Bull. Soc. belge Géol., Paléont., Hydrol. T. 79 fasc. 2 pp. 167-174 Bruxelles 1970 Bull. Belg. Ver. Geol., Paleont., Hydrol. V. 79 deel 2 blz. 167-174 Brussel 1970

MAMMALS OF THE CRAG AND FOREST BED

B. McW1LLIAMs

SuMMARY. In the Red and Norwich Crags mastodonts gradually give way to the southern elephant, large caballine horses and of the Euctenoceros group become common. Large rodents are represented by Castor, Trogontherium and rarely Hystrix; small forms include of Mimomys. Carnivores include hyaena, sabre-toothed cat, leopard, polecat, otter, , seal and walrus. The Forest Bed Series had steppe and forest forms of the southern elephant and the mastodont has been lost. Severa! species of giant deer become widespread and among the many rodents are a. number of voles which develop rootless cheek teeth. The mole is common. Warmth indicators include a monkey, and more commonly hippopotamus. Possible indicators of cold include glutton and musk ox. Rhinoceros is widespread, and it is a time of rapid evolution for the elk. Carnivores include hyaena, bear, glutton, polecat, marten, wold and seal.

The interpretation of mammalian finds from is represented by bones which resemble the the Crags and Forest Bed is not an easy mole remains but are about twice their size. matter. A proportion of the remains have been derived from eatlier horizons, others are Order Primates discovered loose in modern coastal deposits, and early collectors often kept inadequate The order is represented at this period m records. Owing to the uncertain processes of England by a single record of Macaca sp., the fossilisation or inadequate collecting there are distal end of a teft humerus from a sandy many gaps in our knowledge of the ­ horizon of the Cromerian at , ian faunas of these times. Norfolk. Most workers feel grave doubts about the alleged Crag date of the human jaw Order Insectivora found at Foxhall near Ipswich, but as the bone bas been lost for more than a century Three shrews have been recorded from the fully conclusive evidence is not available. Series, one, Sorex savini Hinton, as large as any known or living form, a second, Sorex runtonensis Hinton, Order about the size of the modern pygmy shrew, and the third, Neomys newtoni Hinton, similar In the Lower and early Middle of to our modern water shrew and possibly its East Anglia more than twenty species of predecessor. carnivore have been recorded, though now­ A mole, similar to Talpa europaea L. is here are they common. common in the temperate freshwater beds of The typical Lower Pleistocene hyaena, the Cromerian, and is mainly represented by Hyaena perrieri CROIZET & JOBERT, is found its relatively robust and characteristically in the Crags, and a much larger species, shaped humerus, though other bones and Hyaena brevirostris Aymard, is typical of the mandibles are known. The European desman, early Forest Bed where it is soon replaced by Desmana moschata Pallas, now restricted to the spotted hyaena, Crocuta crocuta Erxleben. south-east Russia and central western Asia, There were two sabre-toothed cats; a larger

