Figures

(Figure 1.1) Sketch map of the principal outcrops of the main stratigraphical units in the British Isles. The pre-Devonian formations, both sedimentary and crystalline, comprise the Precambrian and the three Lower Palaeozoic Systems. The Devonian rocks include both marine and continental (Old Red Sandstone). The Cenozoic, or Tertiary, sedimentary formations are those of the Hampshire Basin, the London Basin and East Anglia. The flood basalts of the North-west Tertiary Igneous Province are marked, but the many intrusive igneous rocks and various older volcanics are omitted for the purposes of this discussion.

(Figure 1.2) A possible scenario of the evolution of the British Isles. (A) Fusion of the continental crustal blocks Laurentia, Avalonia and Baltica in the early to middle Palaeozoic cycle of events. (B) Evolution of middle to late Palaeozoic basins and orogenic uplifts. (C) Mesozoic basin and upland developments. (D) Late Mesozoic-Cenozoic (Alpine) cycle — Atlantic rifting.

(Figure 1.3) A phylogenetic scheme of chordates within the stratigraphical record (after Blieck, 1992). The two known cephalochordates of the Cambrian are Pikaia gracilis from the Middle Cambrian Burgess Shale of Canada and Yunnanozoon lividum from a Chinese Early Cambrian fauna (c. 525 Ma). Thelodonts, as yet unde-scribed, are known from the Ordovician. Acquisition of vertebrate characters: (1) chordate features (mesoderm, notochord, etc.); (2) somitic characters (somites, creatine phosphate, etc.); (3) craniate features (neural crest, cartilaginous endoskeleton); (4) two semicircular canals, dermal ossification, etc. (5) aspidin present; (6) same four-layered structure of dermal bone; (7) paired fins, etc.; (8) gill openings in a slanted line; (9) cellular dermal bone, perichondral bone etc.; (10) heterocercal tail with change of scale ornamentation at caudal peduncle, pectoral tins, concentrated at base, etc. For full discussion see Blieck (1992), also Chen et al. (1995). The Australian group the pituriaspids, the most recently discovered and puzzling, seems to be related to the basic stock from which also came the Chinese galeaspids and the Euramerican osteostracans (see Long, 1995).

(Figure 1.4) A geological history of fishes. The geological periods and the ages of the period boundaries are given at the top. The shaded areas suggest the relative abundance of species within the different classes during each period (after Ommaney, 1963, with data from Benton, 1993).

(Figure 1.5) The Agnatha; fishes lacking jaws. (A) Anglaspis, a heterostracan from the Siluro-Devonian; (B) Pteraspis a heterostracan from the Early Devonian; (C) Hemicyclaspis, an osteostracan from the Late Silurian; (D) Psammolepis, a Late Devonian heterostracan; (E)–(G) Silurian thelodonts, Thelodus, Phlebodus and an unnamed form from Canada; (H) the Devonian anaspid Pharyngolepis; (I) a living lamprey ammocoete larva; (J) an adult living lamprey; (K) the extant adult hagfish. Not shown are the Chinese galeaspid and Australian pituriaspid. All are approximately a third natural size.

(Figure 1.6) Gnathostomata or jaw-bearing fishes. (A) an acanthodian, Parexus, at x 0.5; (B) the antiarch Remigolepis, at x 0.3; (C) the arthrodire Coccosteus, at x 0.3; (D) and (E) fish with bony skeletons: (D) a primitive actinopterygian, at x 0.5; (E) a sarcopterygian, at x 0.1; (F) fish with a cartilaginous skeleton, a hybodont shark, at x 0.1.

(Figure 1.7) Age ranges of Palaeozoic fishes (after Long, 1993).

(Figure 1.8) Cladograms showing the postulated relationships of the main groups of fishes (after Benton, 1993). (A) The Chondrichthyan fishes. Chondrichthyes = cartilaginous endoskeleton with exoskeleton of small scales, sometimes enlarged into head spines or fin spines. Elasmobranchi = a predaceous group with distinctive jaw suspension and quickly replaced characteristic tooth structure; no operculum, gill slits opening directly to the outside. Neoselachii = modern sharks, skates and rays; characteristic vertebrate and fin structures. (B) The major groups of Osteichthyan (bony) fishes. Actinopterygii = ray-finned fishes; Neopterygii = actinopterygians with distinctive separation of cheek and jaw bones from the opercular. Teleosti = advanced bony fishes. Elopocephala = advanced teleosts. Clupeocephala = a group derived from the Elopocephala. Euteleostei = most are characterized by acellular bone and by features of the skull and caudal skeleton. Neoteleostei = most characterized by stiff fin spines and modifications of the positions of the pectoral and pelvic fins and body proportions. Acanthomorpha = spiny teleosts with modifications for swimming concentrated in the caudal fin. Acanthopterygii = further modifications to the bones around the mouth and caudal vertebrae and fin are found in this group.

(Figure 1.9) Fossilization under most circumstances preserved only a very small percentage of the living world in the fossil record. Most of the animals and virtually all the plants that lived alongside these sharks were not fossilized (after Beerbower, 1960).

(Figure 1.10) Under rare conditions exceptional preservation of articulated skeletons and even so-called soft parts occurs. This diagram shows the factors involved in the preservation of a large Eocene biota in a German lake deposit. Many of these factors and pathways played a part in the preservation of complete fishes in the British Middle Devonian, Carboniferous and Jurassic record (after Franzen, 1985).

(Figure 1.11) The basic channels for the flow of materials through a typical ecosystem. This is elaborated in (Figure 1.12) to show the trophic flow in a late Devonian assemblage.

(Figure 1.12) A model of the possible trophic flow and connections in the well-preserved Eusthenodon assemblage in the Late Devonian of the Tula region of Russia (after Lebedev, 1992). This vertebrate community included chondrichthyes, antiarchs, dipnoi and crossopterygians as well as the tetrapod Tulerpeton.

(Figure 1.13) The procedures in stratigraphy and the categories within it (after Holland, 1978). Chronostratigraphy is the central repository for the data derived from the procedures and the phenomena indicated around them. The terms shown in brackets are not often used, but, logically, could be employed to greater extent.

(Figure 1.14) The global stratigraphical column, based on that of the International Union of Geological Sciences (Cowie and Bassett, 1989). In the situation where alternative series and stage names are given, the left hand column is usually favoured. The geochronometric dates are based upon published isotope analyses.

(Figure 1.15) Locality map showing the distribution of the 97 fossil fish sites described in this volume. They are principally grouped upon the ancient sedimentary basins: Caledonian and Orcadian (Devonian), Midland Valley of Scotland (Carboniferous), Anglo-Welsh (Devonian), Oxford-Wessex (Triassic-Jurassic) and the Hampshire and London Basins (Cenozoic). See (Table 1.2) for key to numbers.

(Figure 2.1) Silurian outcrops in Britain.

(Figure 2.2) Early Wenlock palaeogeography of Scotland (after Bassett, 1992).

(Figure 2.3) Location map of the Lower Palaeozoic inliers of the Midland Valley, showing GCR sites 1–6.

(Figure 2.4) Silurian successions in the inliers of the Midland Valley (after Cameron and Stephenson, 1984).

(Figure 2.5) The five basic types of thelodont scales, lateral and basal views and sagittal section to show the typical structure: b, base; cr, non-growing crown. The thelodont Lanarkia bears large spine-like scales as well as small rotund denticles in regular rows (after Turner, 1973).

(Figure 2.6) Thelodont ranges in the Silurian and Devonian of western Europe (after Janvier and Blieck, 1993).

(Figure 2.7) Silurian biozones now of use in international correlation. The vertebrates are largely microvertebrate (e.g. scales of thelodonts and acanthodians) taxa (from Marss, 1986).

(Figure 2.8) The Silurian succession in the Lesmahagow inlier (from Ritchie, 1985). Three vertebrate-arthropod faunal assemblages are distinguished: the jamoytius horizon (lowest), the Ceratiocaris Beds in the Kip Burn Formation, and the (latest) fish beds in the Dippal Burn and Slot Burn Formations. Ainiktozoon is known only from the Jamoytius horizon.

(Figure 2.9) (A) a reconstruction Jamoytius kerwoodi White by Ritchie (1968); (13) anterior end of a specimen from Logan Water with poorly preserved trunk scales; (C) anterior end of specimen with no trunk scales visible; (D) part of trunk of J. kerwoodi showing somites x 1.5 (GLAHM 101 382), Birk Knowes, Upper Llandovery, photographed under water (Photo: courtesy of the Hunterian Museum, Glasgow).

(Figure 2.10) The thelodont Loganellia scotica (Traquair); (A) Traquair's original reconstruction in dorsal view (from Miles, 1971); (B) Turner's (1970) reconstruction in lateral view; (C) Janvier's (1996) representations of Turinia.

(Figure 2.11)A Birkenia elegans Traquair. Well-preserved specimen from the ?Wenlockian of Birkenhead Burn, Shanks Castle, photographed under water, x 2, GLAHM V 8384 (Photo: courtesy of the Hunterian Museum, Glasgow). B Birkenia elegans Traquair. A restoration of this anaspid, the original fossil being c. 10 ems long (after Stetson, 1927).

(Figure 2.12) Map of the geology in the vicinity of Shiel Burn (after Rolfe, 1961).

(Figure 2.13) Stratigraphical sections in the Hagshaw Hills Silurian inlier to show the Fish Bed Formation (after Rolfe, 1961). The Greek letters refer to six divisions distinguished by Rolfe (1961).

(Figure 2.14) A The thelodont Thelodus scoticus (GLAHM V8304), ?Upper Llandovery at Birk Knowes, c. x 1 (Photo: courtesy of the Hunterian Museum, Glasgow). B The thelodont Lanarkia horrida (GLAHM V2302), ?Wenlock at Seggholm, ?Slot Burn; at about natural size (Photo: courtesy of the Hunterian Museum, Glasgow).

(Figure 2.15) A The anaspid Lasanius problematicus Traquair. Specimen from an unidentified horizon, Segholm (Slot Burn), c. x 2, GLAHM V 2286 (Photo: courtesy of the Hunterian Museum, Glasgow). B The anaspid Lasanius problematicus Traquair. Restoration in lateral view and possible feeding and swimming positions as suggested by Parrington (1958).

(Figure 2.16) Ateleaspis tessellata Traquair. (A) Reconstruction in lateral view, c. x 0.3; and of the headshield and front part of the trunk; (B) in dorsal and (C) ventral view, c. x 0.5, (after Ritchie, 1967): cmm, transverse sensory canals; D1 and D2 anterior and posterior dorsal fins; dsf, dorsal sensory field; ifc, infra-orbital sensory canal; lc, lateral sensory canal; lsf, lateral sensory field; o, orbit; pec, left pectoral fin; vlr, ventro-lateral ridge; vrs, ventro-lateral scales.

(Figure 2.17) Geological sketch-map of the coast to the north of Stonehaven (after McGregor, 1978); The Toutties is the area of foreshore south of Cowie Harbour.

(Figure 2.18) Stratigraphical log of the uppermost part of the Downtonian at The Toutties (after MacGregor, 1968).

(Figure 2.19) The ventral disc of Traquairaspis campbelli (Traquair) (holotype, RSM) from the Cowie Harbour Fish Bed, x 1.3. It has the characteristic serrated tubercles now known in related heterostraci throughout the circum-Arctic Siluro-Devonian and may be one of the oldest traquairaspidiforms (from Kiaer, 1932b).

