Trace fossils and pseudofossils from the Wealden strata (non-marine Lower Cretaceous) of southern England

Cretaceous Research 26 (2005) 665e685 www.elsevier.com/locate/CretRes

Trace fossils and pseudofossils from the Wealden strata (non-marine Lower Cretaceous) of southern England Roland Goldring a,*, John E. Pollard b, Jonathan D. Radley c a

Geoscience Building, Postgraduate Research Institute for sedimentology, School of Human and Environmental Sciences, University of Reading, P.O. Box 227, Whiteknights, Reading RG6 6AB, UK b School of Earth, Atmospheric and Environmental Sciences, University of Manchester, Manchester M13 9PL, UK c Warwickshire Museum, Market Place, Warwick, Warwickshire CV34 4SA, UK Received 18 March 2004; accepted in revised form 9 March 2005 Available online 6 September 2005

Abstract The trace fossils of the Wealden (non-marine Lower Cretaceous) of southern England are described. Sixteen invertebrate ichnotaxa include Agrichnium fimbriatus, Beaconites antarcticus, B. barretti, Cochlichnus anguineus, Diplichnites triassicus, Diplocraterion parallelum, Lockeia siliquaria, L. serialis, Monocraterion cf. tentaculum, Palaeophycus striatus, P. tubularis, Planolites montanus, Protovirgularia rugosa, Rhizocorallium isp., Scoyenia cf. gracilis, Unisulcus minutus, insect and root traces. Tetrapod tracks and trackways include tridactyl Iguanodontipus burreyi and other ornithopods, theropod, and tetradactyl sauropod (or possibly ankylosaur), together with extensive dinosaur tramplings. Coprolites are referred to two broad types: spiral, with or without included fish scales (attributable to sharks), and elongate and irregular (possibly produced by reptiles). A skinprint and two types of pseudofossil are also included. Five environmental associations are recognised: (1) lacustrine/lagoonal; (2) brackish incursions (flooding events) into the lacustrine/lagoonal environment; (3) a marginal lacustrine association with fluvial input; (4) a fluvial (lacustrine delta) association; (5) floodplain sediments (seasonal wetlands). These associations are assigned to the fluviallacustrine Scoyenia Ichnofacies and the incursions to Glossifungites Ichnofacies. Ó 2005 Elsevier Ltd. All rights reserved. Keywords: Ichnology; Cretaceous; Wealden; Southern England

1. Introduction Early Cretaceous Wealden sediments in southern England are present in two distinct basinal settings (Fig. 1), collectively referred to as the Weald-Wessex Basin, which display distinct facies, and are referred to as the Weald and Wessex Sub-basins. Correlation of the Wealden of the two sub-basins is tentative (Fig. 2), and distinctive palaeoenvironments are represented (Allen and Wimbledon, 1991). The non-marine Lower Cretaceous Wealden sediments of southern England contain a low * Corresponding author. E-mail address: [email protected] (R. Goldring). 0195-6671/$ - see front matter Ó 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2005.03.001

diversity assemblage of trace fossils that add significantly to an appreciation of Wealden biodiversity. Trace fossils are common in several of the Wealden facies. Vertebrate footprints have attracted most attention, being first recognised in the Wealden of East Sussex by Tagart (1846). Over the years invertebrate traces and roots have been collected (Allen, 1962; Prentice, 1962), particularly in conjunction with insect remains. The traces can be found, often exquisitely preserved, on the soles of thin siltstones in marginal lacustrine/lagoonal facies. These delicate traces mostly occur as positive hyporeliefs, and represent dwelling excavations, feeding and sheltering burrows, and locomotion traces. Several of the more marginal sandy facies are extensively riddled by

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Fig. 1. Map of southern England showing distribution of Wealden and Lower Cretaceous rocks within the Early Cretaceous Wessex-Weald Basin, stratigraphy of the Vectis Formation in the Isle of Wight, and the locations of main sections referred to in the text; 1, Lulworth Cove (to west), Worbarrow and Mupe Bays; 2, Swanage; 3, Compton Chine; 4, Hanover Point; 5, Cowleaze Chine; 6, Shepherd’s Chine; 7, Atherfield; 8, Sandown Bay; 9, Clockhouse Brickworks Pit; 10, Bexhill; 11, Hastings; 12, West Hoathly. (Based on Batten and Lister, 1988.)

finger-size burrows and plant roots, particularly, but rarely, associated with Equisetites. Most of the burrows can be attributed to Beaconites (Goldring and Pollard, 1995), a well-known ichnotaxon in non-marine sediments from the Silurian onwards. These particular burrows have attracted much attention, having been attributed to Ophiomorpha by Kennedy and Macdougall (1969). Ophiomorpha is a pellet-lined burrow that typically occurs in marine or marine-influenced sediments and today represents the work of a callianassid (shrimp) and related crustaceans. Its identification in the Wealden was taken to indicate brackish incursions (e.g., Gallois and Worssam, 1993), though Stewart (1978) demurred. Beaconites is a back-filled burrow, in which taphonomy and diagenesis has often intensified the morphology to give a nodular appearance to the burrow margin, especially with bed-junction preservation. Traced laterally a mud-chip and sand-fill can pass into a wholly sand- or a wholly mud-fill, reflecting the passage of the producer through different lithologies. Although most of the distinctive invertebrate trace fossils in the Wealden sediments were formed endogenically, i.e. below the substrate surface at bedding surfaces, there are a few surface (epigenic) traces which were preserved under quiet hydraulic conditions. But such traces are relatively poorly defined. In general the trace fossils represent only single colonizations, in contrast to the intense bioturbation and complex tiering that is typically seen in marine shelf sediments.

Radley (1994a, c) and Radley et al. (1998a) reported simple borings from gastropod and bivalve shells in the Wessex and Vectis formations of the Isle of Wight. Similar traces also occur in large Viviparus shells in the Upper Weald Clay Formation. Jarzembowski (1990) described beetle larva borings from fossil wood in the Weald Clay. Although insect borings in fossil wood (see below, Paleoscolytus, and Francis and Harland, unpublished manuscript) have been reported from Wealden sediments, and body fossils of termites and wasps are known (Jarzembowski, 1995), no trace fossils of complex nests of social insects or dung beetles are known which characterise the Coprinisphaera Ichnofacies (Genise et al., 2000). Caddisfly larva cases are uncommon in the fossil record. They are regarded as trace fossils, ‘‘aedifichnia/calichnia’’, by Bromley (1996). Caddisfly wings have been found in the Weald Clay at Clockhouse Brickworks pit, near Capel, Surrey, from where many of the traces described below were collected, including several cases. The latter (Jarzembowski, 1995), belonging to several ichnotaxa, are mostly isolated, flattened, tapering tubes up to 35 mm long and 6.5 mm wide, composed of conchostracan valves with some plant material, fish bones, pellets or bivalve fragments. Jarzembowski (1995) also noted a cliff fall from the Ashdown Formation at Hastings which has yielded hundreds of caddis cases. Bioerosion of the vertebrate skeletal elements was recorded by Martill and Naish (2001a). Surface pitting

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2.

3.

4.

5. Fig. 2. Generalized correlation of the marine stages of the Lower Cretaceous with the stratigraphy of the Wealden sediments in the Wessex and Weald Sub-basins. (Based on Allen and Wimbledon, 1991.)

(2e5 mm in diameter) (Martill and Naish, 2001a, fig. 4.1) and asterorhizal borings (0.25e0.5 mm in diameter) (Martill and Naish, 2001a, fig. 4.2) may be attributed to fungal or bacterial attack. Occasional bite marks are also recorded (Naish, 1999). The trace fossils represented here do not constitute a monographic treatment. Most have come from the collections made by E.A. Jarzembowski and A.J. Ross during searches for insects.

