Pleistocene vertebrates in the British Isles 138 139 M iddle and Upper P leistocene interglacial faunas
Table 7.5 Early Flandrian Faunas from Thatcham and Star Carr. fish remains at Star Carr, as indicating that by zone Vertebrae of a pelican Pelecanus sp. were re IV (Fl la) fishes had not yet penetrated the Vale of covered from peat near King's Lynn, Norfolk, Thatcham Star Carr Pickering via the River Derwent. radiocarbon dated to 3 915 ± 120 years B.P. 'Surface MX' at Nazeing, Essex, in the flood (Forbes et al. 1958). Dalmatian pelican Pelecanus MAMMALIA plain deposits of the River Lea, yielded material of crispus is recorded from several localities in the Insectivora common shrew S orex araneus, pika Ochotona pusil Cambridgeshire Fens, and also from the Iron Age Erillaceus europaells L., hedgehog + + Sorex aranells L., common shrew + la, bank vole Clethrionomys glareolus, water vole Lake village at Glastonbury, Somerset (Northcote Talpa europaea L., mole + Arvicola terrestris, northern vole Microtus oeconomus 1980b). and wood mouse Apodemus sylvaticus (Allison et al. The sole record of European pond tortoise E mys Primates Homo sapiens L., man (humerus) + 1952). The pollen spectra from immediately above orbicularis, from the Flandrian of the British Isles, Homo sapiens L., man (artefacts) + + this horizon indicate pine-birch woodland and a is based on remains of three individuals from mere zone V (Fl Ib) age. The fauna is generally consis peats at East Wretham, near Thetford, Norfolk Lagomorpha Lepus sp. + tent with a woodland environment, but the occur (S tuart 1979). Pollen analysis of a peat sample pre Oryctolagus cuniculus (L.), rabbit + rence of Ochotona pusilla - a survival from the Late served with the finds indicate a subzone Vlla (FI Devensian - is interesting as this species now Rodentia lid) age. Castor jiber L., beaver + + occurs only in the Palaearctic steppe (Ch. 5). Northern vole, not recorded from later than the Arvicola terrestris L. , water vole + Bones of wild boar Sus scrofa and red deer Cervus early Flandrian on the mainland, occurs in Bronze Carnivora elaphus are recorded from sands overlying peat and or Iron Age deposits at Nornour, Isles of Scilly Canis luplls L., wolf + + beneath estuarine clays at Belfast Lough; all (Pernetta & Handford 1970). + + Canis (domestic), dog apparently dating from the early Flandrian (Savage Vulpes vlIlpes (L.), red fox + + Written sources provide most of the evidence for Martes martes (L.), pine marten + + 1964, 1966). These finds suggest that both species survival into historical times of many species which Meles meles (L.), badger + + are native to Ireland. subsequently became extinct in Britain (see Table Felis sylvestris Schreber, wild cat + + Vertebrate material dating from later in the Flan 7.6). Perissodactyla drian is available from such deposits as the peats of Equlls ferus Boddaert, horse + the East Anglian Fenland and from archaeological Artiodactyla sites. Many finds are in need of critical reassess Flandrian faunal history SIIS scrofa L., wild boar + + ment, with regard to both identification and strati + + C ervlls elaphus L., red deer graphy. Only the more interesting records will be The history of the wild vertebrate fauna during the Alces alces L., elk + + Capreolus capreollls (L.) roe deer + + considered here. Flandrian (Fig. 7.16) is complicated by factors, not Bos primigenills Bojanus, (extinct) aurochs + + The peats of the Fen margins north of Cam important in earlier interglacials:
AVES bridge (Godwin 1978) date from the second half of (a) isolation of Britain from the Continent early in the Flandrian, i.e. subzone Vlla (FI lld, about the Flandrian - preventing further immigration of Graviiformes 7 000 years B.P.) and later. Most of the bones col terrestrial and freshwater vertebrates; Gavia stellata (Polltoppidall), red-throated diver lected from this area (villages of Burwell, Swaffham (b) extinctions, mainly of large mammals due to Podicipediformes and Reach) during the last century are probably of human activities; and + Podiceps m'status L., great crested grebe zone VIII age (FI lld, early FI Ill). Mammals re Podiceps nljicollis Pallas, little grebe (c) the introduction by man of alien vertebrates to corded include wolf Canis lupus, brown bear Ursus Britain (Ch. 10). Ciconiiformes arctos, otter Lutra lutra, beaver Castor jiber, roe deer The availability of good early Flandrian faunas is cf. Ciconia ciconia L., white stork + Capreolus capreolus, red deer Cervus elaphus and particularly valuable because the faunas of pollen Anseriformes aurochs Bos primigenius. Birds include bittern zone I age from earlier interglacials are almost un Anas plalyrhynchos (L.), mallard + Botaurns stellaris, mute swan Cygnus cygnus, whoop known. Ochotona pusilla, Microtus oeconomus and Anas sp. (small species), a duck + BlIcephala clallgula (L.), golden eye duck + er swan Cygnus olor, white-tailed eagle H aliaetus perhaps Equus ferns as a native species were re Mergus serrator L., red-breasted merganser + albicilla, common crane Grns grns and razorbill stricted to the early Flandrian, in association with Alca torda (Northcote 1980b). F alconiformes regional birch and pine forests, and probably exten Buteo buteo L., buzzard + The record of elk Alces alces, based on droppings sive open habitats. They can be regarded as surviv from peat at Ugg Mere, Cambridgeshire, and als from the Late Devensian. On the other hand, Ralliformes Grus gms L., common crane + + radiocarbon dated to about 3 260 years B.P. (God the earliest Flandrian faunas (zone IV - FI la) win, 1975), is now known to be incorrect. Re already include several taxa nowadays associated Charadriformes examination of the material by A. Lister has shown with the regional temperate forest biotope, and not Vanellus vallellus L., lapwing + that the droppings are too small for A. alces. There extending further north than southern Fennoscan- Passeriformes are no other records of this animal from later than . dia: mammals - Erinaceus europaeus, Talpa euro + Turdus sp., blackbird or thrush, etc. the early Flandrian. paea, Meles meles, Sus scrofa and Cervus elaphus; Pleislocene vertebrates in the British Isles 140 141 Middle and Upper Pleistocene interglacial faunas and birds - Podiceps cristatus, Podiceps rujicollis and The modern fauna Table 7.6. Present day native British fauna of terrestrial vertebrates (*, also occurs in Ireland; t, extinct). Ciconia ciconia. This phenomenon may reflect a rapid amelioration of climate early in the Flan The present-day native vertebrate fauna of the Brit AMPHIBIA (Smith 1969) Rodentia drian. ish Isles comprises about 200 breeding birds, 41 Triturus cristatus (Laurenti), warty newt *S ciurus vulgaris L., red squirrel ?Introduced in The record of Oryctolagus cuniculus from the very non-marine mammals (including 5 species extinct *Triturus vulgaris (L.), smooth newt Ireland Triturus helveticus (Razoumoski), palmate newt early Flandrian is remarkable in that this species is within the last 2 500 years) and 11 breeding bats, tC astor jiber L., beaver. Extinct twelfth century or Bufo bufo (L.), common toad later generally thought to have been restricted to parts