(Figure 2.20) Mesacanthus mitchelli (Egerton) restoration in lateral view, approximately natural size (after Watson, 1937): isp, intermediate spine; with details of squamation of specimen from Tillywhandland (from Young, 1995).

(Figure 3.1) Map of the Downtown Series throughout the Welsh Borders, with schematic summaries of the stratigraphical successions and relationships with the Ludlow Series: UWF, Upper Whitcliffe Formation; DCSF, Downton Castle Sandstone Formation; LBBM, Ludlow Bone Bed Member; PSM, Platyschisma Shale Member; TSF, Temeside Shale Member; LF, Ledbury Formation; PPB, phosphatized pebble beds; PHF, Platyschisma helicites Formation; YDF, Yellow Downton Formation; GDF, Green Downton Formation; RDF, Red Downton Formation; Lud, undifferentiated Ludlow strata; SM, Sandstone Member; TF, Tilestones Formation; u/c, unconformity (after Bassett, Lawson and White, 1982).

(Figure 3.2) Correlation table of Late Silurian-Lower Devonian formations of the Anglo-Welsh province (after Siveter et al., 1984; House et al., 1977).

(Figure 3.3) Přídolí (latest Silurian) palaeogeography of southern Britain (after Bassett et al., 1992). (A) early Přídolí; (B) late Přídolí.

(Figure 3.4) Silurian thelodont denticles from the Welsh Borderland (after Turner, 1973). (A) from the T. pococki zone of the Downtonian: a, b, Goniporus alatus (Gross), c, d, Loganellia ludoviensis (Gross); e, Loanellia cuneata (Gross); f, L. cruciformis; g, Turinia pagei (Powrie); h, Loganellia sp. indet. (B) (Lower) Downton Formation, a, Thelodus costatus, Pander; b, Kataporodus tricavus Gross; c, Goniporus alatus (Gross): (C) Woolhope Limestone, a-b, Thelodus parvidens Agassiz; c-d-e, Loganellia ludoviensis (Gross).

(Figure 3.5) Late Silurian heterostracan vertebrates from the Welsh Borders area. (A) Anglaspis macculloughi in dorsal and lateral views, x 0.75, and, Tesseraspis tessellata in dorsal view, x 0.35; (B) restorations of the carapace of Traquairaspis pococki showing the ornamentation of the dorsal and ventral discs, and of the ventral disc of T symondsi, dorsal view; (C) Traquairaspis symondsi restorations of the carapace in dorsal and ventral views (after Tarrant, 1991); (D) Tesseraspis tessellata part of dorsal carapace, x 0.35 (after Halstead Tarlo, 1964).

(Figure 3.6) Stratigraphical distribution of the Siluro-Devonian agnathans of the Old Red Sandstone continent (England and Wales; after Janvier and Blieck, 1993).

(Figure 3.7) Geological sketch map and stratigraphical section of the Cwar Glas site in the Sawdde Gorge (after Bassett, 1982).

(Figure 3.8) (L) Dorsal and (R) ventral discs of the cyathaspidid Archaegonaspis ludensis from the Ludlovian, x 0.8 (from Lankester, 1864).

(Figure 3.9) Sketch map and stratigraphical section of the site at Ludford Lane, Ludlow (after Bassett et al., 1982).

(Figure 3.10) Fossils from the Ludlow Bone Bed illustrated in a plate from Murchison's Siluria (1872). 1–8, Plectrodus mirabilis, fragments of jaws; 9–12, P. pustuliferus, fragments of jaws; 13, 14, Onchus murchisoni, fin spines; 15–17, O.trenuistriatus, fin spines; 18, Thelodus parvidens, denticles; 19–20, indeterminable fragments; 21–28, coprolites containing invertebrarte fragments; 29, Orthis lunata Sowerby; 30, Pachtheca sphaerica. All approximately natural size.

(Figure 3.11) Thelodonts from the Ludlow Bone Bed (after Turner 1973). (A) Thedolus parvidens; (B) Thelodus bicostatus; (C) Loganellia ludlowiensis head scale; (D) Thedolus pugniformis; (E) Thelodus parvidens?; (F) Thelodus trilobatus.

(Figure 3.12) Ludlow- Přídolí correlation for Europe and North America. Vertical columns not drawn to scale (after Siveter et al., 1989).

(Figure 3.13) Section at Ledbury railway cutting recorded by Symonds (1860). Even more than 100 years later much of this section was still visible; recently, however, much of the western end has become obscured.

(Figure 3.14) Osteostracans from the Ledbury cutting. (A) Hemicyclaspis murchisoni Egerton in rare preservation, one of many such slabs collected over 100 years ago, c. x 0.25, (photograph courtesy The Natural History Museum,London, T05398/A). (B) Restoration of vertebrates found at Ledbury: a, Hemicyclaspis murchisoni; b, Auchenaspis egertoni Lankester; c, Didymaspis grindrodi Lankester (from Blieck and Janvier, in press).

(Figure 3.15) Hemicyclaspis murchisoni Lankester, detail of the dorsal surface of the headshield: p, pineal foramen; o, orbits; nh, nasal hypophysis; lsf, lateral sensory field; msf, median sensory field (after Stensiö, 1932).

(Figure 3.16) Stratigraphical section at Temeside. (After Elles and Slater, 1906).

(Figure 3.17) Sketch map of the GCR site at Tite's Point, and stratigraphical section (after Cave and White, 1971).

(Figure 3.18) Cyathaspis banksi Lankester, x 1.25; a primitive heterostracan, relatively common in Ludlow–Přídolí sites in this region. (A) external surface of ventral disc; (B) internal surface of the ventral disc; (C) cast of the internal surface of dorsal disc; (D) impression of the internal surface of the dorsal disc showing position of the gill pouches (g), semi-circular canals (c), orbits (o) and pineal foramen (p). Specimens from Kington, in the Museum of Practical Geology when figured by Lankester (1868).

(Figure 3.19) Sketch map and section of the outcrop at the cliffs and foreshore at Lydney (after Allen, 1978).

(Figure 3.20) Vertebrate fragments from Lydney. (A) Tesseraspis tessellata Wills, tesserae scattered at many horizons; (B) Sabrinacanthus: (B'), (B") details of ornamentation; al, ascending lamina; ispl first intermediate spine; psp, pectoral spine; ppsp2, 3, paired pectoral spines; sc, scapula. (B) after Miles (1973).

(Figure 3.21) Sclerodus pustuliferus, cephalic shield in dorsal view; apit, anterior pit; cf, central field; dsf, dorsal sensory field; lsf, lateral sensory field; mf, marginal foramen; mpit, median pit; no, nasal opening; o, orbit; po, pineal opening; ppit, posterior pit; approximately natural size (after Forey, 1987, © The Natural History Museum, London).

(Figure 3.22) The stratigraphical section at Tin Mill Race (after Elles and Slater, 1906).

(Figure 4.1) The Old Red Sandstone Continent (Euramerica or Laurussia); a general map, early Devonian to mid-Devonian times.

(Figure 4.2) Stratigraphical sections of the Devonian in the Welsh Borderland–South-west England (after House et al., 1977) with GCR site horizons indicated: A, Afon y Waen; B, Besom Farm; BS, Bedruthan Steps; C, Cwm Mill; D, Devil's Hole; H, Heol Senni; M, Mill Rock; O, Oak Dingle; P, Portishead; PC, Prescott Corner; W Wayne Herbert. Exact stratigraphical positions are not certain.

(Figure 4.3) The Silurian–Devonian succession in the Welsh Borders with faunas and suggested environments (after Allen and Tarlo, 1963).

(Figure 4.4) Fish zones in the Upper Downtonian and Dittonian of the Welsh Borderland and in the Lievin Group in northern France (after Blieck and Janvier, 1989). N.B. a narrow zone of Pteraspis rostrata between those of Rh. crouchi and Protopteraspis is postulated for northern France and may also be justified in Britain.

(Figure 4.5) The sites and stratigraphical distribution of some pteraspidid species in the Welsh Borderland (after Blieck, 1985, with data from Ball and Dineley, 1961). Pteraspidid biozones within the Lower Devonian of the Welsh Borderland. Localities yielding zonal species (mostly akin to Pteraspis rostrata): principal map area: 1, Cradley, near Malvern; 3, Guildings Brook, Trimpley; 4, Wayne Herbert Quarry; 12, Pool Quarry, Walterstone; 14, Goldstop, Newport; 15, Kentchurch Hill, Hereford; 16, Castle Mattock; 17, Pandy, Monmouth; 18, Wern Gwenny, Dorstone; 20, Hopton Brook, Cleobury Mortimer; 22, Hazely Brook West, Cleobury; inset (Brown Clee Hill area): 2, Whitbatch Quarry Ludlow; 5, Ledwyche Brook, Ludlow; 6, Targrove Quarry Ludlow; 7, Oak Dingle, Bouldon; 8, Abdon Brook; 9, Targrove; 10, New Buildings, Holdgate; 11, Rea Bridge, Derrington; 13, Jubilee Brook, Hopton Cangeford; 21, Upton Cressett Quarry. Protopteraspis occurs at: 23, Meadowley Bank, Morville; 24, Monkhopton; 25, The Leath; 26, New Inn; 27, Besom Farm. Rhinopteraspis occurs at: 28, Rea Brook, Silvington; 29, Primrose Hill Quarry; 30, Wilderness Quarry Mitcheldean; 31, Hoel Senni.

(Figure 4.6) Map of the Lower and Upper Old Red Sandstone divisions in the Anglo-Welsh Basin with GCR sites indicated. Key to numbers is provided in Table 1.2.

(Figure 4.7) Modes of occurrence of vertebrates in the Old Red Sandstone (after Dineley and Loeffler, in press). (A) separate, perhaps different, communities within a fluvial basin yield materials that are swept together and mixed downstream; (B) deposition within a meandering river system where vertebrate remains accumulate in a variety of situations. (C) vertebrate remains commonly occur within intraclast conglomeratic layers overlying erosion surfaces cut in siltstones or sandstones.

(Figure 4.8) Stratigraphical section in The Devil's Hole (after M.A. Rowlands, MS). The base of the Main 'Psammosteus' Limestone is mapped as the base of the Ditton Group, which has predominantly pteraspidid faunas.

(Figure 4.9) Traquairaspis (Phialaspis) symondsi (Lankester), restorations of the carapace on the basis of material from Devil's Hole by P.R. Tarrant (1991). (A) carapace in dorsal view, showing ornamentation of tubercles and branchial openings set near the middle of the dorsal surface of the branchial plates; (B) the sensory canal pattern of the dorsal carapace; (C) the carapace in ventral view with characteristic smooth central area; (D) pattern of sensory canals on the ventral side of the carapace; (E) and (F) lateral view of the carapace with sensory canals shown: alp, anterior lateral plate; bcp, branchio-cornual plate; bro, branchial opening; dd, dorsal disc; vd, ventral disc; Ip, lateral place; mop, median oral plate; or, orbit; orp, orbital plate; pi, pineal opening; pip, pineal plate; ro, rostrum; sca, smooth central area; dv, dorsal vane (or spine).

