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Quaternary International 169–170 (2007) 125–136

The ecology of the late middle Pleistocene mammoths in Britain

Katharine Scott�

St Cross College, St Giles, University of Oxford, Oxford OX1 3LZ, UK

Available online 12 December 2006

Abstract

In Britain, most large vertebrate assemblages ascribed to MIS 7 include remains of proboscideans—straight-tusked elephant

Palaeoloxodon antiquus, woolly mammoth Mammuthus primigenius and also a smaller mammoth, unique to this interglacial and often

referred to in Britain as the ‘Ilford’ mammoth. Recent studies of the Eurasian mammoths suggest that the Ilford mammoth, hitherto

described as a primitive form of M. primigenius, is in fact a late form of the steppe mammoth M. trogontherii. This taxonomic change has

significant implications for the interpretation of the MIS 7 fossil assemblages particularly as the small mammoth is the most common

species at many sites. The end of the MIS 7 interglacial and the onset of colder conditions associated with MIS 6 mark a crucial period in

the development of the mammoths in Western Europe with the likelihood that incoming populations of M. primigenius existed

contemporaneously with M. trogontherii. Although M. primigenius is commonly listed among the British vertebrate fauna for MIS 7, in

the light of recently available chronological data for the Late Middle Pleistocene mammoths, this author suggests that these

identifications are erroneous and that woolly mammoths reached Britain too late to have been contemporaneous with the other large

vertebrates common to MIS 7. It is concluded that the only mammoth in Britain during this interglacial was the steppe mammoth and

that the reduction in its size was due to a combination of climate change and competition from a range of other large herbivores.

r 2006 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction: the development of the Eurasian mammoths

The Pleistocene evolution of the Eurasian mammoths isrelatively well documented because mammoth remains areabundantly preserved at many localities and also becausethere has been a significant increase in recent years in thenumber of well-dated specimens. Among the adaptivemorphological changes evident during this period, the mostdiscernible are those affecting the skull and dentition.Three main trends are observed in the molar teeth overtime (Lister and Sher, 2001; Lister et al., 2005). These areparticularly well-expressed in the third (last) molar M3 inwhich (i) the crown doubles in height, (ii) the widely spacedthick plates (lamellae) become thinner until the totalnumber in M3 is doubled and (iii) the thickness of theenamel of each lamella is considerably reduced. This trendappears to correspond to a shift in the distribution ofMammuthus during the Pleistocene from warmer, forestedhabitats to cold, open regions (Lister and Sher, 2001). ThusMammuthus rumanus, appearing in Europe some 3.5

million years ago, is defined by low-crowned, thick-enamelled teeth well suited to browsing deciduous wood-land. The dentally more advanced mammoth Mammuthus

meridionalis, also adapted to a diet of trees and shrubs,eventually replaced M. rumanus in the interval 2.6–2.0Ma.Further advances in dental morphology saw the emergenceof Mammuthus trogontherii, the steppe mammoth, around1 million years ago in a habitat that included bothgrassland as well as deciduous woodland. Finally, thesteppe mammoth was replaced in Europe around 200 kawhen the cool conditions and open habitat of MIS 6evidently favoured the woolly mammoth Mammuthus

primigenius. By the Last Glacial the increased ‘tooth-life’of M. primigenius achieved through increased lamellarfrequency and hypsodonty reached its maximum expres-sion, no doubt reflecting the adaptation of this animal tothe coarse vegetation of the ‘steppe–tundra’ biome.Although the morphological changes that distinguish theEurasian mammoths were broadly incremental, theirevolution appears to have been mosaic rather than linearwith differential development taking place in partiallyisolated populations. In Europe, the absence of a transi-tional form linking M. trogontherii and M. primigenius

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doi:10.1016/j.quaint.2006.10.009

�Corresponding author. Tel.: +441865 880362.

E-mail address: [email protected].

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suggests that local populations of mammoths mademorphological adaptations at different rates in responseto varying environmental conditions. Siberian develop-ments appear to have been consistently ahead of those inthe west and as early as c.800–600 ka, the Siberian fossilsequence includes mammoths that are indistinguishablefrom the later European M. primigenius. Not untilconsiderably later (perhaps late in MIS 7 or early in MIS6) does M. primigenius appear in Western Europe whereM. trogontherii was still extant, thus raising the possibilitythat both forms may have contributed to later populations(Lister et al., 2005).