of 19 primary freshwater fishes, 6 reptiles and 6 *Bufo calamita Laurenti, natterjack Muscardinus avellanarius (L.), dormouse Iberia during the postglacial until its fortunes were amphibians (Table 7.6). *Rana temporaria L., common frog Clethrionomys glareolus (Schreber), bank vole. Table 7.7 demonstrates the way in which the pre dramatically reversed by man from Roman times Introduced in Ireland (probably very recently) onwards, when it was introduced to northern sent-day vertebrate faunas of Britain and Ireland Arvicola terrestris L., water vole Europe. Confirmation of the Thatcham record become progressively impoverished according to Microtus agrestis (L.), field vole would be desirable. It is, however, possible that the their remoteness from Continental Europe. For ex Micromys minutus (Pallas), harvest mouse rabbit became extinct in Britain at some time after ample, voles and snakes are native to Britain but *Apodemus sylvaticus (L.), wood mouse the early Flandrian and was re-introduced in Nor not Ireland, and green lizard Lacerta viridis and gar Introduced in Ireland man or Plantagenet times. den dormouse Eliomys quercinus are absent from the Apodemus flavicollis (Melchior), yellow-necked The single record of Emys orbicularis (three indi British Isles, but are found on the adjacent part of mouse the Continent. This phenomenon is in part due to viduals) of zone Vlla (FI lId) age, at the climatic REPTILIA (Smith 1969) Carnivora optimum of the Flandrian, is particularly signifi climatic factors and a reduced variety of habitats in Anguis fragilis L., slow-worm t*Canis lupus L., wolf. Extinct eighteenth century cant as there are no less than six records from the Ireland, but for terrestrial and freshwater species it Lacerta agilis L., sand lizard *Vulpes vulpes (L.), red fox previous and generally warmer Ipswichian intergla is largely related to the timing of Flandrian land/sea *Lacerta vivipara Jacquin, viviparous lizard t Ursus arctos L., brown bear. Extinct about tenth cial, and numerous Flandrian finds from Denmark level changes. These changes may have isolated Ire N atrix matrix (L.), grass snake century and southern Sweden. The abundance of Scandina land from Britain (formerly connected via south Coronella austriaca Laurenti, smooth snake *Martes martes (L.), pine marten Vipera berus (L.), adder vian records is very probably related to the preva west Scotland) as early as about 9 500 years B.P., *M ustela erminea L., stoat lence of warmer summers further east. and then Britain from the Continent (formerly con M ustela nivalis L., weasel The extinction of a number of wild mammals and nected via the southern North Sea and Straits of M ustela putorius L., polecat *Meles meles (L.), badger birds within the last 2 000 years or so is clearly re Dover) perhaps a thousand years later (Ch. 2). Vertebrate species occurring on the adjacent *Lutra lutra (L.), otter lated to hunting coupled with destruction of the F elis sylvestris Schreber, wild cat forests and later draining of the wetlands by man. Continent at the present day, but absent from the The subject is discussed in Chapter 10. British Isles, are almost entirely those with south MAMMALIA (modified from Corbet & Southern 1977) Artiodactyla ern, or sometimes eastern, distributions in Europe. (Includes species extinct within last 2 500 years, t*Sus scrofa L., wild boar. Extinct probably Presumably they were not abJe to advance suf shown by t) seventeeth century Comparison with Continental faunas ficiently far north, in response to the Flandrian Insectivora *Cervus elaphus L., red deer climatic amelioration, before Britain became iso Sorex araneus L., common shrew Capreolus capreolus (L.), roe deer It is especially interesting to compare the Flandrian lated by the rise in sea level. It is possible, how *Sorex minutus L., pigmy shrew tBos primigenius Bojanus, aurochs. Extinct by Neomysfodiens (Pennant), water shrew 2 500 years B.P.? vertebrate fauna I succession in Denmark and adja ever, that a few such species did manage to reach *Erinaceus europaeus L., hedgehog? Introduced in cent areas, as reviewed by DegerbfiSl (1964), with Britain, but subsequently became extinct due to Ireland that seen in England. Important differences include climatic or other changes, as did Emys orbicularis. Talpa europaea L., mole the records of horse E quus ferus from Swedish de Significantly, the fall in species from France to Lagomorpha posits pollen-dated to zone V (FI Ib), and the sur Britain to Ireland is least marked in the aerial birds Lepus capensis L., brown hare. Introduced vival of elk Alces alces in Denmark until at least and bats, whose distribution is likely to be little recently in Ireland. zone Vllb (early FI Ill). The European pond tor affected by sea barriers. As far as the birds are con *Lepus timidus L., mountain hare toise Emys orbicularis is represented by numerous cerned there are actually more breeding species in fossils from Denmark and southern Sweden, cover Britain than in the rather small area of the Conti buted wholly or partly to human interference roach Rutilus rutilus, tench Tinca tinca); loaches ing pollen zones V (FI Ib) to Vllb (early FI Ill). nent used for comparison, which has a less varied (Ch. la), which complicates the interpretation of Cobitis taenia and Noemacheilus barbatulus; burbot (DegerbfiSl & Krog 1951; Stuart 1979). This dis range of habitats. Similarly, the more restricted natural distribution patterns determined by climat Lota Iota; perch Perca fluviatilis, ruffe Gymnocepha parity between the numbers and stratigraphical availability of habitats, and, to a lesser extent, ic and geographical factors. Species of mammals Ius cernua; and miller's thumb Cottus gobio) now ranges of Scandinavian and English finds reflects climatic conditions account for the rather lower confined in the British Isles to offshore islands found in the British Isles, are only indigenous to the warmer summers in the east, while the species numbers of breeding birds and bats in Ireland com appear to have been introduced by man from the the river catchments bordering the eastern English was probably near the limits of its range in Britain pared with Britain. Continent (see Ch. la). Channel and southern North Sea. The fishes are with its cooler and more cloudy maritime climate The restricted distributions within the British Wheeler (1977) considers that all the primary thought to have entered these rivers when they (Ch. 3). Isles of many native vertebrate species can be attri- freshwater fishes (pike Esox lucius; Cyprinidae (e.g. were formerly joined in the extensive land area con- Pleistocene vertebrates in the British Isles 142
Table 7.7. Numbers of non-marine vertebrate species native to Ireland, England and the adjacent areas of Continental Europe. (Data from Muus & Dahlstrjijm 1971; Wheeler 1977; Arnold & Burton 1978; Heinzel et al. 1972; Sharrock 1976; Brink 1967)
Ireland England Belgium and north-east France