167 species of Crag date, Homotherium sainzelli The boundaries between these species in the Aymard, recorded as far north as Derbyshire Forest Bed are rather blurred. in a cave deposit, and a smaller species, Seal occur throughout the Crags and Homotherium latidens Owen, known in the Forest Bed but in the absence of cranial Cromerian by a few fine canines. A single cat remains little can be said of their specific tooth from the same horizon resembles Felis identity. When limb-bones are compared with lunensis Martelli rather than the modern wild modern North Sea forms they show no cat, and a leopard, Felis pardoides Owen, has recognisable differences. A few walrus tusks been recorded from the Red and Norwich and bones described under the name Triche­ Crag. The lion, Felis leo L., first reached this chus huxleyi LANKES!ER have been found in country in the Forest Bed as a spectacularly both Crags and Forest Bed but three mineral­ large form. ised crania dredged off the Norfolk coast are Severa! mustelids are known from East presumably more recent. Anglian deposits but they are rather rare. A glutton, probably Gulo schlosseri Kormos, Order Proboscidea rather than Gulo gulo L., has been found in Forest Bed strata; unlike the latter species its The central element of the Crag and Forest presence may not indicate cold conditions. Bed fauna are the mastcidonts and elephants, The probable ancestor of the pine marten which were widely distributed and show occurs in the Forest Bed as Martes vetus evolutionary trends in the structure of their Kretzoi, with a large polecat, Pannoniotis teeth which can be matched with environ­ pliocaenica Kormos, first found in a late mental changes which characterise the Pleisto­ Pliocene forest fauna in Hungary. A second cene itself. polecat, Enhydrictis ardea Bravard, has been Two mastodonts survived from the Pliocene recorded from the Red Crag, but polecats of in Europe, but one, Zygolophodon borsoni modern type are first found in the Forest Bed, HAYS, died out early in the Pleistocene and is possibly Mustela putorius L. itself. Remains only represented in the Red Crag by a few of the weasel, or possibly its predecessor very worn molars in the basement bed which Mustela praenivalis Kormos have also been are obivously derived. The second mastodont, found, and in the Norwich Crag there occurs Anancus arvernensis CROIZET & JOBERT, was an otter, Aonyx resvei Newton. the characteristic proboscidean of the early Wolf, Canis lupus L., is undoubtedly present Pleistocene and occurs in both the Red and in the Forest Bed, but earlier records, and Norwich Crags but by the end of the Lower those of a fox from this period of the Crags Pleistocene was extinct in this country and the are rather uncertain. southern elephant, Archidiskodon meridionalis An unusual mammal recorded from three NESTI was common. Molars of the southern localities of the Red Crag is a panda, Parai­ elephant are found in the Red and Norwich /urus anglicus BoYD DA WKINS, evidently Crags, and in the latter are as numerous as closely related to the modern lesser panda those of the mastodont. Limb bones and which is now restricted to bamboo ·forests on vertebrae become better represented in the the slopes of the Himalayas. younger Crag sands and by Forest Bed times Although bear remains, probably are abundant at several localities. It is at this arvernensis CROIZET & JOBERT, are· known period that the species begins to show a from two Red Crag localities it is not until division into presumed forest and steppe forms Forest Bed times that representatives of this which diverge into the temperate straight­ family become relatively common. The fossils tusked elephant, Palaeoloxodon antiquus FAL­ can be separated into two species, Ursus CONER & CAUTLEY, and the steppe mammoth, deningeri REICHENAU, and Ursus savini AN­ Mammuthus trogontherii PoHLIG. The southern DREWS, with the possibility of a third species, elephant ranged through Europe as far as the Ursus spelaeus ROSENMULLER & HEINROTH. Black Sea and North Africa and is