Situated at the northwestern edge of the continent,Britain was thus likely to have been among the lastEuropean regions to see the arrival of M. primigenius in theLate Middle Pleistocene. The effect of this marginallocation is important to the interpretation of the BritishQuaternary vertebrate material in a wider sense as itaccounts for the somewhat fragmentary nature of theBritish fossil record. In the Middle and Late Pleistocene,during the period of development taking place further eastin M. trogontherii and M. primigenius, climatic eventsserved to make Britain periodically uninhabitable (duringmajor ice advances), inaccessible (during periods of highsea-level), or unsuitable for mammoths (during phases ofdense forestation, such as in the Last Interglacial MIS 5e).Thus evolutionary developments in the mammoths thatevidently originated in Siberia and spread westwards bymeans of a complex series of migrations might eventually,but not invariably, have reached Britain. This, of course,applied to all animals, whose appearance in Britain duringthe Pleistocene was likely to have been sporadic, deter-mined not only by the availability of suitable herbaceousvegetation but by ease of access during periods of loweredsea-level. Considering also the destructive effect on thelandscape of fluvial activity during deglaciation, it is notsurprising that deeply stratified sites are uncommon andthat the correlation of isolated assemblages is oftendifficult. Problems of interpretation are further com-pounded by the fact that the majority of British sites wereexcavated or fossils collected from them in the 19th centurywith little concern for stratigraphy, documentation orconservation.

Despite all these potential geographical and historicallimitations, a substantial body of fossil material survivesthat is ascribed to the Late Middle Pleistocene and, morespecifically, to MIS 7. This, as outlined above, is also acrucial period in the development of the mammothstowards the end of which M. primigenius appears inWestern Europe and ultimately replaces M. trogontherii.Since the recognition of the MIS 7 interglacial is relativelyrecent, the identification of the British proboscideans ofthis period has been somewhat equivocal. At most sites,authors have identified one or more of three species: thestraight-tusked elephant Palaeoloxodon antiquus, the typi-cal woolly mammoth M. primigenius and a smaller form ofM. primigenius, also known as the ‘Ilford’ mammoth (after

the site of its discovery). The recent proposal by Lister andSher (2001) that the Ilford mammoth might, on the basis ofits dental morphology, more accurately be referred to as alate form of M. trogontherii rather than M. primigenius

led this author to review the large vertebrate record as awhole and to reconsider the identity of the proboscideansin MIS 7.

2. Elephants and mammoths in Britain during the stage 7

interglacial

Interest in this interglacial has increased significantly inrecent years. Not so very long ago, the existence of theproposed ‘new’ interglacial around 200 000 years ago wasstill a matter of debate (Sutcliffe, 1975; Briggs et al., 1985;Bowen et al., 1989). Today it is widely accepted and,largely due to Schreve’s (1997) study of later MiddlePleistocene vertebrates, some of Britain’s most abundantfossil assemblages are ascribed to this stage (Fig. 1). Thelarge vertebrates of MIS 7 are generally indicative of amixed grassland/deciduous woodland habitat and anabundance of other biological data from recent excavationssuch as at Stanton Harcourt (Buckingham et al., 1996),Aveley (Schreve 1997), and Marsworth (Murton et al.,2001), leaves no doubt as to the fully interglacial nature ofthe climate.It is interesting to compare the large vertebrates recorded

for MIS 7 with those in the climatic stages before and after(Fig. 2). For the purpose of this paper the proboscideansrecorded in Britain during the later Middle and LatePleistocene are listed separately from the large ungulatesmost commonly associated with them. Contemporaneouswith the evolution of the mammoths in northern and

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Fig. 1. British sites with large vertebrate assemblages assigned to MIS 7.

All sites except Pontnewydd Cave and Tornewton Cave have mammoth.

Map redrawn from Schreve (1997) with the addition of Latton (Scott and

Buckingham (2001).

K. Scott / Quaternary International 169–170 (2007) 125–136126

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pyeastern Eurasia during the Middle and Late Pleistocene,the warm climate and deciduous forests of the interglacialsprovided ideal conditions for the migration into Britain ofthe otherwise more southerly-based straight-tusked ele-phant P. antiquus. Straight-tusked elephant was evidentlythe only proboscidean in interglacials MIS 9 and MIS 5e.In MIS 9 it is associated with typical woodland species suchas fallow deer, bear and rhino (Stephanorhinus kirchber-

gensis) although areas of open grassland are indicated bythe presence of horse at some sites, e.g. at Pershore,Purfleet and Grays Thurrock, with the last mentioned alsohaving rhino Stephanorhinus hemitoechus (Schreve 1997).MIS 5e (the Last Interglacial) has a similar large vertebratefauna, although horse is notably absent, perhaps indicativeof heavier forestation than during MIS 9, and hippoappears for the first time in Britain since the early MiddlePleistocene (Stuart 1982).