Fish (primary freshwater species) o 19 24 Amphibians 3 6 16 Reptiles 1 6 8 Birds (breeding species)l 138 182 170+ 8 Mammals (except bats)2 14 30 37 Bats (breeding species) 6 11 13 General graphical record for much of the stage, and all too 1. British Isles 201, Scotland 181 2. Includes species occurring within last 3000 years (see Table 7.6) and man. often one is relying solely on the accuracy of The cold stages are characterized by climates pre radiocarbon dates. dominantly much colder than those of the present Early Devensian deposits are beyond the range of necting England with the Continent in the early land. Lagopus mutus, also recorded from Devensian day, but each covers complex sequences of climatic radiocarbon dating, or near the limits of the Flandrian. The occurrence of many of these prim deposits in England, is now restricted to the Scot fluctuations. As currently recognized, each cold method, but fortunately there exists a fairly con ary freshwater species in other parts of the British tish Highlands (Sharrock 1976). stage lasted several times longer than any inter tinuous pollen record for some of this period, and Isles, is attributed to introduction by man, and the The overall distributions of many birds within glacial. Radiocarbon dates on Devensian material the woodland interstadials can probably be distin only native fishes in these areas, including Ireland, the British Isles are clearly much influenced by show that the duration of this cold stage is at least guished on their pollen spectra (Ch. 2). A further are the secondary freshwater species which are biogeographical factors. Many species are more or 60 000 years, perhaps as much as 90 000 or 100 000 difficulty with cold-stage vertebrate faunas is that at tolerant of saltwater and penetrated the rivers via less coastal, or favour upland or lowland areas, years (see Ch. 2). most open sites the fossils, which often occur in the sea. Fossil evidence is needed to confirm or re while others breed only in, for example, southern Each cold stage covers a period in which climatic very large numbers, are generally not found in the fute this hypothesis. England or conversely only in Scotland, stressing and vegetational conditions varied widely. For datable organic horizons but in the sands and Two secondary freshwater fishes, the arctic charr the importance of climatic controls. The difficulties much of the time, however, the vegetation was gravels above and below. Salvelinus alpinus and whitefishes Coregonus spp., of disentangling such effects, from those produced herb-dominated mainly with grasses and sedges, For the reasons given above it is not possible to are restricted to isolated occurrences in cold upland by man, however, must again be pointed out but there were also milder periods which permitted reconstruct the history of the vertebrate fauna lakes in Scotland, Cumbria, North Wales, and low (Ch. 10). the growth of tree birch and conifers, and, on the through the Beestonian, Anglian or Wolstonian land lakes in Ireland. They also occur in the Alps Amphibians and reptiles with distributions other hand, intensely cold phases with very little stages even in outline. The changes in the mammal and elsewhere. The main present-day distributions which, at least in part, reflect climatic factors, in fauna or flora ('polar desert') may also have occur faunas during the Devensian, however, can be fol of these fishes are, however, in arctic and northern clude natterjack Bufo calamita (restricted to sandy red. Some beetle faunas of Devensian age indicate lowed in some detail. seas, ascending rivers to spawn. The non-migratory coastal and heath areas in England and south-west that there were short periods of warm climate The typical vertebrate faunas of the cold stages, freshwater populations are probably the descen Ireland), sand lizard Lacerta agilis (restricted to which were not accompanied by the establishment excluding periods with woodland interstadials on dants of Late Devensian fishes which became sandy coastal and heath areas in England and of woodland, although perhaps they resulted in a the one hand and polar desert on the other, com isolated when the main distributions shifted north Wales), smooth snake Coronella austriaca (distribu richer herb growth. prise: (a) extinct taxa; (b) taxa now confined to ward with the improved climate of the early tion as in L. agilis, but restricted to southern Eng During the Devensian the main phase of glacia arctic or rarely alpine regions; (c) taxa nowadays Flandrian. A find of salmonid bones and scales, land) and grass snake Natrix natrix (absent from tion appears to have been late in the stage, between characteristic of steppe; and (d) animals now con probably attributable to S alvelinus, from Esthwaite Scotland except the Midland Valley) (Arnold 1973). about 18 000 and 15 000 radiocarbon years B.P. fined to regions further south. Water, Cumbria, in a deposit dating from the end Yalden (1980) has suggested that the disjunct The relative timings of glaciations within earlier These associations are remarkably similar for the of the Late Devensian, lends some support to this ranges of B. calamita, L. agilis and C. austriaca re cold stages can only be estimated very roughly Anglian, Wolstonian and Devensian stages and re interpretation (Freshwater Biological Association sult from initial early Flandrian colonization of (see Ch. 2). There are considerable difficulties in flect climatic and vegetational conditions not repre 1960; Wheeler 1977). large areas of lowland Britain, followed by a drastic dating fossiliferous deposits to or within particular sented anywhere in the world at the present day. Similar relict distributions, paralleling those of reduction of range to coastal areas as forest replaced cold stages older than the Devensian, because There is a rather marked reduction in species certain plants, e.g. dwarf-birch Betula nana, are open herb vegetation. Climatic deterioration re such deposits are well beyond the range of radio diversity in cold-stage faunas in comparison with shown by the mountain hare Lepus timidus and stricted their spread from these refugia when the carbon dating and, unlike interglacials, also lack those of interglacial age. On the other hand, the ptarmigan Lagopus mutus. Lepus timidus, recorded forests were subsequently cleared by man. Of the a continuous pollen or other biostratigraphical abundance of bones, especially of Bison priscus, re from southern England in the Devensian, is now mammals, dormouse Muscardinus avellanarius and record. covered from many deposits of Devensian age indi indigenous only to the Scottish Highlands and Ire harvest mouse M icromys minutus are probably in The vertebrate faunas of the Middle and Upper cates that during certain phases within cold stages land, although introduced to other upland areas. In digenous only to England and Wales, because of Devensian can be dated by radiocarbon and usually the vertebrate biomass may have considerably ex lowland Britain it is replaced by brown hare Lepus climatic limitations. related to fossil floras and beetle faunas. Again, ceeded that of any time within an interglacial capensis, which however is not indigenous to Ire- however, there exists no continuous biostrati- period. Pleistocene vertebrates in the British Isles 144 145 Middle and Upper Pleistocene cold-stage faunas Table 8.1. Mammal faunas from pre-Devensian cold stages. Be, Beeston; Ang, Anglian; Wo, Wolstonian Anglian Stage