2. Environmental and ichnofacies significance of trace fossils Five associations of invertebrate trace fossils are recognised: 1. A lacustrine/lagoonal association where the traces are typically found on the soles of thin calcareous siltstones and sandstones in the Weald Clay Group, Wadhurst Clay and Grinstead Clay formations (Allen and Wimbledon, 1991 and references therein), and Vectis Formation (Stewart et al., 1991). These have been identified as Cochlichnus, Diplichnites,

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Lockeia, Palaeophycus, Planolites and Unisulcus, with rarer Lockeia. Vertebrate footprints may be present with this association. Brackish incursions (flooding events) into the lacustrine/lagoonal environment with Diplocraterion, recorded from the upper Vectis Formation (Shepherd’s Chine Member) (Stewart et al., 1991). A marginal lacustrine association with fluvial input comprising Beaconites antarcticum, Cochlichnus, Scoyenia gracilis, Lockeia and Protovirgularia. This is encountered in the Weald Clay, lower Wessex Formation of Dorset, and the upper Wessex Formation and Vectis Formation (Barnes High Sandstone) of the Isle of Wight (Wach and Ruffell, 1991). Dinosaur footprints may be present with this association (An alternative environmental interpretation for the Barnes High Sandstone, laterally migrating tidal sand bar and channel complex, was suggested by Yoshida et al., 2001). A fluvial (lacustrine delta) association with Beaconites barretti and Planolites in the Ashdown Formation (Allen and Wimbledon, 1991 and references therein; Goldring and Pollard, 1995). Dinosaur footprints may be present with this association. Floodplain sediments and seasonal wetlands (Wright et al., 2000 and references therein), associated with point bars and channel plugs (Stewart, 1981a,b; Insole and Hutt, 1994) of the Wessex Formation of the Isle of Wight, associated with Beaconites antarcticus, B. barretti, Cochlichnus and Lockeia, and dinosaur footprints. Surprisingly the palaeosols of the Wessex Formation of the Isle of Wight (Wright et al., 2000) have not, so far, yielded characteristic trace fossils such as Taenidium (adhesive meniscate burrows; Hasiotis, 2004). The Ardingly Sandstone Member (Lower Tunbridge Wells Sand Formation) of the Weald is inferred to represent a braidplain facies. Beaconites is not recorded.

The invertebrate trace fossils shown in Figs. 3e11 all indicate formation in an entirely aquatic environment. The ichnofacies significance of the Wealden trace fossil associations are restricted to those of aquatic marginal ichnofacies within the diversity of recently defined continental ichnofacies (Bromley, 1996; Buatois et al., 1998; Genise et al., 2000). Associations 1, 3, 4 and 5 belong to the broadly defined archetypal Scoyenia Ichnofacies. Lacustrine/lagoonal Association 1, with surface locomotion and feeding traces of invertebrates preserved on the soles of thin event siltstones and sandstones formed in shallow water, has some elements of the Mermia Ichnofacies. However it is too sparse in ichnogenera and was too ephemeral an environment to fully represent this ichnofacies. Association 3 with marginal lacustrine and fluvial sediments and meniscate Beaconites burrows and probable Scoyenia gracilis, represents classic Scoyenia

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Ichnofacies (Seilacher, 1967; Frey et al., 1984; Buatois and Ma´ngano, 1995). The very limited ichnogenera in Association 4 reflect the ephemeral nature and high sedimentation conditions in a fluvial-lacustrine delta environment (Stewart, 1981a; Goldring and Pollard, 1995). The brackish incursion (flooding) events into the lacustrine/lagoonal environment indicated by Association 2 with protrusive and retrusive Diplocraterion within a nodular ironstone horizon represents an example of Glossifungites Ichnofacies (sensu MacEachern and Pemberton, 1992). However, we prefer to consider this as a Diplocraterion association reflecting a sedimentary hiatus with the establishment of an ephemeral shallow infauna of suspension feeders of more marine affinity, rather than a marine related ‘‘Glossifungites’’ firmground surface (Pemberton and Frey, 1985; Gingras et al., 2001). 3. Invertebrate ichnotaxa Ichnogenus Agrichnium Pfeiffer, 1968 Agrichnium fimbriatus (Ludwig, 1869) Fig. 7D Description. Groups of narrow hypichnial horizontal burrows (0.1 mm wide or less), slightly sinuous and somewhat intertwined. Width of groups about 1 mm. Common on upper and lower surfaces of siltstones and fine sandstones. Distribution. Weald Clay Group. Ichnogenus Beaconites Vialov, 1962 Remarks. Beaconites is a back-filled burrow that is common in non-marine facies. Taenidium Heer, 1877 is a similar back-filled burrow that is generally rather better organised, and found in marine settings. It has taxonomic priority, and has also been applied to nonmarine burrows. At present it is uncertain how the two ichnogenera might be separated. Beaconites antarcticus Vialov, 1962 Fig. 3A, B Fig. 3. A, B, Beaconites antarcticus Vialov, 1962. A, two irregularly meniscate burrows with mixed sand and mud-chip fill, and false branching. Hypichnial positive reliefs of interface burrows; Lower Weald Clay, Clockhouse Sandstone Member, Clockhouse (Butterley) Brickworks pit (TQ 175 385); Booth Museum, Brighton 018890; !0.5. B, hypichnial expression of nodular horizontal burrow resembling Ophiomorpha (a) and vertical burrow with nodular or bioglyph markings on basal rounded terminations (b). Location as for A; !1. C, Beaconites barretti Bradshaw, 1981; horizontal straight to sinuous cross-cutting burrows with regular sand-mud meniscate fill in ripple cross-laminated sandstone; base of Lee Ness Sandstone, Ashdown Formation, shore east of Lee Ness Ledge (TQ 870 112), near Hastings, Sussex. Field photograph; !0.7.

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Description. Finger-sized burrows, mostly beddingparallel or steeply inclined to normal, in cmedm thick beds of fine-grained sandstone. Diameter 5e18 mm, though a smaller range (8e11 mm) is given by Stewart (1978) for occurrences in the Wessex Formation of the Isle of Wight. It is possible that two size classes are present, with a larger burrow averaging 15 mm in width and a smaller about 5 mm wide. Burrows relatively straight and parallel to stratification, or inclined to normal; burrow variably filled, most frequently by small and often angular mud-chips and sand to form coarse menisci, or by mud, or relatively clean sand locally with a discontinuous, thin mud lining. Entrances not clearly observed, but burrows may expand somewhat towards the upper surface of sandstone beds. Terminations rounded, not expanded. No true branching and no networks or boxworks. Distribution. Tilgate Stone facies of lower Wadhurst Clay Formation at Cliff End, Hastings; sand members of the Weald Clay Group, e.g., Horsham Stone; Clockhouse Sandstone, Clockhouse Brickworks pit (TQ 1770 3865) (Goldring and Pollard, 1995, with other localities); Upper Tunbridge Wells Sand Formation, Freshfield Lane Brickworks (TQ 385 261) near Haywards Heath, Sussex; Wessex Formation of south Dorset (Lulworth Cove, Mupe Bay, Swanage Bay); Isle of Wight, Wessex Formation, common at many levels, e.g., Hanover Point Sandstone (Festained), Sudmoor Point Sandstone (taphonomic expression as ‘‘Skolithos’’). Uncommon in mudstones. Top of Wessex Formation (Stewart, 1978), Vectis Formation (White Rock at Cowleaze Chine) (Stewart, 1978). Beaconites barretti Bradshaw, 1981 Fig. 3C Description. The burrows are 10e15 mm in diameter, locally narrowing and expanding, normal but mainly parallel to bedding, and passing in and out of the plane of lamination. In plan, burrows are curved to weakly sinuous for some tens of centimetres. No branching but frequent crossings (often emphasised by contrast in burrow-fill). Margins sharply defined. Fill with marked differentiation, from mud to sandy mud-chip to clean sand. Meniscate structure of fill most prominent in sandy mud-chip fill where it can show regular thin menisci. Distribution. Lee Ness Sandstone, Ashdown Formation, Lee Ness Ledge (TQ 866 109) coastal section, Hastings, Sussex (Goldring and Pollard, 1995). Ichnogenus Cochlichnus Hitchcock, 1858 Cochlichnus anguineus Hitchcock, 1858 Figs. 4A, B, 9A

