scale bar represents 100 mm.


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. crassi

dens 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 mesoeucroco-

dylians, 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 3798. Posteriorly, on the cranial table,

the sculpture comprises relatively large, evenly spaced pits. Between the orbits, how-

ever, these pits become more rostroposteriorly elongated, quickly becoming indis-

tinct 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 addi-

tion to the absence of an interorbital transverse ridge, similarly sized, uniformly

spaced, roughly circular pits occur over most of the cranial table and maxillary ros-

trum. a similar condition also occurs on the skull of one of the ‘G. simus’ specimens

from Bernissart (irsnB r1537; salisbury et al. 1999

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.

Crocodilians324

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

text-FiG. 24.10. 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.


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 osteo-

derms that are longer in a mediolateral direction than they are wide, with a ven-

trolaterally 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 gas-

tral shield comprising interlocking, polygonal osteoderms. While initially iden-

tified 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 con-

tacts 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 pre-

frontal 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

Crocodilians326

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. 1999 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 com-

plete, 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, form-

ing 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 quad-

ratojugal) and the entire maxillary rostrum are uniformly covered in similarly sized,


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 004430; text-fig. 24.5), there is

no clear evidence for a premaxillary fenestra on the preserved portion of the pre-

maxillae of nHMUK r3876. a more complete set of premaxillae from a referred

specimen, iWCMs 2001 446, 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 sur-

face 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 2001 446 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 pre-

maxilla 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 004430, 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

Crocodilians328

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 119453, 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 approxi-

mately 75 per cent of the maximum width of the foramen magnum and approxi-

mately 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 pro-

portions 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

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. 24.13A). 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 50984 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. 80-90%, 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

stevesalisbury

Cross-Out


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 appear

ance. 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 119453 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 24.11

Material. BGS GSM 119453, 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).


locality and horizon. Brook Bay, isle of Wight (between sZ 379837 and sZ 391827),

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 illus-

trate the palatal condition ‘that lies on the boundary line between that characteris-

tic 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 119453 (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 119453 and its catalogue list-

ing (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 379837 and sZ 391827) 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

text-FiG. 24.11. cf. Anteophthalmosuchus, BGs GsM 119453, partial skull and mandible in a, dorsal, B, ventral, C, occipital and d, left lateral aspects. scale bar represents 50 mm.


(Insole et nl. 1998). Although it cannot be confirmed, we therefore assume that BGS GSM 119453 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 119453 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 119453 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 119453 being incomplete and poorly preserved, it displays a number of features that compare well with A. hooleyi. Differences such as its smaller

stevesalisbury

Inserted Text

of the skull and mandible


size and more gracile, less swollen palatines suggest that it represents a distinct

species. alternatively, such differences could be the result of ontogenetic varia-

tion, 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 119453 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


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. 24.12. 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.

Crocodilians334

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. 24.13B).

type locality and horizon. Barnes High, isle of Wight (sZ 434808), ‘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 1979. 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 sub-

sequent 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 some-

time 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 reconstruc-

tion (Buffetaut and Hutt 1980, figs 1–2, 4–5; text-fig. 24.12). 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/termi-

nal 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 sur-

face (text-fig. 24.13B). this feature appears to be unique to V. leptognathus. typical

Crocodilial15 335

t '------L----palrs lat

4-~,::":ermost

car dermost __ ;e;;;:p ,_ , lat

proc art

B car darmost mad D~--p-a-rs-m-ed---~pars lat

TEXT-FIG. 24.13. ,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 10-20 per cent longer than the hindlimbs, whereas in eusuchians they are 10-30 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

Crocodilians336

femur associated with sMns 50984 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 elon-

gated 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 goniopho-

lidids, 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 goniopholid-

ids 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. leptog

nathus 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. 24.13a). 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.


the mandible of V. leptognathus is expanded rostrally, with enlarged and con-

fluent 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 rem-

iniscent 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, 1849–1884, 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 speci-

mens 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 diminu-

tive size of the species compared with other crocodilians known at the time. owen

Crocodilians338

(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 Hateg 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 frag-

mentary 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 follow-

ing 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 rostral-

most extent of the nasals; shallow sulcus on the dorsal surface of the maxillary ros-

trum, 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 mar-

gin of the supratemporal foramen smaller than the orbit throughout ontogeny; lat-

eral 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 den-

tary 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


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 distribu-

tion 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 24.14

material. nHMUK r3697 and nHMUK r4424–31, isolated teeth from the

ashdown and Wadhurst Clay formations of west Fairlight, and possibly Brede,

east sussex (tQ 820180; 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 379837 and sZ 391827; see

section on BGs GsM 119453), 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. 24.14. 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.

