J. S Tuart University of Cambridge Longman London and New York

• J. Stuart University of Cambridge Longman London and New York Pleistocene vertebrates in the British Isles 102 agent in breaking bones, especially as many are far from the Cromerian of West Runton, Norfolk, too large to have been passed through the gut of which has a large bone tumour, with a central pit, another animal, or to have been broken by its teeth. on the antero-medial surface. The lesion is thought The question needs much further investigation in most likely to have been caused by ossification of a view of its implications for the taphonomy of verte sub-periosteal haematoma resulting from injury. brate assemblages (Ch. 4). Pathological elephant molars, showing twisting, Small-mammal bones from the pellets regurgi distortion and eruption at abnormal angles, are fair tated by modern diurnal birds of prey (raptors) show ly common. Bone tumours and direct injury have corrosion, which appears to be of a characteristic been cited as causes of these lesions (McWilliams type and can be matched in Pleistocene fossil 1967). assemblages (Mayhew 1977). Mayhew describes Lesions due to physical injury by weapons have and illustrates a number of recent and fossil exam been described in detail from Star Carr, Yorkshire ples. In vole molars, for example, corrosion is con (early Flandrian), and Blackpool, Lancashire (Late With the exception of one cave assemblage, the imprecise stratigraphical information for many fined to the enamel ridges near the crowns, the rest Devensian). Two scapulae, one of red deer Cervus Lower Pleistocene vertebrates of the British Isles finds, there is little prospect of detecting faunal of the tooth having been protected by the bone of elaphus, the other of elk Alces alces, from Star Carr, come from the predominantly marine crags of East changes within stages for the Lower Pleistocene. the jaw. show partly-healed fractures produced by flint Anglia. The record is therefore not surprisingly The overall faunal history for the period is discuss Material recovered from the pellets of owls shows tipped projectile weapons or barbed points (Noe patchy and unsatisfactory in comparison with the ed in Chapter 9. no such corrosion and little breakage, and is there Nygaard 1975). A complete elk skeleton from 'Villafranchian' faunas of southern and eastern The main sites with Lower Pleistocene vertebrate fore not readily recognizable in fossil assemblages. Blackpool (Ch. 8) bears a variety of unhealed le Europe. faunas are shown in Fig. 6.1 Nevertheless, most of the fossil small-mammal sions resulting from the impact of flint-tipped pro Due to the combination of sparse material and bones and teeth from cave deposits at least almost jectile weapons, together with what appear to be certainly originated from pellets dropped by owls. axe cuts on one of the metacarpals, all dating from Occasionally, discrete patches of fossil small immediately prior to the death of the animal. In WR ER Cm = Sd mammal material have been observed, sometimes addition, however, there are a few lesions with Tm with traces of organic matter, evidently represent accompanying reaction - osteoporosis of the bone Bc ing owl pellets, as at Angel Road, North London due to bacterial infection which must have dated (Devensian). The pellets contained bones and teeth from some time before the elk's death. In particu of arctic lemming Dicrostonyx torquatus in an excel lar, a barbed bone projectile point was excavated in l1li King's Lynn lent state of preservation (Hinton 1912). situ resting in a broad groove in the distal end of the left metatarsal (Hallam et al. 1973) (see Figs 8.6, 8.7). The groove had evidently been eroded by the point which was embedded in the animal's foot, an estimated two to three weeks before death. Noe Palaeopathology Nygaard's (1975) dismissal of the lesions as caused by earth pressure and by damage during excavation Pathological abnormalities are occasionally ob appears to be based on misunderstandings of the served in Pleistocene fossil mammal material. nature of the site and the state of preservation of For example, Wells and Lawrence (1976) describe the bones (Stuart 1976b). a metatarsal of giant deer Megaceros verticornis, o 20 km I I a b Fig. 6.1. Location map of the main Pliocene and Lower Pleistocene vertebrate sites. +, Red Crag Nodule Bed (Pliocene); Red Crag; ., other crags; 0, cave sites. The extent of the Pliocene Coralline Crag (hatched) and the Lower Pleistocene crags (stippled) are indicated, together with the locations of the Ludham and Stradbroke boreholes. Bc, Bacton; Bm, Bramford; Boy, Boy ton; Br, Bramerton; Bu, Butley; Cm, Cromer; DH, Doveholes; EB, Easton Bavents; ER, East Runton; Fa, Falkenham; Fe, Felixstowe; Re, Rendlesham; Sd, Sidestrand; Su, Sutton; Tm, Trimingham; Tp, Thorpe, Norwich; Tr, Trimley' Wa Waldringfield; Wd, Woodbridge; WR, West Runton. ' , Pleistocene vertebrates in the British Isles 104 105 Lower P leislOcene faunas faunas from