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Description. Regularly meandering, smooth, nonbranching burrow of uniform width, resembling sine curve. Larger burrows occasionally convergent on Lockeia (Fig. 4A). C. anguineus is a common trace on upper and lower surfaces of siltstones or sandstones as narrow ridges. Two sizes are present: smaller, with burrow width of ca. 1 mm or somewhat less, wavelength 4e6 mm and amplitude 2e3 mm, and burrow lengths of up to ca. 100 mm are preserved. Larger burrows with width ca. 3 mm and wavelength 14e15 mm. Remarks. Allen (1962, p. 235, pl. 9, fig. A) referred to the aggregations as ‘‘crimping iron’’ traces. In the Silesian (Coal Measures) Eagar et al. (1985) and Pollard and Hardy (1991) recorded a range of size including the above, and also a frequent association with Lockeia. Elliott (1985) also recorded a greater range of wavelength for small forms from the Silesian. Nematodes are generally regarded as the producers of the fine sinuous burrows (Mousa, 1970; Metz, 2000), although insect larvae and worms can produce similar trails (Metz, 1987, 1995). Distribution. Smaller forms in the Weald Clay Group, Wadhurst Clay, Grinstead Clay and Vectis formations; larger forms in the Lower Tunbridge Wells Sand Formation (Ardingly Sandstone Member). Ichnogenus Diplichnites Dawson, 1873 Diplichnites triassicus (Linck, 1943) Fig. 9A Description. Trackway consisting of two parallel rows of simple or crescentic tracks arranged normal or slightly oblique to mid-line of the trackway. Preserved as positive hypichnia on base of a thin sandstone bed. Trackway data: external width 12 mm; internal width 8 mm; tracks 1 ! 2 mm, spaced 2 mm apart. Left track row: length 26.5 mm with 10e12 tracks in the row. Right track row: length 35 mm with 8e9 tracks in a straight row and ca. 6 tracks at a slight offset angle (Z en echelon series). Remarks. This trackway is very similar in form, size and arrangement of simple tracks to Diplichnites [Acripes] triassicus described from Triassic fluvial sediments (Linck, 1943; Bromley and Asgaard, 1979; Pollard, 1985). The trackway width, inferred number of legs in a series (6e9) observed in the en echelon rows, track spacing and mode of locomotion appear similar to tracks produced by crustaceans, most likely notostracans close to Triops (Pollard, 1985). Although body fossils of Triops have not been

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Fig. 4. A, B, Cochlichnus anguineus Hitchcock, 1858. A, C. anguineus apparently converging on Lockeia siliquaria James, 1879 as hypichnia associated with synaeresis cracks; Hastings Group, Lower Tunbridge Wells Sandstone, Hook Quarry, West Hoathly, Sussex (TQ 355 313); Reading University, Geology Archive S38687; !0.7. B, upper surface of thin siltstone biscuit, C. anguineus associated with small depressions (Lockeia) and Unisulcus (grooves), Vectis Formation, Atherfield coastal section, Isle of Wight (SX 450 792); Maidstone Museum MNEMG 2000.3; !1. C, Palaeophycus tubularis Hall, 1847, upper surface of thin siltstone with depressions corresponding with entry of burrows and with many collapsed burrows. Many of the burrows parallel to surface are unfilled and collapsed, but ‘‘levees’’ were formed as animal moved over the mud/silt interface, which was at a shallow depth. Clockhouse Sandstone Member, Lower Weald Clay Formation, Clockhouse (Butterley) Brickworks pit (TQ 175 385); Maidstone Museum MNEMG 2000.7; !1. D, G, Palaeophycus striatus Hall, 1852; hypichnial ridges on sole of siltstone associated with Unisulcus minutus. Lower Weald Clay Formation, Clockhouse (Butterley) Brickworks pit (TQ 175385). D, Maidstone Museum MNEMG 2000.7; G, MNEMG 2000.10; !1. E, F, Palaeophycus striatus, positive hypichnia. Lower Grinstead Clay Member, Philpot’s Quarry, West Hoathly, West Sussex (TQ 355 322); Maidstone Museum MNEMG 2000.15; E, !1.5; F, !0.5.

recorded from Wealden sediments, a form very close to the allied Lepidurus is known from Lower Cretaceous rocks of Turkestan (Chernyschev, 1940; Tasch, 1969). Also the range of environments from which modern

faunas of Triops are known include fluvial and lacustrine habitats in humid sub-tropics similar to the Wealden (e.g. Far East paddy fields; see review in Pollard, 1985).

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Distribution. Upper Weald Clay, Alfold Sand Member, Smokejacks Brickworks, Ockley (TQ 113 373). Ichnogenus Diplocraterion Torell, 1870

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Diplocraterion parallelum Torell, 1870 Figs. 5AeC, 6 Description. Vertical, U-shaped burrow with spreite (the successive margin of a displaced burrow formed as

Fig. 5. AeC, Diplocraterion parallelum Torell, 1870. A, C, broken blocks of ironstone concretions with protrusive and retrusive structure. A, protrusive form, narrowing downwards from omission surface; MNEMG 2000.6. C, with protrusive and retrusive and pellet-filled spreite; MNEMG 2000.5. B, group of retrusive burrows on sole of shell bed (Fig. 6), with mudrock partly removed. Some specimens broken; MNEMG 2000.4. All specimens from Vectis Formation, Atherfield coast section, Isle of Wight (SZ 450 792); !1. D, Cast of small ornithopod footprint (pes); limestone bed 1 (Diplocraterion limestone), Shepherd’s Chine Member, Vectis Formation; at base of cliff between Shepherd’s Chine and Atherfield Point (SZ 448 795); M. Green collection. Scale bar represents 5 cm. E, H, Iguanodontipus burreyi Sarjeant, Delair and Lockley, 1998. E, cast of footprint (pes); Wessex Formation; Hanover Point (SZ 379 837). Dinosaur Isle, Sandown: 5419; H, cast of footprint (pes) in loose block; lower part of White Rock, Vectis Formation, Cowleaze Chine foreshore (SZ 444 801) Isle of Wight. Scale bars represent 10 cm. F, Sauropod or ankylosaur, cast of footprint (pes), Wessex Formation, Brook Bay, Isle of Wight (SZ 382 837). Dinosaur Isle, Sandown: 6508. Scale bar represents 10 cm. G, cast of theropod pes, Vectis Formation, probably Shepherd’s Chine Member, Cowleaze Chine beach (SZ 444 800), Dinosaur Isle, Sandown: 5768. Scale bar represents 10 cm.