(Figure 4.10) Protopteraspis gosseleti Leriche. White's (1950b) Pteraspis (Simopteraspis) leathensis (A)–(C), is now regarded as a junior synonym of this species (D). The lateral line canals are shown by broken lines in C and D. (A)-(B) from White (1950b); (C), (D) from Blieck (1985).

(Figure 4.11) Section through the cyclothem in the Lower Ditton Group at Oak Dingle, Tugord (after Allen, 1964, and M.A. Rowlands MS).

(Figure 4.12) Pteraspis rostrata var. trimpleyensis White, common fragments occur in Oak Dingle, Bouldon, and the adjacent Bouldon ford. This specimen from Bouldon figured by White (1961).

(Figure 4.13) The succession at Cwm Mill, Abergavenny (after Allen, 1964).

(Figure 4.14) Cephalaspids from Cwm Mill (after White and Toombs, 1983). (A) Cephalaspis cradleyensis Stensiö, restoration of headshield; (B) Cephalaspis cwmmillensis White, holotype in dorsal impression and counterpart, with restoration: nsf, nerve canals to lateral sensory field; cv, rostral vein; sof, supra-oral field; (C) Cephalaspis abergavenniensis White, external dorsal aspect of the headshield and part of the thorax with counterpart and restoration; (D.) Cwmaspis (Cephalaspis) billcrofti White, part of headshield in dorsal aspect, with restoration (after White, 1963).

(Figure 4.15) Rhinopteraspis crouchi (Lankester) from Cwm Mill (after White, 1973; Blieck, 1980). (A) R. crouchi carapace in dorsal view with sensory canal system shown by broken lines: b, branchial openings; o, orbits; (B) R. dunensis (Roemer) carapace in dorsal view, the extreme elongated form found in the higher Dittonian strata: (C), (D) dorsal discs; (E), (F) rostral plates.

(Figure 4.16) Pteraspids from Wayne Herbert Quarry, after White (1935). (A) Pteraspis rostrata var. toombsi, dorsal view of the carapace with parts of the lateral line system shown; (B), (C) ventral views of the rostrum and oral appratus, NHM P17488 x 0.4 and NHM P17487 x 1; (D) Pteraspis waynensis (P 16524) x 0.5, outline of the ventral disc with lateral line canals indicated; (E) Pteraspis rostrata var. toombsi (P16789) x 0.33, almost complete individual preserved as an external mould, ventral view of carapace trunk and tail; (F) the same specimen preserved in counterpart, x 0.33, seen in dorsal view; (G) Pteraspis jackana (P17628) x 0.5 approximately, dorsal disc with traces of lateral line canals in dorsal view; (H)—(1) Pteraspis rostrata var. toombsi, external lateral impressions of tails with squamation (NHM P 17488, NHM P17521, NHM P 17477), at c.x 0.25.

(Figure 4.17) (A) Poraspis sericea Lankester from Wayne Herbert Quarry: a dorsal aspect of the dorsal disc of the holotype (NHM P.4117); (B) median dorsal, lateral, ventro-lateral and median ventral scales of Poraspis; (C) lateral profile of the dorsal disc of Poraspis; Br, position of the branchial opening between margins on the disc and the branchial plate; O, position of the orbit; R, rostrum; (D) Nikolivia milesi, denticles from the trunk showing overlap (NHM P 53902). (From Turner, 1982a).

(Figure 4.18) Acanthodians from Wayne Herbert Quarry (after Miles, 1973; Young, 1995). (A) Ptomacanthus anglicus Miles x 0.25; (B) Vernicomacanthus waynensis Miles, x 0.45; (C) Uraniacanthus spinosus Miles, x 0.66, with detail (x 2) of scale type in vicinity of the pectoral spine.

(Figure 4.19) Section in the old quarry at Besom Farm, Wheathill (after M.A. Rowlands MS).

(Figure 4.20) Restorations by Blieck and Janvier (1989) of vertebrates occurring in the Lower Devonian at Artois, France. The same species (A, B, C, D, E) and Pattenaspis occur in South Wales and the Welsh Borderland. (A) Turinia pagei (Powrie); (B) Rhinopteraspis crouchi (Lankester); (C) Kujdanowiaspis sp.; (D) Pattenaspis artesensis (Agassiz); (E) Protaspis (Europrotaspis) cremulata White, the holotype (NHM P28801) an imperfect dorsal shield c. x 0.66. br, Branchial plate; bro, branchial opening; brd, branchial duct; coc, cornual contact surface on branchial plate; cop, cornual plate; do, dorsal disc; or, orbital plate; ora, orbit; ro, rostrum; sp, shells of spirorbis attached to undersurface of dorsal disc (after White, 1961).

(Figure 4.21) Althaspis senniensis:(A) rostral plate from Hoel Senni Quarry; (B) a ventral disc from Hoel Senni Quarry (after Loeffler and Thomas, 1980), both approximately natural size.

(Figure 5.1) Palaeogeography of the Early Devonian in northern Britain (after Bluck et al., 1992). (A) palaeogeography of Lochkovian time, c. 408 Ma; (B) palaeogeography of Late Pragian–Emsian time, c. 400 Ma. Continuing movement of the fault-bounded blocks dominated the area of Scotland throughout this interval, with uplift continuing to the north and south of the Midland Valley.

(Figure 5.2) Lateral variation in the Lower Devonian strata of the Strathmore Syncline in the northeast of the Midland Valley of Scotland (after Armstrong and Paterson, 1970).

(Figure 5.3) Cephalaspis lyelli Agassiz. (A), (B) Dorsal and (C) lateral views of the headshield and trunk region of the lectotype (NHM P 20087) from Glamis, Angus (after White, 1958b): lsf, lateral sensory field; msf, median sensory field; n, nasal hypophysis; o, orbit; p, pineal; pa, pectoral appendage.

(Figure 5.4) Cephalaspids from the Early Devonian of Scotland (from Stensiö, 1932, © The Natural History Museum, London). (A) Cephalaspis powriei Lankester headshield in dorsal view after specimen RSM 138, x 0.8; (B) C. spinifer headshield after the holotype RSM 1891.92.149, x 0.8; (C) C. spinifer complete animal in dorsal view, x 0.7.

(Figure 5.5) New restorations of Scottish Early Devonian thelodonts by Turner (1992): (A) Turinia pagei (Powrie); (B) Lanarkia spinosa Traquair; (C) Loganellia scotica (Traquair), in which post-pectoral fin and tail are hypothetical. All are c. x 0.75.

(Figure 5.6) Restorations of acanthodians from the Early Devonian of Scotland. (A) Mesacanthus mitchelli (Egerton), x 2.0; (B), (C) Euthacanthus macnicoli Powrie x 0.8 in lateral and ventral views; (D) Ischnacanthus gracilis (Egerton), x 1.25 approx.; (E) Mesacanthus sp. approximately natural size (taken with permission from J. Long, 1995); (F) Parexus sp., approximately natural size. From various sources.

(Figure 5.7)A Tillywhandland Quarry: the succession (after Trewin and Davidson, 1996). B Tillywhandland Quarry: the recorded positions of the fauna and flora within the fish bed (after Trewin and Davidson, 1996).

(Figure 5.8) Characteristic acanthodian scales from Tillywhandland Quarry. (A) Ischnacanthus gracilis (Egerton), NHM P 62266, scales rhombic, sub-rhombic or polygonal, relatively flat to gently convex and smooth; (B) Mesacanthus mitchelli (Egerton), P 140 and 10892, scales smooth with flat crowns, sharp posterior point; (C) Euthacanthus macnicoli Powrie, scales with simple ribs, very flat crowns with five to ten well-spaced grooves. (Courtesy V.T. Young.)

(Figure 5.9) A restoration of the margins of 'Lake Forfar' as proposed by Trewin and Davidson (1996). A vagrant benthos of cephalaspids, eurypterids and other invertebrates occupied the shallows; acanthodians lived in the open waters. Bottom conditions were probably oxygen-poor. Terrestrial arthropods inhabited the vegetated land adjacent to the lake. From nearby volcanoes quantities of fine ash periodically drifted over the lake, while there was throughout a seasonally variable input of fine fluvial sediment.

(Figure 5.10) Aberlemno Quarry, stratigraphical section (after Armstrong, 1978).

(Figure 5.11) Cephalaspids from Wolf's Hole Quarry. (A) Cephalaspis scotica White, imperfect headshield is dorsal aspect, holotype RSM Powrie collection 1981.92.135, x 2. (B) Securiaspis waterstoni White, imperfect headshield in dorsal aspect, holotype, Perth Museum, unregistered, x 1. (C) Securiaspis caledonica White, external impression of headshield. Area to left of dotted line is restored. Perth Museum, unregistered, slightly less than natural size. (From White, 1963)

(Figure 5.12) Pteraspis mitchelli Powrie from Wolfs Hole Quarry (A), (B) Negative and positive counterparts of a small dorsal shield. Lectotype, Bridge of Alan, RSM Powrie Collection, 1891.92.118 and 117, x 1. (C) External impression of imperfect dorsal shield, Bridge of Alan, Perth Museum, unregistered, x 1. (D) Very imperfect dorsal shield figured by Lankester (1868), Bridge of Alan, RSM Powrie Collection, 1892.92.119, x 1. (E) External impression if imperfect dorsal shield. Specimen figured by White (1935). Bridge of Alan, NHM P 16808, x 1. (F) external impression of small dorsal shield. RSM Powrie Collection, 1892.92.122 x 1. (All figures from White, 1963.)

(Figure 6.1) Palaeogeography of the Middle Devonian, 380–375 Ma, of Scotland and adjacent North Sea (after Bluck et. at, 1992).

(Figure 6.2) Stratigraphical sections of the Middle Old Red Sandstone of Scotland, Orkney and Shetland, with GCR sites shown. A, Achanarras Quarry; B, Banniskirk Quarry; C, Cruaday Quarry; D, Dipple Brae; E, Blackpark; F, Edderton; G, Gamrie, Den of Findon, Tynet Burn; H, Holburn Head Quarry; J, John o'Groats; M, Melby; P, Papa Stour; Py, Pennylands; S, Spital Quarry, Su, Sumburgh Head; W Weydale Quarry; X, Exnaboe (after Mykura (1991), Donovan et al. (1974) and others).

(Figure 6.3) Ranges of the common fossil fishes in the Middle Old Red Sandstone of the Orcadian Basin compared with the Eastern Baltic vertebrate biozones (largely after Donovan et al., 1974; Dineley and Loeffler, 1993; and Mark-Kurik, 1978). Z, Biozones distinguished by Donovan et al., (1974): 1, Thursius macrolepidotus; 2, Coccosteus cuspidatus; 3, Palaeospondylus gunni; 4, Dickosteus threiplandi; 5, Asmussia murchisoniana; 6, Millerosteus minor; 7, Watsonosteus fletti. A, Asterolepis; P, Psammosteus; Sch, Schizosteus.

(Figure 6.4) Diagrammatic reconstruction of the marginal environments around the Middle Devonian Orcadian Basin of Scotland and the North Sea at a time of high stable water level (after Trewin, 1986). The shallow-water well-oxygenated zone (A) provided a variety of habitats for the fish, which were ultimately preserved in the deeper deoxygenated zone (B).

