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Volume 10: Fossil Reptiles of Great Britain
 

Figure 1.1
Lower end of the thigh bone of Megalosaurus, from Cornwell, Oxfordshire: one of the first fossil reptile bones to be illustrated from Britain, and the oldest recorded figure of a dinosaur (from Plot, 1677).

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Figure 1.2
The skull patterns, in side view, of the major lineages of reptiles. The anapsid pattern (A) is plesiomorphic (primitive), being present also in fishes and amphibians, while the diapsid (B) and synapsid (C) patterns define two major clades of amniotes, the Diapsida (thecodontians, dinosaurs, pterosaurs, crocodiles, birds) and the Synapsida (mammal-like reptiles and mammals). The euryapsid pattern (D) may have arisen more than once, in different marine groups, and appears to be a derivative of the diapsid pattern (after Benton, 1990a).

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Figure 1.3
Cladogram of the major groups of reptiles, based on recent analyses (after Benton, 1990a).

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Figure 1.4
Evolutionary tree of the main groups of reptiles, with proposed relationships based on recent cladistic analyses, and the stratigraphical distributions based on global data. The width of the ‘spindles’ represents diversity of the groups (after Benton, 1990a).

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Figure 1.5
Table showing the classification of the major groups of reptiles, based on the cladograms summarized in Figure 1.3. Symbols: † extinct group; * paraphyletic group (after Benton, 1990a).

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Figure 1.6
(A) Map of Great Britain showing the distribution of the 50 GCR fossil reptile sites; (B) The outcrop pattern of Permian, Triassic, Jurassic, Cretaceous and Tertiary rocks in Great Britain. After Benton (1988).

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Figure 1.7
Generalized stratigraphic column showing the major British fossil reptile sites in sequence, and comparable sites elsewhere in the world.

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Figure 3.1
Map showing the distribution of Permian rocks in Great Britain. GCR Permian reptile sites: (1) Middridge; (2) Cutties Hillock; (3) Masonshaugh.

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Figure 3.2
Reptile specimens from the Late Permian Marl Slate of Middridge Quarry, County Durham: (A) Protorosaurus speneri Meyer, 1830, part of the backbone (after Hancock and Howse, 1870b); (B) Adelosaurus huxleyi (Hancock and Howse, 1870), partial skeleton (after Evans, 1988b).

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Figure 3.3
Part of the worked face at the east end of the main quarry, Cutties Hillock, showing cross-bedding. The fossil reptile remains were recovered from the foot of the cross-bedded units. (Photo: M.J. Benton.)

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Figure 3.4
Reptile specimens from the Late Permian Cutties Hillocks Sandstone Formation of Cutties Hillock Quarry, Morayshire. Skulls of (A) Gordonia; (B) Geikia; and (C) Elginia, all drawn to the same scale. After Benton and Walker (1985).

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Figure 3.5
Reptile footprints from the Late Permian Hopeman Sandstone Formation of Masonshaugh Quarry, Morayshire: (A) small prints; (B) medium prints, Chelichnus megacheirus Huxley, 1877; and (C) large prints. After Benton and Walker (1985).

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Figure 4.1
Map showing the distribution of Triassic rocks in Great Britain. GCR Triassic reptile sites: (1) Grinshill Quarries; (2) Coten End Quarry; (3) Guy’s Cliffe; (4) Sidmouth coast section; (5) Otterton Point; (6) Lossiemouth East Quarry; (7) Spynie; (8) Findrassie; (9) Bendrick Rock; (10) Aust Cliff; (11) Slickstones (Cromhall) Quarry; (12) Durdham Down; (13) Emborough Quarry; (14) Tytherington Quarry.

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Figure 4.2
The stratigraphy of the British Triassic reptile faunas. Correlations of the standard Triassic divisions and the German Triassic sequence with the British Triassic, as proposed by Hull (1869) for the ‘classical’ British succession, and by Warrington et al. (1980) for currently recognized lithostratigraphical units. Skull symbols indicate the levels of the main tetrapod faunas, and asterisks denote palynological evidence of relative age. Age dates (Ma ± 5) after Forster and Warrington (1985). From Benton et al. (1993).

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Figure 4.3
The Grinshill localities. The map is based on published maps of the British Geological Survey (BGS 1:63, 360 scale Geological Sheet 138, Wem) and on field observations by M.J.B. I. Bridge Quarries; II. working quarry (ECC Quarries Ltd).

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Figure 4.4
The operational quarry on Grinshill: view of the north face, showing the massive cross-bedded Helsby Sandstone Formation at the bottom, and the softer, more thin-bedded Tarporley Siltstone Formation above. (Photo: M.J. Benton.)

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Figure 4.5
Rhynchosaurus articeps, the only member of the Grinshill skeletal fauna: typical fossil remains (A–C) and restorations (D–H). (A) Partial skeleton lacking the tail and the limbs of the left side, in ventral view (BMNH R1237, R1238); (B) dorsal vertebrae, ribs, and right forelimb in posteroventral view (SHRBM 6); (C) pelvic region, right leg with ankle bones, presacral vertebrae 22–25, sacral vertebrae 1 and 2, and caudal vertebrae 1–8 (BATGM M20a/b); (D)–(G) restoration of the skull, based on SHRBM G132/1982 and 3 and BMNH R1236, in lateral (D), dorsal (E), ventral (F), and occipital (G) views; (H) restoration of the skeleton in lateral view in walking pose. All based on Benton (1990a).

