the evolution of the guild of large carnivora of the british isles during the middle and late...

JOURNAL OF QUATERNARY SCIENCE (2009) 24(8) 991–1005Copyright � 2009 John Wiley & Sons, Ltd.Published online 1 May 2009 in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/jqs.1278
The evolution of the guild of large Carnivoraof the British Isles during the Middle and LatePleistoceneALAN TURNER*

School of Biological and Earth Sciences, Liverpool John Moores University, Liverpool, UK

Turner, A. 2009. The evolution of the guild of large Carnivora of the British Isles during the Middle and Late Pleistocene. J. Quaternary Sci., Vol. 24 pp. 991–1005. ISSN0267-8179.

Received 12 August 2008; Revised 13 February 2009; Accepted 25 February 2009

ABSTRACT: Recent refinements to the dating of a number of Middle and Late Pleistocene localitiesand fossil mammal assemblages from the British Isles, and their correlation with Marine Isotope Stages(MIS), permit a more detailed assessment of the evolution of the guild of larger Carnivora. By around0.4–0.5 Ma (MIS 13–11) or just after, the more archaic elements of the guild, such as the sabretoothedcat Homotherium latidens, the medium-sized pantherine Panthera gombaszoegensis and the large dogCanis lycaonoides, had disappeared and had been replaced, or were being replaced, by extant speciessuch as lion (Panthera leo), spotted hyaena (Crocuta crocuta) and the wolf (Canis lupus), although
claims for a last-glaciation appearance – or reappearance – of Homotherium require assessment.During MIS 5a this guild seems to have been temporarily replaced by a more restricted one of wolf inconjunction with a very large brown bear (Ursus arctos), together with a reduced prey fauna of bisonand reindeer. Final extinction of Crocuta and lion in these islands took place towards the end of MIS 3,arguably prior to the Last Glacial Maximum. Copyright # 2009 John Wiley & Sons, Ltd.
KEYWORDS: large Carnivora; guild evolution; Middle Pleistocene; Late Pleistocene; British Isles.

Introduction

The British Isles have one of the best European sequences ofMiddle–Late Pleistocene assemblages of fossil mammals, withthe potential to inform us about evolution, dispersal patternsand the effects of isolation. The larger Carnivora are particularlywell represented here (Turner, 1995a), and afford the mostcomplete picture of the evolution of the European predatorcommunity from the Middle Pleistocene onwards. They wereamong the most widely distributed of the Pleistocene species,and played an important ecological role as top predators, whileas denning animals and bone collectors many helped provideus with much of the fossil record and thus potential evidence ofpredator–prey interactions (Turner, 1990, 1992, 1995c). Thisperiod also covers the timespan of occupation of these islandsby humans (Parfitt et al., 2005), and they and the largerCarnivora are likely to have had shared interests in access toprey and to carcasses in the landscape (Turner, 1992, 1999c;Anton et al., 2005).

Several assessments of the Carnivora of Africa (Turner, 1990;Turner and Anton, 1999) and Europe (Turner, 1992, 1995a–d)have mentioned Britain, while closer attention has focused onmaterial from specific localities here (Turner, 1999b, 2000), but

* Correspondence to: A. Turner, School of Biological and Earth Sciences, Liver-pool John Moores University, Liverpool L3 3AF, UK.E-mail: [email protected]

despite the importance of the material the guild, in whole or inpart, has not been studied since the 1860s (Dawkins andSanford, 1866–8), beyond evidence for contact with theEuropean mainland (Turner, 1995a). Even Yalden’s (1999)History of the British Mammals gives relatively little attention topre-Holocene matters. Recently improved chronology of anumber of Middle and Late Pleistocene localities (Table 1) andmammalian assemblages (Currant and Jacobi, 2001, 2004;Schreve, 2001a,b; Currant, 2004a,b; Jacobi et al., 2006;Gilmour et al., 2007), together with the possibility of a longersurvival of the sabretoothed cat Homotherium latidens thanpreviously suspected (Reumer et al., 2003; Proctor et al., 2005),suggest it is now time to assess the evolution and movement ofthe European Carnivora using Britain as the focal case study.