168 guished by its large, gently-curving tusks, wide systematics of this family are rather confused. molars with short, broad grinding surfaces Rolled teeth of Hipparion cf. gracile KAUP enclosing relatively few widely spaced lamel­ occur in the Red Crag, obviously derived lae. The adults reached a shoulder height of from an older horizon. over 12 ft. and have correspondingly massive limb bones. Order Artiodactyla The forest form of the southern elephant appears to have become a distinct species by Pig fossils from the Red Crag are scarce and Forest Bed times. Palaeoloxodon antiquus may belong to Sus strozzii MENEGHINI. The FALCONER & CAUTLEY has rather narrow deep first satisfactory remains corne from the Forest molars, the lamellae in the grinding surface of Bed of Norfolk. These include a fine skull and the lower molars in particular tend towards mandible from one individual which appears a lozenge-shaped pattern of wear with rela­ to be a large form of the modern wild hog, tively thick, coarsely folded enamel. The tusks Sus scrofa L. are large and gently curved. The hippopotamus which reached this Forms transitional between the southern country by Forest Bed times has often been elephant and the steppe mammoth, Mam­ thought of as a separate species from the muthus trogontherii PoHLIG, are first found in modern Hippopotamus amphibius L., but the the Forest Bed. Molars of this species have a only difference is in size, the early form being relatively large number of lamellae which tend rather larger than the modern . Hippo­ towards a division into three parts in the potamus is absent until the last interglacial, posterior part of the grinding surface. At its the lpswichian, where it occurs in some acme this species attained a shoulder height abundance. of over 14 feet and later in the Pleistocene gave Deer remains form an important part of rise to the woolly mammoth, Mammuthus the fauna of the Crags and Forest Bed but are primigenius BOJANUS. a difficult family to work on as most identifi­ cations are based on the antlers. Other parts Order Perissodactyla of the skeleton are grouped with antlers mainly on size because associated bones are rare. A number of tapir teeth described under the Where similar sized deer occur together the species Tapirus arvernensis CROIZET & JOBERT problems are formidable. have been found in the nodule bed of the Four species of the genus Euctenoceros have Red Crag, these may have been derived from been separated on the basis of the forms of the a Pliocene source. antlers, which are quite large and have a few There are a few records of rhinoceros from openly arranged slightly curved tines. Eucte­ the Crags, but not until Forest Bed times are noceros ctenoides NEsTI is rare, possibly remains common, where a single species, present in the Norwich Crag and recorded a Dicerorhinus etruscus F ALCONER is present. few times from the Forest Bed. Euctenoceros This rather small rhinoceros extended over tetraceros DAWKINS with four widèly spaeed most of southern Europe and was close to its tines is better known in the Forest Bed, though northern limit in Norfolk. Most Forest Bed it has been recorded from the Norwich Crag localities of Norfolk and Suffolk have yielded and possibly also the Red Crag. Euctenoceros mandibles of which more than twenty are falconeri DAWKINS is a smaller species found preserved in Norwich Castle Museum. in the Red and Norwich Crags but absent in Equine remains occur throughout the Crags later deposits. The largest of this group, and more commonly in the Forest Bed. Cabal­ Euctenoceros sedgwicki FALCONER has mark­ line horses, Equus robustus PoMEL, are distin­ edly ftattened tines each of which is divided guished by their large size compared with the into three or four points. A fine antler from zebrine horse, Equus stenonis Coccm, which the Forest Bed at Bacton indicates a total span may be absent in the Forest Bed, but the of over nine feet. This species has also been

