Cranium, jrg. 9, no. 2, pag. 77-84, augustus 1992
The Pleistocene Vertebrate Faunas of West Runton, Norfolk,
England
A.J. Stuart
Summary
the Norfolk is the fossil vertebrate the Cromer Forest Bed For- West Runton, on coast, most important single locality of
marine fauna mation. The Lower Pleistocene (Pre-Pastonian and Pastonian stages) ’crags’ have yielded a sparse inclu- ding Mammuthus meridionalis, Mimomys pliocaenicus, and M. blanci.
The fluvial of the West Runton Cromerian have organic deposits Freshwater Bed, type site, produced an exceptionally rich early MiddlePleistocene vertebrate assemblage, comprising: 8 fish taxa; 5 amphibians; 3 reptiles; several birds; and
43 mammals. The mammals include: voles Mimomys savini and and Pliomys episcopalis; extinct beaver Trogontherium cuvieri; monkey Macaca sylvanus, mammoth Mammuthus cf. meridionalis, rhinoceros Dicerorhinus etruscus, and giant
deer Megaloceros verticornis.
Pollen and macrofossils show that this fauna lived in association with forest in the first half of plant regional temperate
the temperate stage.
Samenvatting
West Runton, aan de Norfolk kust in Engeland, is de meest belangrijke vindplaats van vertebratenfossielen afkomstig uit
de Cromer Forest Bed Formatie. In de Vroeg Pleistocene (Pre-Pastonien en Pastonien stages) mariene ’crags’ zijn een
klein aantal M. relatief zoogdierresten van o.a. Mammuthus meridionalis, Mimomys pliocaenicus en blanci verzameld.
de Freshwater hebben De organische, fluviatiele afzettingen van West Runton Bed, type-lokaliteit van het Cromerien,
Midden vertebratenfauna 8 5 een buitengewoon rijke vroeg Pleistocene opgeleverd met soorten vissen, soorten amphi-
biën, 3 soorten reptielen, meerdere vogels en 43 soorten zoogdieren. De zoogdierfauna bevat o.a. woelmuizen Mimomys
en de bever de de mammoet savini Pliomys episcopalis, uitgestorven Trogontherium cuvieri, aap Macaca sylvanus, Mam-
cf. meridionalis, neushoornDicerorhinus muthus de etruscus en het reuzenhertMegaloceros verticornis.
Pollen dat de in de helft en plantaardige macroresten tonen aan faunaheeft geleefd een bosrijke omgeving tijdens eerste
van de gematigd warme periode.
Introductie and Pastonian of Lower Pleistocene age, and the Bees-
tonian and Cromerian dating from the early Middle The Cromer Forest Bed Formation (C.F.B.F.), exposed Pleistocene. Some deposits assigned to the Beestonian along the coasts of northeast Norfolk and Suffolk, has date from distinct Lower Pleistocene cold may a stage. been famous for its fossil mammal remains since the Bed also Recent work suggests that the Forest covers early part of the last century (GREEN, 1842; OWEN, one or more early MiddlePleistocene temperate stages 1846). The classic nineteenth century study of the stra- which are post- Cromerian, but still pre-Anglian tigraphy and palaeobotany of these complex deposits by (MEIJER & PREECE, 1992; STUART, 1992). Clement Reid (REID, 1882; 1890), was matched by ex- work the vertebrates cellent on by E.T. NEWTON (e.g. The Forest Bed mammal "localities" are shown in Fig. 1.
beach 1882a; 1886; 1891). Forest Bed fossils found on the at Eccles, appear contributions Other major on the vertebrates include to have been washed out of deposits below low tide le- last those made century by OWEN (e.g. 1846) and vel. Stray, probably wave-transported, finds occur on and earlier this the southeast DAWK1NS (e.g. 1883); century by HIN- the beaches even further to at Sea Palling
and should be stressed TON (1908; 1911; 1926), AZZAROLI (1953). More and Winterton. It that many of the
includes Bed recent work MCWILLIAMS (1967); STUART Forest 'localities', as recorded in museum labels fossil (1975; 1982; 1992); STUART & WEST (1976); and LISTER and catalogues, are not specific sites, but merely indication of where (1992a; 1992b). give a general specimens were
in relation the coastal found to nearest village or town. WEST'S (1980) massive reinterpretation of the stratigra- in fossils found the Moreover, many cases were loose on phy and of the C.F.B.F., utilizing palyno- palaeobotany been washed of beach, having out deposits in the cliffor
- - a technique unknown to Reid provided an inva- logy it is difficult foreshore, so that or impossible to assign luable framework for sorting out the sequence of these units. to particular stratigraphic Exposures were vertebrate faunas. According to West the C.F.B.F. co- better in the generally past, as nowadays most are obs- the vers several climate-based stages, from oldest to the cured by sea defences, but some localities are still avai- Pastonian youngest: Pre-Pastonian (cold); (temperate); lableand important finds continue to be made. Beestonian (cold); and Cromerian (temperate), the de-
posits of which are succeeded by tills and outwash of The most important single locality where Forest Bed hiatus the Anglian Glaciation. A major stratigraphical is vertebrateshave been found in situ is the cliff section at
within the with the Pre-Pastonian the present sequence, West Runton (Figs. 2,3), stratotype for the Cromeri-
77 in situ. Fig. 1: Forest Bed Vertebrate Localities. Open circles indicate localities where Forest Bed deposits do not occur Corton, Pakefield and
Kessingland are in the County of Suffolk; the rest are in Norfolk.