Carnivore guilds and the present enquiry

The term ‘guild’ has often been used in the literature inconnection with the large carnivores (Van Valkenburgh, 1985,1988; Turner, 1990, 1995c) as a useful shorthand expression. Itwas originally employed by Root (1967) to describe a groupof species that exploit a resource in a similar way, in the presentcase by catching, killing and eating other mammalian speciesand in particular the smaller and medium-sized ungulates. Themost diverse example occurs today in Africa, where the guild

Table 1 British Middle and Late Pleistocene mammal sequence showing probable age (Ma), MIS, named terrestrial stages and mammal assemblagezones (MAZs), together with an indication of important localities

Ma

0.8–0.6 MIS 19–15 ‘Cromerian Complex’ – localities include West Runton Freshwater Bed, Corton, Bacton, Norton Subcourse,Pakefield–Kessingland

0.5 MIS 13 – (Warm)Boxgrove, Happisburgh, Westbury

0.45 MIS 12 – Anglian (Cold)0.4 MIS 11 – Hoxnian (Warm) – Swanscombe MAZ

Hoxne, Swanscombe, Clacton, Ingress Vale0.34 MIS 9 – Purfleet (Warm) – Purfleet MAZ

Purfleet, Grays Thurrock, Belhus Park, Cudmore Grove, Cauliflower Pit0.22 MIS 7 – Avely (Warm) – Ponds Farm/Sandy Lane MAZs

Aveley, Ilford, Oreston, Crayford and Erith, Bleadon, Hutton, Bielsbeck, Pontnewydd and numerous others0.15 MIS 6 – (Cold)

Minchin Hole, Bacon Hole (lower), Waterhall Farm (lower), Balderton0.125 MIS 5e – Ipswichian (Warm) – Joint Mitnor MAZ

Hoe Grange, Joint Mitnor, Kirkdale, Tornewton Bear Stratum (in part), Waterhall Farm, Barrington, Victoria Cave0.087 MIS 5c – (Temperate) Bacon Hole MAZ

Bacon Hole (upper cave earth), Tornewton Hyaena Stratum0.0.8 MIS 5a – (Temperate with cold-stage mammals) Banwell Bone Cave MAZ

Banwell, Windy Knoll, Steetley Quarry, Pickens Hole layer 5, Wookey Hole Hyaena Den, Brean Down lowermost deposit,and numerous others

0.07 MIS 4 – (Cool) – As yet unnamed MAZLikely to be based on fauna like that in uppermost assemblage from Brean Down

0.05 MIS 3 – Devensian (Cold) – Pin Hole MAZNumerous localities include Coygan, Creswell (Pin Hole, Church Hole, Robin Hood’s Cave), Kent’s Cavern, Wookey HoleHyaena Den upper deposit, Sandford Hill, Uphill

Sources: Currant (1989, 2004a,b; Lister and Brandon (1991); Currant and Jacobi (2001); Schreve (2001a,b, 2004); Stuart and Lister (2001); Lewis et al.(2004); Stuart et al. (2004); Parfitt et al. (2005); Higham et al. (2006); Gilmour et al. (2007).

992 JOURNAL OF QUATERNARY SCIENCE

consists of medium and large-sized cats, hyaenas and dogs,animals that variously chase or ambush their prey, often singlybut sometimes by means of group action, and who may at timesscavenge the remains originally taken either by conspecifics orby other species (Kingdon, 1997; Turner and Anton, 1997,2004; Turner et al., 2008; Wang and Tedford, 2008). Thismodern-day guild, together with those rather more restrictedguilds to be found on other continents, can afford us insightsinto structure and function of past guilds and their constituentfossil species, or at least help us chart our way through thetaphonomic fog of an imperfect fossil record (Dennell et al.,2008). However, it is worth stressing that my intention in usingmodern guild comparators is simply to have a guide to whatvarious animals do and can do. Ecologists find it useful to thinkin terms of niche partitioning and character displacement infeatures such as tooth sizes, with co-evolutionary pressuresover time frequently cited as a cause of such separations (Dayanand Simberloff, 1996; Meiri et al., 2005), but a marked featureof carnivore behaviour is flexibility and adaptability, both inprey selection and capture and in interactions with other guildmembers, (Estes, 1992; Mills and Biggs, 1993; Kingdon, 1997)rather than strict operation within some abstract and compart-mentalised framework. It is also clear that things such as theevolution of large, bone-crushing and carcass-destroyinghyaenas and the development of large sabre-like teeth in themachairodont cats, frequently thought to have exerted co-evolutionary pressure upon each other, actually developedover quite different timescales and owe little to suchinterfamilial influence (Werdelin and Turner, 1996a,b; Salesaet al., 2005; Turner et al., 2008). Thus, instead of seeking toestablish a rigid framework from which past behaviours andinteractions can be read off in any simple manner, I insteadmake reference to characteristics of appropriate modernspecies, or to the guilds within which they operate, at variouspoints as in the text as necessary.