A crushed skeleton of a suslik Spermophilus sp. was .s:: c: co found in freshwater silts (,Arctic Bed') immediately o "0 Q) C ID I beneath the till in the cliff at Mundesley, Norfolk :::l c: :.= ~ Q) a: o Q) (Newton 1882b). These silts are now referred to the ti Q) c: E ti Q) E >< o Q) Q) o o o Early Anglian Cold Stage, and the pollen spectra ~ co I I as indicate open grassland and a cold climate (West ~Wo Stage/substage Be Be e-Ang ?Ang ?Ang I-Ang e-Wo Wo Wo I-Wo Bd 1980a). Unfortunately, this constitutes the only known fossil mammal definitely dated to the begin PRIMATES ning of this stage. The association of Spermophilus Homo sapiens L., man + 11 WI with unwooded vegetational conditions is consistent RODENTIA with the ecology of this genus at the present day. S permophilus sp., suslik + The Wallingford Fan Gravels occur, as remnants Lemmus lemmus (L.), Norway lemming + of a formerly more extensive sheet, near Benson, Arvicola cantiana (Hinton), extinct Oxfordshire. The deposits, about 6 m thick, main water vole + ly comprise chalky flint gravels, thought to have Microtus oeconomus (pallas), been soliflucted (coombe deposits), and angular northern vole + Fig. 8.1. Location map of Beestonian, Anglian and flint gravels and sands with seams of sands, cal Microtus gregalis (pallas), tundra Wolstonian vertebrate sites. Beestonian; careous silts and clays, largely deposited under vole + Anglian; ., Wolstonian; (cave sites shown by open fluviatile conditions (Horton & Whittow, in symbols). Bd, Brandon; Be, Beeston; BH, Baker's Shephard-Thorn & Wymer 1977). The stratifica CARNIVORA Hole, Northfleet; Brm, Broome Heath; Hm tion has been contorted by cryoturbation, and ice Crocuta crocuta Erxleben, spotted hyaena + Homersfield; Hx, Hoxne; Mu, Mundesley; TN, wedge casts also testify to permafrost conditions Tornewton Cave; WH, Water Hall Farm; WI, during the deposition of these beds. The Walling PROBOSCIDEA Wallingford; Wo, Wolston, WR, West Runton. cf. Mammuthus trogontherii Pohlig, ford Fan Gravels appear to grade to the level of the extinct elephant + Winter Hill or Black Park Terrace of the River Mammuthus primigenius Blumenbach, Thames, which suggests an Anglian age, but this is mammoth + + + + by no means certain. Beestonian Stage In Hall's Pit near Benson, Dr A. Horton found PERISSODACTYLA in the calcareous silt unit (unit ii) non-marine Mol Equus ferns Boddaert, horse + + + + At present only four fossil mammal specimens can lusca and rare vertebrate remains including frog Coelodonta antiquitatis Blumenbach, be confidently assigned to the Beestonian Cold woolly rhino + Rana sp. and/or toad Bufo sp., Norway lemming + + Stage. A mandible of an elephant apparently in Lemmus lemmus, extinct water vole Arvicola can ARTIODACTYLA termediate between Archidiskodon meridionali~ and tiana, tundra vole Microtus gregalis, horse Equus Megaceros giganteus Blumenbach, giant deer + M ammuthus trogontherii, was found in situ in the ferns, and reindeer Rangifer tarandus. Cervus elaphus L., red deer + cliff at Beeston, Norfolk, in marine beach sands A single well-dated record of red deer Cervus Rangifer tarandus L., reindeer + + + and gravels (Bed 0), assigned to the Beestonian elaphus from the Late Anglian is known from Hox Indet. deer .+ Stage, and resting on an eroded surface of Pasto ne, Suffolk. Three foot bones were found resting Bison sp. or Bos sp., bison or nian marine silts (West 1980a). Pollen spectra from on the surface of the Anglian till in lacustrine clay aurochs + + + + Bed 0 consist mainly of grasses, birch and pine, mud and marl (Bed F) containing pollen indicative but with some pollen of thermophilous trees prob of the Anglian late-glacial (Spencer, in West 1956). ably reworked from the Pastonian. A cool climate The pollen points to open herb-dominated vegeta Pre-Devensian Cold Stages ly common, but in general are assumed to be of and grassy vegetation with stands of birch and pine tional conditions with sea buckthorn Hippophae Devensian age in the absence of clear stratigraphical can be tentatively inferred. scrub. This gives a clear association of C. elaphus (Fig. 8.1, Table 8.1) evidence. At West Runton, Norfolk, the topmost bed of with unwooded vegetational conditions (Ch. 5). The paucity of faunas and the impossibility of the pre-West Runton Freshwater Bed (Cromerian) In the valley of the River Waveney, which is the The very few available 'cold' faunas for which dating deposits precisely within the stage, pre succession (Bed i) consists of sands and silts with boundary between Norfolk and Suffolk, two dis a pre-Devensian age can be demonstrated are cludes any attempt at present to reconstruct faunal marine shells assigned to the Beestonian Stage tinct terraces occur above pollen-dated Ipswichian very similar to typical faunas of Devensian age. histories for the Beestonian, Anglian and Wolsto (West 1980a). Bison and a deer are recorded from deposits, at Wortwell, Norfolk (see Ch. 2) which The implication is that the former are perhaps fair- nian cold stages. this bed. are at approximately the level of the modern flood- Pleistocene vertebrates ill the British Isles 146 147 Middle and Upper Pleistocene cold-stage faunas plain (Sparks & West 1968). The higher, and there gravels yielded a very interesting fauna of fresh characteristic Ipswichian fauna, suggests that both include Mammuthus trogontherii, Coelodonta anti fore presumably older, Homersfield Terrace surface water fishes (Osborne & Shotton 1968). According to the Glutton and Bear Strata are of Wolstonian age; quitatis, Rangifer tarandus and Bison priscus. is at approximately 8 m above the floodplain. In the H. Greenwood is Osborne & Shotton (1968), the most probably dating from near the end of the At Steinheim (Murr), West Germany, the 'Main early 1950s mammalian remains were collected species present include pike Esox lucius, gudgeon stage. Unfortunately, however, there are strong Mammoth Gravels' overlying the gravels with a from sands and gravels of this terrace at Gobio gobio, possibly minnow Phoxinus phoxinus, suspicions that the faunal remains from the Glutton Holsteinian fauna (Ch. 7) and therefore of probable Homersfield, Suffolk, by Prof. B. Funnell, and the chub Leuciscus cephalus, roach Rutilus rutilus, Stratum are mixed with material of interglacial age. Saalian age, have yielded a fauna which includes fauna comprises mammoth Mammuthus primigenius, various other undetermined cyprinids, perch Perca Mammals recorded from the Glutton and Bear elephants referred to both M. trogontherii and M. woolly rhino Coelodonta antiquitatis, horse Equus fluviatilis and three-spined stickleback Gasterosteus Strata are bear Ursus sp., wolf Canis lupus, red fox primigenius, C. antiquitatis, M egaceros giganteus ferus, reindeer Rangifer tarandus, giant deer aculeatus. The beetle fauna is stated to include some Vulpes vulpes, lion Panthera leo, badger Meles meles, and B. prise us (Kahlke 1975a). Megaceros giganteus and a large bovid Bison or Bos, southern forms and lacks arctic stenotherms, and clawless otter Aonyx antiqua Blainville, glutton Similar faunas assigned to the 'Rissian' (Saalian, and is thus a