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producer shifts broadside through sediment) connecting the arms. Two modes of preservation are present in the Shepherd’s Chine Member (upper Vectis Formation) of the Isle of Wight: 1. (Figs. 5B, 6) Hypichnial preservation as prominent, short tongues normal to the sole of a thin coquina, with a compressed elliptical cross-section, rarely a somewhat dumb-bell outline. Structure filled with fragmentary bioclastic material, with a slight indication of retrusive lamination. No preferred orientation and burrows frequently cross-cut. Width 10e20 mm, thickness 4e5 mm, maximum depth of sole structure, l5 mm. 2. (Fig. 5A, C) In association with a 50e70-mm-thick clay ironstone concretionary layer, representing firmground colonization; protrusive and retrusive, or partly protrusive, partly retrusive, vertical to steeply inclined. Ironstone with irregular, penecontemporaneously eroded upper surface, overlain (but not encrusted) by a shaley layer rich in oysters. Size more variable than mode 1, with U-burrow width of up to 60 mm. Depth originally greater than thickness of concretionary layer. Diameter of marginal tube 4e6 mm. Burrows mudfilled and generally rather compressed. Thickness variable because of compaction of muddy, faecal-rich spreite. (Viewed from the side, weathered out faecal pellets may resemble scratch marks.) Remarks. Diplocraterion is a useful indicator of brackish to marine environments, though occurrences of mode 1 appear to be associated with low salinity. Occurrences in the Vectis Formation probably mark flooding surfaces over a firm substrate. Distribution. Shepherd’s Chine Member, Vectis Formation, Isle of Wight. Mode 1, ‘‘Diplocraterion’’ limestone, associated with theropod footcasts, approximately 17 m above base of member, intermittently exposed from the cliff top south-east of Cowleaze Chine (SZ 4444 8000) to a low bank south-east of Shepherd’s Chine (SZ 4491 7953) and in the sides of Shepherd’s Chine (Radley et al., 1998a). Poorly preserved examples occur in the correlative bed at Compton Bay (Radley and Barker, 2000). Mode 2, ‘‘Diplocraterion ironstone’’, approximately 10.5 m below the top of Vectis Formation at Atherfield (SZ 4491 7954) (Radley, 1994b), and in a similar position at Sandown Bay (Radley, 1994a) and Compton Bay (Radley and Barker, 1998). Ichnogenus Lockeia James, 1879 Lockeia siliquaria James, 1879 Figs. 4A, 7C

Fig. 6. Diagrammatic representation of the theropod trackway horizon, limestone bed 1 ‘‘Diplocraterion limestone’’; Shepherd’s Chine Member, Vectis Formation, Isle of Wight. The footprint was cast soon after formation by washed-in Viviparus. The mud substrate was then colonised by the Diplocraterion-producer before colonization was curtailed and then overwhelmed by a Filosina-rich ‘‘event’’ bed. (After Radley et al., 1998a.)

Description. Ovoid protruberances on soles of sandstones or siltstones corresponding to the infaunal crypt of a bivalved animal. Length approximately 20 mm. Occasionally seen in vertical sections of sandstone as superimposed (stacked) down-bent laminae (Allen, 1975, pl. 3, fig. B). Also, very small (ca. 1 mm) protubarances on siltstone soles and depressions on upper surfaces (which may represent the activity of ostracods). Remarks. Allen (1962, pl. 9) illustrated two instances of probable Lockeia with converging Cochlichnus, similar to an association interpreted by Hardy (in Eagar et al., 1985, p. 129, pl. 12, fig. F) as small worms attracted to an autochthonous decaying bivalve. In the Barnes High Sandstone Member the crypts can be attributed to unionid and corbiculid bivalves and are associated with adjustment (escape) structures (Radley et al., 1998b, fig. 2). Distribution. Lower Tunbridge Wells Sand Formation of the Weald and Barnes High Sandstone Member of Vectis Formation. A smaller form is present in the Wadhurst Clay Formation. Possible ostracod or conchostracan resting traces are common in the Weald Clay and Vectis Formation. Lockeia serialis Seilacher and Seilacher, 1994 Fig. 7C Description. Serially-aligned ovoid, almond-shaped protuberances of L. siliquaria on sole of bed. Remarks. These serially-aligned crypt casts are attributed to bivalve crawling activity (see also Protovirgularia rugosa). Distribution. Top of Barnes High Sandstone Member of Vectis Formation, associated with corbiculid and unionid bivalves and viviparid gastropods (Radley et al., 1998b). Ichnogenus Monocraterion Torell, 1870

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Fig. 7. A, B, Protovirgularia rugosa (Miller and Dyer, 1878). A, epichnial grooves leading to ovoid cubichnion (Lockeia siliquaria) (upper centre). B, detail with chevron ridges; approximately 1 m below top of Ardingly Sandstone Member, Lower Tunbridge Wells Sand Formation; Philpots Quarry, West Hoathly, West Sussex (TQ 355 322), A, Maidstone Museum MNEMG 2000.12; !0.5; B, MNEMG 2000.13; !1. C, Lockeia siliquaria James, 1879 (solitary hypichnia) and Lockeia serialis Seilacher and Seilacher, 1994 (serially-aligned hypichnia). Ripple-marked underside of quartz arenite slab, derived from uppermost 2 m of the Barnes High Sandstone Member, Vectis Formation, shore between Shepherd’s Chine and Cowleaze Chine (SZ 445 799); field photograph, scale bar represents 5 cm. D, Agrichnium fimbriatus (Ludwig, 1869); hypichnia with trace cutting tool-marked surface. Lower Weald Clay Formation, Clockhouse (Butterley) Brickworks pit (TQ 175 385), north-east of Horsham, Surrey; Maidstone Museum MNEMG.2000.2; !2. E, Rhizocorallium isp., ‘‘Tilgate stone bed’’, Upper Wadhurst Clay Formation, Freshfield Lane Brickworks, Sussex; Maidstone Museum MNEMG 2000.11; !1.

Monocraterion cf. tentaculum Torell, 1870 (cf. Jensen, 1997, pp. 62e64, figs. 40, 41) Fig. 9C Description. Shallow, conical, circular, positive hypichnion with two concentric rings, faint radial markings and a central nipple-like protruberance. External diameter 10 mm, central ‘‘nipple’’ ca. 1.5 mm, depth ca. 5 mm. Remarks. This specimen is similar to small buttonshaped positive hypichnia on soles of thin-bedded sandstones from the Lower Cambrian of Sweden

(Jensen, 1997) and Upper Carboniferous of central England (Eagar et al., 1985) who referred them to Mammilichnis Chamberlain, 1971 or Monocraterion. The nipple-shaped central protuberance distinguishes the specimen from Bergaueria Prantl, 1946 and most other plug-shaped burrows except perhaps Mammilichnis, which has a central protuberance and may or may not have radial ridges and concentric constrictions (Crimes et al., 1981; Pemberton et al., 1988). However, following Jensen we interpret the ‘‘nipple’’ to represent the broken vertical burrow below a shallow outlet funnel rather than a resting trace of an anemone-like organism.