Copyright � 2009 John Wiley & Sons, Ltd.

As for the details of the British guild, although mention ofspecific material and of size changes, taxonomic niceties andthe like is made from time to time, the intention is not to repeatsuch details as can be found elsewhere in the literature. The aimin this paper is instead to provide an overview of the diversity,structure and predator–prey relationships of the British guild oflarger Carnivora over the past 800 ka or so, including changesin proportions of flesh-eaters, carcass destroyers and both groupand solitary predators in the guild, the range of prey and preysizes and the changing patterns of predator–prey interactions.The guild is taken to include the families Hyaenidae, Ursidaeand larger members of the Felidae and Canidae – what couldessentially be considered the top predators. As such, it excludeslynx and smaller felids and also foxes. Discussion begins with asummary of the current consensus on stratigraphic frameworkfor the fossil-bearing localities, an essential basis from which toinvestigate first and last appearances, changing patterns ofassociation of faunal elements and likely predator–preyrelationships. This is followed by a bestiary of the guildmembers, outlining the major characteristics of each family andits member species and the known time ranges based on thestratigraphic framework, and then by an overview of thechanging prey fauna. In the final section changes in thestructure of the guild over time, including the question of finalextinctions, are considered against this background of changesto the physical and biotic environment.

Chronology and stratigraphic framework

In recent years a better understanding of the sequence andchronology of Middle and Late Pleistocene cold and warmstages in the British Isles, and their likely correlations with the

J. Quaternary Sci., Vol. 24(8) 991–1005 (2009)DOI: 10.1002/jqs

BRITISH PLEISTOCENE CARNIVORA 993

Marine Isotope Stages (MIS), has developed. This has beendriven in part by a clearer understanding of the terrestrialstratigraphic framework (Bowen, 1999), but has also in largepart been led, and strongly supported, by detailed studies of thebiostratigraphic evidence from fossil mammal assemblages(Currant, 1989, 2004a,b; Currant and Jacobi, 2001; Schreve,2001a,b, 2004; Stuart and Lister, 2001, 2007; Stuart et al.,2004; Higham et al., 2006; Jacobi et al., 2006; Gilmour et al.,2007), which have led to suggestions for allocation to mammalassemblage zones (MAZs). But numerous questions remain.Relatively few of the older sites can be given absolute dates(Pike et al., 2005), and assignment of assemblages such asCrayford in the lower Thames Valley continue to poseproblems; Sutcliffe (1995) favoured placement in cold MIS 6,while Schreve (2001b) has favoured warm MIS 7. The numberand age of phases within the ‘Cromerian Complex’ (Stuartet al., 2004), and the allocation of various assemblages tothem, continues to exercise stratigraphers, palaeontologists andarchaeologists alike, not least because of the effect of anydecisions on ideas about earliest human presence here (Lewiset al., 2004; Stuart et al., 2004; Parfitt et al., 2005). For mypurpose here I have placed assemblages variously argued tocome from MIS 19–15 within the heading ‘CromerianComplex’. More recently, Gilmour et al. (2007) have suggestedthat assignments of mammalian assemblages to MIS 5 and 4,the Joint Mitnor, Bacon Hole and Banwell MAZs requireadjustment and realignment based on new data derived fromthermal ionisation mass spectromatic (TIMS) dates. Table 1therefore presents a model of the stratigraphic and biostrati-graphic framework for localities and assemblages derived fromthe various studies, and forms the basis for the allocations ofpredator and prey species in Tables 2 and 3. I make no claimsthat it will be the last word on the subject.