Crocodilians340

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 1980. 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, 1883.

lectotype. irsnB r46, an almost complete, mounted skeleton. in addition to the

axial skeleton (which lacks only the posteriormost portion of the tail), the appen-

dicular 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 coal-

seam 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


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 mate-

rial 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).

Crocodilians342

Bernissartia sp. indet.

text-figure 24.15

material. nHMUK r37712, a partial skeleton from the Hastings Group (most

probably the ashdown Formation) at Hastings, east sussex (between tQ 831095

and tQ 853105; owen 1851, 1849–1884, vol. 2; Woodward 1885, 1886; lydekker

1888a; Buffetaut and Ford 1979; text-fig. 24.15a–C); BMB 018419, 018420, 018421,

018422, isolated teeth from the lower Weald Clay Formation at Keymer tileworks,

Burgess Hill, West sussex (tQ 323193; Cook 1995; Cook and ross 1996, fig. 9C);

nHMUK r9296 and MnHn 1978–2, about 40 isolated teeth from the Wessex

Formation in an area mid-way between Compton Grange Chine and Hanover

Point (sZ 377840) 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 1851. 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. 24.15. 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.


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. 24.15a–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 37712 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 (1849–1884, vol. 2,

index, p. vi) subsequently named the specimen Crocodilus sauli. Woodward (1885)

was the first to suggest that nHMUK 37712 was similar to Bernissartia; a view sup-

ported by lydekker (1888a, p. 77). Buffetaut and Ford (1979), after identifying a sin-

gle Bernissartia-type tooth among the preserved elements, assigned nHMUK 37712

to Bernissartia. Buffetaut and Ford (1979) also assigned about 40 teeth from the

Wessex Formation to Bernissartia sp. indet. isolated teeth and osteoderms assign-

able 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, mesio-

distally 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 ref-

erence 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 poste-

riormost 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

Crocodilians344

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 verti-

cal 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 hylaeo-

champsid Iharkutosuchus makadii (osi et al. 2007; osi 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 (osi and

Weisampel 2009); the more rostrally positioned teeth of I. makadii also appear to

deviate from the typical neosuchian shape, being incisiform (osi 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 323193).

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, single-

keeled 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, pho-

lidosaurids 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


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 rec-

tangular, with considerably smaller and fewer pits (salisbury 2001).

the double-keeled osteoderms from Keymer are remarkably similar to the par-

avetebral 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 lack-

ing 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 gas-

tral 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 eusuch-

ian 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 compel-

ling 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.

Crocodilians346

type species. Leiokarinosuchus brookensis.

diagnosis. as for type species.

Leiokarinosuchus brookensis sp. nov.


derivation of name. old english, broc, 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 verte-

brae and associated ribs, and five paravertebral osteoderms.

locality and horizon. Brook Bay, isle of Wight (between sZ 379837 and sZ 391827),

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 (autapomor-

phies 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 geof

froyi, 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 max-

illary 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. hoo

leyi 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, lack-

ing a sagittal keel and an articular process, with the lateral part inclined at approxi-

mately 40 degrees to the medial part (shared with B. fagesii and some eusuchians);

cervical vertebrae amphicoelous.

remarks. nHMUK 28966 was originally part of the Hastings Collection, purchased

by the British Museum in 1855. other than that it comes from the “Wealden of

Brook, isle of Wight” (lydekker 1888a, p. 88), little else is known about its collec-

tion history. as with other specimens from this locality, we assume that it was found

at Brook Bay (between sZ 379837 and sZ 391827), presumably from the Wessex

Formation (see section on cf. Anteophthalmosuchus, BGs GsM 119453).

although he never figured it, lydekker (1887a, p. 311) provided a brief descrip-

tion of nHMUK 28966 and referred it to Pholidosaurus meyeri dunker, 1843–1844,


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 1843–1844; Meyer 1846). lydekker’s assignment of nHMUK

28966 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 1849–1884, vol. 3). salisbury

(2002) placed all the previously recognized Purbeck pholidosaurids (Steneosaurus

text-FiG. 24.16. 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.