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Distribution. Lower Tunbridge Wells Sand Formation (Ardingly Sandstone), Valanginian, Hook Quarry, West Hoathly, West Sussex (TQ 335 313). Ichnogenus Palaeophycus Hall, 1847 Palaeophycus striatus Hall, 1852 Fig. 4DeG Description. Straight to slightly curved hypichnial ridges with distinct thin muddy margin, sometimes with longitudinal striations. Silt-filled burrow, width l.5e 2 mm. Remarks. Some specimens resemble Scoyenia, a common non-marine filled burrow, but none of the Wealden material examined displays a meniscate backfill. Distribution. Weald Clay Group, Lower Grinstead Clay Formation and Vectis Formation. Palaeophycus tubularis Hall, 1847 Fig. 4C Description. Straight to curved hypichnia and epichnia with smooth margin and often with evidence of collapse of incomplete passive fill (Allen, 1975, pl. 3, fig. A). Width 2e3 mm. Distribution. On thin siltstones and very fine sandstones in the Weald Clay Group and Vectis Formation. Ichnogenus Planolites Nicholson, 1873 Planolites montanus Richter, 1937 Fig. 8B Description. Typically, short lengths of straight to curved hypichnial semireliefs or detached, sand- or siltfilled burrows; less commonly as epichnial structures. Width 1.5e3.0 mm, occasionally with compactional striations parallel to length of burrow or with weak annulation. Ranging into dense intertwining ramifications of burrows on soles of thin sandstones and siltstones, often with a slight protrusive structure in cross-section, and somewhat variable burrow diameter. In one instance loop-like aggregations on sole are suggestive of Fuersichnus Bromley and Asgaard, 1979. Distribution. Widespread, particularly on soles of thin sandstones or siltstones, or ‘‘riddling’’ muddy siltstones and sandstones in the Weald Clay Group and Vectis Formation. Ichnogenus Protovirgularia McCoy, 1850

Fig. 8. A, Unisulcus minutus Hitchcock, 1858; hypichnial ridges on sole of mm-thick siltstone associated with Palaeophycus isp. Lower Weald Clay Formation, Hauterivian; Clockhouse (Butterley) Brickworks pit (TQ 175 385); Maidstone Museum MNEMG 2000.9; !2. B, Planolites montanus Richter, 1937. Lower surface hypichnia of composite and partly bioturbated siltstone with muddy partings; Lower Weald Clay Formation, Clockhouse (Butterley) Brickworks (TQ 175 385); Maidstone Museum MNEMG 2000.8; !0.7.

Protovirgularia rugosa (Miller and Dyer, 1878) [taphonomic preservation Chevronichnus imbricatus Hakes, 1976] Fig. 7A, B Description. Shallow epichnial grooves bordered by ridges with variable morphology along a gently curved course. On ripple crests, ridges variable and often strongly asymmetrical. Weak chevron ridges in troughs. Grooves may lead to ovoid cubichnion as in Lockeia siliquaria. Remarks. This trace is probably attributable to the ploughing action of a bivalve leading to burial. Seilacher and Seilacher (1994) considered this ichnotaxon to occur only on the soles of ‘‘sandy tempestites’’ (event beds), and that it identified escape structures from a Lockeialike resting burrow. In the Wealden of Sussex and the Isle of Wight the long shallow grooves resemble those described and

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Fig. 9. A, Diplichnites triassicus Linck, 1943. Short biserial trackways preserved as hypichnia. Short lengths of Cochlichnus anguineus also present. Upper Weald Clay Formation (Barremian), Smokejacks Brickpit, Ockley, Surrey (MMEMG 2003.26); scale bar represents 2 cm. B, Scoyenia cf. gracilis White, 1929. Horizontal burrow in hyporelief with segmentally arranged scratch marks. Collapsed cross-section of similar vertical burrow (top left). (From Kennedy and Macdougall, 1969, pl. 88, fig. 1. Weald Clay Group, No. 7 sand), Nutbourne Brickworks, Hambledon, Surrey; scale bar represents 1 cm. C, Monocraterion cf. tentaculum Torrel, 1897 (cf. Jensen, 1997). Hypichnial preservation of vertical burrow. Lower Tumbridge Wells Sand Formation (Ardingly Sandstone), Valanginian, Hook Quarry, West Hoathly, West Sussex (TQ 335 313); scale bar represents 5 mm.

figured as Chevronichnus imbricatus by Hakes (1976) and Ma´ngano et al. (2002) from the Upper Carboniferous of North America. They were formed either epigenically or, more likely, just below a thin mud with the burrow passing from a repichnion to deeper burial and a cubichnion. The variable morphology reflects substrate variation with the troughs being wetter (and the sediment too liquid for penetration), and the ripple crest

probably emergent (with the sand too loose for burrowing). Only as the water table fell did the substrate become sufficiently dilatent for burrowing. Distribution. Weald Clay Group and Barnes High Sandstone Member, Vectis Formation. Ichnogenus Rhizocorallium Zenker, 1836

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Rhizocorallium isp. Fig. 7E Description. Small bedding-parallel U-burrows with closely spaced silt-filled tubes (3 mm in diameter; separation 3 mm) but apparently without spreite. Remarks. This trace, though smaller, resembles examples of R. jenense (where the spreite is often not present) as described from the tops of clay lenses intercalated between fluvial cross-bedded sands and gravels of the Upper Freshwater Molasse (Upper Miocene) of southern Germany (Fu¨rsich and Mayr, 1981). These examples were ascribed to excavations by mayfly larvae. Distribution. Base of a ‘‘Tilgate Stone bed’’, Upper Wadhurst Clay Formation. Ichnogenus Scoyenia White, 1929 Scoyenia cf. gracilis White, 1929 Fig. 10B Description. Horizontal burrows, unbranched, preserved as semi-relief hypichnia with longitudinal, commonly segmentally arranged scratch marks. Burrow diameter 5e10 mm. Vertical burrows of the same diameter may show similar bioglyph marks (Goldring and Pollard, 1995, fig. 3b). Internal meniscate burrow-fill not seen. Remarks. The specimens of these burrows originally described as ‘‘Ophiomorpha’’ or Spongeliomorpha by Kennedy and Macdougall (1969, text-fig. 1; pl. 87, fig. 1; pl. 88, fig. 1) are now lost and so could not be reexamined in this study. Goldring and Pollard (1995)

pointed out the close similarity of these burrows to type specimens of Scoyenia gracilis (White, 1929; Haubold, 1982; Frey et al., 1984) and how they may be related to meniscate-filled B. antarcticus. Distribution. Weald Clay Group, base of sandstone beds, especially Topley’s (1875) ‘‘No. 7 sand’’, Nutbourne Brickworks, Hambledon, Surrey (SU 973375). Ichnogenus Unisulcus Hitchcock, 1858 Unisulcus minutus Hitchcock, 1858 Figs. 4G, 8A Description. Narrow, hypichnial ridges, almost detached, or epichnial ridges, unlined, smooth (though with occasional faint ridges parallel to burrow length), straight to winding with tendency to form apparently irregular networks. Over-crossing at very similar depths produces false branching at moderate to high angles. Burrow fill of medium to coarse silt (contrasting markedly with lag on sole of bed). Definite terminations not recognised and burrow typically emerges from sole of bed. Rarely, narrow muddy streak within fill. Width less than l mm. Remarks. This fine trail or burrow is very similar to Helminthoidichnites tenuis (Buatois et al., 1996; Metz, 1995). However the size, predominance of straight elements and frequent tangled over-crossing (Fig. 4D, G) were features mentioned by Hitchcock (1858, p. 151, pl. 26, fig. 3). More meandering elements may approach Helminthopsis Metz (1995, pl. 4, fig. 1), and Prentice (1962, p. 182, pl. 7) commented on the similarity of this trace to Paleodictyon. The tendency to form a network

Fig. 10. Paleoscolytus sussexensis Jarzembowski, 1990. Wadhurst Clay/Ashdown Formation, Crowborough, East Sussex. Scale bar represents 5 cm.

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may resemble more regular nets referred to ?Protopaleodictyon (Eagar et al., l985, pl. 6D), but the apparent nets are too irregular and small and the trace cannot be regarded as an agrichnion. Fill passive with silt derived from overlying sediment.

traces cross Beaconites isp., and in the Vectis Formation, spectacularly at the top of the White Rock.

Distribution. Weald Clay Group and Vectis Formation.