the Ludham and Stradbroke boreholes Upper Pliocene strongly suggest that the Red Crag was deposited (Red Crag Nodule Bed) largely or entirely within the first stage of the Brit ish Pleistocene, the Pre-Ludhamian (Funnell & The Red Crag occurs over much of south-east Suf West 1977). Most fossil terrestrial vertebrates re folk and as small outliers in north-east Essex. At its corded from the Red Crag probably originate from base at a number of localities, (notably Wood the Red Crag Nodule Bed. A few specimens appear bridge, Boy ton, Sutton and Butley, all in Suffolk) from their less mineralized condition, andlor is found a condensed deposit - comprising flint where they were found, to have come from the Red pebbles and phosphatic nodules. Vertebrate fossils Crag proper (Spencer 1964, 1966) and are there from this horizon are generally heavily mineralized, fore probably of Pre-Ludhamian age, if not re and often well rolled and polished, imparting an worked from older strata. Taxa represented include attractive glass, so that specimens were much beaver C. fiber, gomphothere Anancus arvernensis, prized by nineteenth-century, and later, collectors. extinct elephant Archidiskodon meridionalis, horses Some of the fossil vertebrate material has clearly including both Equus cf. stenonis and a 'caballine' been reworked from much older strata. In particu species Equus bressanus, a deer Eucladoceros sp., lar, remains of Coryphodon and Hyracotherium were a gazelle Gazella sp., and albatross Diomedea derived from Eocene London Clay which occurs sp. Marine vertebrates are also recorded (Newton over much of the area. Comparison with Continen 1891). tal faunas suggests that the bulk of the vertebrate material is of broadly Upper Pliocene ('Lower Vil lafranchian') age. The Pliocene age strictly puts the Red Crag Nodule Bed fauna outside the scope of this chapter, but it is nevertheless briefly consid Antian and Baventian Stages ered here as a prelude to the faunas of the Lower (N orwich Crag, in part) Pleistocene. Little modern work has been done on this material, which was, however, extensively de It is uncertain to what extent the Ludhamian and scribed and figured by Newton (1891). Thurnian stages, which occur in the Ludham and Taxa recorded include porcupine sp., Hystrix Stradbroke boreholes, can be recognized in surface beaver Castor fiber, a small beaver-like rodent Tro exposures of deposits assigned to the Red or N or gontherium minus, a bear-like carnivore Agriotherium wich Crag. Vertebrate records from the Norwich sp., extinct hyaena Hyaena perrieri, an extinct pan Crag so far available can be assigned to one or more da Parailurus anglicus, gomphothere mastodont of the stages from the Antian to the Pre-Pastonian. Anancus arvernensis, true mastodon Zygolophodon The cliff section at Easton Bavents, north of Fig. 6.2. Shelly marine sands ('crag') of Antian/early Baventian age at Easton Bavents, Suffolk. borsoni, extinct elephant Archidiskodon meridionalis, Southwold, Suffolk, includes at least 2.8 m of Nor horse E quus sp., extinct three-toed horse H ipparion wich Crag (Fig. 6.2), overlain by more than 2 m of sp., extinct tapir Tapirus arvernensis, extinct rhi laminated grey and blue clays (Funnell & West Marine fish material is common in the crag, but noceros Dicerorhinus megarhinus and an extinct Bramertonian Stage and 1962). Pollen analyses of silt bands within the basal remains of non-marine vertebrates are sparse, and bovid cf. Leptobos sp. metre of crag indicates temperate forests with hem much of our knowledge results from patient collect Pre-Pastonian a Substage Marine vertebrate remains also occur abundant lock Tsuga, and the deposits are assigned to the ing over many years by Mr T. H. Gardner. Taxa ly. Taxa represented include fishes, turtles, a (N orwich Crag , part) Antian Stage with Easton Bavents as the type local recorded from the rarely exposed 'Lower Shell sirenian, a walrus and several whales (Newton ity. A single pollen spectrum obtained from the Bed', of probable Antian age, include Equus cf. ste The pit at Bramerton Common, near Norwich, Nor 1891). upper part of the crag is of intermediate character nonis and Archidiskodon meridionalis. Large mam folk, has long been regarded as the type section for and was assigned to the early part of the Baven mals from the 'Upper Shell Bed' of early Baventian, the Norwich Crag, and dates from the early tian (zone Lp4a of Ludham). The crag therefore or late Antian age, include E. cf. stenonis, A. meri nineteenth century. A later excavation, Blake's Pit, ranges from the Antian to the early Baventian in dionalis, Eucladoceros falconeri, Eucladoceros sp. and 300 m to the east, was first noted in 1870. The strati age. Anancus arvernensis. Teeth of the extinct water vole graphy, pollen, foraminifers and marine molluscs Pre-Ludhamian Stage (Red Crag) The marine mollusc faunas (Norton & Beck M imomys pliocaenicus were described by Carred:.

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