typical cold-stage fauna, identical to the pollen spectra indicate meadow birch woodland Gulo gulo, mountain hare Lepus timidus, horse Wolstonian) Cold Stage are recorded from the cave faunas of Wolstonian or Devensian age. Recent Kelly (1968). Gobio gobio and L. cephalus do not E quus ferus, rhinoceros cf. C oelodonta sp., reindeer sequences of Southern France (Lumley 1976). The work by Coxon (1979) has shown that at nearby occur further north than southern Scandinavia at Rangifer tarandus and a small bison or aurochs southern location, however, is reflected in the Flixton, the Homersfield Terrace deposits are in the present day. (Sutcliffe & Zeuner 1962). Small vertebrates in occurrence of such temperate species as fallow deer timately associated with chalky till. This and the The general picture is of a cool but not arctic cli clude fishes, frogs and/or toads, common hamster Dama dama and aurochs Bos primigenius in many of height relationships to the type Hoxnian intergla mate, suggesting interstadial conditions within the Cricetus cricetus, a small hamster assigned to the ex these faunas and the proximity to mountainous cial lake deposits at Hoxne, also in the Waveney Early Wolstonian. tinct AUoericetus bursae, steppe lemming Lagurus areas by the many finds of ibex Capra ibex and cha Valley area, suggest an Anglian age for this terrace. At Northfleet, Kent, in a small tributary valley lagurus, arctic lemming Dicrostonyx torquatus, N or mois Rupicapra rupicapra. Previously the terrace had been tentatively ascribed of the River Thames, coombe rock, frost-heaved way lemming Lemmus lemmus and northern vole to the Wolstonian Stage by Sparks and West. chalk, soliflucted gravels and loess underlie fresh Microtus oeconomus (Kowalski 1967; Sutcliffe & water silts, attributed to the Ipswichian Interglacial Kowalski 1976. The status of the records of snow on the basis of temperate non-marine molluscan vole NI ierotus nivalis from this site have been discuss faunas. Further periglacial deposits occur above ed in Chapter 3. Birds identified from these beds Devensian stage (Kerney & Sieveking, in Shephard-Thorn & by Harrison (1980) include ducks, kestrel Falco tin W olstonian Stage Wymer 1977). Excavations at the Baker's Hole pit nunculus, eagle owl Bubo bubo, cross bill Loxia cf. An outline of the stratigraphy, climate, vegetation early in the present century (Smith 1911) yielded a curoirostra, carrion crow Corous corone and raven and history of the beetle faunas of this stage is The surface of the Broome Terrace of the River few large-mammal remains (Table 8.1) from the Corvus corax. A fragment found in the Glutton given in Chapter 2. Waveney lies about 3 m above the floodplain. earlier periglacial deposits, which appear to be of Stratum is the sole basis for the description of the Teeth of mammoth Mammuthus primigenius, and Wolstonian age - again probably from late in this extinct patridge Alectoris sutcliffei Harrison. Con bones of a bovid Bison sp. or Bos sp., have been stage. firmation of this record with more material would Sites and faunas (Fig. 8.2, Table 8.2) found in the sands and gravels of this terrace at Lacustrine stonewort Chara marls at Water Hall be desirable. Broome Heath, and organic deposits at this site Farm, Hertfordshire, occur beneath sands and The Glutton and Bear Strata faunas show a mix The vertebrate faunas of the Devensian are far bet have yielded a full-glacial flora of plant macrofos gravels containing Hippopotamus amphibius and ture of steppe, boreal-arctic, and temperate com ter known than those of earlier cold stages, and sits. The discovery of reworked Hoxnian plant other mammals indicative of the Ipswichian inter ponents as is seen in many Devensian faunas, have been found in most areas of the British Isles. remains at this site by Coxon here confirms the glacial. The marls are penetrated by one or more ice although the steppe contribution, including hams Open-site records are mostly from fluviatile de Wolstonian age previously suggested by Sparks wedge casts which indicate a Wolstonian age, most ters and Lagurus lagurus, is especially marked. The posits, but some of the best-preserved material and West (1968). probably near the end of this stage. Small verte presence of the distinctly temperate Meles meles, comes from lacustrine sediments. Most assemblages Large-mammal remains have been collected from brates from the deposit include pike Esox lucius, and if correct of Loxia curoirostra, a species depen from caves are also of Devensian age. Unfortunate pits in the fluviatile Baginton-Lillington Gravel in frog Rana sp. and/or toad Bufo sp. and northern dent on coniferous trees, however, either indicates ly, however, the majority of finds are neither accur the vicinity of Wolston, Warwickshire (Shotton vole Microtus oeconomus. interstadial conditions or may reflect contamination ately stratified nor dated. 1953, 1968). These deposits are the lowest member In Tornewton Cave, Torbryan, Devon, a series with interglacial material. of the type sequence of the Wolstonian, and most of deposits with vertebrate fossils represents parts Early Devensian (pre-SO 000 radiocarbon year B.P.) probably date from the very early part of this stage. of the Wolstonian, Ipswichian, Devensian and Early Devensian vertebrate faunas are known so far The fauna is given as: spotted hyaena Crocuta ero Flandrian Stages (Sutcliffe & Zeuner 1962) (see Comparison with Continental faunas almost entirely from only two localities, both in cuta, mammoth Mammuthus primigenius, straight Fig. 7.11). The oldest fossiliferous bed, termed the Norfolk (Table 8.2). tusked elephant Palaeoloxodon antiquus, woolly Glutton Stratum, thought to have been soliflucted Some of the faunas from central Europe (e.g. In the early 1960s excavations for a flood-relief rhino Coelodonta antiquitatis, horse Equus ferus, into place, contains stalagmite apparently shattered Stninska Skala, Vertesszollos, Mauer) which have channel along the Fenland margin at Wretton, reindeer Rangifer tarandus and bison Bison sp. or by freeze-thaw processes, and has yielded arctic been ascribed to the Elsterian (Anglian) (Kahlke Norfolk, revealed an extremely complex sequence aurochs Bos sp. The record of P. antiquus, if cor mammals. The immediately overlying Bear Stra 1975a), could be more convincingly correlated of Ipswichian and Devensian deposits of the River rect, is unexpected in a cold-stage fauna. tum has much the same fauna but lacks evidence of with the Cromerian Interglacial (Ch. 7). Faunas of Wissey (Sparks & West 1970; West et al. 1974) At Brandon, Warwickshire, fossiliferous silts periglacial conditions. The stratigraphical position probable Elsterian age listed by Kahlke, e.g. (Fig. 8.3). The Devensian sediments record deposi occupying a channel within the Baginton-Lillington of these beds, beneath the Hyaena Stratum with its Bornhausen and Neuekrug (Harz), West Germany, tion in a wide range of environments. Much of the Table 8.2. Mammal faunas from well-stratified/dated Devensian open sites. Circled crosses indicate radiocarbon-dated records. Other Middle and Upper Devensian records are more broadly associated with the radiocarbon dates given.