The term ‘British Isles’ is taken here to refer to what some mayprefer to call the islands of Britain and Ireland, and whateverterm we use it is clear that the Pleistocene mammalian fauna ofIreland itself is much more restricted in time and in diversity –indeed, as pointed out in a recent overview, there is at best only‘limited evidence for a mammalian fauna older than 40 ka’(Woodman et al., 1997). Whether refinements to acceleratormass spectrometry (AMS) radiocarbon dating such as thosereported by Jacobi et al. (2006) will alter the chronologicalframe of this picture of Irish mammalian history remainsunclear, but they will do little to extend the relatively restrictedrange of species known from there. Whether the reduced faunareflects difficulties of access or destruction of evidence by laterice advances – or both – is unclear, but the extent and frequencyof land bridges between Ireland and Britain during thePleistocene seem far from clear (Devoy, 1995). I shall discussaspects of the Irish fauna at relevant points below. As for Britain,evidence for Pleistocene mammals comes in fact from sites thatwere for the most part to the south of the last ice sheet, and thusmainly from cave and open-air localities in England and somecave localities in Wales (Stuart, 1982, Fig. 2.2).

It will be apparent from Table 1 that we generally know moreabout the fauna of the warmer phases, the interglacials or atleast interstadials, than the colder or glacial phases, with theobvious exception of the last cold phase. It may be thought thatthe colder phases made for a less hospitable environment,although the extensive evidence for occupation during at leastpart of the Devensian cold stage tends to argue against thatsimplistic view. The very fact of possible correlations with theMIS sequence raises the question of changing sea levels and therecurrent isolation of the British Isles, and of Ireland fromBritain, a major factor in the dispersal and thus distribution ofterrestrial mammals. It seems generally accepted that until theAnglian (MIS 12) the British Isles were essentially joined to the


mainland of Europe, of which they formed a north-westernpeninsula, and had been since the end of the Pliocene(Gibbard, 1995), although Britain and Ireland seem to havebeen separated at that point from each other. Once the Straits ofDover were formed by rapid downcutting sea level thencontrolled insularity, but what seems less clear is the degreeand frequency of isolation during the Early Pleistocene andPliocene, although Funnell (1995) argues for an earlier seawayuntil the latest Pliocene. The paucity of terrestrial mammalianrecord for the Pliocene and Early Pleistocene of the British Islesscarcely helps any attempt to look at a longer timescale ofoccupation and movement, as Stuart (1982) pointed out.

The British guild of large Carnivora: thebroad pattern of appearance and diversityby family

A small but diverse large-mammal fauna may be broadlyassigned to the late Pliocene and perhaps Early Pleistocene inBritain (Stuart, 1982; Lister, 1998; Gentry, 1999), althoughallocation to cold and warm stages and hence MIS is difficult.However, the only clear evidence of larger Carnivora before theMiddle Pleistocene is the occurrence of the sabretoothedHomotherium latidens and an undetermined hyaena at theDerbyshire locality of Dove Holes associated with the Pliocenegomphothere Anancus arvernensis (Spencer and Melville,1974), the appearance of the large hyaena Pliocrocuta perrieriand a large ursid in Red Crag deposits (Newton, 1891), thecheetah Acinonyx pardinensis in the Norwich Crag at EastonBavents (Turner, 1995a) and the large hyaena Pachycrocutabrevirostris in the Siliceous Member at Westbury-sub-Mendip(Bishop, 1982). The paucity of evidence for this earlier periodmeans that we have no clear way of establishing presence orabsence, or of assessing the Early Pleistocene guild structure,hence the emphasis here on the Middle and Late Pleistocene.Nevertheless, it will be important to take some account of thewider guild structure known for the earlier period fromcontinental Europe, from which the guild of the British Islesmust derive and where we may see something of the transitionthat produced the Middle Pleistocene carnivore fauna. It is alsonecessary to consider here a small number of important guildspecies for which there is no record in the British Isles, and toconsider whether this may be a true picture of distributions.

The general distribution of guild species from the ‘CromerianComplex’ onwards is shown in Table 2, where some of the mostimportant localities are indicated. Particular attention is givento sole known occurrences of some species and to periodsduring which a more extensive record from a number of sitesallows us to infer actual absence from the British Isles. I stressthis latter point since absence from the fossil record is always adifficult thing to deal with and interpret. Some of the MiddlePleistocene patterns of presence and absence are based onsparse remains from such a small number of particular kinds ofsites that chance might well have left recorded speciesunknown and turned up the odd specimen of others so farunrecorded. In contrast, absences in some other periods arebased on a complete lack of specimens from a substantialnumber of diverse localities with very reasonable numbers ofother fossils. That of course is the nature of palaeontology, andwe must deal with what we have. In this context I have notincluded species recorded from Ireland in the tables either aspresent or absent, and will simply refer to those that are knownas relevant.