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Figure 4.6
Typical elements of the Warwick fauna. (A) Left postero-external angle of the skull of ‘Cyclotosaurus pachygnathus’ (Cyclotosaurinae incertae sedis) in lateral view (WARMS Gz13); (B) part of the snout of ‘Stenotosaurus leptognathus’ (Stenotosaurinae incertae sedis) in palatal view (WARMS Gz38); (C) posterior portion of a left lower jaw of ‘Stenotosaurus leptognathus’ in lateral view (WARMS Gz35); (D) scattered bones of cf. Macrocnemus (Rhombopholis scutulata) (WARMS Gz10); (E)–(H), assorted remains of Rhynchosaurus brodiei: (E) anterior part of the skull in lateral view (WARMS Gz6097/ BMNH R8495), (F) anterior part of a dentary in medial view (WARMS Gz950), (G) mid-dorsal vertebra in right lateral view (WARMS Gz17), and (H) interclavicle in ventral view (WARMS Gz34); (I) right ilium of Bromsgroveia walkeri in lateral view (WARMS Gz3). After various sources; from Benton et al. (1993).

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Figure 4.7
Map of the coastal outcrop of the Otter Sandstone Formation between Sidmouth and Budleigh Salterton, Devon. The major Triassic formations are indicated, together with mean fluvial palaeoflow directions, and principal tetrapod localities. From Benton et al. (1993).

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Figure 4.8
The Otter Sandstone Formation exposed in the cliffs west of Sidmouth, view looking east. Fish and reptile remains have been found at various horizons. (Photo: M.J. Benton.)

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Figure 4.9
Larger elements of the Otter Sandstone Formation fauna of Devon. (A) Spine of an unknown vertebrate, possibly a dorsal neural spine of a ctenosauriscid archosaur (EXEMS 60/1985.88); (B) fragment of the skull roof of Mastodonsaurus lavisi in dorsal view (EXEMS 60/1985.287); (C) posterior portion of a right mandible of an unknown capitosaurid, in lateral view (EXEMS 60/1985.78); (D) incomplete skull roof of Eocyclotosaurus sp., in dorsal view (EXEMS 60/1985.72); (E)–(I) remains of Rhynchosaurus spenceri: (E) left humerus in ventral view (EXEMS 60/1985.282), (F) restored skull in right lateral view (EXEMS 60/1985.292), (G) right maxilla in ventral view (EXEMS 60/1985.292), and (H) right dentary in lingual view (BMNH R9190). (I) Vertebra of an archosaur (Bristol Univ. unnumb.); (J) the neopterygian (‘palaeonisciform’) fish Dipteronotus cyphus (EXEMS 60/1985.293). After various sources; from Benton et al. (1993).

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Figure 4.10
Smaller elements of the Otter Sandstone Formation fauna of Devon. Right dentaries (A, C) and a left maxilla (B) of a procolophonid, all in lateral view (EXEMS 60/1985.311, 3, and 154); (D) dentary fragment of an unknown small pleurodont reptile, showing pits for teeth, in lingual view; (E) tooth of ?Tanystropheus, showing small accessory cusps (EXEMS 60/1985.143); (F), (G) recurved teeth of two kinds of unknown archosaurs (Bristol Univ. unnumb.); (H) unidentified insect wing (Bristol Univ. unnumb.); (I), (J) carapaces of the conchostracan Euestheria (Bristol Univ. unnumb.). After various sources; from Benton et al. (1993).

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Figure 4.11
Imaginary scene during Mid Triassic times in Devon, based on specimens recovered from the Otter Sandstone Formation between Sidmouth and Budleigh Salterton. A scorpion (mid-foreground) contemplates a pair of procolophonids on the rocks. Opposite them, a hefty temnospondyl amphibian has spotted some palaeonisciform fishes, Dipteronotus, in the water. Two Rhynchosaurus stand in the middle distance and, behind them, a pair of rauisuchians lurk. The plants include Equisetites (horsetails) around the waterside and Voltzia, a coniferous tree. Drawn by Pam Baldaro, based on her colour painting.

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Figure 4.12
The distribution of the Permo–Triassic beds around Elgin, Morayshire. The formations are indicated by shading, and the main reptile and footprint localities are named. From Benton and Walker (1985).

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Figure 4.13
Lossiemouth East Quarry: view of heavily jointed dune cross-bedded sandstones at the eastern end of the site. Reptile skeletons were found at the base of the quarry. (Photo: M.J. Benton.)

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Figure 4.14
The reptiles of the Late Triassic Lossiemouth Sandstone Formation, near Elgin, Morayshire. Skulls of (A) Stagonolepis; (B) Hyperodapedon; (C) Ornithosuchus; (D) Erpetosuchus; (E) Leptopleuron; (F) Brachyrhinodon; (G) Scleromochlus. From Benton and Walker (1985).