(Figure 6.5) Fossil fishes from Westerdale Quarry (A) The lungfish Dipterus valenciennesi Agassiz (Photo: courtesy The Natural History Museum, London, T00826/A), approximately natural size.Fossil fishes from Westerdale Quarry (B) D. valenciennesi in restoration by Ahlberg and Trewin (1995); (C) the osteolepid Thursius (Photo: courtesy The Natural History Museum, London, T00448/A), c. 12 cm; (D) T. macrolepidotus (Sedgwick and Murchison) (from Jarvik, 1948a).

(Figure 6.6) The section of Achanarras Quarry, after Trewin (1986). Numbers 1–6 are the main fossiliferous horizons.

(Figure 6.7) Achanarras Quarry (photo: D.L. Dineley).

(Figure 6.8) Particularly fossiliferous horizons with well-preserved fishes may result from mass mortality events induced by planktonic blooms. The axonic conditions extend throughout the shallow marginal areas (A); following this, carcasses drift into deeper water in a bloated conditions (B); after further decay (C), they sink through the thermocline and are preserved in laminites in the anoxic hypolimnion (D). The depth of the thermocline may have been no more than a few tens of metres (after Trewin, 1986).

(Figure 6.9) Common fishes at Achanarras Quarry: Pterichthyodes rnilleri (Miller) reconstructions in (A) dorsal, (B) ventral and (C) lateral aspects, x 0.33 approximately. (D) Palaeospondylus gunni Traquair in characteristic preservation with the head and anterior end in dorsal view but the posterior part of the vertebral column in lateral view, after Moy-Thomas (1940). (E) Common fishes at Achanarras Quarry: Pterichthyodes milleri (Miller). Photograph GLAHM V7015 showing typical compression preservation, c. x 0.8 (Photo: courtesy of Hunterian Museum, Glasgow). (F) Common fishes at Achanarras Quarry: Pterichthyodes milleri (Miller), T05399/A, with an early model of the animal, x 0.5 (Photo: courtesy The Natural History Museum, London). (G) Common fishes at Achanarras Quarry: Palaeospondylus gunni Traquair in characteristic preservation with the head and anterior end in dorsal view but the posterior part of the vertebral column in lateral view, c. x 6 (Photo: courtesy of the Hunterian Museum, Glasgow). (H) Common fishes at Achanarras Quarry: Dipterus valenciennesi Agassiz Photograph GLAMH V3656 showing the commonly well-preserved nature of this fossil lungfish, c. x 0.75 (Photo: courtesy of the Hunterian Museum, Glasgow).

(Figure 6.10) Geological sketch map of GCR Site Cruaday Quarry, Mainland, Orkney, based on the Geological Survey Orkney sheet.

(Figure 6.11) The Sandwick Fish Bed (after Trewin, 1976), showing division of the fish-bearing laminites into an upper and a lower leaf. Fish distribution is shown in the histograms for the upper leaf. 'Larger acanthodians' are mostly Mesacanthus and Cheiracanthus.

(Figure 6.12) Acanthodian species from Cruaday Quarry. Restoration of Diplacanthus crassisimus Duff: (A) lateral view; (B) ventral view, approximately natural size. (C), (D) Scales of D. crassisimus, posterior to the right: (C) exterior view; (D) side view, c. x 60. (E), (F) Scales of Cheiracanthus murchisoni; (E) interior view; (F) exterior view, c. x 55 (scales from Denison, 1979). (G) 'Rhadinacanthus'Diplacanthus longispinus Agassiz scales with well-defined ribs and scalloped posterior margin; (H) Diplacanthus striatus Duff, scales with fine transverse grooves, (G) and (H) c. x 50 (courtesy of V.T. Young).

(Figure 6.13) Osteichthyan species from Cruaday Quarry (A)—(C) The osteolepid Osteolepis macrolepidotus (Sedgwick and Murchison), restoration in lateral, dorsal and ventral views respectively; (D) Gyroptychius agassizi Traill, restoration in lateral view. (After Jarvik, 1948a.)

(Figure 6.14) The stratigraphical section in the brook at Edderton. The fish are confined to units 2, 3 and 4. (After Peach et al., 1912.)

(Figure 6.15) The Edderton ptyctodont Rhamphodopsis threiplandi Watson. (A) restoration of the skeleton in lateral view; (B) restoration of the skull in lateral view; (C) restoration of pectoral girdle in ventral view (after Miles, 1967).

(Figure 6.16) Correlation of the area Strathpeffer–Black Isle–Tarbat Ness, showing the central position of the Nodular Fish Beds and the Edderton Fish Beds (after Donovan, 1978).

(Figure 6.17) Den of Findon section (from Trewin and Kneller, 1987).

(Figure 6.18) Fishes from the Den of Findon. (A), (B) Cheirolepis trailli Agassiz, restorations of lateral and ventral views respectively (from Pearson and Westoll, 1979). (C)–(E), Coccosteus cuspidatus Agassiz: (C) restoration of the fish head in lateral view; (D) restoration of the head and trunk shields in dorsal view; (E) restoration of the skull in anterior view (after Miles and Westoll, 1968). Fishes from the Den of Findon. (F) Cheirolepis trailli Agassiz, a more or less complete specimen in lateral view, T00382/A, x 0.75 (Photo: courtesy The Natural History Museum, London).

(Figure 6.19) Tynet Burn, map of the GCR site and section through the fish-bearing beds.

(Figure 6.20) Fossil fishes from Tynet Burn. (A) Cher acanthus latus Egerton, a restoration based upon NHM P 15286 with scales from below and above the lateral line; (B) Cheiracanthus murchisoni Agassiz, a restoration based upon NHM P 6189 with scales from above and below the lateral line ((A) and (B) from Young, 1995); (C) T04134R of Cheiracanthus x 2 (Photo: courtesy The Natural History Museum, London); (E), (F) Reconstructions based on species from Tynet Burn: (E) skull roof with surface ornamentation and lateral line canal grooves; (F) in outline with radiation centres shown. (D) Fossil fishes from Tynet Burn. (D) photograph GLAHM V3573, of Mesacanthus pusillus (Agassiz) x 2.5, specimens preserved in nodules.

(Figure 6.21) Stratal settings of the Melby Fish Beds (after Mykura and Phemister, 1976).

(Figure 6.22) Fishes from Melby: (A) dorsal view of the head and pectoral shield of the arthrodire Homosteus (after Moy-Thomas and Miles, 1971); (B) the osteichthyid Gyroptychius in dorsal and lateral view (after Jarvik, 1948a).

(Figure 6.23) Stratigraphical section through Dipple Brae Fish Bed (based on data from M.A. Rowlands MS).

(Figure 6.24) (A) The coccosteid arthrodire Dickosteus threiplandi Miles and Westoll; outline of the skull roof and cheek bones laid out in a single plane, based on the holotype RSM 1962.4, (after Miles and Westoll, 1963). (B) Cephalaspis magnifica Traquair, cephalic shield in ventral view at x 0.23.

(Figure 6.25) Sketch map of the geology of Holborn Head, near Thurso (after Hamilton and Trewin, 1994).

(Figure 6.26) Distribution and abundances of fossil fish in Holborn Head Quarry (after Hamilton and Trewin, 1988).

(Figure 6.27) The arthrodire Millerosteus minor (Miller). (A) Reconstruction in lateral view of the head and thoracic region of this species from the Mey and Ackergill Beds, Thurso Flagstone Group, of Caithness and Orkney (after Desmond, 1974); md, median dorsal plate; nu, nuchal plate; x, rostrum. (B) median dorsal plate, external aspect; (B') median dorsal plate, internal aspect. (C) Nuchal plate, external aspect.

(Figure 6.28) Locality map of the John o'Groats area (after Trewin and Hurst, 15 )93).

(Figure 6.29) Sketch map of the geology of south-east Shetland, including Exnaboe and Sumburgh Head (after Mykura, 1976).

(Figure 6.30) Fossil fish found at Exnaboe, Shetland. (A) Stegotrachelus finlayi Woodward and White, an early actinopterygian from Exnaboe; (B) Pentlandia macroptera Traquair, x 0.5; (C) Microbrachius dicki Traquair, reconstruction in dorsal view of the carapace of the smallest antiarch, based largely on RSM 1877.22.4 from John o'Groats and DMSW P 513 from Deerness; (D) Watsonosteus fletti (Watson) from Deerness (after Miles and Westoll, 1963); outline drawing of NHM P 11732 in dorsal view; avl, anterior ventro-lateral plate; pmv, posterior ventro-lateral plate; pvl, posterior ventro-lateral plate; r, rostrum.

(Figure 6.31) The antiarch Asterolepis thule Watson from Sumburgh Head, c. x 0.2. Reconstruction after Janvier (1996).

(Figure 7.1) The palaeogeography of southern England and Wales during Late Devonian time (c. 370–365 Ma), based upon outcrop and borehole evidence (after Bluck et al., 1992).

(Figure 7.2) Distribution of Mid and Late Devonian outcrops and GCR sites in England and Wales (site numbers as in Table 1.2).

(Figure 7.3) Simplified non-palinspastic fades section of Devon and Cornwall (after House, 1975).

(Figure 7.4) Sketch map and section through the Late Devonian near Portishead (after Pick, 1964).

(Figure 7.5) Fossil fishes from the Late Devonian of Woodhill Bay, near Portishead. (A) Reconstruction of the large porolepiform Holoptychius spp. (after Janvier, 1996); (B) Holoptychius scales, c. x 0.5; (C) Bothriolepis maxima Agassiz, restoration in dorsal view, c. x 0.5; (D) Asterolepis maxima Agassiz, x 0.3; (E) Glypopomus kinnairdi Woodward restoration after Jarvik.

(Figure 7.6) Groenlandaspis from Portishead and other localities. Reconstruction of Groenlandaspsis (taken with permission from J. Long, 1995), length of animal c. 25 cm. (A) Dorsal view of cranial roof plates; (B) plates of the trunk armour in lateral view; (C) trunk armour in ventral view (after Ritchie, 1974), (A)–(C) x 0.5; (D) reconstruction (after Long, 1995); (E) reconstruction of Groenlandaspsis showing ventral surface, by Ritchie (1971).

(Figure 7.7) Vertebrates from the Upper Old Red Sandstone at Prescott Corner and nearby, Shropshire. (A) Pseudosauripterus anglicus (Woodward) cast of the right entoperygoid, x 0.75; (B) Eusthenopteron farloviensis White, right cleithrum, Prcl, process of cleithrum, x 0.5. Specimens from Church Quarry, figures by White (1961).

(Figure 7.8) Stratigraphical section in the stream at Afon y Waen (after Hall et al., 1973).

(Figure 8.1) Upper Old Red Sandstone outcrops and locations of GCR sites; 0, Oxendean; HH, Hawk's Heugh; Bhl, Boghole; Sc, Scaat (Scat) Craig.

(Figure 8.2) Late Devonian palaeogeography of Scotland, c. 370–365 Ma (after Bluck et al., 1992).

(Figure 8.3) Upper Devonian stratigraphical successions in the Cheviot, Midland Valley and Moray Firth areas of Scotland (after Westoll, in House et al., 1977).

(Figure 8.4) Stratigraphical ranges of widespread and relatively common Mid- and Late Devonian vertebrates in Euramerica (after Blieck et al., 1988). P, Phyllolepis; B, Bothriolepis; R, Remigolepis; G, Groenlandaspis; Pt, Pterichthyodes; A, Asterolepis (all x 0.25); M, Microbrachius, (x 1.0).

(Figure 8.5) Sketch map of Oxendean Burn area.