In view of their association with deciduous woodlandelements, one might thus expect straight-tusked elephantsand steppe mammoths to have been in Britain during theinterglacial periods and woolly mammoths in the cooler,open grassland conditions of the cold stages. Although nolarge vertebrate assemblages are reliably attributed to MIS8, M. primigenius indeed appears to have been the only

proboscidean in the cold stages from MIS 6 onwards, beingespecially abundant among fossils of Last Glacial age.Although woolly mammoth is not certainly in Britain byMIS 4, by MIS 3 (c. 60–30 ka) a large number of sitesdenote a strong association between woolly mammoth,woolly rhino, horse and reindeer. Rather than being thearctic climate envisaged in earlier literature, this period wasevidently relatively mild (van Andel, 2003) with extensiveherbaceous vegetation supporting herds of large ungulatesand dependent large predators—lion, wolf and spottedhyaena.It is possibly the result of having more data for MIS 7

than for the other two interglacials that it appears also tohave been more complex, but it certainly differs from theother interglacial stages. On the basis of her study of theBritish material, Schreve (1997) proposed an earlier,forested phase of MIS 7 characterised by straight-tuskedelephant, fallow deer and other woodland species (butwithout mammoth), and a later phase characterised by amore open habitat with deciduous woodland and straight-tusked elephant, the typical woolly mammoth M. primi-

genius and the smaller ‘Ilford’ mammoth. It has yet to beestablished how these two ecosystems relate to thenumerous warm episodes identified in the marine isotope

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Fig. 2. Large vertebrates commonly associated with mammoths and elephants in the Middle and Late Pleistocene at British sites. The diagram depicting

‘sawtooth’ climate change derived from ice core and oxygen isotope data was modified from the University of London (Royal Holloway) website. The

faunal data for marine isotope stages 9 through 5e in the first and third columns is summarised from Schreve’s (1997) review of the British Middle

Pleistocene large vertebrates. Other studies complete the list for the Last Glacial (Stuart, 1982; Scott, 1986; Currant and Jacobi, 1997).

K. Scott / Quaternary International 169–170 (2007) 125–136 127

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record for MIS 7 (e.g. Reille et al., 2000; Fig. 2) but it isinteresting that the majority of vertebrate assemblages (26out of 30) ascribed by Schreve to this interglacial fall intothe latter category. Recent excavations at Latton (Scottand Buckingham, 2001; Lewis et al., 2006) add yet anotherto this group. As mammoth is absent from Schreve’s earliergroup of MIS 7 sites, the rest of this paper is concernedwith large vertebrates represented in the later category.

The proboscideans are notable for several reasons. Forthe only time in the British fossil record straight-tuskedelephant P. antiquus and mammoth M. primigenius areoften recorded as contemporaneous in these stage 7assemblages. Frequent reference is also made to thepresence of the Ilford mammoth which is unique to thisinterglacial and might be regarded as a stratigraphicmarker for MIS 7 (Schreve, 1997). There has long beensome uncertainty about the taxonomy of the Ilfordmammoth and it has been referred to variously as ‘a smallwoolly mammoth’, ‘Mammuthus primigenius (Ilford type)’,‘a primitive’ or ‘early form’ of M. primigenius, a ‘smallmammoth intermediate between M. trogontherii andM. primigenius’ and ‘a mammoth of the Mammuthus

trogontherii–primigenius lineage’. Recently, Labe andGuerin (2005) have resurrected the name Mammuthus

intermedius to represent a late Middle Pleistocene Eur-opean mammoth intermediate in form between M.

trogontherii and M. primigenius. The fossils in questionmight, however, be alternatively interpreted as a mixture ofsmall, late M. trogontherii and early M. primigenius. Thispossibility and the question of how they compare with theBritish material will be addressed elsewhere (Lister andScott, in prep.). It is important to note that it was notsimply size that differentiated the Ilford mammoth fromthe typical woolly mammoth (Fig. 3) but its dental

morphology. Thus this author concurs with Lister andSher (2001) who proposed that the combination of smalltooth size, thicker enamel, and relatively low lamellarfrequency in the Ilford mammoth (about 19–21 in M3 asopposed to 24 or more in the Late Pleistocene woollymammoth) warrants its consideration as ‘a late form ofM. trogontherii’ rather than an ‘early M. primigenius’ or aform intermediate between the two.Changing the identity of the Ilford mammoth from M.