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Figure 4.15
Imaginary scene at Elgin, Morayshire, in Late Triassic times, showing reconstructions of reptiles with typical Late Triassic plants. Three Hyperodapedon feed on seed-ferns in the foreground. Behind them, an Ornithosuchus runs towards the armoured Stagonolepis which is looking over its shoulder. Behind Stagonolepis, two Erpetosuchus feed on a small carcass. On the rocks in the left foreground are two Leptopleuron, a tiny Brachyrhinodon and a small bipedal dinosaur, Saltopus. To the right of the rocks is the tiny Scleromochlus at the side of the pond. In and around the pond there are horsetails, cycads and ferns, and there are tall lycopods in the distance. Based on a colour painting by Jenny Halstead. From Benton and Walker (1985).

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Figure 4.16
The main quarry at Spynie, looking northwards along the east face. Blasting has just taken place, leaving broken blocks that have, from time to time, yielded fossil reptile remains. (Photo: M.J. Benton.)

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Figure 4.17
Aerial view of a bedding plane on the foreshore at Bendrick, covered with three-toed dinosaur footprints, named Anchisauripus. Each small depression is a footprint. Width of field of view is about 5 m. (Photo: M.J. Benton.)

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Figure 4.18
The Bendrick Rock footprints. (A) Sedimentary log showing the horizon at which footprints occur; (B) the two footprint types from the locality, each of which is ascribed to Anchisauripus. After Tucker and Burchette (1977).

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Figure 4.19
The Rhaetian at Aust Cliff. (A) Geological map of the Aust Cliff area; (B) the broad anticlinal structure of Aust Cliff, showing the Lias (1); the Rhaetian (Penarth Group) (2); the ‘Tea Green Marls’ (3); and the ‘Keuper Marls’ (4). Both after Hamilton (1977).

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Figure 4.20
Aust Cliff: view on the north-eastern side of the Severn Bridge, looking south-east. The red sediments of the Mercia Mudstone Group extend about four-fifths of the way up the cliff, capped by the Penarth Group (latest Triassic). The Blue Lias of the Jurassic lies at the very top, in the vegetation line. Vertebrate remains are found in lenses of ‘Rhaetic’ Bone Bed, at the base of the Penarth Group. (Photo: G.W. Storrs.)

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Figure 4.21
Reconstruction of the Rhaetian sea floor, based on fossils from Aust Cliff and neighbouring localities. The fishes are Saurichthys, at top right, Birgeria at top left, and a hybodont shark in front of it. The marine reptiles include ichthyosaurs (mid-top) and placodonts (lower left). After Duff, McKirdy and Harley (1985).

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Figure 4.22
Map showing the distribution of Carboniferous Limestone and of tetrapod-bearing GCR fissure sites in south-west England. After Fraser (1985).

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Figure 4.23
Slickstones (Cromhall) Quarry: view of the north face, at the uppermost level, of this working site. A bone-bearing fissure is seen extending down the middle of the photograph. The fissure fills are late Triassic in age, and they occupy caves dissolved into uplifted Carboniferous marine limestones. (Photo: M.J. Benton.)

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Figure 4.24
Typical reptiles from the Late Triassic fissures in South Wales and around Bristol. Skeletal reconstructions of (A) the sphenodontid Clevosaurus; (B) the crocodilomorph Saltoposuchus; (C) the prosauropod dinosaur Thecodontosaurus; and, (D) the gliding diapsid Kuehneosaurus. After various sources; in Fraser (in press).

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Figure 4.25
Reconstruction of the faunas of the Bristol region in the late Triassic, based on fossil remains found in several fissure deposits. The prosauropod dinosaur Thecodontosaurus stands in the background, while the gliding diapsid Kuehneosaurus passes over a sphenodontid (bottom left), the possibly lizard-like Variodens (bottom middle), and the early mammal Haramiya (bottom right). After Duff, McKirdy and Harley (1985).

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Figure 4.26
Tytherington Quarry: view taken in 1981. Fissures containing Triassic sediment occur in the upper levels. (Photo: R.J.G. Savage.)p

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Figure 5.1
Map showing the distribution of Jurassic (Lower, Middle and Upper) rocks in Great Britain. GCR Jurassic reptile sites: (1) Lyme Regis; (2) Whitby; (3) Loftus; (4) Eigg; (5) New Park Quarry; (6) Stonesfield; (7) Huntsman’s Quarry; (8) Shipton-on-Cherwell Quarry; (9) Kirtlington Old Cement Works; (10) Furzy Cliff, Overcombe; (11) Smallmouth Sands; (12) Kimmeridge Bay; (13) Encombe Bay; (14) Chawley Brickpits; (15) Roswell Pits, Ely; (16) Isle of Portland; (17) Bugle Pit, Hartwell; (18) Durlston Bay.

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Figure 5.2
Summary of Jurassic stratigraphy, showing global standards and some major British formations (based on Harland et al., 1990).

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Figure 5.3
The reptile-bearing Lower Jurassic of Lyme Regis. (A) Map of the coastal section from Pinhay Bay to Charmouth, showing the major units, and indicating areas that have yielded fossil reptiles in the past. (B) The rock succession of the Blue Lias, based on the work of W.D. Lang. From House (1990).

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Figure 5.4
The Lower Lias sediments at Lyme Regis: view of the succession below Ware Cliffs, showing interbedded limestones and shales. (Photo: G.W. Storrs.)