(Figure 8.6) The antiarch Bothriolepis hayi Miles (1968) from the Oxendean Beds of Oxendean Burn. (A) restoration of the trunk-armour in ventral aspect based on RSM 11967.34.43 and FR 1867, x 0.2; (B) restoration of the dorsal trunk armour, somewhat flattened, based on RSM 1967.34.16, x 0.2 (after Miles, 1968); (C) reconstruction of Bothriolepis (after Long, 1995), x 0.42.

(Figure 8.7) Geological sketch map and section of the Hawk's Heugh (GCR site) coastline (after Greig and Davies, 1978).

(Figure 8.8) The antiarch Remigolepis. (A) Restoration after Ritchie (1986); (B) the carapace in dorsal aspect (after Miles, 1968).

(Figure 8.9) Sketch map of Muckle Burn–Whitemire GCR site area.

(Figure 8.10) Suggested correlation of the Upper Old Red Sandstone of the Moray–Nairn area with the successions in Greenland, Spitsbergen, Belgium and the Baltic Province (after Miles, 1968; Westoll, 1977).

(Figure 8.11) Scaat Craig GCR site sketch map.

(Figure 8.12) Psammosteids from Scaat Craig. (A) Restored dorsal view; (B) a restoration of Psammolepis, both x 0.5 (after A. Bistrova).

(Figure 8.13) (A) Holoptychius nobilissimus Agassiz, a restoration from Scaat Craig, x 0.2; (B) Holoptychius specimen from Classbinnie, Perthshire, figure in ventral aspect, x 0.3 (after Murchison, 1872).

(Figure 8.14) Fossil tetrapod bone fragments from Scaat Craig (from Ahlberg, 1995). (A), (B) Elginerpeton pancheni Ahlberg, holotype RSM G 1967.17.1, in ventro-lateral and dorsal views. (C), (D) Reconstruction of the mandible in lateral and mesial views: brackets indicate parts of the jaws preserved in different specimens. (E), (F) the premaxilla, OUM Geol. Col D796, anterior part in dorsal and ventral views. (G)–(I) the posterior part of the premaxilla, NHM P9776, in lateral, ventral and dorsal views (vertical hatching = broken bone).

(Figure 9.1) Maps of Carboniferous palaeogeographies: (A) Late Tournaisian; and (B) Late Viséan; and (C) Namurian.

(Figure 9.2) Carboniferous stratigraphy and correlation (after MacGregor, 1960).

(Figure 9.3) Carboniferous outcrops in Britain: present GCR sites are limited to the south of Scotland and to the Border country and the south-east Pennines. Horizons previously yielding fossil fish occur at Oreton (Shropshire), in the Bristol Avon gorge and the west coast of Devon. Midland Valley sites: 54, Wardie; 56, Cheese Bay; 57, Inchkeith; 58, Ardross Castle; 59, Abden. Glasgow area site: 61, Bearsden. Berwickshire sites: 53, Foulden; 55, Glencartholm. Derbyshire: 60, Steeplehouse Quarry.

(Figure 9.4) Basal actinopterygian fish morphology (after Lauder and Liem, 1983). (A) Moythomasia nitida, Late Devonian, with stout body, large eyes and long jaws, dorsal ridge scales behind the dorsal fin; (B) Cheirolepis canadensis, Mid-Late Devonian, elongate with tiny scales; (C) Aduella blainvillei, Early Permian, stout body, large scales, arrow points to 'chondrostean hinge', large eyes, suborbital and preopercular bones in head, small mouth; (D) Andriochthys tuberculatus, Early Carboniferous, deep body, small tail and paired fins, large scales and prominent dorsal ridge scales; (E) a characteristic early actinopterygian scale bearing a small peg that articulates with the scale immediately above it.

(Figure 9.5) Cladogram of the relationships of the different groups of early actinopterygians (from Gardiner and Schaeffer, 1989). It is based on a preliminary cladogram of a visual comparison of the nasal-temporal bones of the head and also upon a computer analysis of a further large data matrix.

(Figure 9.6) Carboniferous chondrichthyans: (A) a cla-doselachid persisting from the Late Devonian, x 0.16; (B) a symmoriid, Denea, x 0.16; (C) the Late Carboniferous eugenodont Fadenia, x 0.16; (D) the Early Carboniferous xenacanthid Diplodoselache, x 0.08; (E) the xenacanthid Xenacanthus, x 0.08. Note the differences in cranial and vertebral ossification and fin structure (after Carroll, 1988).

(Figure 9.7) Geological sketch map of southern Scotland and northern England with the positions of the Carboniferous GCR fish sites. Key to localities: A, Abden; B. Bearsden; CB, Cheese Bay; EK, East Kirkton; F, Foulden; G, Glencartholm; I, Inchkeith; R, Ardross Castle; W, Wardie.

(Figure 9.8) Sketch map of the GCR site at Foulden.

(Figure 9.9) Stratigraphical section at the Foulden site with the fish bed detailed (after Wood and Rolfe, 1985).

(Figure 9.10) Foulden acanthodians: (A) Acanthodes ovensi White, with a series of immature forms showing the progressive spread of squamation forward during growth (from Forey and Young, 1985): (B) White's (1927) figure of the holotype (NHM P 13137) of A. ovensi.

(Figure 9.11) Foulden sarcopterygians: (A) preliminary reconstruction of ?Strepsodus aulaconamensis sp. with scale cover; (B) ?Strepsodus aulaconamensis Andrews, with scales omitted to show lepidotrichia and axial skeleton (after S.M. Andrews, 1985) Many of the head and axial skeleton bones are largely conjectural. (C) Coelacanth flank scales (from Forey and Young, 1985).

(Figure 9.12) Foulden actinopterygians (after White, 1927). (A) Aetheretmon valentiacum White; (B) Phanerosteon mirabile White; (C) Strepheoschema fouldenensis White.

(Figure 9.13) Sketch map and geological section at the Wardie GCR site, East Lothian (after Wood, 1975).

(Figure 9.14) Actinopterygians in nodules from the Wardie Shale, Wardie, (A)–(D) are all c. X 0.66. (A) Cosmoptychius striatus Agassiz, specimen in nodule; (B) Elonichthys robisoni Hibbert; (C) Rhadinichthys ferrox Traquair; (D) Nematoptychius greenocki Traquair; (E) Gonatodus punctatus Agassiz, restoration from Gardiner (1967a); (F) Nematoptychius greenocki Traquair, restoration of the head by Gardiner (1963), c. x 0.5.

(Figure 9.15) Wardie elasmobranchs; (A) Diplodoselache woodi Dick, restoration of the skeleton, after Dick (1981); (B,a) scales from the anterior part of the trunk; (B,b) scales from the anal fin; (B,c) flank scales; (B,d) scales from head, body and tail; (C) Tristychius arcuatus Agassiz restoration of the skeleton (after Dick, 1978); (D) Onychoselache traquairi Dick, restoration (after Dick, 1978); (E) The Wardie tetrapod Lethiscus stocki Wellstead, a restoration of the dorsal surface of the skull (after Wellstead, 1982); (F) the Acanthodes sulcatus Agassiz restoration after Moy-Thomas and Miles (1971).

(Figure 9.16) Sketch map of the Glencartholm GCR site, Berwickshire.

(Figure 9.17) Some of the relatively more common Glencartholm actinopterygians: restorations in lateral view by Moy-Thomas and Bradley Dyne (1938). (A) Rbadiniththys fusiformis Traquair; (B) R. canobensis Tranquair; (C) Canobius ramsayi Traquair; (D) C. elegantulus Traquair.

(Figure 9.18) Glencartholm actinopterygians: restorations in lateral view by Moy-Thomas and Bradley Dyne (1938). (A) Cycloptychius concentricus Traquair; (B) Mesopoma politum Traquair; (C) Mesopoma crassum Traquair.

(Figure 9.19) Glencartholm actinopterygians: restorations in lateral view by Moy-Thomas and Bradley Dyne (1938). (A) Holurus parki Traquair; (B) the elongated Tarrasius problematicus Traquair, once thought to be a crossopterygian (after Moy-Thomas, 1937b); (C) Platysomus superbus Traquair; (D) Cheirodopsis geikiei Traquair. (C) and (D) are typical deep-bodied actinopterygians.

(Figure 9.20) Glencartholm elasmobranchs. (A) Sphenacanthus costellatus Traquair, restoration (after Moy-Thomas, 1936); (B) Goodrichichthvs eskdalensis, Moy-Thomas restoration; (C) Tristychius arcuatus Agassiz, isolated teeth; a, lingual; b, lateral; c, labial views of RSM 1972.276.461A; d, labial view of RSM 1972.27.460B.

(Figure 9.21) Glencartholm holocephalians. (A) Chondrenchelys problematica Traquair in lateral restoration after Patterson (1965). (B) Deltoptychius armigerus Traquair, restoration of the headshield in dorsal view based on NHM P 11372 (after Patterson, 1966); anterior at top, incompletely fused tesserae make up the central part of the shield. (C) Dentition restored as if seen in front of wide-open mouth. (D) Deltoptychius, restoration of the fish courtesy of the Hunterian Museum, Glasgow; overall length of the specimen from Bearsden (q.v.) c. 60 cm.

(Figure 9.22) Section of the Cheese Bay GCR site.

(Figure 9.23) Sketch map of Inchkieth.

(Figure 9.24) Ardross Castle site fossil actinopterygians: (A) Watsonichthys pectinatus Traquair, skull in lateral view (after Gardiner, 1963); (B) Rhadiniscus wrighti White, skull in lateral view; (C) the coelacanth Rhabdoderma, restoration in lateral view (after Forey, 1981).

(Figure 9.25) Sketch map of the GCR site at Abden.

(Figure 9.26) Steeplehouse Quarry elasmobranchs. Teeth of the hybodont selachian Lissodus wirksworthensis Duffin: (A) NHM P 60741 in labial view; (B) P 60739 in occlusal view; (C) holotype P 60740 in occlusal view; (D) in lingual view; (E) labial view; (F) P 60752 in occlusal view; (G) labial view; (H) lingual view; (I) P 60748 in occlusal view; (J) labial view (after Duffin, 1985).

(Figure 9.27) (A) Sketch map of the Bearsden area, Glasgow, with the stratigraphical section (B) found in the Manse Burn Formation (after Wood, 1982).

(Figure 9.28) Bearsden actinopterygians: (A) Mesopoma carricki Coates in lateral restoration (after Coates, 1993); (B) Frederichthys musadentatus Coates, a composite restoration after the holotype GLAHM V 8286 (from Coates, 1993). (Continued on p. 309.) (Figure 9.28) — contd. Bearsden actinopterygians: (C) Melanecta annaea, GLAHM V 8255, x 2, photographed in UV light; (D) Mesopoma carricki, GLAHM V 8254, x 1.5, photographed in toluene; (E) Cheirodus sp., GLAHM, X 1.25 (Photos: courtesy of Hunterian Museum„ Glasgow).

(Figure 9.29) Bearsden fishes: (A) the acanthodian Acanthodes sp., GLAHM unnumbered photo in toluene and figured as a restoration in lateral view; (B) Deltoptychius, GLAHM unnumbered photo in toluene at x 0.3; (C) Stethacanthus sp. (the 'Bearsden Shark') GLAHM V8246 at x 0.25 (Photos: courtesy of Hunterian Museum„ Glasgow).Bearsden fishes: (D) reconstruction of the Stethacanthus skeleton (after Zangerl, 1981); (E) the Bearsden shark Stethacanthus sp., c. x 0.1.