Tab

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Bri

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Copyright � 2009 John Wiley & Sons, Ltd. J. Quaternary Sci., Vol. 24(8) 991–1005 (2009)DOI: 10.1002/jqs


Table 3 British ungulate prey fauna for large Carnivora, showing known presence by MIS. Species are grouped into four broad size classes, from small(S) (ovicaprid or roe deer) through medium (M), large (L) and very large (VL) (rhino and above) based in part on sizes in living European cervids (Lister,1988)

MIS 19-15 13 11 9 7 6 5e 5c 5a 3

ElephantidaeMammuthus meridionalis (VL) X XMammuthus trogontherii (VL) X XMammuthus primigenius (VL) X X XPalaeoloxodon antiquus (VL) X X X X X X XRhinocerotidaeStephanorhinus hundsheimensis (VL) X XStephanorhinus kirchbergensis (VL) X X XStephanorhinus hemitoechus (VL) X X X X XCoelodonta antiquitatis (VL) X XEquidaeEquus altidens (L) XEquus sussenbornensis (L) XEquus ferus (L) X X X X X X XEquus hydruntinus (M) X XCervidaeCervus elaphus (M–L) X X X X X X X X XDama dama (M) X X X X XCapreolus capreolus (S) X X X X X X XMegaloceros dawkinsi (M) X XMegaloceros verticornis (L) X XMegaloceros savini (M–L) XMegaloceros giganteus (L) X X X XAlces latifrons (L) XCervalces sp. (L)Rangifer tarandus (M) X X X XBovidaeBison schoetensacki (L) X XBison priscus (L) X X X X X XBos primigenius (L) X X XBovidae sp. (L) X XOvibos moschatus (M–L) X X XSaiga tartarica (S) XSoergelia elisabethae (M–L) cf.Caprini indet. (S) XHippopotamidaeHippopotamus amphibius (VL) XSuidaeSus scrofa (S–M) X X X X X X

Sources: Currant (1989, 2004a,b); Gentry (1999); Currant and Jacobi (2001); Schreve (2001a,b, 2004); Stuart and Lister (2001, 2007); Stuart et al.(2004); Higham et al. (2006); Jacobi et al. (2006); Gilmour et al. (2007).


Hyaenidae

Hyaenas are important guild members from the mid Mioceneonwards in Africa and Eurasia (Werdelin and Turner, 1996a,b;Turner et al., 2008). The living species are cursorial predatorsthat may chase their prey for some distance before bringing itdown and killing by eating it. Group-hunting abilities meanthey take medium and small prey and therefore sample much ofthe ungulate fauna, while their numbers and the carcass-destroying capabilities of the large species help themappropriate kills and affect the availability of food to would-be scavengers. They can accumulate bone, and suchassemblages may hold much information about predator andprey activities.

The spotted hyaena, Crocuta crocuta, is recorded in thelower unit TD 3/4 of the Spanish site of Atapuerca, below theBrunhes/Matuyama boundary (Garcia and Arsuaga, 2001). It isalso found in latest Villafranchian assemblages from the karstfissures at Selva Vecchia in Italy (Turner, 1995a), and these maywell be the earliest records of the species in Europe. In Britain it


first appears in the Freshwater Bed of the Cromerian typelocality at West Runton in Norfolk as well as in ‘CromerianComplex’ deposits at Palling and the rootlet deposits at CortonCliff (Newton, 1883; Lewis et al., 2009). Small numbers occurin the latest Calcareous Member at Westbury in the Mendips(Turner, 1999b), but it has a patchy and sparse distribution untilthe truly enormous numbers at Ipswichian and Devensianlocalities (Turner, 1981, 1995a). There are indications, basedon dental morphometrics, that the British Ipswichian andDevensian hyaenas were separate populations (Turner, 1995a),and refinements to the stratigraphic position of several hyaena-less assemblages from Mendip in particular that may now fallinto the intervening period of time (Currant, 2004a,b; Gilmouret al., 2007) support this view but it needs further examination(see below and Table 1). Crocuta is one of the few speciesrecorded from last cold stage deposits in Ireland.