Early Middle Late
0 ~ 0 0 ~. Q) C\J a O'lco C') 0 '"~ en I ::;: CO ::::: ..0 0 CO ~ 0 C') 0 ::J - "E
CARNIVORA Canis lupus L., wolf + + + Alpex lagopus CL.), arctic fox + Ursus arctos L., brown bear + (±) +
PROBOSCIDEA Mammuthus primigenius Blumenbach, mammoth + + + + + + + + +
PERISSODACTYLA Equus ferns Boddaert, horse + + + + + + + + + + Coelodonta antiquitatis Blumenbach, woolly rhino + + + + + + + + + ARTIODACTYLA Cervus elaphus L., red deer + + Rangifer tarandus L., reindeer + + G) G) + + + + Megaceros giganteus CBlumenbach), giant deer + + + Alces alces CL.), elk G) G) Bison priscus Bojanus, extinct bison + + G) G) + + + Ovibos moschatus Zimmerman, musk ox +
1. Could be slightly older
...... -1::.. \Q
~ 5..: \::l... ~ I::l ;::: \::l... ~ '\::i- ...,~ "v ~ a ~'" ~ '"Cl ~ '" ~ ~ l:: ~ Pleistocene vertebrates in the British Isles 150 151 Middle and Upper Pleistocene cold-stage faunas
Fx, Flixton; Fs, Fisherton; GC, Gough's Cave; B~ HF, High Furlong, Blackpool; Is, Isleworth; Kb, Kirkby-on-Bain; KC, Kent's Cavern Ki, Kilmaurs; Kw, Kew; Ns, Neasham; Nz, Nazeing; Ox, Oxbow; PH, Picken's Hole; Sh, Shandon Cave; occ Kb SH, Sun Hole; Sp, Sproughton; TC, Tattershall o rc' Castle; TN, Tornewton Cave (Reindeer Stratum); Tw, Twickenham; UW, Upton Warren; Wr, Wretton.
sequence comprises fluviatile sands, silts and gravels, with fine clastic and organic sediments in small enclosed basins thought to have been formed by the melting of ice mounds. Eight local pollen zones A to H, representing three herb substages and two woodland substages (interstadials) were recognized on the basis of pollen spectra. The later interstadial is thought to be equivalent to the Chel ford (Cheshire) woodland substage, and the earlier designated as the Wretton woodland substage. These may correlate with the BrliSrup and Amers Ca) foort interstadials respectively recognized In Denmark, North Germany and the Netherlands. Bones of wolf Canis lupus have been found in situ in beds with pollen spectra of herb substage I type (Fig. 8.3b). The majority of the Wrettop mammal material was found loose in the mounds of exca vated deposits, but all or most of it probably comes from the beds of herb substage I age; a conclusion corroborated by the uniformity of the fauna. The abundant fossils, collected by Dr K. A. }oysey, are now being studied by him in collaboration with the author (Table 8.2). The extinct bison Bison priscus is by far the best represented taxon, followed by reindeer Rangifer tarandus. The other taxa, Canis lupus, arctic fox Alopex lagopus, bear Ursus sp., mammoth Mam muthus primigenius, woolly rhinoceros Coelodonta antiquitatis and horse E quus ferns, are mostly repre sented only by one or two specimens each. The pic ture of large herds of grazing bison and reindeer is entirely consistent with the palaeobotanical evi dence for a rich herbaceous vegetation. The pre Fig. 8.2. Location map of Devensian vertebrate dominance of male shed antlers at Wretton indi sites. Ill, Early Devensian; Middle Devensian (or cates that reindeer herds were in the area mainly in uncertain); ~, Late Devensian; (cave sites shown the winter (see Ch. 5). by open symbols). AH, Aveline's Hole; BD, Brean Cb) Down; Bd, Brandon; Bg, Ballaugh; Bk, Beckford; Beetle remains recovered from zones A and B of Fig. 8-.3. Early Devensian river terrace deposits at Wretton, Norfolk. (a) general view of flood-relief El, Ballybetagh; Bp, Brimpton; BS, Barnwell Wretton herb substage I were interpreted as sug channel looking eastwards towards Wretton Fen Bridge. The high banks on either side consist of excavated Station, Cambridge; Bsh, Bishopbriggs; Ca, gesting open, rather marshy, grassland with sandy spoil in which many bones were found. In situ deposits are exposed in sections above and below water level. Castlepook Cave; Cor, Corstophine; CC, Creswell soils, and a cool but not arctic climate (Coope, in (b) section in south bank showing sands and silts, overlain by organic deposits yielding herb substage I Crags (Robin Hood's Cave, Pinhole Cave); Cf, Cefn West et al. 1974). There is, however, a marked dis pollen spectra (see text). A partial skeleton of wolf Canis lupus was found in the sands and silts. The scapula Cave; CH, Clifford Hill; Cs, Coston; Fd, Fladbury; agreement between the beetle faunas and the pollen and another bone are visible in the photo to the left and above the trowel.