primigenius to M. trogontherii has significant implicationsfor the interpretation of the British fossil material. For thefirst time, it acknowledges the presence of the steppemammoth in Britain during MIS 7 thereby making sense ofthe rest of the biological data at the many sites where itoccurs. While it was considered to be a ‘primitive’ form ofwoolly mammoth M. primigenius, it was always somethingof an enigma that a species usually associated with evidencefor cool climate and open habitat was consistentlyassociated in Britain with evidence for a warm, partlywooded environment. At Ilford itself, more than a hundredyears ago, it was noted that a small form of mammothoccurred with warm-climate animals (Adams, 1877–1881)and at Aveley, the fact that remains of ‘Ilford type’mammoth directly overlay remains of elephant P. antiquus

was interpreted as evidence of a forested landscape rapidlyreplaced by cold climate and open grassland (Sutcliffe,1995). When interglacial deposits at Stanton Harcourt werefirst reported (Briggs et al., 1985), the anomaly was thepresence of mammoth. Extensive excavations at StantonHarcourt between 1990 and 1999 not only confirmed thepresence of mammoth remains in a fully interglacialcontext but yielded the biggest assemblage of Ilfordmammoth in Britain (Buckingham et al., 1996; Scott,2001). In preparing the Stanton Harcourt material forpublication, this author considered that a possible ex-planation for the occurrence of woolly mammoths amongstotherwise interglacial elements at Stanton Harcourt andother MIS 7 sites might be climatic: that the onset ofinterglacial conditions and rising sea-levels sometimeduring MIS 7 had prevented herds of woolly mammothsalready in Britain from dispersing north-eastwards intomainland Europe and that their reduction in size was aconsequence of being stranded in an unsuitable environ-ment. At various times in the Pleistocene, elephants andmammoths have undergone (sometimes extraordinary) sizereduction in island situations viz. Columbian mammothson the California Channel Islands (Agenbroad and Nelson2002), elephants on Mediterranean islands (Palombo 2001)and mammoths on Wrangel Island, north of Siberia(Tikhonov and Vartanyan (2000). In recognising theBritish material as M. trogontherii, the possibility that sizereduction was a response of the woolly mammoths to anunsuitable habitat is no longer relevant as the associatedpalaeoenvironmental data is entirely consistent with theexpected environment for steppe mammoths. It thenremains to question whether size reduction in the steppemammoth was an effect of insularity. There are two

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Fig. 3. Above: lower right M3 of mammoth M. trogontherii (lingual view)

from Stanton Harcourt c. 200 ka. Below: lower left M3 of mammoth M.

primigenius (buccal view) from Sutton Courtenay (Last Glacial).


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reasons to argue that it was not. Firstly, sea-level dataindicate that Britain might only temporarily have beenisolated from the Continental mainland during MIS 7,perhaps during the warmest phases of the interglacial(Gibbard, 1995; Keen, 1995). Secondly, the dating evidencesuggests that size reduction in M. trogontherii was a processthat began on the Continental mainland before MIS 7. InEurope, the Early Middle Pleistocene M. trogontherii wasextremely large and became successively smaller betweenapproximately 600 and 200 ka (Lister et al., 2005) with theBritish MIS 7 specimens being the smallest apart fromthose at Ehringsdorf, Germany (c. 200 ka). The Europeandates, combined with the fact that there are no MIS 7 fossilassemblages in Britain with large or intermediary forms ofM. trogontherii, indicate that the small mammoths spreadinto Britain from populations already established furtherwest. The factors that brought about size reduction in theLate Middle Pleistocene M. trogontherii are probablycomplex but, in general, dwarfing is seen as a response todiminishing resources (e.g. Guthrie, 1984; Haynes, 1991;Roth, 2001). Insularity was clearly not a factor in Europe,but perhaps fluctuating climatic and vegetational changesgradually brought about the diminishment of the preferredrange of the steppe mammoth.

Despite the implications of size reduction in the steppemammoth, its abundant representation in fossil assem-blages indicates an adequate environment during at leastthe latter part of MIS. It was evidently also a suitablehabitat for the straight-tusked elephant. It remains todiscuss the possible presence of the woolly mammoth, alsorecorded at some sites. Apart from the fact that theenvironmental data suggest an unlikely habitat for thewoolly mammoth for most of MIS 7, the chronologicaldata on the Eurasian mammoths (Lister et al., 2005)indicate that M. primigenius did not reach Europe untilafter 200 ka, probably early in MIS 6. Its presence inBritain during MIS 7 would therefore be unexpected. It issuggested here that the misidentification of the Ilfordmammoth as a small form of M. primigenius led to thefurther misinterpretation of the mammoth remains aswoolly mammoth. It is concluded that, with the possibleexception of one site, wherever M. primigenius is recordedfor this interglacial, it is actually the Ilford mammoth—M. trogontherii.