The large, short-faced hyaena Pachycrocuta brevirostrisoccurs at a number of Villafranchian/early Galerian faunal sitesin continental Europe (Turner and Anton, 1996; Turner, 2001;Turner et al., 2008). In East Anglia it occurs in the Cromer



Forest-bed Formation (CF-bF) at Bacton, Mundesley andSidestrand, but presence in the Freshwater Bed of theCromerian ss is based only on coprolites, as it is at NortonSubcourse (Lewis et al., 2004). It is also present in thelowermost (Siliceous Member) pre-Cromerian deposits atWestbury (Bishop, 1982; Gentry, 1999; Turner, 1999b). Earliestknown appearance at the early Pliocene locality of Olivola inItaly and the apparent hiatus in the CF-fb (Lister, 1996; Stuartet al., 2004), implies that specimens from East Anglia musttherefore belong either to the ‘Cromerian Complex’ or to thevery earliest Pleistocene, although the Westbury specimen mayfall within the intervening period. Co-occurrence with Crocutatherefore seems unlikely in the British Isles, but needs furtherinvestigation.

A third large European bone-crushing hyaena, Pliocrocutaperrieri, disappeared from Continental records at the end of thePliocene after a lengthy and widespread occupation (Turner,1995c). It is included in Red Crag deposits of Suffolk (Lydekker,1886; Newton, 1891), which probably equate with the latePliocene Praetiglian of the Netherlands (Stuart, 1982, 1988;Gibbard et al., 1991; Stuart et al., 2004). The species thenreappeared in the earlier part of the Middle Pleistocene atPetralona in Greece, Mauer and Mosbach 2 in Germany, and atL’Escale and Lunel Viel in southern France, where it waspreviously identified as a large striped hyaena, Hyaena prisca(Turner, 1990). But it is not known in Britain during the MiddlePleistocene, nor in Italy. However, it may have reached theIberian Peninsula, contrary to earlier suggestions (Turner,1995c), in view of the presence there of what has beenidentified as Hyaena prisca at the Portuguese Late Pleistocenelocality of Furninha (Cardoso, 1992). Based on Cardoso’sdescriptions, measurements and illustrations of this material itappears to be identical to other specimens of P. perrieri (Turneret al., 2008). What prevented recolonisation of Britain?

A fourth species should be mentioned, although it isunrecorded from the British Isles. This is Chasmaportheteslunensis, often referred to as the ‘hunting hyaena’ because of itsrelatively gracile build (Anton et al., 2007; Turner et al., 2008).The animal was widespread throughout the earlier part of theVillafranchian (Turner et al., 2008), with its last Europeanknown appearance at the German Early Pleistocene site ofSchernfeld (Turner, 1995a), and appears to have been animportant component of the fauna as a cursorial predator.

Felidae

Felids are essentially soft-tissue consumers. They hunt using acombination of teeth and claws to capture their prey and killbefore eating, and while the size and strength of larger speciesmeans they may take large prey, even groups of lions may lose acarcass to spotted hyaenas (Turner and Anton, 1997). Lionsapart, living species tend to be solitary hunters, but it is possiblethat some of the extinct machairodont species (the variouslyknown sabretooths and dirktooths) may also have tended tooperate in groups (Turner and Anton 1999; Anton et al., 2005).It is worth pointing out here that the larger felids are unknownfrom Ireland except for lynx, otherwise unconsidered here,which occurs with a date of around 9 ka BP in Kilgreany Cave(Woodman et al., 1997). The lion-sized sabretoothed Homo-therium latidens is known throughout the Villafranchian faunalspan and into the Middle Pleistocene of continental Europe(Anton et al., 2005). The animal occurs in various British EastAnglian deposits of the CF-bF – it is recorded from Bacton (a left


upper canine, Natural History Museum collection M17926),Kessingland/Pakefield (a right mandible; Stuart, 1982) andSidestrand (a fragment of the crown of a right upper canine,Natural History Museum collection M6084) – and is probablythe larger of the two previously unidentified large felids in theFreshwater Bed deposits at West Runton (Turner, 1999b; Lewiset al., 2009). Its last established occurrence is in the latest (MIS13) deposits at Westbury (Turner, 1999b), although canines ofthe species have been claimed to derive from Devensian-agecave earth deposits at Kent’s Cavern in Devon (Proctor et al.,2005) and the authenticity of a find of similar apparent age atRobin Hood’s Cave in Derbyshire remains unclear (Kurten,1968). However, a mandible from the North Sea has recentlybeen AMS dated to around 28 ka BP (Reumer et al., 2003); theimplications of such a late occurrence would be profound(Anton et al., 2005), and I return to this question below in thesection on guild evolution.