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Figure 5.5
Typical reptiles from the Lyme Regis section. Skeletons of (A) Ichthyosaurus; (B) Plesiosaurus; (C) Dimorphodon; (D) Scelidosaurus. (A) and (B) from various sources; (C) from Padian (1984); (D) from Coombs et al. (1990).

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Figure 5.6
The reptile-bearing Lower Jurassic (Toarcian) of Whitby. (A) Map of the Upper Lias (Jet Rock Member and Alum Shale Member) exposed on the foreshore between Whitby Harbour and Saltwick Bay. (B) North-east Yorkshire with fossil reptile localities marked. The coastal outcrop of Lias rocks is stippled and the area shown in (A) is outlined. (C) The Lower Toarcian sequence at Whitby, showing ammonite zones and subzones, formations, bed numbers from Howarth (1962), and thicknesses for sections near Whitby (after Cope et al., 1980a). The terminology used by earlier authors is also indicated. From Benton and Taylor (1984), after Howarth (1962).

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Figure 5.7
The Upper Lias sediments east of Whitby at Saltwick Nab, showing the fossiliferous rocks on the wave-cut platform. (Photo: C. Little.)

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Figure 5.8
Marine reptiles from the Lower Jurassic Alum Shale Member of Whitby. (A) The crocodile Steneosaurus gracilirostris Westphal, 1961, type specimen (BMNH 14792); (B) the ichthyosaur Temnodontosaurus longirostris (Mantell, 1851), type specimen (BMNH 14566); (C) the pliosauroid plesiosaur Rhomaleosaurus cramptoni (Carte and Baily, 1863), type specimen (NMI F8785), skull and skeleton.

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Figure 5.9
The pterosaur Parapsicephalus purdoni (Newton, 1888) from the Lower Jurassic Alum Shales Member of Loftus, Yorkshire. (A), (B) and (C) skull in lateral and ventral views; (D) and (E) brain cast in left lateral and dorsal views. From Westphal (1976).

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Figure 6.1
Distribution of British Bathonian tetrapod localities. Dorset: Long Burton (1), Watton Cliff (1a), Swyre (1b); Somerset: Closworth (2); Avon: Bath (3); Wiltshire: Avoncliff (4), Bradford-on-Avon (5), Frankley (6), Box Tunnel (7), Atford (8), Malmesbury (9), Leigh Delamere (9a); Gloucestershire: Minchinhampton (10), Sapperton Tunnel (11), Avening (12), Cirencester (13), Tarlton Clay Pit (13a), Sevenhampton (14), Chedworth (15), Stanton (16), Bibury (17), Naunton (18), Kyneton Thorns (19), Huntsman’s Quarry (20), Eyeford (21), New Park Quarry (22), Oakham (23), Longborough Road Quarry (24); Oxfordshire: Chipping Norton (25), Sarsden (26), Over Norton (27), Sharp’s Hill (28), Temple Mills Quarry (29), Enstone (30), Stonesfield (31), Slape Hill (32), Glympton (33), Bladon (34), Hanborough (35), Enslow Bridge (36), Bletchingdon Station (37), Shipton Quarry (38), Kirtlington (39), Hampton Common (40), Fritwell (41), Littlemore (42), Woodeaton (43), Ardley (44), Stratton Audley (45); Buckinghamshire: Stony Stratford (46), Olney (47); Northamptonshire: Blisworth (48), Cogenhoe (49), Northampton (50), Kingsthorpe (51), Rushden (52), Thrapston (53), Ilchester (54), Oundle (55); Leicestershire: Essendine (56), Belmesthorpe (57); Cambridgeshire: Peterborough (58), Botolph’s Bridge (59), Orton Longueville, Peterborough (60), Stilton (61); Yorkshire: Scarborough (62); Hebrides: Eigg (63), Muck (64), Elgol, Skye (65a, b), Bearreraig, Skye (66). Based on information in Evans and Milner (in press), Metcalf et al. (1992), and original.

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Figure 6.2
Stratigraphy of the British Bathonian (after Cope et al., 1980b), with ages of the localities listed in Figure 6.1 indicated.

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Figure 6.3
Hugh MillerÂ’s Bone Bed on Eigg, showing collecting operations in 1972. (Photo: R.J.G. Savage.)

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Figure 6.4
Exposure of the Chipping Norton Limestone Member in New Park Quarry. Reptile bones were recovered from the top of the underlying Hook Norton Limestone Member, in the floor of the quarry. (Photo: M.J. Benton.)

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Figure 6.5
Scene in Early Bathonian times, showing a small lake in Gloucestershire surrounded by seed ferns and conifers. Fishes (Lepidotus) live in the water, and frogs (Eodiscoglossus) disport themselves around the sides. Dinosaurs include some of the earliest stegosaurs and maniraptorans (?), plated and small carnivorous dinosaurs respectively. A carcass of the large sauropod, Cetiosaurus, is rotting in the water, and Megalosaurus scavenges. Lizard-like animals, crocodiles, pterosaurs, mammals and tritylodont mammal-like reptiles complete the scene. Based on a restoration painting by Pam Baldaro, showing the scene at Hornsleasow Quarry, Gloucestershire. Reproduced with permission of the University of Bristol.