Vertebrate footprints have been recorded from most formations of the Wealden and were first recorded from the Hastings area in 1846 (Woodhams and Hines, 1989). Tridactyl footprints attributable to iguanodontid dinosaurs (if not with greater confidence to Iguanodon) have been recorded from several horizons in the Hastings Beds of Sussex (Woodhams and Hines, 1989), the Weald Clay of Surrey (Allen, 1975) and in the Wessex and Vectis formations of the Isle of Wight (Radley, 1994a,b; Radley et al., 1998a; Martill and Naish, 2001a,b). Other occurrences of footprints include the Wessex Formation of Swanage and Worbarrow bays, Dorset (MansellPlaydell, 1888, 1896; Delair, 1989). In the Wealden the footprints are mostly preserved on the soles of sandstones and may be seen, for instance, at outcrop close to the base of the Lee Ness Sandstone, lower Ashdown Beds, at Lee Ness near Hastings (Lake and ShephardThorn, 1987, pl. 3; Woodhams and Hines, 1989; Parkes, 1993) and among Wessex Formation sandstones at Hanover Point, Isle of Wight. Most relate to floodplain sand sheets. Four-toed tracks attributed to ankylosaurs and/or sauropods occur in the Wessex Formation on the Isle of Wight (Radley, 1994b,c). Theropod and ornithopod footprints (Fig. 5D) on the sole of a Filosina-Viviparus shell bed (Radley, 1994b, fig. 3c; Radley et al., 1998a, fig. 3f) are from the Shepherd’s Chine Member of the Vectis Formation (Isle of Wight) and indicate the shallowness of the lagoonal environment. Locally, the strongly disorganised nature of the mudstones of the upper Wessex Formation and lower part of the White Rock (Vectis Formation), suggest extensive dinosaur trampling (dinoturbation).

4. Insect traces Jarzembowski (1990) described the remarkable beetle borings (Fig. 10) into coniferous wood from the Wadhurst Clay/Ashdown Formation, originally reported, but not figured, by Blair (1943). In addition to the radiating pattern (Paleoscolytus sussexensis Jarzembowski, 1990) attributed to scolytid borings, winding borings are also known; see also Rhizocorallium isp. (above). Scott (1992, fig. 12.1), also figured these beetle borings in gymnosperm bark, but recorded that Chaloner et al. (1991) disagreed with Jarzembowski (1990) about the causal organism, suggesting that they were produced by weevils. Insect damage to Wealden plants has been recorded by Watson et al. (2001) from plant debris beds of the Wessex Formation of Worbarrow Bay and Mupe Bay, Dorset. The cuticle of leaves of Ginkgoites weatherwaxiae Watson et al. show repair to damage, possibly caused by phytophagous bugs, as well as post-mortem damage attributed to litter feeding beetles. The latter, approximately circular holes in the leaves were referred to the ichnogenus Phagophytichnus (Watson et al., 2001, p. 43). Francis and Harland (unpublished manuscript) have described a pyrite infilled chamber in fossil wood, from a plant debris bed in the Wessex Formation at Hanover Point, Isle of Wight, which they interpreted as a termite boring containing hexagonal faecal pellets. Surprisingly, dung beetles have not yet been recorded, and the fate of iguanodont faecal matter is an unsolved mystery.

5. Root traces Poorly preserved roots traces are common in the fluvial and lacustrine facies as slender, essentially upright strands of carbonaceous sand or silt, extending downwards and branching from the top of a sand bed, and often crosscutting burrows in the bed. Examples are widespread, in many alluvial and lake-margin facies of both sub-basins. In the Wessex Formation of the Isle of Wight (e.g., sandstones at Hanover Point), Allen (1946, 1959) described many localities, often associated with rhizomes and stems of Equisetites lyellii. In the Wessex Formation of the Isle of Wight (e.g., Hanover Point Sandstone) root

6. Tetrapod tracks and trackways

Ichnogenus Iguanodontipus Sarjeant, Delair and Lockley, 1998 Iguanodontipus burreyi Sarjeant, Delair and Lockley, 1998 Fig. 5E, H Description. Large tridactyl footprints with thick digits and bluntly rounded or pad-like toe imprints. Heel impressions typically wide and rounded but can vary considerably in preserved form depending on the nature of the substrate (see Woodhams and Hines, 1989; Parkes, 1993). Dimensions. Length 0.25e0.50 m (mean 0.38 m) (Lee Ness; Woodhams and Hines, 1989; Parkes, 1993); 0.50e 0.56 m (mean 0.53 m) (Cooden; Woodhams and Hines,

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1989), 0.7 m (Hanover Point; Radley, 1994b). Width: 0.18e0.5 m (mean 0.34 m) (Lee Ness; Woodhams and Hines, 1989; Parkes, 1993); 0.45e0.51 m (Cooden; Woodhams and Hines, 1989). Trackway data. Pace, 0.87e1.38 m; stride, 1.76e 2.76 m (Lee Ness, Parkes, 1993); pace, ca. 1e1.3 m; stride, ca. 2.5 m (Cooden, Woodhams and Hines, 1989, fig. 32.4). Remarks. The precise form and size of iguanodont footprints and the trackway dimensions can vary considerably (see References for details). Locally heavily trampled ‘‘dinoturbated’’ bedding surfaces are known (Radley, 1994c). Distribution. Iguanodontid footprints are widespread throughout the Wealden succession (see above). Major recently analysed tracksites are: East Sussex, Cooden beach, Bexhill, Tunbridge Wells Sand (Delair and Sarjeant, 1985; Woodhams and Hines, 1989); Lee Ness, Hastings, lower Ashdown Beds (Woodhams and Hines, 1989; Parkes, 1993). Isle of Wight, Wessex Formation, Yaverland, Wessex Formation (Radley, 1994b), Chilton Chine, Wessex Formation (Blows, 1978; Delair, 1985); Cowleaze Chine, White Rock, Cowleaze Chine Member and Barnes High Sandstone Member, Vectis Formation (Radley et al., 1998a). An inferred Iguanodon manus print has been collected from the White Rock, Cowleaze Chine by D.M. Martill (University of Portsmouth collection).

Wight; Wessex Formation, Yaverland; Shepherd’s Chine Member, Vectis Formation, Shepherd’s Chine. ?Sauropod or ankylosaur footprints Fig. 5F Description. Broadly sub-triangular footcasts often expanding rapidly from the heel region towards four blunt digits. Dimensions. Length, 0.3 m (Brook Bay, Isle of Wight; Radley, 1994b); 0.55 m (Yaverland; Radley, 1994c); 0.29 m, 0.16 m (Brook Bay; Radley, 1997). Remarks. Some of these casts are similar to the prints of the hind feet (pes) of sauropods figured from the Middle Jurassic deltaic sediments of Yorkshire (Whyte and Romano, 1993, 1994; Romano and Whyte, 2003) and ‘‘Wealden-age’’ tracksites in the Cameros Basin of northern Spain (Moratalla et al., 1994; Wright et al., 1998). In Spain, sauropod footprints form a small percentage of ichnofaunas dominated by theropods and ornithopods, including rare quadripedal Iguanodon. Distribution. Larger examples, almost certainly attributable to sauropods, have been discovered in the uppermost Wessex Formation near Cowleaze Chine, Isle of Wight.