1: Forest Bed. de Forest bed niet in situ voorkomen. Fig. Vindplaatsen van gewervelden van Open rondjes geven plaatsen aan waar afzettingen
Corton, Pakefield en Kessingland liggen in Suffolk, de andere in Norfolk.
Pre-Pastonian and Pastonian few an Temperate Stage, and designated a geological stages (WEST,1980). A
S.S.S.I. (Site of Special Scientific Interest) by English large-mammal remains have been found in the fore-
Natural EnvironmentResearch a molar of the Nature (formerly Coun- shore exposures, including extinct ele- cil). Although threatened on more than one occasion, phant Mammuthus meridionalis from the Stone Bed this Vole material prime palaeontological locality is still allowed to (Pre-Pastonian a substage) (WEST, 1980). erode less and thus remains from more or naturally, available was obtained by sieving crag dug beneath the for study. beach by mechanical excavator during the construction
wall of a concrete sea immediately to the east of West Two of and faunas in groups deposits widely separated Runton Gap (MAYHEW & STUART, 1986). this The Lower Pleis- time are present at locality. older,
which tocene, assemblages need more detailed study and are The vole fauna, comprises Mimomys pliocaeni- considered only briefly here. cus, Mimomys reidi, Mimomys newtoni, Mimomys blanci
and Mimomys pitymyoides, is very different to that of Cold and Pastonian Crag (pre-Pastonian Stage Tempe- Freshwater the West Runton Bed, which occurs in a rate Stage) stratigraphically higher position, but is similar to those
East Runton recorded from (MAYHEW & STUART, Overlying the Upper Chalk, which can be seen exposed A 1986). desman upperP4 from the same deposits has on the foreshore at low tide, are Lower Pleistocene ma- been referred to al. rine Galemys kormosi by HARRISON et deposits; the Stone Bed and crag ("Weybourne The above material is in the Universi- which (1988). preserved Crag") occur both on the foreshore and beneath ty Museumof Zoology, Cambridge. the modern beach. Pollen assemblages obtained from
silt horizons show the interdigitated that the crag covers
78 Fig. 2: Schematic section of cliff and foreshore deposits at West Runton, Norfolk (based on WEST, 1980). Length of section approximately 500 metres; depth 22 metres. The V-shaped structure is an ice-wedge cast.
Sediment Environment Stage
8. tilland outwash sands and gravels, glacial Anglan Cold Stage
occasional ice wedge casts penetrate
the underlying sands and silts permafrost Anglian Cold stage
7. laminated silts and sands (below), marine Cromerian Temperate Stage, sands (above) tidal substage III, IV
6. gravel ("Monkey Gravel") marine Cromerian TemperateStage, substage III palaeoasol subaerial
5. West Runton Freshwater Bed freshwater,fluvial Cromerian TemperateStage, substage Ib-IIb
4. Calcareous silt freshwater,lacustrine Late Beestonian Cold Stage to early Cromerian substage la
3. Gravels and sands marine Beestonian Cold Stage
2. Crag and silts marine Pre-Pastonian Cold Stage and Pastonian Temperate Stage
1. Stone Bed on frost-shattered chalk marine Pre-Pastonian Cold Stage
The Pleistocene deposits rest on Chalk (Upper Cretaceous, Campanian).
2: Schematisch de klif de de kust West Norfolk de door- Fig. profiel van en afzettingen voor van Runton, (gebaseerd op WEST, 1980). Lengte van 500 snede is ongeveer m.; diepte 22 m. De V-vormige struktuur is een vorstwig.