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Figure 6.6
The Stonesfield Slate mines. Map based on ground surveys and studies of historical records by Aston (1974).

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Figure 6.7
View of the entrance to an adit at Stonesfield, re-opened in 1980 in an operation funded by the Nature Conservancy Council. (Photo: W.A. Wimbledon.)

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Figure 6.8
Bathonian reptiles from Stonesfield. (A) The rhamphorhynchoid pterosaur Rhamphocephalus bucklandi (Meyer, 1832), anterior part of the lower jaw; (B) the type specimen of Megalosaurus bucklandi Meyer, 1832, a partial lower jaw, seen from the inside; (C) Stereognathus ooliticus Charlesworth, 1855, reconstructed right upper molar, showing posterior and crown views. (A) After Wellnhofer (1978); (B) after Buckland (1824); (C) after Simpson 1928).

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Figure 6.9
Exposure of the Eyford Member, or ‘Cotswolds Slates’, at Huntsman’s Quarry. Reptiles occur as isolated bones at various levels in the succession. (Photo: M.J. Benton.)

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Figure 6.10
The quarries around Shipton-on-Cherwell. Up to seven quarries (detailed in the text) appear to have yielded fossil reptiles from the White Limestone and the Forest Marble formations. Based on old Ordnance Survey maps.

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Figure 6.11
Bathonian reptiles from Shipton-on-Cherwell. (A) The crocodile Steneosaurus boutilieri Deslongchamps, 1869, skull in dorsal and ventral views; (B) the stegosaur Lexovisaurus vetustus Huene, 1910, right femur in lateral and anterior views.

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Figure 6.12
Bathonian amphibians and reptiles from Kirtlington Old Cementworks Quarry. (A) The frog Eodiscoglossus oxoniensis Evans, Milner, and Mussett, 1990, right maxilla in medial view; (B) the salamander Marmorerpeton kermacki Evans et al., 1988, atlantal centrum in ventral and anterior views; (C) the lepidosauromorph Marmoretta oxoniensis Evans, 1991, reconstructed skull in lateral and dorsal views; (D) the choristodere Cteniogenys oxoniensis Evans 1990, reconstructed skull in dorsal view; teeth of: (E) rhamphorhynchoid pterosaur; (F) pterodactyloid pterosaur; (G) goniopholidid crocodile; (H) atoposaurid crocodile; (I) fabrosaurid dinosaur; (J) megalosaurid dinosaur; (K) maniraptoran dinosaur. All after Evans and Milner (in press).

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Figure 7.1
The rapidly eroding exposure of Oxford Clay at Furzy Cliff, Overcombe. Fossil reptile bones came from the Jordan Cliff Clays, at the bottom of the sequence. (Photo: M.J. Benton.)

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Figure 7.2
The pliosaur Pliosaurus brachyspondylus (Owen, 1840), an associated skeleton found in 1889 in the Kimmeridge Clay of Roswell Pits, Ely. (A) The lower jaw in crown view; (B) tooth; (C) cervical vertebra in anterior and ventral views; (D) vertebral column, and associated elements, in dorsal view; (E) reconstructed shoulder girdle. After Tarlo (1959a).

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Figure 7.3
The ornithopod dinosaur Camptosaurus. (A) Partial restoration of the skull of C. prestwichii (Hulke, 1880) showing the known fragments of bone; (B) restored skull of the North American C. dispar Marsh, 1879; (C) restoration of the skeleton of C. prestwichii (Hulke, 1880): the bones present include parts of the skull, much of the vertebral column, forelimbs and hindlimbs. After Galton and Powell (1980).

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Figure 7.4
The Kimmeridgian of Kimmeridge Bay, showing dipping limestone and shale units, facing south. (Photo: M.J. Benton.)

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Figure 7.5
Kimmeridgian reptiles from Kimmeridge Bay, Dorset. (A) The elongate slender snout of the marine crocodile Steneosaurus megarhinus (Hulke, 1871) in ventral view; (B) lower jaw of Colymbosaurus trochanterius (Owen, 1840) in crown view; (C) skeleton of the ichthyosaur Nannopterygius (Ichthyosaurus) enthekiodon (Hulke, 1871). (A) after Hulke (1871b); (B) after Owen (1861); (C) after Hulke (1871a).

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Figure 7.6
(A) Locality map and vertical section of the Swyre Head–Chapman’s Pool Kimmeridge Clay site on the Isle of Purbeck, Dorset. The beds dip gently southwards, and the shales and mudstones are punctuated by distinctive limestone beds (‘stone bands’) which have been named. These may also be matched with the (B) tabulation of the ammonite zones of the Kimmeridgian. Abbreviations: (42) Blake’s Bed 42; (BS) Blackstone; (FS) Freshwater Steps Stone Band; (r.N) rotunda Nodules; (SB) Stone Band; (W) White Stone Band; (YL) Yellow Ledge Stone Band; (in the zonal chart, B, L, M, U and T refer to basal, lower, middle, upper and topmost parts of the zones). After Taylor and Benton (1986); based on Cope (1967, 1978); Cope et al. (1980b); Cox and Gallois (1981).

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Figure 7.7
The plesiosauroid Kimmerosaurus langhami Brown, 1981, from the Upper Kimmeridge Clay of Egmont Bay. (A) Restoration of the skull in lateral view; (B) lower jaws viewed from above. After Brown (1981).