The earliest European appearance of the lion, Panthera leo, haspreviously been cited as the southern French Early Pleistocenesite of Le Vallonnet (de Lumley et al., 1988; Turner, 1995a), butit is now apparent that the identification was made on what isprobably the canine of P. gombaszoegensis (Moulle, 1998) andthat the earliest known lion from the Continent is from theItalian early Middle Pleistocene site of Isernia la Pineta (Sala,1990; Coltorti et al., 2005). It is first seen in Britain in‘Cromerian Complex’ deposits, with the best specimen being adamaged mandible from Pakefield in Suffolk (Turner 1995a;Lewis et al., 2009). A large, rolled felid distal humerus, foundon the beach at Cromer and held at Norwich Castle Museum,also appears to be of lion (Turner, 1995a). The ‘undeterminedlarge cat’ from the West Runton Freshwater Bed listed by Stuart(1996) in fact represents two other felid taxa – Homotherium(see above) and probably P. gombaszoegensis (see below) – buttwo further felid specimens in the West Runton collection at theNatural History Museum are much more likely to be of lion,Panthera leo; M17904, is a first phalanx, M17887 a secondphalanx.

Lion is common in MIS 13 deposits at Westbury, and thenappears consistently but in small numbers in later deposits ofMIS 11, 9 and 7 before becoming more numerous in Ipswichian(MIS 5e) and Devensian (MIS 3) localities, particularly in theMendips (Turner, 1995a, 1999b), although whether this impliesa continuous, indigenous population is unclear.

The leopard, Panthera pardus, has been thought to be firstclearly seen in Europe in early Middle Pleistocene deposits atMauer and Mosbach in Germany, and to have dispersed fromAfrica with the lion (Turner, 1992). However, the first Europeanrecord of leopard may be close to the Jaramillo event at LeVallonnet (de Lumley et al., 1988; Turner, 1995a), althoughHemmer (2001) suggests that this may also be a representativeof his Puma pardoides (see below). Having seen the materialI tend to favour the earlier identification as leopard; the bulge atthe rear of the M1 appears more in keeping with that species.Leopard is never a common member of fossil assemblages,except at the Italian Late Pleistocene locality of Equi Cave (DelCampana, 1954), although its known range in Europe is nowextensive. In the British Isles it is absent from ‘CromerianComplex’ deposits and Westbury (Turner, 1999b), and asidefrom isolated material from localities in the Mendips (Dawkinsand Sanford, 1866–8) – probably the MIS 7 deposits at BleadonCave (Currant and Jacobi, 2001) – the only other Britishmaterial is from the contemporaneous Intermediate Complexand Lower Breccia deposits at Pontnewydd in North Wales,dated to ca. 225 ka (Currant, 1984). How one interprets thisdistribution is less clear. Leopards are solitary and secretive cats,and even in Africa they are scarce as fossils, with the exception ofthe South African site of Swartkrans (Turner and Anton, 2004).



Panthera gombaszoegensis, a large cat intermediate in sizebetween lion and leopard, is first recorded in the earliest part ofthe Early Pleistocene at Olivola and in what seems to have beenthe uppermost levels at Tegelen in the Netherlands (Turner,1995c; O’Regan, 2002; O’Regan and Turner, 2004). It isrepresented by several teeth at Westbury (Bishop, 1982), and isprobably the smaller of two large felids from the type Cromeriandeposits at West Runton mentioned above (Turner, 1999a). Itmay also be present at Pakefield based on a distal right humerusidentified from a series of three colour photographs providedvia A. J. Stuart by a private collector who retains the item. Likethe very much larger lion humerus from Cromer beachmentioned above, it has all the attributes of a pantherine catand yet appears to be too small to be a lion (Lewis et al., 2009).A further probable occurrence is in MIS 11 deposits atSwanscombe in Kent, where lion is also clearly present, whichmay be the species’ latest European record (Turner, 1999a).

The European cheetah, Acinonyx pardinensis, together withHomotherium latidens, is one of the longest-lived of theVillafranchian taxa. It is best represented in late Pliocenedeposits at St Vallier in the Rhone Valley, but occurs verypatchily until perhaps the last record at Mosbach (Turner, 1995a).It is known from the British Isles from a single specimen, a small,broken P3 (Natural History Museum Collection M20227)previously identified as Felis pardoides (Turner, 1995a), fromlate Pliocene cliff deposits at Easton Bavents in Suffolk.