* d P leistocene vertebrates in the British I sies 152 153 Middle and Upper Pleistocene cold-stage faunas spectra for zones F and G. According to Coope, the the gravels has yielded a fossil beetle fauna of arctic beetles indicate a severe arctic climate, whereas the type and radiocarbon dates of 44 300 ± ~ ~~~ and pollen shows the development of pine Pinus and 42 100 ± ~ igg years B.P. (Lab. no. Birm-408, spruce Picea forest. This discrepancy has yet to be 398. A higher silt band up to 2 m thick has, in resolved. contrast, yielded a temperate beetle fauna, and Organic deposits in sands and silts of the second radiocarbon dates of 43 000 ± i ~gg and terrace of the River Cam at Sidgwick Avenue, Cam 42 200 ± 1 000 years B.P. (Birm-341, 409). bridge, have been attributed to the Early Deven Numerous large-mammal bones have been found at sian (Lambert, Pearson & Sparks 1962). Plant mac this site, of which as at Wretton the majority are of rofossils indicate an open unwooded landscape and Bison priscus, with Rangifer tarandus in second a climate colder than today. A skull of Bison priscus place (Rackham 1978). Bones of both these species was recovered from these deposits. have been found in situ in the upper silt bed, but Organic sands and silts beneath gravels at Coston most finds are not stratigraphically located and may near Barnham Broom, Norfolk, have yielded ma cover several thousands of years, within which croscopic plant remains, molluscs and vertebrate there may have been considerable changes in cli remains. Preliminary pollen analyses by P. L. Gib mate. The presence of abundant shed male antlers bard indicate spruce Picea/pine Pinus forest giving at the site indicates that Rangifer tarandus was pre way to an unwooded environment dominated by sent in the winter months (Ch. 5). grasses and sedges. A correlation with the end of Of particular interest is the occurrence of numer the Early Devensian Chelford interstadial seems ous dung beetles in the upper silt bed coleopteran very probable. Ipswichian interglacial deposits also fauna, indicating the presence of large herbivores in occur in the gravel pit and a hippopotamus Hippo the area during the summer months as well. potamus amphibius tooth has been recorded from Numerous large-mammal remains were dis the site, although not in situ. covered in the late 1950s and early 1960s at Will The majority of large-mammal bones from Cos ment's Pit, Isleworth, Middlesex, and collected by ton are of Bison priscus. Other taxa, represented by J. W. Simons. The deposits, laid down by the Riv a few specimens each, comprise Coelodonta anti er Thames, comprise a sequence about 5 m thick of quitatis, Mammuthus primigenius, Rangifer tarandus, gravels, sands and organic silty clays, the latter giv and red deer Cervus elaphus. All of these species ing a radiocarbon date of 43 140 ± i ~~~ years B.P. were recorded in situ in the organic deposits, (Birm-319) (Coope & Angus 1975). The vertebrate although many unstratified finds have also been fauna (Table 8.2) includes fishes, amphibians, and made. Small-vertebrate remains collected in situ by the voles Microtus oeconomus and Microtus gregalis the author include: three-spined stickleback Gaster from the dated silty clay bed. Of the far more abun osteus aculeatus, frog Rana sp. and/or Bufo sp., dant large-mammal remains, bear Ursus sp., rein toad, short-tailed vole Microtus agrestis and extinct deer Rangifer tarandus and bison Bison priscus are water vole Arvicola cantiana. definitely recorded from the silty clay bed, while A basal antler fragment of C. elaphus has been the majority of finds are from the gravels and found in deposits attributed to a post Chelford in sands, and are therefore less precisely dated. terstadial at Brimpton, Berkshire (Bryant & The vertebrate assemblage is dominated by the Holyoak 1980). remains of Bison priscus. Remains of Rangifer taran dus are also conspicuous, though much less abun Middle Devensian (50 000 to 26000 radiocarbon dant than Bison; but this impression is exaggerated years B.P.) since they include many shed antlers, and each Middle Devensian vertebrate faunas are available reindeer produces several pairs of antlers in its life from a number of open sites in association with time. The other taxa are each represented by one or radiocarbon dates, beetle faunas and pollen (Table a few specimens only. 8.2). The organic silty clay bed has produced a rich At Tattershall Castle, Lincolnshire, several metres beetle fauna, described by Coope and Angus, with Fig. 8.4. Section showing sequence of Middle Devensian deposits at Tattershall Castle, Lincolnshire. A of fluviatile gravel, deposited by the River Witham, fossiliferous organic rich channel fill (dark) near the base, is succeeded by· cross-bedded sands and gravels representatives of aquatic pool, riparian and terrest (the holes were made by sand martins). overlie fossiliferous organic deposits dated by pol rial environments. The latter were interpreted as len analyses to the Ipswichian interglacial (Girling comprising an adjacent marshy area with lush her 1974) (Fig. 8.4). A thin silt band near the base of baceous vegetation, and patchy vegetation includ- Pleistocene vertebrates in the British Isles 154 155 M iddle and Upper Pleistocene cold-stage faunas ing heath on higher and drier ground. More recent with insects and pollen and a radiocarbon date of Table 8.3. Devensian radiocarbon-dated mammal records based on bone collagen. (Data from pollen analyses, by P. L. Gibbard, show typical 38 600 ~ : ~~~ years B.P. (NPL-163B) (Gaunt, Radiocarbon 1959-79; Camp bell 1977) Devensian herb pollen spectra dominated by grass Coope & Franks 1970). The pollen indicates a thin es and sedges. According to Coope and Angus, the herbaceous plant cover with patches of bare Lab. and assay Date in radiocarbon beetle fauna consists of temperate species now ground, and the beetles also point to arctic condi no. years S.P. found in Britain plus a smaller number of more tions. southern or eastern distribution, indicating mean River sands and gravels at Fladbury, Worcester Homo sapiens, man July temperatures °of 18 QC or more, i.e. higher shire, include a peat horizon radiocarbon dated to Paviland, Glamorganshire (,Red Lady') BM-374 18 460 ± 340 than at the present day. The absence of trees was 38 200 ± 600 years