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Figure 7.8
Some typical marine reptiles of Late Jurassic times in southern England. (A) The pliosaur Liopleurodon, one of the largest marine predators of all time at 12 m long. (B) The ichthyosaur Ichthyosaurus, which was 3–4 m long. (C) The plesiosauroid Cryptoclidus, which was 4 m long. These animals occur typically in the Oxford Clay and Kimmeridge Clay faunas. Drawn by John G. Martin, copyright City of Bristol Museums and Art Gallery.

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Figure 7.9
Nicodemus Nob on the eastern side of the Isle of Portland, showing the partly overgrown quarried cliffline. Upper parts of the Portland sequence are exposed. (Photo: M.J. Benton.)

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Figure 7.10
Broadcroft Quarries on the Isle of Portland, showing large blocks quarried for building stone. Fossil reptiles have been found in most of the inland and cliffline quarries. (Photo: M.J. Benton.)

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Figure 7.11
Turtles from the Portlandian of the Isle of Portland. (A) Plesiochelys planiceps (Owen, 1842), skull in partially restored ventral view; (B) Portlandemys mcdowelli Gaffney, 1975, partial skull in ventral view. In both cases, the toothless jaws are directed to the top, and the palate and braincase extend to the bottom. The top of both skulls is missing. After Gaffney (1975a).

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Figure 7.12
The latest Jurassic and earliest Cretaceous sequences in Durlston Bay, showing marine limestones in the northern part of the section. (Photo: J.L. Wright.)

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Figure 7.13
Cliff profiles of Durlston Bay showing the type section of the Durlston Beds (after Strahan, 1898).

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Figure 7.14
Sedimentary log of the reptile-bearing units at Durlston Bay. Bone and footprint symbols indicate fossiliferous horizons. Supplied by W.A. Wimbledon.

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Figure 7.15
A small selection of the Purbeck menagerie from Durlston Bay: turtles, sphenodontid, and lizards. (A) The skull of the cryptodire turtle Mesochelys durlstonensis Evans and Kemp, 1975, in dorsal and lateral views; (B) the skull of the cryptodire turtle Dorsetochelys delairi Evans and Kemp, 1976, in dorsal and lateral views; (C) the sphenodontid ?Homoeosaurus, partial left lower jaw; (D) the lizard Paramacellodus oweni Hoffstetter, 1967, left lower jaw in lateral and medial views; (E) the lizard Becklesius hoffstetteri (Seiffert, 1973), left lower jaw in lateral and medial views; (F) the lizard Saurillus obtusus Owen, 1854, anterior end of right lower jaw in lateral and medial views; (G) the lizard Pseudosaurillus becklesi Hoffstetter, 1967, right lower jaw in lateral and medial views; (H) the lizard Dorsetisaurus purbeckensis Hoffstetter, 1967, left lower jaw in lateral and medial views. (A) after Evans and Kemp (1975); (B) after Evans and Kemp (1976); (C) after Boulenger (1891); (D)–(H) after Estes (1983), based on various sources.

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Figure 7.16
A small selection of the Purbeck menagerie from Durlston Bay: crocodilians and dinosaurs. (A) Skull of the crocodilian Goniopholis simus Owen, 1878, in dorsal view; (B) skull of the crocodilian Nannosuchus gracilidens Owen, 1879, in dorsal view; (C) skull of the crocodilian Theriosuchus pusillus Owen, 1879, in dorsal view; (D) elongate cervical vertebra of the pterosaur Doratorhynchus validus Owen, 1870, dorsal view; (E) the ornithopod dinosaur Echinodon becklesi Owen, 1861, partially restored snout region, and detail of lower jaw; (F) jaw fragment of the theropod dinosaur Nuthetes pusillus Owen, 1854, in lateral and medial views. (A)–(C) after Joffe (1967); (D) after Howse (1986); (E) after Galton (1978); (F) after Owen (1854).

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Figure 8.1
Map showing the distribution of Cretaceous (Lower and Upper) rocks in Great Britain. GCR Cretaceous reptile sites: (1) Hastings; (2) Black Horse Quarry, Telham; (3) Hare Farm, Brede; (4) Smokejacks Pit, Ockley; (5) Brook–Atherfield Point, Isle of Wight; (6) Yaverland; (7) Wicklesham Pit, Faringdon; (8) East Wear Bay, Folkestone; (9) Culand Pits, Burham; (10) St James’s Pit, Norwich.

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Figure 8.2
Summary of Cretaceous stratigraphy, showing global stage nomenclature and some major southern British formations. Based on Harland et al. (1990).

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Figure 8.3
The Wealden of the Weald. (A) Summary stratigraphic succession, showing the relative temporal position of the bone beds; (B) map of some key Wealden reptile sites. Courtesy of E. Cook.

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Figure 8.4
The cliff at Cliff End, east of Hastings, with the Cliff End Bone Bed near the top of the section. Fossil footprints and reptile bones have been found at, and in the vicinity of this locality. (Photo: E. Cook.)