(Figure 10.1) Map showing the outcrop of Permian rocks in Great Britain, with the Middridge GCR site indicated (1).

(Figure 10.2) Palaeogeographical map showing Britain during the early part of the English Zechstein Cycle 1 and equivalents, c. 255 Ma (after D.B. Smith, in Cope et al., 1992). Symbols indicate various invertebrate fossils.

(Figure 10.3) Correlation of part of the Permian succession of northern England with that in North Germany and Holland (after Smith, 1989).

(Figure 10.4) The Zechstein Sea, showing the major sub-basins and the land masses (after Smith, 1992). Sea depth was nowhere more than 30 m; the English Basin was less than 250 mm deep.

(Figure 10.5) The palaeoniscid Palaeoniscus freieslebeni Blainville: (A) restored lateral view (after Aldinger, 1937); (B) reconstruction of the head (after Westoll, 1941).

(Figure 10.7) (A) the semionotid Acentrophorus restored in lateral view (after Gill, 1925), with specimens, (B), (C) from the Marl Slate, Middridge (natural size) (figured by King, 1850).

(Figure 10.6) (A) Dorypterus hoffmanni Germar, restoration in lateral view (after Westoll, 1941); (B) Platysomus striatus Agassiz (from King, 1850).

(Figure 10.8) The holocephalian janassa bituminosa (Schlotheim) in ventral view, and showing the petalodont dentition (after Shaumberg, 1978).

(Figure 10.10) Wodnika striatula Munster, reconstruction by Schaumberg (1978).

(Figure 10.9) (A) Coelacanthus granulatus Agassiz, a restoration in lateral view (after Moy-Thomas and Westoll, 1935); (B) specimen from the Marl Slate, Middridge, figured by King (1850).

(Figure 10.11) Components of the fossil fish fauna from the German Kupferschiefer, probably closely comparable to that at Middridge. Palaeoniscids make up more than 90% of the assemblage, while the larger actinoptery-glans and the coelacanth constitute less than 5% at the top of the trophic pyramid (after Schaumberg, 1978).

(Figure 11.1) Palaeogeography of Britain in late Triassic time (after Bradshaw et al., 1992). Connection with the Tethyan ocean lies via the east and south, with the Boreal ocean via the north.

(Figure 11.2) British Triassic stratigraphy (after Warrington et al., 1980).

(Figure 11.3) Map showing main outcrop of Triassic strata in England and Wales, with GCR sites at Sidmouth and Aust Cliff indicated.

(Figure 11.4) The diversity of fishes in the Triassic and Jurassic periods (after McCune and Schaeffer, 1986). The width of each band at the middle of a division reflects the number of genera known from that division.

(Figure 11.5) Map of coastal outcrop of the Otter Sandstone Formation between Sidmouth and Budleigh Salterton, South Devon (from Benton and Spencer, 1995).

(Figure 11.6) View eastwards of Sidmouth cliffs, exposing the Otter Sandstone Formation in which fish and tetra-pod remains occur at various levels (photo: M.J. Benton).

(Figure 11.7) Otter Sandstone Formation fish: Dipteronotus cyphus Egerton.

(Figure 11.8) Aust Cliff, north of the Aust Service Area: the pale band of the Penarth Group high in the cliff face contains lenses of Rhaetic Bone Bed (photo: G. W Storrs).

(Figure 11.9) The Rhaetian at Aust Cliff: (A) geological map; (B) stratigraphical section (both from Benton and Spencer, 1995, after Hamilton, 1977).

(Figure 11.10) Fossils from the Rhaetic Bone Bed, Aust Cliff: (A) idealized characteristic tooth plate of Ceratodus latissimus Agassiz (c. 6 cm long) in occlusal aspect; (B), (C) Polyacrodus cloadcinus (Quenstedt), idealized tooth (c. 2.5 cm in antero-posterior length) in lingual and apical aspects; (D) Gyrolepis alberi Agassiz, idealized scale (c. 0.5 cm long; (E), (F) Nemacanthus monilifer Agassiz, reconstructed fin spine (c. 12 cm long) in transverse section and right lateral aspects; (G), (H) typical Lissodus minirnus (Agassiz) tooth (c. 0.3 cm in antero-posterior length, in lingual and apical aspects; (I), (J) Severnichthys acuminatus (Agassiz) neotype, dentary, BRSMG 7976 (c. 10.5 cm long) in left lateral aspect and medial aspect; (K) fin spine of the rare Palaeospinax rhaeticus Duffin (c. 7 cm long). (All figures from Storrs, 1994.)

(Figure 11.11) Representative fish remains from the Westbury Formation, Aust Cliff. (A)–(D) rostropre-maxillary of Severnichthys acuminatus (Agassiz) BRSMG Cd222, c. 3 cm long, in platal, anterodorsal and left lateral aspects. (E) idealized ?lower tooth of Pseudodalatius barnstonensis (Sykes), c. 0.3 cm long. (All figures from Storrs, 1994.)

(Figure 12.1) Map of the outcrop of Jurassic rocks in Britain (after Benton and Spencer, 1995) with GCR sites for fossil fishes indicated.

(Figure 12.2) Table of British Jurassic stratigraphy (modified from Benton and Spencer, 1995).

(Figure 12.3) Early Jurassic palaeogeography (from Bradshaw et al., 1992).

(Figure 12.4) The Lower Liassic shales and cementstones in Ware Cliffs, north-west of Lyme Regis harbour (photo: G.W Storrs).

(Figure 12.5) (A) Map of the coastal outcrop of the Lower Lias, Charmouth to Lyme Regis (after Benton and Spencer, 1995); (B) rock succession (after House, 1993).

(Figure 12.6) Liassic fishes from Lyme Regis: (A), Dapedium politus (Leach), x 0.5 (Photo: courtesy The Natural History Museum, London, T00144/A). Liassic fishes from Lyme Regis: (AI) (restoration), Dapedium politus x 0.2; (13) Chondrosteus acipenseriodes Egerton, x 0.1, restoration by Woodward (1895); (C) Chondrosteus aeipenseroides (Egerton) skeleton and body outline preserved, x 0.2; (D) the teleostPholidophorus sp., x 0.1, restoration by Woodward (1895); (E) the coelacanth Holophagus gulo Egerton, x 0.25, restoration by Woodward (1895); (F) the palaeoniscoid Coccolepis bucklandi Agassiz, x 0.5; (G) the elasmobranch Hybodus, x 0.1, restoration by Maisey (1982); (H) teeth of the elasmobranch Hybodus, isolated individual teeth and part of the dental array, x 1.0. Figures from Woodward (1889–1901) © The Natural History Museum, London. (Continued on p. 368.) (Figure 12.6) — contd. Liassic fishes from Lyme Regis: (I) array of Acrodus teeth, x 1.0 (from Woodward, 1895, © The Natural History Museum, London).

(Figure 12.7) Sketch map of the area around the Blockley Old Station Quarry GCR site, which is to the north-east of Blockley.

(Figure 12.8)A Blockley Station Quarry, close-up of north-west faces of the Upper Clays in the Lower Lias, with basal limestones (photo: K. Page). B Blockley Station Quarry, north-west faces of the Upper Clays in the Lower Lias, with basal limestones (photo: S. Metcalf)

(Figure 12.9) The elasmobranch fishes from Blockley Old Station Quarry: (A) restoration of Synechodus by Duffin and Ward (1983b); (B) denticles (x 30) and (C) teeth of Synechodus from Duffin and Ward (1983b); (D) views of tooth (holotype NHM P 60788) of Agaleus dorsetensis Duffin and Ward (1983b).

(Figure 12.10) (A), (B) Sketch map of the geology near Whitby with (C), rock succession in the local Upper Liassic (after Benton and Spencer, 1995). Fish bearing localities are marked on (A).

(Figure 12.11) Saltwick Bay, Whitby: the cliff at the left is of the Alum Shale, topped by Dogger Sandstone and boulder clay. The distant Nab headland is also of Alum Shale; the Jet Shale crops out on the lower tidal area of the beach beyond the Nab. Both formations have yielded fossil fishes. (Photo: BGS no. A5519, Crown Copyright reserved.)

(Figure 12.12) Upper Liassic osteichthyan fishes from the coast at Whitby: (A) Pachycormus curtus Agassiz, x 0.4, restoration of the head in lateral view by Lehman 1966; (B) Lepidotes semiserratus Agassiz, x 0.6, restoration by Jaekel 1929; (C) lateral views of Lepidotes semiserratus after Woodward 1895a; 1901, x 0.7; (D) Ptycholepis sp. x 0.25.

(Figure 12.13) Palaeogeography of the Middle Jurassic (after Bradshaw et al., 1992).

(Figure 12.14) Map of the mines and workings in the Stonesfield Slate around Stonesfield (after Aston, 1974).

(Figure 12.15) The Bathonian succession at Stonesfield, where the 'slate' occurs within the Taynton Limestone Formation (after Boneham and Wyatt, 1993).

(Figure 12.16) Vertebrate remains from the Stonesfield Slate were well known when Phillips published this plate in his Geology of Oxford and Thames Valley (1871): (A) all at x 1; a and b, Hybodus jugosus Phillips; c, H. polyprion Agassiz; d-1 H. grossiconus Agassiz; g, Strophodus magnus Agassiz; h, S. lingualis Phillips; i, S. tenuis Agassiz; j, Acrodus sp. (B) all at x 0.5; a, Hybodus dorsalis Agassiz; b, H. apicalis Agassiz; c, Nemacanthus brevis Phillips; d, Ischyodus sp.; e, granulated surface; _f, H. marginatus Agassiz; g, Asteracanthus tenuistriatus Agassiz; Leptacanthus striatus Agassiz; i, L. semistriatus Agassiz; j, Pristacanthus securis Agassiz. Several of these names are now invalid — the specimens belong to pre-established taxa. (Continued on p. 388.)Vertebrate remains from the Stonesfield Slate (C) Pholidophorus sp. in lateral view showing rare excellent preservation of fish in this facies, x 1.0 (Photo: courtesy The Natural History Museum, London, T00848/A).

(Figure 12.17) Fishes from the Stonesfield Slate. (A) The chimaeroid Ischyodus sp. in lateral view, x 0.5; (B) Allothrissops, x 0.3, restoration in lateral view (after Taverne, 1977).

(Figure 12.18) Photograph of the Kirtlington section, north face (photo: R Cottle).

(Figure 12.19) Amphibian remains from Kirtlington; the albanerpetonid Celtedens, restoration (after McGowan and Evans, 1995).

(Figure 12.20) (A) Sketch map of the vicinity of the Watton Cliff (West Cliff) GCR site, Upper Bathonian of Dorset; (B) stratigraphical section through the Forest Marble (after Holloway, 1985). (Continued on p. 396).(C) View of Watton Cliff facing west (photo: S.J. Metcalf).

(Figure 12.21) Elasmobranchs from the Bathonian: (A) Protospinax, x 0.1 restoration after Woodward 1919b. (B) Lissodus pattersoni Duffin (NHM P60706); a, occlusal view; b, labial view. L. wardi Duffin (holotype NHM P58701): c, occlusal view; d, lingual view; e, labial view; f, lateral view (from Duffin, 1985).