West Ronton Freshwater Bed beds fluvial muds and (Cromerian Temperate c-e, shelly organic sands with nu-
small Stage) merous reworked marl lumps and pebbles (mainly
subzones - flint), with pollen spectra essentially of Cr lb This famous deposit, known since the early nineteenth Cr Ila. The clast size decreases upwards and the upper has of the richest century, produced one Quaternary of the bed muds part deposit, f, comprises detritus vertebrate faunas in Europe. So far it has escaped being (shelly in part), mainly of subzone Cr lib age. The alte- obscured coastal defences and by sea important new red top of the bed, interpreted as a palaeosol, indicates including new species continues to be material, records, substantial of mari- a period subaerial exposure prior to found. basal ne deposition of the overlying gravel ("Monkey Freshwater The West Runton Bed occupying a broad Gravel") and thin bedded silts and sands attributed to channel about 200m in section and about across up to Cromerian the second half of the Interglacial. The top 2m thick, cut in sands and gravels attributed to the of the Freshwater Bed is extensively bored by the mari- Beestonian Cold out at the base of the cliff Stage, crops shells ne mollusc Mya. Occasional can be seen in life east of West Runton Gap (Woman Hythe) (Fig. 2,3). position. and Details of the stratigraphy, pollen plant macro as-
and semblages are given by WEST (1980). The channel filling The Freshwater Bed pollen plant macro assembla- is for the Cromerian record designated the stratotype Stage. ges (WEST, 1980) (Fig. 4) changes from wood-
land with pine, elm, birch, extensive herbaceous vegeta- The basal bed of the channel filling, bed a (of WEST, tion and restricted fen and reedswamp (Cr lb), to a in Goss' 1980), occurs only a limited area near Gap more diverse woodland with oak and other thermophi- It near the eastern end of the exposure. comprises lous trees more persistent local habi- widespread, open freshwater marls with pollen assemblages assigned to tats and fen and reedswamp (Cr Ila, Cr lib). the late Beestonian Cold Stage and subzoneCr la of the
Cromerian (Table 1, Fig. 4). On its eroded surface rest
79 PISCES PISCES (provisional identifications) (STUART,(STUART, 1975; AVES (according to CJ.O. HARRISON, 1979)1979)
1988; STINTON, 1985)1985) Bewick 1. cf. Cygnus bewickii, swan
1.1. Esox luciuslucius L., pikepike 2.2. Anser anser, grey-lag goose
2. Gasterosteus aculeatusaculeatus L., three-spinedthree-spined sticklebackstickleback 3. AnasAnasplatyrhynchos,platyrhynchos, mallard
3. eel 4. Anas Anguilla anguilla (L.), common 4. Anaspenelope, wigeon teal 4. TincaTinea tincatinea (L.), tenchtench 5. Anas crecca, teal
5. Scardinius erythropthalmuserythropthalmus (L.), ruddrudd 6. Netta rufina,rufina, red-crested pochard
6. Rutilusrutilus (L.), roach 7. AythyaAythyaferina,ferina, pochard
7. Perca fluviatilisfluviatilis L., perch 8. Aythya fuligula, tuftedtufted duck mandarin 8. Gymnocephalus cernuacemua (L.), rufferuffe 9. Aix galericulata,galericulata, mandarin
Somateria thick eider 10. Somateriagravipes, leggedlegged (extinct spespe cies, described by Harison) AMPHIBIA (HOLMAN,(holman, CLAYDENclayden & STUART, 1988; 11. Bucephala clangula, goldeneye HOLMAN, 1989) 12. Mergus albellus,albellus, smewsmew
red-breasted 13. Mergus senator,serrator, red-breastedmerganser 1. Triturus vulgaris L., common newt 14.14. Gallinula chloropus, moorhen 2. Bufo bufo (L.), common toad 15.15. Turdus Imerula,?merula, ?blackbird 3. Rana arvalis Nilsson, moor frog 16.16. SturnusStumus ?vulgaris, ?starling?starling 4. Rana ’esculenta’'esculenta' Rana edible ?vulgar\s, or ridibunda, edible frog or 17. Garulus ?glandarius, ?jay?jay marsh frog
5. Rana temporaria L., common frog
MAMMALIA (sources include: AZZAROLI, 19531953; HIN-
TON, 1908; 1926; NEWTON, 1882a; 1909; STUART, 1975; REPTILIA (HOLMAN, CLAYDEN & STUART, 1988; HOL- 1982;1982; 1988; 1992; D L HARRISON & BATES, 1984);1984) W1L-WIL- MAN, 1989) LEMSEN, 1990; LISTER, 1992a)
1. Anguis fragilis L., slow worm Chiroptera 2.2. Natrix natrix (L.), grass snake 1.1. Nyctalus noctula Schreber, noctule bat 3. Vipera berus L., adder
remains and the In review of Small-vertebrate are fairly common, a major European Quaternary otters, WIL-
have been the the West majority found by wet-sieving sediments. LEMSEN (1990) has referred Runton material
remains are much but the the extinct Lutra Large-mammal rarer, over to species simplicidens. Carnivores, are in the years numerous bones and teeth have been collected, clearly underrepresented collections considering
fossils the available especially when the have been exposed after a wide variety of prey. Hopefully, more ma-
combination of gales and high tides has eroded the terial will be discovered in the future.