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Figure 8.5
Fossil reptile remains from the Early Cretaceous Hastings Beds of Hastings. (A) Sequence of dorsal vertebrae of the crocodilian Heterosuchus valdensis Seeley, 1887; (B) iguanodontid footprints; (C) theropod footprints from the foreshore. (A) After Seeley (1887c); (B) and (C) after Woodhams and Hines (1989).

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Figure 8.6
Excavation of a partial Iguanodon skeleton at Smokejacks Brickpit in summer, 1992. (Photo: M.J. Benton.)

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Figure 8.7
Dr Glenn Storrs consolidates an Iguanodon vertebra in Smokejacks Brickpit, part of the partial skeleton excavated in summer, 1992. (Photo: M.J. Benton.)

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Figure 8.8
Sedimentological process models for the formation of the Wealden of the Weald. (A) Arenaceous formations; (B) argillaceous formations; (C) regional setting. Uplift of the London horsts, to the north of the basin of deposition, produced an area of high relief and an extensive source of sediment (A). Braided alluvial sand plains expanded southwards from the uplands, and the lowlands supported diverse floras and faunas, including dinosaurs (A). Downfaulting and denudation of the London horsts reduced relief and the rate of sediment supply (B), and the Weald area became a brackish–freshwater lagoonal–alluvial mudplain. Again, abundant vegetation grew around the lakes, and a diverse fauna of fishes, insects and reptiles inhabited the area.

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Figure 8.9
The most famous recent British dinosaur discovery, the enigmatic theropod Baryonyx walkeri Charig and Milner, 1986 (‘Superclaw’), from the Early Cretaceous Weald Clay of Smokejacks Pit, Ockley, Surrey. (A) Skull bones as preserved; (B) normal digit, presumably from the hand; (C) the claw; (D) restoration of the skeleton; (E), imagined life appearance. (A) and (C) after Charig and Milner (1990); (B), (D) and (E) after Milner (1987).

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Figure 8.10
A reconstructed scene in the Wealden of southern England, combining elements of the faunas from the Weald and from the Isle of Wight. The small ornithopod Hypsilophodon (bottom left) looks up at its larger relative Iguanodon, just behind. A mammal and a turtle stand in the bottom right, while behind them the theropod Baryonyx prepares to eat a fish. Behind it, a small theropod runs towards a small herd of the sauropod Pelorosaurus. Based on a painting by Graham Rosewarne in Benton (1989). Reproduced with permission of Quarto Publishing plc.

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Figure 8.11
Summary sedimentary log through the Wealden beds (the Wessex and Vectis formations) of the south-western coast of the Isle of Wight between Sudmoor Point and Atherfield Point. Known reptile bone-bearing horizons are noted (R), as are footprint beds (F), and the numbers 7–12 match those used in the text in the locality descriptions. After Stewart (1981b).

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Figure 8.12
Maps and diagrammatic cliff views of the coastal section from Compton Chine to Atherfield Point, on the south-western coast of the Isle of Wight. Fossil reptile localities are indicated as 1–11, corresponding to the sites described in the text. After Stewart (1981b).

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Figure 8.13
The Hypsilophodon Bed at Cowleaze Chine, high in the Wealden sequence. Stephen Hutt points to the horizon from which several complete skeletons of Hypsilophodon have been excavated. (Photo: M.J. Benton.)

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Figure 8.14
Typical non-dinosaurian reptiles from the Early Cretaceous Wealden of the south-western coast of the Isle of Wight. (A) The crocodilian Goniopholis crassidens Owen, 1841, skull in dorsal view; (B) the crocodilian Vectisuchus leptognathus Buffetaut and Hutt, 1980, restored skull and lower jaws in dorsal view; (C) teeth of the crocodilian Bernissartia sp., in crown and side views; (D) the crocodilian Hylaeochampsa vectiana Owen, 1874, skull in dorsal and ventral views; (E) the pterosaur Ornithodesmus latidens Seeley, 1901, restoration of skull, humerus, wrist, and femur. (A) After Hooley (1907); (B) after Buffetaut and Hutt (1980); (C) after Buffetaut and Ford (1979); (D) after Clark and Norell (1992); (E) after Wellnhofer (1978), based on several sources.

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Figure 8.15
Typical dinosaurs from the Early Cretaceous Wealden of the south-western coast of the Isle of Wight. (A) The theropod dinosaur Calamospondylus oweni Fox, 1866, sacrum and pubis in dorsal and lateral views; (B) the theropod dinosaur Thecospondylus horneri Seeley, 1882, natural cast of the sacral cavity; (C) the sauropod dinosaur Pelorosaurus hulkei (Seeley, 1870), a dorsal vertebra in anterior view, a coracoid, and the pubis and ischium; (D) the large ornithopod Iguanodon atherfieldensis Hooley, 1925, skull and skeleton; (E) the large ornithopod Iguanodon bernissartensis Boulenger, 1881, skull and skeleton; (F) the small ornithopod Hypsilophodon foxii Huxley, 1869, skull and restored skeleton; (G) the ankylosaur Polacanthus foxi Hulke, 1881, skeleton. (A) After Seeley (1887c); (B) after Seeley (1882a); (C) after Hulke (1880b, 1882d), Seeley (1882); (D) and (E) after Norman (1980, 1986); (F) after Galton (1974); (G) after Blows (1987).