3. Mammoths and other large vertebrates at British MIS 7

sites

This section considers the reasons for concluding thatthe only mammoth in Britain during MIS 7 was the steppemammoth M. trogontherii. It also reviews the representa-tion of other large vertebrates associated with themammoths.

Of the 27 sites attributed to MIS 7 (Fig. 1) onlyPontnewydd Cave and Tornewton Cave (Otter Stratum)lack mammoth. Taphonomic factors are relevant here: theOtter Stratum consists of a small mammal fauna very likely

brought in by a fox or bird of prey whereas thePontnewydd material is a debris flow and it will have beena matter of chance what became incorporated into the cavedeposits. The deposits in Hutton and Bleadon Caves arealso debris flows but in these cases mammoth and elephantremains happen to have come in with the other fauna. Theremaining MIS 7 large vertebrate assemblages are fromfloodplain and fluvial deposits. The species represented ateach site are given Table 1 (omitting species that occur veryrarely, e.g. wild cat, leopard and various mustelids) andthese form the basis of further discussion here. A fewgeneral observations may be made: that mammoth ispresent at every site; that half also have straight-tuskedelephant; that horse is almost as commonly represented asmammoth; and that those with forest or woodlandpreferences such as straight-tusked elephant and red deerfeature fairly consistently. The overall picture is of a mixedgrassland and deciduous woodland habitat in a temperateclimate.More informative than species representation, however,

is the relative abundance of the various vertebratesrepresented (Table 2). The proboscideans are among themost common. The straight-tusked elephant occurs atapproximately half the sites but mammoth is presentthroughout. It is interesting to consider the representationof the mammoths which, for the purposes of this paper aredivided into three categories. The first two denote sites atwhich authors specify the presence either of ‘woollymammoth’ or of ‘Ilford mammoth’ on the basis ofdentition. In the third category (mammoth unidentified tospecies) are those remains where the absence of teeth madethat distinction impossible. In the literature, these mam-moth remains are assigned to M. primigenius. It is clearfrom Table 2 that, at most sites where there was dentition,the mammoths were recognised as of ‘Ilford’ type (i.e. M.

trogontherii). At only two sites are there references to thepresence of teeth of the ‘evolved’ mammoth (i.e. M.

primigenius). The provenance of much of the Crayfordmaterial is highly dubious and the assemblage includesspecies of distinctly ‘cold’ habitat preference (Currant,1986). It would not therefore be possible to statecategorically that woolly mammoth was truly contempora-neous with the rest of the fauna believed to be of MIS 7age. As regards the second site, the lower channel atMarsworth, M. primigenius dentition was reportedly inreliable context associated with MIS 7 material (DavidParish, pers. comm.; Murton et al., 2001) and maythus denote the arrival of woolly mammoth in Britain,perhaps towards the end of the interglacial. Apart fromMarsworth, the absence of conclusive evidence for M.

primigenius strongly suggests that the ‘unidentified mam-moth’ at the remaining sites is also the steppe mammothM. trogontherii. The author plans a comparative studyof the mammoth post-cranial remains from the MIS 7sites that might clarify this matter and a quantitativestudy of the mammoth dentition is underway (Lister andScott, in prep.).

ARTICLE IN PRESSK. Scott / Quaternary International 169–170 (2007) 125–136 129

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Table

1

RepresentationoflargevertebratesatBritish

MIS

7sites

Aveley

upper

channel

Biels-

beck

Bleadon

Cave

Brun-

don

Cray-

ford

brick-

earths

Great

Yeldham

Hark-

stead

Hindlow

Cave

Hutton

Cave

Ilford

Ittering-

ham

Latton

Lexden

Mars-

worth

lower

channel

North-

fleet

A

North-

fleet

B

North-

fleet

F

Oreston

upper

Cave

Selsey/W

.

Wittering

Sible

Hedding-

ham

Stanton

Harcourt

Stoke

bone

bed

Stoke

Goldington

upper

channel

Stutton

Upper

Strensham

West

Thurrock

Her

biv

ore

s

Straight-tusked

elephant’

’’

’’

’’

’’

’’

?’’

’

Pa

laeo

lox

odo

na

nti

qu

us

Woollymammoth

’’

’’

’’

’’

’’

’’

’’

’’

’’

’’

’’

?’’