cliffs. Although known to palaeontologists for more than a
The list of vertebrates from The Freshwater Bed has century and a half the Freshwater Bed continues to sur-
new the been significantly extended in the last few years (STU- prise us with discoveries; most recent of which
ART, 1975; 1981; 1982; 1988b; 1992). J. A. Holman has is the most spectacular ever found at this site and in- studied the reptile and amphibian material and disco- deed in the entire Cromer Forest Bed Formation. Until
vered two taxa not recognized by NEWTON (1882), Rana recently only very sparse material of elephant was avai-
the arvalis, moor frog and Rana ridibunda or R. ’esculenta’, lable from Freshwater Bed. However in December
after and marsh or edible frog (HOLMAN, CLAYDEN & STUART, 1990, a gale high tide, Mr and Mrs H. Hems discovered the of 1988). pelvis an adult male elephant. A year
Recent erosion additions to the mammal list include: Nyctalus later, further exposed several more bones which found Sinclair. noctula, noctule bat (see D. HARRISON & BATES, 1984); were by R. A controlled excavation in
Pliomys episcopalis, extinct vole (first recognized by A. January 1992 recovered the mandible, most of the verte- bral humerus P. Currant, on the basis of material collected by J Clay- column, right and ulna, several ribs and a
a small number of other bones den); hamster, species not yet determined of Mammuthus cf. meridionalis
(tooth collected by J. Clayden) and Megaloceros savini, (Figs. 4,5). More precise identification awaits the remo-
deer J. val of the mandible giant (antler collected by Lightwing and identi- from its plaster jacket. The finds so
A. M. fied by Lister). In addition Macaca sylvanu, ma- far comprising about a quarter of the entire skeleton,
is Freshwater in the Castle caque monkey, now recorded from the are Museum, Norwich. The presence of
Bed on the basis of an incisor tooth collected by P. two hyaena coprolites found in situ with the elephant
the find of from West characteristic toothmarks Bergdahl; previous macaque bones, plus on a phalanx,
Runton from the show was "Monkey Gravel" (HINTON, 1908), that hyaenas fed on the carcass. The evidence for
80 Insectivora Carnivora
2. Erinaceus cf. europaeus L., hedgehog 24.24. Canis lupus L., wolf Sorex extinct shrew 3. Sorex runtonensis Hinton, extinct shrew 25.25. Ursus deningeri von Reichenau, extinct bear
4. Sorexsavini Hinton, extinct shrew 26.26. Mustela nivalis L., weasel
5. extinct water shrew 5. Neomys newtoni Hinton, water shrew 27.27. Martes sp.,sp., marten mole 6. Talpa europaea L., common mole 28.28. PannonictisPannonictis sp., extinct mustelid
7. Talpa minorFreudenberg, extinct mole 29. Lutra simplicidens Thenius, extinct otter
8. Desmana moschata (Palla), Russian desman 30. Crocuta crocuta (Erxleben), spotted hyaena
31. Felis cf.cf. lunensislunensis Martelli, extinct cat Primates undetermined 32. undeterminedlarge felid, probably lion oror sabre- 9. Macaca sylvanus (L.), Barbary Barbary macaque tooth
Lagomorpha Proboscidea 10. Lepus sp., a hare sp., 33. Mammuthus cf. meridionalis(Nesti), extinct ele-
phant.phant. Rodentia
11. SciurusSciunts whitei Hinton, extinct squirrell Perissodactyla 12.12. Trogontherium cuvieri Fischer, exti 34. Equus sp. (caballine), aa horsehorse
35. Dicerorhinus etruscus (Falconer), extinct 'Etruscan' 13. CastorCastorfiberfiber L., beaver rhinoceros 14. Cricetus cricetus hamster (L.), commoncommon
small 15. hamster, species not yet determined Artiodactyla 16. ClethrionornysClethrionomys glareolus Schreber, bank volevole 36. Sus scrofa L., wild boar 17. Pliomys episcopalis Mehely, extinct vole 37. Megaloceros verticornis (Dawkins), a giant deer 18. Mimomys savini Hinton, extinct water vole Mimomys deer 38. Megaloceros savini (Dawkins), a giant 19. Pitymys arvaloides Hinton, extinct pine vole 39. Dama dama (L.), fallow deer 20. Pitymys gregaloides Hinton, extinct pine vole 40. Cervus elaphus L., red deer 21. Microtus arvalisarvalis vole cf. (Pallas), common vole 41. Alces latifrons Johnson, extinct elkelk (moose) 22. Microtus oeconomus (Pallas), northern vole 42. Capreolus capreoluscapreolus (L.), roe deer 23. Apodemus sylvaticus (L.), wood mouse 43.43. Bison schoetensacki Freudenberg, extinctextinct bisonbison