B.P. (GrN-1269). The verte attributed to the climatic amelioration having b~en brate fauna from the gravels comprises only Mam Ursus arctos, brown bear too short for trees to immigrate. The possibility of muthus primigenius, Coelodonta antiquitatis and horse Sun Hole, Somersetshire BM-524 12 378 ± 150 grazing pressure by the large herbivores, prevent Equusferus (Coope 1962). The beetles from the peat Kent's Cavern, Devonshire GrN-6203 14 275 ± 120 ing tree growth, was also mentioned. Tree growth again indicate an arctic environment. Robin Hood's Cave, Derbyshire BM-602 28 500 ± ~ ~~ may, however, have been restricted by severe win At Kirkby-on-Bain (Tattershall Thorpe) Lincoln Kent's Cavern, Devonshire GrN-6202 28 720 450 ters (West 1977b). shire, only some 4 km from Tattershall Castle in ± It is not known whether Bison priscus was present the valley of the River Witham, several metres of Mammuthus primigenius, mammoth at Isleworth throughout the year. Provisional study sands and gravels (Fig. 8.4) have produced a num Cae Gwyn Cave, Flintshire Birm-146 of the numerous Rangifer tarandus antlers from the ber of large-mammal bones. A silt band, radiocar 18 000 ± ~ igg English Channel off Sussex site suggests that most of the shed antlers are from bon dated to 34 800 ± 1 000 years B.P. (Birm- Gif.-UlO 19 300 ± 700 Castlepook Cave, County Cork males, indicating presence of the herds in the win 250), has yielded an arctic beetle fauna (Girling D-122 33 500 ± 1 200 ter months (see Ch. 5). 1974). Rackham (1978) has listed the fauna (Table Little Rissington, Gloucestershire Birm-466 34 500 ± 500 As at Tattershall Castle the large and varied 8.2) and pointed out the striking differences when Oxbow, near Leeds NPL-162B 38 600 ± ~ ~~g dung-beetle fauna indicates that large herbivores compared with the earlier sites. At Kirkby Bison were present during the summer. Carrion beetles priscus is virtually absent, and no remains of Ran Coelodonta antiquitatis, woolly rhino are also recorded. gifer tarandus have been found. The faunal assem Ogof-yr-Ichen, Caldey Island, Pembrokeshire Birm-340 22 350 ± 620 At Upton Warren, Worcestershire, many bones blage is here dominated by remains of mammoth Bishopbriggs, Lanarkshire GX 27 550 ± :~~g and teeth of large mammals were found in terrace Mammuthus primigenius and rarer woolly rhinoceros Kent's Cavern, Devonshire GrN-6201 28 160 ± 435 deposits of the River Salwarpe (Coope, Shotton & Coelodonta antiquitatis. Leadenhall Street, London GrN-4630 29450 ± 350 Strachan 1961). Silt bands within the sands and At Brandon, Warwickshire, organic silts, gravels yielded pollen spectra indicative of treeless radiocarbon dated to about 28 200 years B.P., E quus ferus, horse herb-dominated vegetation. One of these silt bands occur at the base of gravels of the River Avon num Robin Hood's Cave, Derbyshire BM-604 10 590 ± 90 gave a radiocarbon date of 42 100 ± 800 years ber 2 terrace (Shotton 1968). They contain plant Kent's Cavern, Devonshire GrN-6324 38 270 ± : i~ B.P. (GrN-1245). Only a few remains of three and insect fossils (Kelly 1968; Coope 1968a) both of spined stickleback Gasterosteus aculeatus were, which indicate an open, herb-rich, treeless environ Megaceros giganteus, giant deer however, actually found in this dated silt. Bones ment. Dwarf birch Betula nana and dwarf willow Kent's Cavern, Devonshire GrN-6204 12 180 ± 100 assigned to common frog Rana temporaria, arctic S alix herbacea were also present. The beetles imply Brandesburton, Yorkshire (reworked) Birm-55 12 850 ± 250 lemming Dicrostonyx torquatus and a vole Microtus a severe arctic climate. The single vertebrate record sp. were recovered from the other silt bands. The from the silt deposits is of arctic lemming Dicros Rangifer tarandus, reindeer large-mammal bones were mostly found by work tonyx torquatus. Dead Man's Cave, Anston, Yorkshire BM-440b 9 750 110 men in the sands and gravels so that the majority of ± Organic silt bands within terrace sands and Rodden's Port, County Down records are only broadly associated with the LJ-658 10 250 ± 350 gravels of the Carrant Brook at Beckford, Worces Ossoms Cave, Staffordshire radiocarbon date. Mammoth Mammuthus pri tershire, gave a radiocarbon date of 27 650 ± 250 GrN-7400 10 590 ± 70 Cattedown Cave, Plymouth, Devonshire migenius is much better represented than in the years B.P. (Birm-293) (Briggs, Coope & Gilbertson BM-729 15 125 ± 390 Coygan Cave, Camarthenshire faunas from Isleworth and Tattershall Castle, 1975). Non-marine molluscs from the silt were BM-499 38 684 ± ; ~i! although both Bison priscus and Rangifer tarandus thought to suggest marshy ground and exposed Kilmaurs, Ayrshire Birm-93 >40000 are still very abundant. open environments with lack of continuous grass cf. Bison priscus, extinct bison From the abundance of both dung and carcass cover. The beetle assemblage is of arctic tundra beetles, Coope et al. suggested that the pools on the type, again suggesting sparse vegetation. One spe Broadway, Worcestershire Birm-656 26 600 ± ~~~ floodplain, represented by fossiliferous silt bands, cies of carcass beetle occurs. According to White Kent's Cavern GN-6325 27 730 ± 350 were used as waterholes by large mammals. head (in Shotton, 1977c) the large-mammal bones Ovibos moschatus, musk ox An organic silt horizon at Oxbow near Leeds from the sands and gravels, and therefore only yielded a tusk of M ammuthus primigenius together broadly associated with the radiocarbon date, in- Clifford Hill, Northamptonshire BM-725 18 213 ± 310 Pleistocene vertebrates in the British Isles 156 157 Middle and Upper Pleistocene cold-stage faunas
Table 8.4 Mammalian faunas of selected Middle and Upper Devensian cave deposits. Thousands of radiocarbon ~c, years B.P. ~c, ....QI ~c, ~o IJ)<::- Qlc, 0.: 0.: ,IJ) . C,U ~u cri cri ,IJ) ~~ IJ) ~ ~ (f) ~ «-. ~. ~. Q/' ca co Flandrian I 0) 0) >. 0)>' 1.. 0) 0) E I.{') >0 10 ~~OO-----=I~·------~------co-> ::J COo ~ O) ~~ 01.{') o~ "0 ... °E cC/) 0-.::1' ~C') 15 ------, ------•• -- c ::J I IC') oC') o .... .8(tj (f)- 0""'" Late Main glaciation Cl. - ~ "- ~~ "~I.{') "c C') 0)"0 Devensian 20------f---??-----Q------0)- 0)"0 0) .... 0) .... -c cC/) c c ~O) ,:::t:.0) .... ~ u>' u>' 0- o'a; .- co .- co ~~ I-W 1-0: Cl...... J Cl...... J 0..= 25 <> ,+9+ 9 9 +9' Insectivora 30 Talpa europaea L., mole + 35 <>t 9 Primates Middle