’’

Ma

mm

uth

us

pri

mig

eniu

s

Woollyrhino

’’

’’

Co

elo

do

nta

an

tiq

uit

ati

s

Rhino(M

erck’s)

’’

’

S.

kir

chb

erg

ensi

s

Rhino(narrow-nosed)

’’

’’

’’

’’

’’

’’

’

S.

hem

ito

ech

us

Horse

’’

’’

’’

’’

’’

’’

’’

’’

’’

’’

’

Eq

uu

sfe

rus

Wildboar

’’

’’

’

Su

ssc

rofa

Giantdeer

’’

’’

’

Meg

alo

cero

sg

iga

nte

us

Red

dee

r’

’’

’’

’’

’’

’’

’’

’’

’’

Cer

vus

ela

ph

us

Roedeer

’’

’’

’

Ca

pre

olu

sca

pre

olu

s

Bison

’’

’’

’’

’?

?&?&

?&’

’’

’

Bis

on

pri

scu

s

Aurochs

’’

’’

’’

’’

’’

?&?&

?&’

’’

’’

’

Bo

sp

rim

igen

ius

Ca

rniv

ore

s

Spotted

hyaena

’’

’’

Cro

cuta

cro

cuta

Bear

’’

’’

’’

’’

’’

Urs

us

arc

tos

Wo

lf’

’’

’’

’’

’’

’

Ca

nis

lup

us

Lion

Pa

nth

era

leo

’’

’’

’’

’’

’’

’’

Data

forStantonHarcourt

andLatton:K.Scott;forNorthfleet:Wenban-Smith(1995);forallother

sites:Schreve(1997).

?’Indeterminate

proboscideanpost-cranial.

?&Distinctionnotmadebetweenbosandbison.

?Identified

ascf.

bis

on

pri

scu

s.


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Table

2

RepresentationoflargevertebratesatBritish

MIS

7sites(numbersofidentified

specim

ens)

Aveley

upper

channel

Biels-

beck

Bleadon

Cave

Brundon

Cray-

ford-

brick-

earths

Great

Yeldham

Hark-

stead

Hindlow

Cave

Hutton

Cave

Ilford

Ittering-

ham

Latton

Lexden

Mars-

worth

lower

channel

North-

fleet

A

North-

fleet

B

North-

fleet

F

Oreston

upper

Cave

Selsey/W

.

Wittering

Sible

Hedding-

ham

Stanton

Harcourt

Stoke

bone

bed

Stoke

Golding-

ton

Stutton

Upper

Strensham

West

Thurrock

Her

biv

ore

s

Straight-tusked

elephant

15

75

11

641

12

33

42

44

1

P.

an

tiq

uu

s

Woollymammoth

++

+

M.

pri

mig

eniu

s

Ilford

mammoth

136

259*

146*

8441

170

18

193

18

10

515

6737

77

26

24

Mammoth

3+

19

125

12

++

(un

iden

tifi

edto

spec

ies)

Woollyrhino

66

34

21

C.

an

tiq

uit

ati

s

Rhino(M

erck’s)

523

4

S.

kir

chb

erg

ensi

s

Rhino(narrow-nosed)

13

614

41

161

18

13

141

16

S.

hem

ito

ech

us

Horse

43

11

124

301

14

57

462

75

26

293

25

14

290

25

31

160

81

Eq

uu

sfe

rus

Wildboar

311

217

12

Su

ssc

rofa

Giantdeer

73

25

19

M.

gig

an

teu

s

Red

deer

23

532

22

42

82

106

134

139

24

76

19

2

C.

ela

ph

us

Roedeer

1101

32

1

C.

cap

reo

lus

Bison

43

34

534

142

2?3

?1?3

111

2+

1

Bis

on

pri

scu

s

Aurochs

19

10

16

17

72

718

3548

49

?3?1

?320

35

120

116

Bo

sp

rim

igen

ius

Ca

rniv

ore

s

Spotted

hyaena

12

93

G1

Cro

cuta

cro

cuta

Bear

22

212

320

31

18

911

1

Urs

us

arc

tos

Wolf

35

118

1188

344

51

17

Ca

nis

lup

us

Lion

380

336

88

416

37

21

24

319

5

Pa

nth

era

leo

Data

forStantonHarcourt

andLatton:K.Scott;forNorthfleet:Wenban-Smith(1995);forallother

sites:Schreve(1997).

+Recorded

aspresent,nonumbersgiven.

*Severalteethof

M.

pri

mig

eniu

s(andpresumably

somepost-crania)included

inthistotal.

?Distinctionnotmadebetweenbosandbison.

GGnawed

mammoth

bone.

Poorlyprovenancedvertebrates.