(Figure 12.22) The shales and cementstones of the Kimmeridgian at Kimmeridge Bay, facing south (photo: M.J. Benton).

(Figure 12.23) Fish from the Kimmeridgian: (A) Hypsocormus, sp. x 0.25 restoration by Woodward (1891, © The Natural History Museum, London); (B) Thrissops sp. x 0.5 restoration after Cox et al., 1988.

(Figure 12.24) The uppermost Jurassic and Lower Cretaceous shales and limestones in the northern part of cliff section at Durlston Bay (photo: J.L. Wright).

(Figure 12.25) (A) Sketch map of the northern part of Durlston Bay (after Clements, 1993); letters and numbers refer to Clements' labelling of the beds. (B) Cliff profile at Durlston Bay (after Clements, 1993; Benton and Spencer, 1995).

(Figure 12.26) Stratigraphical section through the Purbeckian at Durlston Bay (after Benton and Spencer, 1995, and based on Wimbledon and Hunt, 1993). Numbers are those of Clements (in Torrens, 1969a).

(Figure 12.27) Fossil fishes from the Middle Purbeck Beds of Durlston Bay. (A) Hybodus ensis Woodward, teeth x 1; (B) Lepidotes minor Agassiz, x 0.3; (C) Holophagus purbeckensis Woodward, x 0.2; (D) Macromesodon daviesi (Woodward), x 0.5; (E) Proscinates (Microdon) radiatus Agassiz, x 0.5; (F) Ophiopsis dorsalis Agassiz x 0.3; (G) Amiopsis damoni (Egerton), x 0.3; (H) Histionotus angularis Egerton x 0.5. Figures from Woodward (1889–1901) © The Natural History Museum, London.Fossil fishes from the Middle Purbeck Beds of Durlston Bay. (I) Pachythrissops laevis Woodward, x 0.25; (J) Caturuspurbeckensis Woodward, head x 0.5; (K) Asteracanthus verrucosus Egerton fin spine x 0.5; (L) Lepidotes sp. x 0.5 (Photo: courtesy The Natural History Museum, London, T00562/A).

(Figure 13.1) Map of outcrop of Cretaceous with GCR fish sites indicated (from Benton and Spencer, 1995).

(Figure 13.2) Cretaceous stratigraphy (from Benton and Spencer, 1995).

(Figure 13.3) (A) Map of the Lower Cretaceous Hastings Formation (Beds) and Weald Clay Formation of the Weald area of southeast England (after Cook, 1995); (B) geological map of Isle of Wight.

(Figure 13.4) Cliff End, Hastings, exposing Wealden sediments. Looking to the east. (Photo: S.J. Metcalf.)

(Figure 13.5) Stratigraphical log of Cliff End section, near Hastings (after Lake et al., 1987).

(Figure 13.6) Wealden fossil fish: (A), (B) Hybodus parvidens Woodward anterior and posterior teeth in (a) labial, (b) lingual and (c) medial view, NHM P 46930 31; (C)–(E) Lissodus rhizion 'teeth' in (a) lingual, (b) labial, (c) medial, (d) occlusal and (e) basal view: (D) the holotype NHM P 47144 (from Patterson, 1966); (F) Coelodus mantelli Agassiz, oral view of the right splenial bone with teeth, x 1, Wealdon, Hastings; (G) Coelodus mantelli left splenial, x 1, Wealdon, Hastings. Figures from Woodward (1889–1901) © The Natural History Museum, London.

(Figure 13.7) Brook–Atherfield Point section with locality details (after Benton and Spencer, 1995).

(Figure 13.8) Location map, Brook-Atherfield area (from Benton and Spencer, 1995).

(Figure 13.9) Wealden fossil fishes from Brook-Atherfield and other localities (after Woodward, 1917): (A) Hybodus ensis Agassiz, fin spine, x 0.5; (B) Coelodus multidens Woodward, right splenial with teeth, x 1.0; (C), (D) Pachythrissops vectensis Woodward: (C) crushed parts of right opercular region, x 0.5; (D) right lateral view of skull and axial skeleton, x 0.25; (E), (F) Hybodus basanus Egerton, restoration of the cranium in dorsal view and right lateral view of the skull with jaws, about half natural size. Figures from Woodward © The Natural History Museum, London.

(Figure 13.10) Speeton Cliffs, North Yorkshire, looking north-north-west. The Red Chalk lies near the base of the Lower Chalk and crops out in the foreshore below the beach in the foreground. Cliffs of Middle Chalk in the middle distance show small landslips. Fossil fish remains have been found in both of these formations. (Photo: BGS no. A5467, Crown Copyright reserved.)

(Figure 13.11) The Upper Cretaceous Chalk Group succession, northern and southern provinces (Wood and Smith, 1978; Owen, 1975). Correlation is provisional because of uncertainties in the biostratigraphical scales: a, belemnite; b, bivalve, c, ammonite; d, echinoid; e, crinoid; f, brachiopod.

(Figure 13.12) Photograph of Blue Bell Hill upper chalk pit: north-east face (photo: S.J. Metcalf).

(Figure 13.13) Fossil fishes from Burham Blue Bell Hill Pits (all from Woodwood, 1895a): (A) Cantioscyllium decipiens Woodward, anterior part of the skeleton; hy, hyoid arch; md, mandible; pct, pectoral arch; pq, pala-to-quadrate, x 0.75; (B), (C) Scyliorhinus antiquus ( = Scyllium antiquum) Agassiz, teeth in labial and lingual views, x 4.0 and x 0.5 respectively; (D) Scylliorhinus antiquus ( = Scyllium antiquum) Agassiz, dermal den-tides, c. x 10; (E) Ischyodus incisus Newton, left mandibular dental plate; outer, inner and upper views, x 1.5; (F), (G) Pachyrbizodus basalis Dixon; (F) part of the right premaxilla with teeth, x 1; (G) right dentary in lower and inner aspects, x 0.25; (H) Pachyrbizodus dibleyi Woodward, left maxilla in outer view, x 0.3.Fossil fishes from Burham Blue Bell Hill Pits (all from Woodwood, 1895a): (I) Neorhombolepis excelsus Woodward (from Kent), imperfect head and anterior abdominal region (N. punctatus) Woodward, (known from scales in Burnham area), X 0.6; (J) Enchodus pulchellus Woodward, holotype (NHM P1703) left-hand side of the head and opercular area„ x 0.9; (K) Sardinioides illustrans Woodward, holotype (NHM P3977) imperfect crushed specimen in ventral view, x 0.6; (L) Ichthyodectes tenuidens Woodward, right side of head and jaw, x 1; (M) Ichthyodectes sp., two vertebral centra, x 1.

(Figure 13.14) Totternhoe Chalk stratigraphical log (after P.J. Smart).

(Figure 13.15) Fossil chondrichthyan teeth genera from the Chalk at Totternhoe (after Cappetta, 1987). (A) Protosqualus sp., x 24; (B) Leptostyrax sp., x 1.25 in distal view; (C) Synodontaspis (Carcharias) striatula, x 2 in distal view; (D) Synechodus sp. x 5; (E) Paraorthacodus sp., x 4; (F) Squatirhina sp., x 10, in labial and distal views.

(Figure 13.16) Map of chalk pits around the Mt Caburn area, Lewes (after Lake et al., 1987).

(Figure 13.17) Chalk sections in the Machine Bottom Pit and Southerham Grey Pit (after Lake et al., 1987).

(Figure 13.18) Lithology and biostratigraphy of Chalk pits near Lewes (after Lake et al., 1987).

(Figure 13.19) Fossil fishes from the Chalk of the Lewes area (after Woodward, 1895b). All distorted head and jaw parts; (A), (A') Apateodus striatus Woodward, distorted head, right and left lateral views, both are c. x 1.0; (B) Hoplopteryx lewesiensis Woodward, restoration x 0.5; (C) Thrissopater megalops Woodward, head with left pectoral arch, x 0.6; (D) Tomognathus mordax Dixon, imperfect skull and left mandible, x 1; (E) Elasmodectes willetti Newton, right side of head and part of trunk, x 0.8.(F) Edaphodon agassizi (Buckland), left mandibular dental plate in labial, lingual and upper views, x 1; (G) Aulolepis typus Agassiz, incomplete head and trunk in left lateral view, x 3; (H)–(j) Cimolichthys lewesiensis Leidy: (H) head and opercular region in dorsal and left lateral views x 0.35, (I) damaged trunk and tail parts x 0.35, (J) labial aspect of left dentary x 0.9.(K) Macropoma mantelli Agassiz, crushed skull in right lateral view, x 0.6; (L) Enchodus lewisiensis Mantell, head and anterior part of trunk in lateral view, x 0.9; (M), (N) Halec eupterginus (Dixon): (M) head and abdominal region in lateral view, x 0.9, (N) head in lateral view, x 1.

(Figure 13.20) Photograph of the eastern face of Southerham Grey Pit section (photo: S. J. Metcalf).

(Figure 13.21) Photograph of Lime Kiln Quarries, the marly chalk seen (above the talus) in the left (north) face of the quarry is in the Zone of Holaster subglobosus; that in the lowest 3 m or so of the farther (east) face is in the Zone of Schloenbachia varians. (Photo: BGS no. 2962; Crown Copyright Reserved).

(Figure 13.22) Chondrichthyan fishes from the Chalk at Southerham Lime Kiln Quarry (A) Plicatolamna crassidens x 1.0; (B) Ptychodus decurrens Agassiz, a restoration of the lower jaw x 1.0; (C) Paranometodon angustidens x 2.0. (From Longbottom and Patterson, 1987.)

(Figure 13.23) Photograph of the southern face of Boxford Chalk Pit (photo: S.J. Metcalf).

(Figure 13.24) Fossil elasmobranchs from the Chalk at Boxford Chalk Pit. (A) Pararhinocodon angustidens, X 16; (B), (C) Paratriakis sp., rostral teeth, x 2.5; (D) Ganopristis sp. (after Cappetta, 1987), oral tooth, x 10.

(Figure 14.1) Palaeogeographical maps of southern Britain and adjacent mainland Europe during part of the Palaeogene (Early Cenozoic) (after Curry, in Duff and Smith, 1992).

(Figure 14.2) Summary table of Early Cenozoic stratigraphy in southern England (based on Curry, in Duff and Smith, 1992), ages from Cowie and Bassett (1989).

(Figure 14.3) The section of Thanet Beds overlying eroded Chalk exposed in the Pegwell Bay cliffs (after Shephard-Thorn, 1988).

(Figure 14.4) (A) Sedimentary log of a generalized vertical section of the Palaeocene and Lower Eocene in the Herne Bay Cliffs and foreshore (after Ward, 1978a). (B) London Clay exposed in Beltinge Cliff, Herne Bay, view to the west from Bishopstone Glen (photo: BGS no. A7933; Crown copyright reserved).

(Figure 14.5) Fossil fishes from Herne Bay, Upnor and Abbey Wood. (A) Notidanodon sp., lingual view of lower anterior tooth, x 0.65, Herne Bay (from Cappetta, 1987); (B) elasmobranch teeth from Abbey Wood, Odontaspis (Synodontaspis) macrota (Agassiz) x 1.3; (C) O. striata (Winkler) (after White, 1931) x 2. (Continued on page 496.)Fossil fishes from Herne Bay, Upnor and Abbey Wood. (D) Palaeogaleus vincenti (Hooker and Ward) x 1; (E) Galeorhinus lefevrei Gunn, x 0.1 (after Cappetta, 1987).