169 recorded from the Norwich Crag and may be of A/ces gallicus AzzAROLI in the Norwich present in the Red Crag. Crag, by Forest Bed times considerable size The true red deer, Gervus elaphus L., has variation had taken place and two further been doubtfully recorded from the Forest Bed species, Alces latifrons JOHNSON and the very on the basis of a few antlers, some of which large A/ces reynoldsi AzzAROLI, are distin­ are beach specimens. Unfortunately the guished. The antlers have long gently curved diagnostic terminal tines are invariably absent. beams which end in a small palmation edged The commonest deer of Forest Bed times with small tines. The trend in this group is were the giant deer, two species of which can towards a large palmation and a shorter beam be traced back into the Crags. In Praeme­ which is gently twisted along its length. gaceros verticornis DAWKINS the antlers spread Bovids are first represented in this country almost horizontally out on either side of the by Gazella anglica NEWTON from the Red and skull after the first tine which is almost Norwich Crags. The musk ox, Ovibos circular in section and curved abruptly moschatus L., may be present in the Forest forwards and downwards. A number of antiers Bed, @ut most of the ovibovine remains are have a small extra tine just below this first tine, now referred to Praeovibos priscus Staudinger. though often only one antler out of a pair There is no good evidence to suggest that possesses this additional feature. The beam is musk ox was at this time adapted to cokl long and stout, and throws off a second for­ conditions. The characteristic in thie ward facing tine followed by one at the rear, Forest Bed is the large steppe Bison priscus then the beam spreads out upwards into a BOJANUS but a small species, Bison schoeten­ large :fiat palmation which is rarely found sacki FREUDENBERG, probably a woodland intact. A peculiarity of the Megaceros group, animal, has also been found. which is especially marked in M. verticornis, is a strange thickening of the bone of the Order Rodentia lower jaw, and to a lesser extent of the skull. This has been thought of as a mark of Fossi! rodents have a potentially high value degeneration or a pathological character; in stratigraphical studies because they are perhaps the extra bone may have acted as a found in varied habitats and exhibited rapid source of calcium required when new antlers evolution. About twenty species are known were growing. savini DAWKINS from Forest Bed deposits, perhaps only five has widely spread antlers in which the beam from the Crags. Systematic collecting of small is flattened but does not terminate in a large mammal remains in this area would tell us palmate structure as in other giant deer. The much of the details of past environments. first tine is short, almost fan-shaped, and is A squirre!, Sciurus whitei HINTON, probably very close to the origin of the beam. There is ancestral to the modern red squirre!, has been a second rather small tine and a third back­ found in the West Runton Cromerian deposts. ward-facing tine after which the beam curves Two types of beaver occur in the Crags and upwards to a terminal fork. Megaceros Forest Bed; a large form, Trogontherium dwakinsi NEWTON is common in the Forest cuvieri FISCHER, and the then Jess common Bed but there are no Crag records. The antlers Castor fiber L., to which is attributed a mass are backward-pointing and spread rapidly into of gnawed branches in the Forest Bed near a broad fan with six or seven fiat tines. Bacton, Norfolk, part of a fossil dam. A An early fallow deer. nestii MAJOR, porcupine, Hystrix sp., has been recognised has been collected from both Crags but there from the Red Crag, and a second unusual are doubts about its presence in the Forest rodent, a large hamster, Cricetus cricetus Bed as Dama clactoniana FALCONER may also runtonensis H1NTON is known from West be present. The roe deer, Capreolus capreolus Run ton. L., first occurs in Forest Bed deposits. Amongst the voles six species of the Elk first reached this country in the form Mimomys group have been described from the

170 Lisr of of the Crag and Forest Bed

R.C. N.C. 1 W.C. & C.F.B.S.

SPECIES Insectivora Sorex savini Hinton. X Sorex runtonen!iis Hinton. X Neomys newtoni Hinton. X Talpa europaea L. X Desmana moschata Pallas. X

Primates Macaca sp. X

Carnivora Hyaena perrieri CROIZET & JOBERT. X X Hyaena brevirostris AYMARD. X Crocuta crocuta ERXLEBEN. X Homotherium sainzelli AYMARD. X Homotherium latidens ÜWEN. X X Pelis cf. !unensis MARTELLI. X Pelis pardoides ÜWEN. X X Pelis leo L. X Gulo schlosseri KoRMOS. X Martes velus KRETZOI. X Enhydrictis ardea BRAVARD. X Pannonictfa pliocaenica KoRMOS. X Mustela cf. putorius L. X Mustela praenivalis KORMOS. X Aonyx reevei NEWTON (Pomel) X Canis lupus L. ? X Parai/urus anglicus BoYD DAWKINS. X REICHENAU. X Ursus savini ANDREWS. X Ursus spelaeus ROSENMULLER & HEINROTH. ? Ursus cf. arvernensis CROIZET & JOBERT. X Phoca sp. X X X Trichechus hux!eyi LANKESTER X X X Proboscidea Anancus arvernensis CROIZET & JOBERT. X X Archidiskodon meridionalis NEsn. X X X Palaeoloxodon antiquus FALCONER & CAUTLEY. X Mammuthus trogontherii PoHLIG. X

Perissodactyla Tapirus arvernensis CROIZET & JOBERT. X Dicerorhinus etruscus F ALCONER. X Rhinoceros sp. X X Equu; stenonis Coccm. X Equus robustus POMEL. X X X Hipparion cf. gracile KAuP. X