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The next group to consider is the perissodactyls (rhinosand horse). There is evidently a strong correlation betweenhorse and mammoth at all the sites corroborating otherbiological evidence for extensive grassland during thisinterglacial. The representation of the rhinos is somewhatmore ambiguous. The woolly rhino Coelodonta antiquitatis

is conventionally a ‘cold stage’, open landscape indicator.Although its presence at 5 sites could be interpreted asindicative of the onset of the ensuing cold stage (Stuart1976), the comparative rarity of woolly rhino (except atCrayford) and the poor stratigraphic resolution at the siteswhere they are recorded (especially in the case of Crayford)argue against including this species as a contemporarymember of the fauna. As regards the interglacial rhinos, thefrequent occurrence of the narrow-nosed rhino S. hemi-

toechus at many sites further emphasises the openherbaceous nature of the landscape. However, the absenceof Merck’s rhino S. kirchbergensis from all but three sites iscurious given the strong presence of other forest browsers(red deer, roe deer, aurochs and elephant) in theseassemblages. It is also interesting that the rhinos occuronly at sites in the south and east of Britain and are absentfrom the central region. This is unlikely to be a samplingbias especially in the cases of Marsworth, StantonHarcourt and Latton which are all large faunal assem-blages, and must therefore be related either to differencesin regional habitat or to differences in the time ofdeposition of these accumulations.

The occurrence of wild boar and the various cervids(giant, red and roe deer) appropriately denote woodland toa greater or lesser extent in the vicinity of some sites. Lessclear is the relative representation of bison and aurochs. Inthe European Pleistocene Bos primigenius is restricted tothe interglacials and is generally associated with forestdwelling species. Bison priscus, however, is known fromboth cold and interglacial stages, associated with herbac-eous vegetation but also with evidence for open forestconditions. For example, it is common in MIS 3 inassociation with M. primigenius, C. antiquitatis and othergrazers at Sutton Courtenay (Scott and Eeles, in prep.), atTuckwell’s Pit (? MIS 5a) in association with R. tarandus ina cool temperate, herbaceous and boreal forest habitat(Buckingham and Scott, in prep.), and at Stanton Harcourtit is the most common species after mammoth in fullyinterglacial conditions characterised by grassland anddeciduous woodland (Table 2; Scott, 2001). The presenceof both aurochs and bison at almost half the sites confirmsthe picture provided by the rest of the fauna—that of amosaic of forest and grassland. Despite the fact that theextant European bison Bison bonasus is an inhabitant ofdeciduous forest, it derives 90% of its forage by grazingclearings of open grassland within the forest (Krasinski,1993). The similar dentition of all three bovids suggests thelikelihood that the Pleistocene bison and aurochs were alsoprimarily grazers, alternating between forest cover andopen terrain. The abundance of other large grazers in MIS7 (such as mammoths and rhinos) is perhaps an indication

that the herbaceous areas were more extensive than inMIS 5e or in the current forest range of Europeanbison. The fact that some of the MIS 7 sites have eitheraurochs or bison may be evidence (as discussed for therhinos) of subtle habitat differences across the region ormay indicate that some of these accumulations representdifferent phases of the interglacial. The latter explanationmay more logically account for the bovid representation atsites whose proximity suggests that local vegetationaldifferences are unlikely to account for species differences.At Marsworth, for example, the large bovid is predomi-nantly aurochs; only 50 k west at Stanton Harcourtaurochs are absent but bison is common and, at Latton,a little further up the River Thames, there are no bovidsat all.A landscape so rich in herbivores would undoubtedly

have attracted a range of predators. At most sites there aresmall numbers of wolf, bear and a very large lion (Table 2).Excluded from this table are the smaller carnivores such asfox and badger and rare occurrences such as wild cat andleopard. Spotted hyaena is recorded at a few MIS 7 sites(Table 2) but, in this review, its presence is regarded aserroneous. The poor provenance of much of the Crayfordcollections has already been noted; the other occurrences(so few as to be insignificant) are from caves, none of whichhas reliable stratigraphic data and all reportedly containingdebris flows that indicate a likely admixture of material ofvarying ages. This would support Turner’s (1995) view thatthe spotted hyaena Crocuta crocuta disappeared fromBritain at the end of the Cromerian (c.450 ka) and did notreappear here until MIS 5e.Although the large herbivores are of broadly interglacial