(Figure 14.6)A Upnor sand pit, the eastern face exposing the upper part of the Oldhaven Formation (Photo: S.J. Metcalf.).B Exploratory trench in the Blackheath and Oldhaven Formation in Abbey Wood, Blackheath, (Photo: S.J. Metcalf, 1994).

(Figure 14.7) Fossil fishes from the London Clay at Bognor Regis. (A) labial, mesial and lingual views of Isistius triangularis (Probst), lower lateral tooth (after Capetta, 1987), x 9; (B) Isurolamna affinis (Casier): (B') anterior tooth, labial and lingual views x 2; (B") antero-lateral tooth, labial view x 2; (C) Myliobatis dixoni Agassiz median tooth, basal view, x 2; (D) Otodus obliquus Agassiz: (D') upper lateral tooth labial view x 1; (D") lower anterior tooth, lingual and labial views, x 1. (B)-(D) from Kemp et al., 1990).

(Figure 14.8) The Isle of Sheppey: (A) sketch map, (B) section.

(Figure 14.9) Common elasmobranch fossils from the London Clay, as at the Isle of Sheppey (after Kemp et al., 1990). (A) Squatina prima (Winkler), lateral tooth (left) and anterior tooth (right), x 2.5. (B) Physogaleus secundus (Winkler) female antero-lateral tooth, x 2, lingual and labial views (left) and male antero-lateral tooth, lingual and labial views (right). (C) Carcharias hopei (Agassiz), lower anterior tooth, x 1.23, labial lateral and lingual views (left) and upper lateral tooth, lingual view (right). (D) Aetobatus irregularis (Agassiz), single tooth from lower dentition, x 1.2, basal and occlusal views. (E) Burnhamia daviesi (Woodward), tooth, x 1.25, occlusal, basal and lateral views.

(Figure 14.10) Uncommon teleosts from the London Clay at Sheppey (after Casier, 1966, © The Natural History Museum, London). (A), (B) Ampheristus toliapicus Konig, reconstruction of the skull, right side and dorsal views; (C) Brychaetus muelleri Woodward, right side of head; (D) Eocoelopoma hopwoodi Casier, reconstruction of skull in dorsal and right side views; (E) Eothynnus salmonens Woodward, reconstruction of head in right side view; (F) Promegalops signeuxae Casier, right lateral view of skull. All figures c. x 0.5. (Continued on page 514.)Uncommon teleosts from the London Clay at Sheppey (after Casier, 1966, © The Natural History Museum, London). (G) Percostoma angustum Casier, reconstruction of skull in right side view; (H) Sciaenurus bowerbanki Agassiz, head in right profile; (I) Rhinocephalus planiceps Casier, skull in left side view; (j) Wetberellus cristatus Casier, incomplete skull in left profile. All figures c. x 0.5.

(Figure 14.11) 'The Cliff' section, looking east, soil on London Clay, Burnham-on-Crouch, (photo: S.J. Metcalf).

(Figure 14.12) Chondrichthyes from Sheppey and Burnham-on-Crouch. Sheppey: (A) Notorhynchus serratissimus upper anterolateral tooth, x 5. Burnham-on-Crouch: (B) Hexanchus agassizi, lower lateral tooth, x 5; (C) Hypotodus verticalis, lower anterior tooth, x 4; (D) Odontaspis winkleri, lower lateral tooth, x 4; (E) Xiphodlamia ecocaena, x 3; (F) Megascyliorhinus cooperi, x 10; (G) and (H) Triakis wardi: (G) lower lateral tooth, X 2; (H) upper lateral tooth, X 2. (All figures after Cappetta, 1987.)

(Figure 14.13) Sketch map of the foreshore at Bracklesham Bay, West Sussex (after Curry et al., 1977).

(Figure 14.14)A Exposure of the Barton Clay, looking east from below Highcliffe, in the vicinity of Barton-on-Sea, Hampshire (photo: S.J. Metcalf). B Section from Mudeford to Milford-on-Sea.

(Figure 14.15) Headon Beds at Hordle Cliff, Hampshire (after Cray, 1973).

(Figure 15.1) Cladogram of the fish–tetrapod transition in Late Devonian–Early Carboniferous times (after Ahlberg and Milner, 1994). The taxa are grouped on the basis of shared derived characters. The characters shared by the panderichthyids and the tetrapods appear to be adaptations to shallow-water habitats. Devonian tetrapods retained lateral line systems and so must have been at least semi-aquatic. The black dots represent taxa known from a single locality; bars show the stratigraphical range of taxa known from several levels. The osteolepiformes originated in Middle Devonian times, perhaps some 10 Ma earlier.

(Figure 15.2) Devonian tetrapods: (A) the sarcopterygian Panderichthys (after Janvier, 1996); (B), (C) Ichthyostega; (D) Acanthostega (both after Coates and Clack, 1995); (E) Tulerpeton (after Lebedev, 1990; Janvier, 1996). These animals were all up to c. 1 m long.

(Figure 15.3) Map of GCR sites and localities yielding amphibians mentioned in the text below. C, Coton End, Warwickshire; EK, East Kirkton, Edinburgh; G, Guy's Cliffe, Warwickshire; H, Headon Hill, Isle of Wight; Ho, Hordle Cliff, Hampshire; K, Kirtlington, Oxfordshire; S, Stonesfield, Oxfordshire; Sc, Scaat Craig, Moray; Si, Sidmouth Cliffs, Devon.

(Figure 15.4) Vertebrates from Dora in a reconstruction by Janvier (1996; based on Milner et al., 1986). Namurian tetrapods (A) Crassigyrinus and (B) Spathicephalus are supposedly primitive, the others are anthracosaurs: (C) Protogyrinus and Eoherpeton. Also present is the acanthodian (F) Gyracanthus, primitive actinopterygians (E) and lungfish (D).

(Figure 15.5) Sketch map of the GCR Site at East Kirkton Quarry (after Rolfe et al., 1994).

(Figure 15.6) Succession in the top 250 m of the West Lothian Oil Shale Formation (after Whyte, 1994).

(Figure 15.7) Fossil acanthodian and actinopterygian fishes from East Kirkton. (A) A very small acanthodian (RSM G 1993.6.1) from the Little Cliff Shale: Br, branchiostegal rays; dsp, dorsal spine; 11, lateral line; m, mandible; oto, otoliths; pcg, pectoral girdle; pcsp, pectoral spine; pq, palatoquadrate; pvsp, pelvic spine; sc, scapula; at, anchylosed tooth. (B) An articulated specimen of Eurynotus with head bones, tooth plates and scales (after Coates, 1994): cl, cleithrum; d, dentary; mx, maxillary; ptp, pterygoid; scl, supraclei-thrum; sop, suboperculum, tp, tooth plate. (C) a diagrammatic representation of the total acanthodian and actinopterygian fauna, based on scales and drawn to estimated proportional sizes: Ac, acanthodian material; Ac?, possible climatiid tesserae; a, actinopterygians; b, isolated scales; c, spp. of Elonichthys; d, spp. of Mesopoma; e, Cosmoptychius; f, eurynotid material; j, juvenile specimens.

(Figure 15.8) Balanerpeton woodi Milner and Sequiera; photographs under UV light (x 1) of (A) the skull and anterior part of the axial skeleton in dorsal view (GLAHM V2051) and (B) posterior part of the axial skeleton, hind limb and squamation in dorsal view (GLAHM V 2052). From the Upper Viséan, Bathgate. (Photographs courtesy of the Hunterian Museum, Glasgow)

(Figure 15.9) Balanerpeton woodi Milner and Sequiera (based on GLAHM V 2051): (A) the dorso-ventrally compressed skull in palatal view; (B) restoration of the skull in dorsal view showing ornamentation on the outer surface of bones; (C) the skull restored in palatal view with marginal teeth and fangs and surface ornamentation indicated; (D) restoration of the incomplete skeleton in dorsal view; (E) the animal restored in dorsal view (after Milner and Sequiera, 1994).

(Figure 15.10) Ophiderpeton kirktonense Milner, holotype RSM G 1988.3.1; part of skull and vertebral column, the vertebrae 5–10 are restored from counterpart; ds, dorsal scales; h, hyoid; o, orbit (after Milner, 1994).

(Figure 15.11) Silvanerpeton miripedes Clack; an interpretative drawing of the skeleton (specimen UMXC V1317; after Clack, 1994).

(Figure 15.12) Eldeceeon rolfei Smithson; the skull and most of the post-cranial skeleton in a provisional restoration, based mainly on the holotype (RSM G 1990.7.1) from the East Kirkton Limestone (after Smithson, 1994).

(Figure 15.13) Westlothiana lizziae Smithson and Rolfe; reconstruction of the skull and skeleton based on both known specimens. (A) Skull in dorsal aspect; (B) skull in palatal aspect; (C) skull in lateral aspect; (D) restoration of incomplete skeleton (after Smithson et al., 1994.)

(Figure 15.14) Reconstruction of the environments represented in the vicinity of the East Kirkton Lake (after Durant, 1994). 1, intermittent rainfall; 2, recently expired volcano with lava flows; 3, erupting basaltic volcano with associated heavy rainfall; 4, fumaroles and hot springs emitting mineral-rich waters; 5, intermittent stream in flood transporting volcanic detritus from flanks of volcanoes; 6, terrestrial fossils incorporated in sediment during transport; 7, exposed lake shallows or floodplain with limestone and stromatolite debris; 8, deposition of graded units in lake.

(Figure 15.15) Reconstruction of the hot springs environment that is indicated in the East Kirkton Limestone. The temnospondyl Balanerpeton is in the foreground while an Eoherpeton-like anthracosaur (Eldeceeon) is in the middle distance, next to a fumarole. The eurypterid Hibbertopterus is at the water's edge and the pteri-dosperm Sphenopteris is shown in the bottom-left corner. (From Milner et al., 1986; Courtesy of the Hunterian Museum/Modern Geology)

(Figure 15.16) Section through the Headon Hill Formation at Headon Hill, Isle of Wight (after Cray, 1973).

Tables

(Table 1.1) Classification of fishes and amphibians (living = t; paraphyletic exceptions = *).

(Table 1.2) Fossil fish sites described in this volume. See also (Figure 1.15).

(Table 3.1) Section of the Ledbury Group, described as 225 m of alternating thick red mudstones and red to purple micaceous sandstones with abundant vertebrates in the lowest 120 m

(Table 3.2) Comparison of faunal assemblages at Ludlow railway cutting, Temeside and Ledbury.

(Table 4.1) Thelodont faunas in the Upper Silurian-Lower Devonian Old Red Sandstone of the Anglo-Welsh Basin. They are based on scale species (largely after Turner, 1973): recent discoveries, as yet unpublished, show that other fish groups are also represented by scales and may be of similar stratigraphical value.

(Table 11.1) Aust Cliff section (based on Reynolds, 1946; Hamilton, 1977; Warrington et al., 1980).

(Table 12.1) Designated stratal units in the Purbeck Limestone Group of the Durlston Bay section. Discoveries of fossils, including vertebrates, can be referred to precise horizons in this unbroken sequence, and related to the ostracod zonation.

(Table 14.1) Table of Palaeogene formations and environments in southern Britain.

References