Crocodilians348

purbeckensis Mansel-Pleydell, 1888, Pholidosaurus decipiens Watson, 1911 and P. lae

vis 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 differ-

ence 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 formal-

ized, then P. schaumburgensis Meyer, 1841 is the name that would have priority.

nHMUK 28966 shows none of the characters typical of P. purbeckensis, P.

schaumburgensis and P. meyeri. as noted by Clark (1986), Pholidosaurus is essen-

tially a long/slender-snouted version of Goniopholis with reduced pitting on the

cranial table. in P. purbeckensis and P. schaumburgensis, the cranial table is approxi-

mately 1.4–1.5 times broader than it is long, and in dorsal aspect the lateral mar-

gins 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

text-FiG. 24.17. 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.


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

1849–1884, vol. 4, pl. 18, fig. 1).

the preserved portion of the skull of nHMUK 28966 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 28966 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 inconsisten-

cies 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 character-

istic 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

28966 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 28966 was most likely square or slightly rectangular, with its long axis

aligned mediolaterally. the latter feature is shared with P. purbeckensis, P. schaum

burgensis, 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 supratem-

poral foramen with the orbit, as neither has a complete margin. the posterior por-

tion 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 por-

tions 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.

Crocodilians350

in common with most neosuchians, the frontoparietal suture on nHMUK 28966

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 28966 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 pos-

torbital 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 28966 are similar in size, shape and posi-

tion to those of A. hooleyi (nHMUK r3876) and cf. Anteophthalmosuchus (BGs

GsM 119453). the rostral margin of each secondary choana is located just ros-

tral 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 sub-

orbital 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 28966 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

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 119453; Text-fig. 24.11D). 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 28966 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

Crocodilians352

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 15–20 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 dis-

tinct 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 occip-

ital 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 rep-

resents 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 24.18

holotype. nHMUK r177, an imperfectly preserved posterior portion of skull from

a crocodilian that was once 1.5–2 m long, lacking most of the maxillary rostrum,


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 379837 and sZ

391827), presumably from the Wessex Formation.

remarks. the holotype of Hylaeochampsa vectiana was collected by William Fox

some time prior to 1874. 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 119453). as with other crocodilian fossils collected

from this part of the isle of Wight (e.g. BGs GsM 119453 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

text-FiG. 24.18. 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.

Crocodilians354

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 cov-

ered 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 croco-

dilians, 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/slender-

snouted 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 posteri-

orly. With the secondary choana situated well within the pterygoid, the palate of

H. vectiana conformed to the pattern considered characteristic of eusuchia, the sub-

order that included all extant crocodilians (Huxley 1875). But with the lateral open-

ings 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 ‘trans-

palatine’ 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 osi et al., 2007, a closely related hylae-

ochampsid from the late Cretaceous Csehbánya Formation of iharkút, western

Hungary, by osi et al. (2007) and osi (2008) confirmed Clark and norell’s (1992)

reinterpretation of palatal ‘foramina’ of H. vectiana as enlarged alveoli. the pos-

teriormost maxillary alveoli of I. makadii are also greatly enlarged, each housing


distinctive multi-cusped teeth. each tooth has a primary mesiodistally-aligned row

of cusps surrounded by many radially orientated rows of smaller cusps (see osi et al.

2007, fig. 3; osi 2008, figs 6C, 7; osi 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 sug-

gested 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 cur-

vature 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 (osi 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 (osi 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) con-

clusion 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

Crocodilians356

(osi and Weisampel 2009). Both taxa also have an unusual lacrimal that is consider-

ably 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; osi et al. 2007; turner and Buckley 2008), and its possession of other

characters support this placement (e.g. the basisphenoid is expanded rostroposte-

riorly ventral to the basioccipital and is also broadly exposed ventral to the basi-

occipital 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 independ-

ently (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. 2009 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 eusuch-

ian, 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 24.19

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 831095 and tQ 853105; 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


sZ 379837 and sZ 391827), 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 300256; seeley 1887a); nHMUK 36527, an isolated vertebra, possi-

bly 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 113375).

remarks. Procoelous crocodilian vertebrae have been collected from several hori-

zons in the Wealden supergroup. the best-known example is nHMUK 36555, a

series of 12 articulated vertebrae from the Hastings Group (text-fig. 24.19). 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 36555 were sufficient

text-FiG. 24.19. cf. eusuchia, nHMUK 36555, an articulated series of 12 procoelous cervi-cal, 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.