character, they reflect considerable intra-site variability(Fig. 4). For the purpose of this discussion, the caveassemblages have been omitted (for reasons stated above)as have a few sites with very small samples. The remainingsites reflect abundant herbaceous vegetation with a varyingdegree of woodland nearby, but the non-uniformity of therepresentation of grazing species from site to site isstriking. It might have been anticipated that where thesteppe mammoth is most common there would be fewerforest species and vice versa. In fact, although there is aslight tendency for mammoth numbers to be lower wherethe cervids and elephant are more common, almost anycombination of woodland and open grassland species maybe seen. It seems less likely that these illustratedassemblages represent marked habitat variety across arelatively small area (Fig. 1) than that they are of varyingages within MIS 7. Their context may offer someexplanation in this respect for, while they differ in manyrespects, these assemblages have in common the fact thatthey are from floodplain and fluvial channel deposits.While this could imply poor context, at all the sites wherethe condition of the material is recorded, it is noted that thepreservation of the bones is generally good with relativelylittle abrasion, indicating that they had not been trans-ported any distance. Such ‘almost in situ’ assemblages are

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discussed with reference to a number of Upper Thamessites (Scott and Buckingham, 2001) where it is argued thatburied remnant channels containing synchronously depos-ited biological material can, in some circumstances, bedifferentiated from derived material that has been trans-ported by rivers or glacial outwash. Buckingham (thisvolume) discusses the conditions under which vertebrate

remains, insects, molluscs and vegetation can accumulateand be preserved in fluvial environments. The importanceof these channel assemblages (which were subsequentlyburied by gravel and other material of ensuing cold stages)is that they possibly each preserve a snapshot of theimmediate environment during a relatively brief period ofthe MIS 7 interglacial. The oxygen isotope data indicate

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Fig. 4. The Ilford mammoth (M. trogontherii) shown as a percentage of other large herbivores at British MIS7 sites.


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that there were numerous climatic fluctuations during MIS7 that may have resulted in a number of relatively short-lived ecosystems characterised by the faunal variation seenin the fossil record. It is interesting that there are more ofthese assemblages known from MIS 7 than for any otherinterglacial which perhaps suggests that debris on thelandscape has a better chance of preservation aroundmeandering rivers in open country than in a denselywooded landscape where the course of the river is moreconfined and debris are constantly carried away with thepressure of flow.

4. Conclusion

In reviewing the large vertebrates recorded for MIS 7,the main purpose was to define the landscape in which theIlford mammoth lived. The fossil record illustrates avariety of environments to which the mammoth evidently

adapted even to the point of being the most numerouslyrepresented of all species. It appears to have been mostcommonly associated with other large grazers in apredominantly grassland habitat and somewhat lessnumerous where cervids and elephant indicate denserforestation. Its disappearance from the British fossil recordcoincided with the onset of colder conditions and moreopen terrain associated with MIS 6 and perhaps also withthe arrival of M. primigenius. However, its demise mayhave begun well before this point. Although the palaeoen-vironmental data for MIS 7 suggest ideal conditions for thesteppe mammoth, it was during this period that thesemammoths were smaller than any mammoths in Britainbefore or since. The sea-level data and the presence ofincreasingly smaller forms of M. trogontherii in Europebetween c. 600 and 200 ka (i.e. earlier than in Britain)signify that insularity cannot be invoked to account for sizereduction.

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Fig. 4. (Continued)


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It is likely that a combination of factors resulted in itbeing advantageous to the mammoth to be smaller.Fluctuations in climate may have resulted in less thanoptimum habitats while, simultaneously, a range of largeherbivores were competing with the mammoth forresources. In as far as the mammoth browsed in woodland,its only serious rival would have been the much largerstraight-tusked elephant. However, in more open terrain,where it would have needed to acquire the bulk of itsforage, competition would have been greatest in the formof rhino, aurochs, bison and horse. It may be significantthat aurochs, bison and horse were substantially largerduring MIS 7 than at any time during the Middle and LatePleistocene. Size reduction on the part of the steppemammoth might have been an initial response to competi-tion from such a range of very large grazers. Perhapsequilibrium between all these herbivores was established inthe latter part of MIS 7 but the balance might have been afine one, tipped eventually by the combined effects ofcooler climate, vegetational changes, and the appearance ofanother even larger herbivore in the form of the woollymammoth.

Acknowledgements

I am indebted to Danielle Schreve whose study of theMIS 7 vertebrates made this review possible. My thanksare due to Dr. Christine Buckingham and Profs. AdrianLister and Tony Stuart for many valuable discussions andfor their helpful comments on an earlier draft of this paper.I also appreciate the input from two anonymous reviewersof the submitted manuscript. A British Academy travelgrant that enabled me to attend the second World ofElephants Congress in South Dakota is gratefully acknowl-edged.

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