1 1 Species recorded so far only from marine gravelgravel over-over-
Freshwater Bed lying West Runton Freshwater Bed (probably rewor-
ked)
Table 1: Faunal List West Runton Freshwater Bed (West, Runton, Norfolk, England). Cromerian Interglacial (stratotype), substages Cr Ib-IIb
burial of the skeleton much faster evidence rapid implies deposi- Taking the palaeontological together, one can tion of the Freshwater Bed them had rich in and been assumed pre- picture a slow-flowing river, aquatic plants
wide of viously. fringed by fen, supporting a variety fishes, such
small-vertebrate have in lowland Sparse remains, also been found as is found a typical English river today, plus in the marine which "Monkey Gravel", rests directly on frogs and toads, grass snake, waterfowl, and mammals the Freshwater Bed. According to WEST (1980), this ho- of waterside habitats, including Russian desman, bea- rizon to Cromerian Cr III. The extinct beaver otter and belongs substage verte- ver, Trogontherium cuvieri, ex- which include brates, Mimomys savini (see above), are tinct water vole Mimomys savini.
those of the and very similar to Freshwater Bed, are Much of the fauna, is consistent with forest probably derived partly or entirely from it. However, temperate indicated the Sciurus as by pollen spectra (WEST, 1980); one species the squirrel whitei is recorded, on e.g. fallow deer the basis of far wild boar, macaque, and woodmouse. How- a single upper P4, so only from the the abundant remains of "Monkey Gravel". ever, grassland voles Microtus and indicates the of spp. Pitymys spp. presence at least To 43 17 date mammalian taxa, birds, 5 3 local areas ofherb while the amphibians, vegetation, presence of elk
and at least 8 fishes are recorded from the West of reptiles and two species giant deer, in which the males car-
Runton Cromerian. Some of the bird in ried identifications, enormous outspread antlers, together with horse,
the of an extinct of eider indicates more extensive habitats particular diagnosis species (c open than are sugge- be the of HARRISON, 1979) may questioned on grounds sted by the pollen spectra. The activities of large herbi- insufficient fish identifications should material. The be - vores, especially elephants possibly present in substan-
The material is at - regarded as provisional. present un- tial herds would have probably promoted and der by B. maintained within study Irving. more open areas the forest.
81 dark above Fig. 3: West Runton cliffs east of West Runton Gap (Woman Hythe). The West Runton Freshwater Bed can be seen as a band the top
of the beach.
Bed is zien als Fig. 3: West Runton kliffen ten oosten van West Runton Gap (Woman Hythe). Het West Runton Freshwater te een donkere band
boven de top van het strand.
Pinus Carpinus PiceaAbiesAlnus Betula UlmusQuercusTilia Corylus
4: on WEST, - the Fig. Schematic pollen diagram for the Cromerian Temperate Stage (based 1980a). Cr Ia Cr IVc are pollen substages of Crome-
rian. The diagram also includes the end of the preceeding Beestonian Cold Stage and the beginning of the succeeding Anglian Cold Stage.
4: Schematisch van warme Cromerien s.str. WEST, - Fig. pollendiagram het gematigd (gebaseerd op 1980a). Cr Ia Cr IVc zijn pollen stages van
het Cromerien. Het van de koude het Beestonien diagram bevat ook het laatste deel voorafgaande periode, en het begin van het Anglien, de kou-
die de periode volgt op het Cromerien.
82 Fig. 5: Elephant mandible in situ, January 1992. (Courtesy of Eastern Caonties Newspapers).
Fig. 5: Onderkaak van een olifant in situ, januari 1992.
The large amount of material from the West Runton
Freshwater Bed is divided between several museums,
notably: Natural History Museum, London; Norwich
Castle Museum; Harrison Museum, Sevenoaks; and
University Museum of Zoology, Cambridge.
Adress of the author:
A.J. Stuart
Castle Museum
Norwich NR1 3JU
England
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