Artiodactyla Sus strozzii MENEGHINI. X

171 Sus scrofa L. X Hippopotamus amphibius L. X Euctenoceros ctenoides NESTI. ? X E. tetraceros DAWKINS. ? X X E. Jalconeri DAWKINS. X X E. sedgwicki F ALCONER. X X X Cervus cf. elaphus L. X Praemegaceros verticornis DAWKINS. ? X X Megaloceros savini DAWKINS. ? X X Megaceros dawkinsi NEWTON. X Dama nestii MAJOR. X X X Dama clactoniana F ALCONER. ? Capreolus capreolu~ L. X A/ces gallicus AzzAROLI. X X A/ces latifrons JOHNSON. X Alces reynoldsi AzzAROLI. X Gazella anglica NEWTON. X X Praeovibos priscus STAUDINGER. X Ovibos moschatus L. ? Ovis savini NEWTON. X Bison prfacus BoJANUS. X Bison schoetensacki FREUDENBERG. X

Rodentia Sciurus whitei HINTON. X Castor /iber L. ? X X Trogontherium cuvieri FISCHER. X X X Hystrix sp. X Cricetus c. runtonensis HINTON. X Clethrionomys cf. g[areolus ScHREBER. X Mimomys newtoni MAJOR. X X Mimomys reidi HINTON. X M. savini HINTON. X M. intermedius NEWTON. X M. pliocaenicus MAJOR. X X M. majori HINTON. X Arvicola bactonensis HINTON. X A. greenii HINTON. X Pitymys gregaluides HINTON. X P. arvaloides HINTON. X Microtus arvalinus HINTON. X Microtus nivaloides FoRSYTH MAJOR. X M. nivalinus HINTON. X M. ratticepoides HINTON. X Apodemus cf. sylvaticus L. X Lepus sp. ? ?

Cetacea include Balaena spp. X X X Megaptera sp. X Balaenoptera spp. X X Monodon monoceros L. ? X Delphinus sp. X X Phocaena sp. X

172 Forest Bed, two of which also occur in the Order Cetacea Norwich Crag. During the early Middle Pleistocene the Arvicola group emerged with Many species of cetaceans are known from species of Microtus and Pitymys, characterised the nodule bed of the Red Crag but most are by their rootless high-crowned cheek teeth heavily mineralised, rolled and polished, the which grow throughout life. signs of derived fossils. Cetacean finds thought The common field mouse, Apodemus sylva­ to be contemporaneous with the Red Crag are ticus L., is known from the Forest Bed at not as common as in the Norwich Crag where West Runton where a few cheek teeth very lightly mineralised bones have been referred similar to the modern species have been found. to species of dolphin, fin and toothed whales. In addition to these the Forest Bed has yielded porpoise and narwhal.

BIBLIOGRAPHY

AzzAROLI, A. (1953) The Deer of the Weybourne East Anglian Crags. Trans. Su.If. nat. Soc. 12, Crag and Forest Bed of Norfolk. ( Brit. Mus.) 262-266. CARRECK, J.N. (1967) Microtine remains from the SPENCER, H.E.P. (1963b) The Contemporary Norwich Crag (Lower Pleistocene) of Eastern Mammals of the Crags. Trans. Su.If. nat. Soc. Bavents, Suffolk. P.G.A. (77) 491-496. 12, 333-334. H!NTON, M.A.C. (1926) British Voles and Lem­ SPENCER, H.E.P. (1966) New Mammalian fossils mings ( Brit. Mus.) from the Red Crag. Trans. Su.If. nat. Soc. 13, KuRTEN, B. (1968) Pleistocene Mammals of 154-156. Europe. Weidenfeld & Nicolson, London. SPENCER, H.E.P. (1970) A contribution to the McWILLIAMS, B. (1967) Fossil Vertebrates of the Geological History of Suffolk, Part 4. Trans. Cromer Forest Bed in Norwich Castle Museum. Su.If. nat. Soc. 15, 148-196. ( Norw. Mus.) W&<;T, R.G. (1968) Pleistocene Geology and Bio­ SPENCER, H.E.P. (1963a) Prehistoric Deer of the logy. Longmans, London.

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