7. Vertebrate skinprint Theropod footprints Fig. 5G Description. Tridactyl footprints with generally narrow digits that have sharply pointed toe impressions, sometimes with terminal claws. Heel imprints are usually more angular in shape than those of iguanodont footprints. Theropod footprints are generally smaller than iguanodont, but larger prints occur frequently. Dimensions. Length, 0.23e0.45 m (mean 0.34 m) (Lee Ness; Woodhams and Hines, 1989; Parkes, 1993); 0.23 m (Yaverland, Isle of Wight; Radley, 1994c); 0.27 m (Shepherd’s Chine, Isle of Wight; Radley et al., 1998a). Width, 0.20e0.42 m (mean 0.31 m) (Lee Ness: Woodhams and Hines, 1989; Parkes, 1993); 0.25 m (Yaverland, Isle of Wight; Radley, 1994c); 0.27 m (Shepherd’s Chine, Isle of Wight; Radley et al., 1998a). Trackway data. Pace 0.82 m; stride, 1.64 m (Lee Ness; Parkes, 1993). Distribution. Theropod footprints have been recognised at most Iguanodontipus sites. East Sussex; Hastings Beds; lower Ashdown Beds, Fairlight Cove. Isle of

Although strictly to be regarded as a body fossil, a skinprint was figured by Allen (1975, pl. 3, fig. C) from the Lower Tunbridge Wells Sand Formation, West Hoathly, West Sussex. It is possibly of an iguanodon since skeletal remains were found nearby.

8. Vertebrate coprolites Although coprolites were first collected by Gideon Mantell from the sandstone of Tilgate Forest in the 1820s and figured by Buckland (1829), they have been rarely recorded and little studied subsequently. No collection has been made that is comparable to the famous Wealden coprolites of Bernissart, Belgium, originally assigned to Iguanodon but now regarded as theropod or crocodile faeces (Bertrand, 1903; Abel, 1935; Casier, 1960; Thulborn, 1991). However, recent collecting in brick pits in the Weald Clay of Surrey and Sussex (Jarzembowski, 1991; A.J. Ross, pers. comm. 1991; Cooke and Ross, 1996; Ruffell et al., 1996), and Wessex and Vectis Formations of the Isle of Wight (Martill and Naish, 2001b), and a reawakened interest in the nature

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and interpretation of vertebrate coprolites (Thulborn, 1991; Hunt et al., 1994; Chin et al., 1998) has enabled several distinct forms to be recognised and described. At present vertebrate coprolites are generally described in terms of their overall shape, surface morphology, internal structure, internal contents and chemical composition. Their naming is highly controversial. Some workers, especially those concerned with invertebrate coprolites, apply ichnogeneric and ichnospecific names (e.g., Ha¨ntzschel, 1975; Bischoff, 1990), while most workers on vertebrate coprolites do not, on account of the variability in size, shape, preservation, and uncertainty of the producer (Hunt et al., 1994). Here they are described in terms of morphotypes that probably reflect different producers. Two broad morphotypes are recognised among Wealden coprolites: Type 1 comprises spiral coprolites, frequently with delicate surface markings and containing fish scales. These were probably produced by piscivorous aquatic predators with a spiral intestinal valve, most likely sharks. Type 2 is elongate and irregular in shape with only poor spiral structure, or is segmented; some specimens have polygonal surface cracking and rarely contain fish scales. These are likely to have been produced by reptiles, possibly carnivorous dinosaurs or crocodiles. These morphotypes described here from Wealden sediments of Sussex are similar to those from the Isle of Wight (Martill and Naish, 2001b, pp. 319e321) but markedly different from those recently described from the underlying Purbeck Group (Ensom, 2002). Heteropolar spiral coprolites Fig. 11A, B Description. Sub-ovoid coprolite with strong, overlapping, spiral folds concentrated towards one end. The surface of the folds has distinct transverse or dendritic markings. Length in excess of 30 mm (broken end, common); width: 25 mm maximum. Four or five spiral folds are present and appear to have been separated originally by a membrane. Internally, phosphatic material including fish scales and fin rays is densely packed across the folds and reflected in the surface markings. Remarks. This morphotype crudely resembles the ‘‘larch-cone’’ type of Buckland (1829, pl. 31, figs. 1e11) but lacks the fine ‘‘bract-like’’ layering within the folds that are characteristic of Chalk coprolites. The producer was a piscivorous predator with a spiral intestinal valve, probably a shark such as Hybodus or other chondrichthyan. Although similarly heteropolar and spirally folded to the coprolite ichnogenus Heteropolacoprus (Hunt et al., 1998), the major differences of shape, arrangement and sculpture of folds suggest that ichnogeneric assignment is unwise.

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Distribution. Weald Clay Group and Vectis Formation (Martill and Naish, 2001b, pl. 44, fig. 1), probably in offshore lacustrine or lagoonal facies. Amphipolar spiral coprolites Fig. 11CeE Description. Ovoid medium-sized coprolites with one or two indistinct, adherent spiral folds in the medial part of the coprolite. Length in excess of 57.5 mm; maximum width, 37 mm. Surface smooth or with faint transverse spiral lines; a few isolated fish scales project from the surface. Shape irregular (Fig. 11C): length, 32 mm; width 21.5 mm; thickness 12.5 mm, compressed; contains large scales of Lepidotes (8 ! 6.5 mm) and other densely packed fish debris crudely layered parallel to compression. Remarks. This is a common form of spiral coprolite usually assigned to sharks, which are quite well known from Wealden sediments (Patterson, 1966; Cooke and Ross, 1996). Mantell’s original Wealden coprolite (Buckland, 1829, pl. 31, fig. 18), which was crudely spiral, phosphatic in composition and contained fish scales, was probably of this type. Distribution. Cuckfield Stone of Tilgate Forest; Weald Clay Group and Vectis Formation (Martill and Naish, 2001b, pl. 44, fig. 1). In plant debris beds of Wessex Formation, Isle of Wight, lacustrine/lagoonal facies of Vectis Formation. Elongate irregular coprolites Fig. 11FeH Description. Elongate, cylindrical, irregular-shaped coprolites with only a crude spiral or segmented structure. Shape is often sausage-like or sinuous with both ends evenly rounded (isopolar; Fig. 11G) or with one end rounded, one pointed (anisopolar; Fig. 11H) (terminology of Thulborn, 1991). Where the form is crudely spiral (Fig. 11H) there are no internal folds or separation surfaces. Length 30 to O65 mm, width ca. 25 mm. Annulated or constricted forms (Fig. 11F) commonly break into segments, so one end is often missing. Surface is usually irregular, although it can have polygonal cracking or adherent clay with carbonaceous matter. Internally some coprolites show septarian cracking with calcite veins or cavities. There are no fish scales but the matrix is phosphatic claystone. Remarks. The irregularity, lack of spiral structure, occasional segmental constrictions, and occasional surface and internal cracking (desiccation), suggest that these coprolites may have been extruded subaerially, possibly by reptiles (Thulborn, 1991; Hunt et al., 1994). The small

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Fig. 11. A, B, heteropolar spiral coprolite of crude ‘‘larch-cone’’ form. A, upper end is broken off but four distinct spiral folds with delicate surface markings are visible. B, rear view shows smooth internal surface separating the spiral folds, Weald Clay Group, Hamsey Brickworks, South Chailey, Sussex (TQ 399 160); Booth Museum, Brighton BMB 023035 (collection M. Zdrzalek); !1.5. C, compressed irregular spiral coprolite with large Lepidotes scales; Weald Clay Group, Leybrook Brickworks, Thakenham, Sussex (TQ 118 188); Booth Museum, Brighton BMB 023036 (collection A.J. Ross); !1.4. D, E, amphipolar spiral coprolite with prominent central fold and delicate spiral markings. Rear view (E) shows that the lower end is missing and the fractured internal surface reveals a dark brown to black phosphatic matrix with scattered fish scales. Weald Clay Formation (Bed 3), Rudgwick Brickworks, Rudgwick, Sussex; Booth Museum, Brighton BMB 023037, collection A.J. Ross; !1. F, elongate ‘‘sausage’’ shaped coprolite with four crudely spiral segments separated by constrictions; lower end broken off along a constriction; composed of buff phosphatic claystone with small ovoid internal cavities ca. 1 mm in diameter (gas bubbles?); Weald Clay Group, Keymer Brick and Tile Works, Burgess Hill, Sussex (TQ 324 193); Booth Museum, Brighton. BMB 023038 (collection A.J. Ross); !1.5. G, slightly sinuous elongate coprolite with a bluntly pointed end and four weak segments. The irregular surface of the coprolite shows primary cracks forming rough polygons 3e4 mm across, suggesting desiccation of the surface, perhaps subaerially. Horizon and locality as F; Booth Museum, Brighton BMB 023039 (collection A.J. Ross; !0.8). H, elongate unsegmented coprolite with a distinct spiral twist at the upper end (proximal end on discharge?). Smooth surface with adherent small carbonaceous fragments. Horizon and locality as F; Booth Museum, Brighton BMB 023040 (collection A.J. Ross); !1.5.