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Figure 8.16
The skull cap of the oldest pachycephalosaurid, Yaverlandia bitholos Galton, 1971, from the Early Cretaceous Wessex Formation of Yaverland, Isle of Wight, in (A) dorsal and (B) lateral views. Skull of the Late Cretaceous pachycephalosaurid Stegoceras in (C) dorsal and (D) lateral views, for comparison. Abbreviations: f, frontal; n, nasal; p, parietal; pf, postfrontal; po, postorbital; sq, squamosal. After Galton (1971c).

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Figure 8.17
The Gault clays at East Wear Bay, Folkestone. (Photo: D.J. Ward.)

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Figure 8.18
Reptiles from the mid-Cretaceous Gault of Folkestone. (A) The elasmosaur Mauisaurus gardneri Seeley, 1877, tooth and dorsal vertebra; (B) the pterosaur Ornithocheirus daviesi (Owen, 1874), top of snout in medial and ventral views. (A) After Seeley (1877); (B) after Owen (1874a).

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Figure 8.19
The rather overgrown Upper Culand Pit, Burham, showing the Middle and Upper Chalk, source of several specimens of fossil turtles, marine lizards, pterosaurs and plesiosaurs. (Photo: M.J. Benton.)

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Figure 8.20
Typical reptiles of the Late Cretaceous Middle Chalk of the Culand Pits, Burham, Kent. (A) and (B) The turtle Chelone (Cimolochelys) benstedi (Mantell, 1841), carapace in dorsal and ventral views; (C) the elongate marine lizard Dolichosaurus longicollis Owen, 1850, crushed skull and anterior part of skeleton; (D) the pterosaur Ornithocheirus compressirostris (Owen, 1851), skull in lateral view; (E) O. cuvieri (Bowerbank, 1851), anterior part of snout in lateral and crown views; (F) O. giganteus (Bowerbank, 1846), anterior part of snout, right and left sides. (A)–(C) After Owen (1851b); (D)–(F) after Wellnhofer (1978), from various sources.

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Figure 9.1
Map showing the distribution of Tertiary rocks in Great Britain. Only major divisions are indicated, and an enlargement of the Hampshire and Isle of Wight areas is given. GCR Tertiary reptile sites: (1) Warden Point; (2) Barton Cliff; (3) Hordle Cliff; (4) Headon Hill and Totland Bay; (5) Bouldnor and Hamstead Cliffs.

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Figure 9.2
Summary of Tertiary stratigraphy, showing global standards and some major British formations. Based on Curry et al. (1978).

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Figure 9.3
The London Clay, exposed at Warden Point, Isle of Sheppey, showing collapsed cliffs and fossil-bearing material on the foreshore. (Photo: D.J. Ward.)

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Figure 9.4
Typical reptiles of the Eocene London Clay of Sheppey. (A)–(D) The turtle Puppigerus camperi (Gray, 1831), skull in (A) dorsal and (B) ventral views, (C) carapace in dorsal view, (D) plastron in ventral view; (E) Chrysemys bicarinata (Bell, 1849), partial carapace in dorsal view; (F) Platemys bullocki Owen, 1841, plastron in ventral view; (G) and (H) the crocodile Crocodilus spenceri Buckland, 1837, skull in (G) dorsal and (H) lateral views; (I) the snake Palaeophis toliapicus Owen, 1841, thirty dorsal vertebrae in side view. (A)–(F) After Owen and Bell (1849); (G) and (H) after Owen (1850b); (I) after Owen (1850c).

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Figure 9.5
The turtle Argillochelys athersuchi Moody, 1980, from the Late Eocene of Barton Cliff, partial skull in (A) lateral and (B) ventral views. After Moody (1980b).

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Figure 9.6
The Lower Headon Beds of Hordle Cliff, looking towards Becton Bunny. (Photo: D.L. Harrison.)

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Figure 9.7
Typical reptiles of the Late Eocene Lower Headon Beds of Hordle Cliff. (A) the turtle Trionyx henrici Owen, 1849, carapace in dorsal view; (B) The turtle Ocadia crassa (Owen, 1849), internal views of plastron elements; (C) and (D) the crocodilian Diplocynodon hantoniensis (Wood, 1844), skull in (C) dorsal and (D) lateral views; (E) the snake Paleryx rhombifer Owen, 1850, mid-body vertebra in posterior, anterior, lateral, and dorsal views. (A) and (B) After Owen and Bell (1849); (C) and (D) after Owen (1850b); (E) after Owen (1850c).

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Figure 9.8
Alum Cliff, at the southern end of the Headon Beds outcrop on Headon Hill, Isle of Wight. (Photo: M.J. Benton.)

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Figure 9.9
Typical reptiles and amphibians of the Late Eocene Lower Headon Beds of Headon Hill and Totland Bay. (A) A palaeobatrachid frog, fragmentary atlas; (B) the limbless lizard Ophisaurus sp., scute and trunk vertebra in ventral view; (C) the snake Paleryx rhombifer Owen, 1850, mid-trunk vertebra in dorsal, lateral, and anterior views; (D) the snake Vectophis wardi Rage and Ford, 1980, mid-trunk vertebra in dorsal, lateral, and anterior views. (A), (C) and (D) After Rage and Ford (1980); (B) after Meszoely and Ford (1976).

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