Crocodilians358

grounds upon which to name it, as no crocodilian with this type of vertebral mor-

phology 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 ver-

tebrae collected by Mantell that seeley (1887a) assigned to He. valdensis included

nHMUK 36525 and 36527 (from the ‘tilgate Forest’) and nHMUK 36524 (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 syno-

nym 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 par-

tial 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 48244 (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 36524 and 36525) from the Wessex

Formation at Brook Bay may belong to Theriosuchus. as discussed above, isolated

teeth (nHMUK r4424–31 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 procoe-

lous vertebrae, it seems reasonable to assume that the isolated procoelous vertebrae


from the Wessex Formation could pertain to this taxon. nevertheless, only the ante-

riormost cervical vertebrae of T. pusillus are unequivocally procoelous; the anterior-

most 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. 103–104). Considering that nHMUK 36524 and 36525 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 mey

eri, but subsequently had its label changed to that of an eocene dyrosaurid from

nigeria, Rhabdognathus rarus swinton, 1930. some of the thoracic vertebrae associ-

ated 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 identifi-

cation cannot be established.

although it seems likely that, based on multiple occurrences of isolated procoe-

lous 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 mate-

rial was originally part of the Fox Collection, purchased by the British Museum in

1882. 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 iso-

lated 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

Crocodilians360

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 simi-

lar 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 rea-

son, are rarely visited by researchers. despite recent advances in our understand-

ing 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 gen-

era 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 indetermi-

nate 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

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 004430, 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 001876) 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 004430, 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 004430 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 2001.446, 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 119453, 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.

Crocodilians362

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spec

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niss

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(lyd

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ov. [

hol

otyp

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7 H

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[h

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cf. e

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, 188

7a (

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4418

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7a (

lyde

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Mes

oeu

croc

odyl

ia in

det.

nH

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4418

Het

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sis

seel

ey, 1

887a

(ly

dekk

er 1

888a

)cf

. eu

such

ia

nH

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188

cf. H

eter

osuc

hus

vald

ensi

s se

eley

, 188

7a (

lyde

kker

188

8a)

lab

elle

d in

itia

lly a

s P

holid

osau

rus

mey

eri d

un

ker,

1843

–184

4, b

ut

subs

equ

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ange

d to

Rha

bdog

nath

us

raru

s sw

into

n, 1

930

cf. e

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1008

0 n

o pr

evio

us

assi

gnm

ent

in a

pu

blic

atio

n, b

ut

labe

lled

as

Het

eros

uchu

s va

lden

sis

seel

ey, 1

887a

cf. e

usu

chia


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 3.5-4 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 WessexWeald 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 brooken

sis. 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

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 24.2. 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.


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


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. interme

dius 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 utiliz-

ing 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 croco-

dilians 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

Crocodilians368

Theriosuchus (usually all three, but sometimes just two), also occur in the follow-

ing 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 pres-

ence 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.


Acknowledgements. thanks to the following people who helped with the exami-

nation 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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* Wealden News can be found at: http://www.kentrigs.org.uk/wealden.html

PALAEONTOLOGICAL ASSOCIATION

FIELD GUIDE TO FOSSILS: Number 14

English Wealden fossils

Edited by

DAVID J. BATTEN

THE PALAEONTOLOGICAL ASSOCIATION

LONDON 2011

© The Palaeontological Association, 2011

ISBN 978-1-4443-6711-9 PB ISSN 0962 5321

Series Editor Philip D. Lane School of Earth Sciences and Geography

Keele University, Keele Staffordshire ST5 5BG, UK

Front cover Images taken from chapters 3, 28, 32 and 25 in this volume: cliffs at Rock-a-Nore,

Hastings, East Sussex, exposing the lower part of the Wadhurst Clay Formation at the top and sandstones of the Ashdown Formation beneath; insets from top

downwards: Pelorosaurus conybeari, caudal vertebra in posterior view; Weichselia retiClllata, rachis with attached pinnae and pinnules; Istiodactylus latidens, rostrum

tip in anterior view showing dentition and symphysial pseudotooth of dentary.

Printed in Great Britain by Henry Ling Limited at the Dorset Press, Dorchester DTl IHD

24. crocodilians...24. crocodilians by steven w. salisbury and darren naish“having burned my...

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