size of the Wealden specimens (length 30e60 mm) when compared to the Bernissart (Belgian) forms (80e 120 mm), suggests that the producers could have been crocodiles or small theropod dinosaurs rather than large megalosaurs (Bertrand, 1903; Thulborn, 1991). In common with the Belgian coprolites, all those collected are composed of phosphatic material produced by predators (Hill, 1976; Chin and Gill, 1996), not comminuted plant material indicative of herbivores, possibly ornithopods (Hill, 1976). No coprolites have been recognised that can be assigned to Iguanodon, the most abundant skeletal

remains and footprint producer (see above). This anomaly was discussed by Thulborn (1991), who emphasised the higher preservation potential through rapid diagenesis, of phosphatic coprolite material and the likely rapid breakdown and dispersal of herbivore droppings unless deposited in peat swamps or rapidly buried in anoxic and fine-grained sediment or permineralised in semi-arid palaeosols (Chin and Gill, 1996). Such favourable preservation conditions would have been rare in Wealden environments. Alternatively, iguanodont faeces may have been reworked and destroyed by the action of dung beetles

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as has been recorded for herbivorous dinosaur faeces preserved by permineralisation in the Upper Cretaceous Two Medicine Formation of Montana (Chin and Gill, 1996). These favourable preservation conditions would have been rare in Wealden environments. Distribution. Fluvial-lacustrine sediments at several localities in the Weald Clay, and in plant debris beds in the Wessex Formation, Isle of Wight (Martill and Naish, 2001b, pl. 44, figs. 3, 4). 9. Pseudofossils When palaeoenvironmentally significant fossils are absent or uncommon there is a natural tendency to search even more diligently. Structures that superficially resemble trace fossils are quite common. Proving a biological origin can be difficult and it is generally useful first to attempt to eliminate primary physical and chemical processes. The large spiral concretions in the Wadhurst Clay Formation near Hastings were attributed to moulds of gigantic gastropods and given the name Dinocochlea ingens by Woodward (1922) before Thomas (1935) reported their inorganic origin. Load casts on the underside of sandstones in the Upper Tunbridge Wells Sand Formation and the Wessex Formations often resemble dinosaur footprint casts. In the Wessex Formation some of these may grade into recognizable tridactyl casts, and are thus plausibly interpreted as such, but others are highly suspect. Fig. 12 shows two sedimentary structures that resemble Beaconites (Fig. 3), a back-filled (meniscate) burrow, which is not uncommon in Wealden sandstones of marginal lacustrine facies. Fig. 12A is part of a large block of sandstone in the rockstore at Clockhouse Brickworks (TQ 175 385) with a strongly convex lower surface and planar top forming a section of an erosional (channel)-fill structure. The horizon in the quarry (Lower Weald Clay Formation) from which it came is unknown but is unlikely to have been from the Clockhouse Sandstone Member. On one side of the block are cross-sections of two upward-directed meniscate structures that run up one margin of the scour. That these are not trace fossils is evident from the manner in which the width of each expands rapidly from the base, where some menisci also pass laterally into primary lamination in the sandstone. Further changes in width occur higher in the block. The structures are a result of gas/water escape from underlying sediment that locally heaved up the laminae of the sandy fill. Where such escape took place in more uniform sediment the domed laminae have generally collapsed. In this case collapse was probably prevented by the muddy wall of the scour. Gas generated by plant decomposition is likely, and the impression of a log (20 cm diameter) is present on the opposite side of the scour-fill.

Fig. 12. A, complex structure of upwardly deformed and expanding meniscate laminae superficially resembling large burrows of Beaconites barretti; interpreted as a gas or water escape structure expanding upwards through cross-bedded sandstone at margin of a channel fill. Sandstone in Lower Weald Clay Formation, Clockhouse Brickworks rockstore (TQ 175 385), Capel, Surrey; field photograph; scale bar represents 5 cm. B, weathered surface of grey-green sandstone showing a structure resembling meniscate burrow-fill of B. barretti, cross-cut by vertical sand-filled burrows with thin mud walls; interpreted as weathering of rib and furrow structure in a gully-fill in trough crossbedded sandstone. Top of Lee Ness Sandstone (Ashdown Formation), Lee Ness Ledge (TQ 866 109), near Hastings, Sussex; field photograph. Scale bar represents 5 cm.

Fig. 12B is from the upper part of the Less Ness Sandstone (Ashdown Formation) in the Hastingse Fairlight section of the Hastings Beds (Lake and Shephard-Thorn, 1987). The sandstone exhibits crosssections of meniscate-like structures; these are gently curving and either more or less parallel to the stratification (dipping at a low angle westwards on the west limb of the anticline) or lie at a steeper angle, parallel to inclined heterolithic stratification. The unit has been intensely reworked by cut and fill and subsequently burrowed by Beaconites antarcticus and penetrated by thin roots. The structures differ from meniscate trace fossils in that laterally they can be traced into small-scale cross-lamination forming a sheet rather than a gutter. The margins are also far more ragged than in a burrow, owing to uneven avalanching and deposition of the fine-grained sand during deposition. The structures are examples of ‘‘rib and furrow’’ structure (e.g., Collinson and Thompson, 1989, figs. 6.9, 6.11) produced by advancing current ripples.

10. Conclusions The ichnofossils recovered so far include 16 invertebrate ichnotaxa, insect and root traces, three- and four-toed vertebrate footprints, a skinprint, and three

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broad types of vertebrate coprolite. Two types of sedimentary structure that have been previously attributed to trace fossils are described. New discoveries are being constantly made in the stone and brick quarries in the Wealden of the Weald Basin, and in the sea cliffs of the Wessex Basin, either in situ or as eroded material. With the relatively small number of ichnogenera we can only place the assemblages in the broadly defined Scoyenia Ichnofacies within the formally recognised continental ichnofacies, and the brackish incursion association within the Glossifungites Ichnofacies.

Acknowledgements Many of the specimens from the Weald Basin have been collected in the past few years by Ed Jarzembowski (Maidstone Museum), Andrew Ross (Natural History Museum, London) and colleagues, often during search for fossil insects. Jon Radley acknowledges the assistance of Martyn Munt and Steve Hutt (Dinosaur Isle, Sandown) and Mick Green (Brighstone, Isle of Wight) for assistance in the field during the production of the Jurassic-Cretaceous Boundary Conservation Review volume, and for allowing access to collections in their care. He also acknowledges the support received during the production of the Wealden Conservation Review volume, under the direction of Perce Allen (University of Reading) and Bill Wimbledon (Countryside Council for Wales). Permission from the Palaeontological Association to reproduce Fig. 9B is acknowledged.

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