All of these latter three species are likely to have been solitaryanimals, at least beyond the size of a family group of femaleswith adolescent cubs, and simulation studies of extinctions ofsolitary cats (O’Regan et al., 2002) suggest that they coulddisappear quite quickly in an area the size of the British Isleseven if one allows a reasonably large initial population, so thattheir scant traces in the fossil record here are not especially odd.

One of the long-standing puzzles of European felid taxonomyhas been the medium-sized cat from the late Pliocene of StVallier in France, originally named Panthera schaubi by Viret(1954) and later Viretailurus schaubi by Hemmer (1964). Theanimal has been tentatively claimed at Early Pleistocenelocalities of Villarroya, Puebla de Valverde and Cueva Victoriain Spain (Turner, 1995a). Hemmer (2001) and Hemmer et al.(2004) have most recently transferred all of this material, alongwith many other specimens from Eurasia and Africa, to theAmerican genus Puma, represented by the modern-daymountain lion, with the Eurasian material named Pumapardoides. However, one should note that only St Vallierprovides substantially complete material and thus a compellingargument for this identification, and that many of the otherreferrals over the years, whether to Viretailurus or now to Puma,have essentially been made by default, rather than by positivematching with known characters of the material from St Vallier.

Mention should also be made of another species as yetunknown from Britain, the dirktoothed Megantereon. Thismedium-sized, strongly built machairodont cat has probablythe most restricted geographical distribution among Europeanguild members, leading Kurten (1968) to suggest that it waslargely a circum-Mediterranean animal. As such, it may trulynever have reached the British Isles, although the latestoccurrence is now in Jaramillo age deposits at Untermassfeld inThuringia, eastern Germany (Hemmer, 2001).

Canidae

As pack hunters the canids would have been important guildmembers (Wang and Tedford, 2008), able to take larger prey


but probably more prone than hyaenas to losing it to com-petitors. Masini and Torre (1990) identify a small, jackal-sized,wolf-like animal variously referred to Canis etruscus orC. mosbachensis in the Early Pleistocene Olivola fauna ofItaly, and this appears to be the common small canid during theearly part of the Middle Pleistocene (Sotnikova, 2001). Towardsthe end of the Middle Pleistocene, from MIS 7 onwards, there amarked increase in the size of specimens, which are identifiedas the living Holarctic wolf, Canis lupus (Turner, 1999b;Sotnikova 2001). In between these events lies the firstappearance of a second large canid previously referred toXenocyon lycaonoides but now also placed within Canis oreven Lycaon (Martınez-Navarro and Rook, 2003).

The earliest appearance of the smaller Canis mosbachensis inBritain is based on material from CF-bF deposits at Sidestrand(Stuart, 1982, 1988; Gibbard et al., 1991). It is also present inthe Freshwater Bed at West Runton, and is similar in size tospecimens from Westbury (Turner, 1999b) and Boxgrove(Roberts, 1990) in MIS13, from Swanscombe (Turner, 1999b) inMIS 11 and Grays Thurrock in MIS 9 (Turner, unpublisheddata). Representatives of the larger Canis lupus are seen innumerous deposits of Devensian and last interglacial age atDevon sites such as Kent’s Cavern, Joint Mitnor and theTornewton Cave Hyaena Stratum (Turner, 1995a), and it isknown from Ireland in pre-Last Glacial Maximum deposits(Woodman et al., 1997). A small number of specimens from theunderlying Bear/Glutton Stratum at Tornewton Cave, togetherwith the MIS 7 material from Crayford, also appear large,although one specimen from the Bear Stratum at Tornewtondoes fall within the range of Westbury measurements (Turner,1999b). Currant (1984) describes fragmentary material of Canisfrom the MIS 7 Intermediate Complex at Pontnewydd whichfalls within the larger size range. The large, Holarctic wolf-sized Canis lycaonoides is known in Britain only from the MIS13 deposits at Westbury (Bishop, 1982), where it makes animportant addition to the sparse European record of the speciesbut provides little by way of additional understanding of itsecological significance (Turner, 1999b).

Ursidae

Bears are often considered as omnivores, but they can beeffective hunters too and will drive other, smaller predatorsfrom kills. As large animals they can consume considerablequantities of food, but they are essentially territorial and (exceptfor mothers with cubs up to 2 years old) solitary. Maximumobserved and estimated densities of 55 animals of all ages per100 km2 are recorded from North America, but these occuronly in areas with abundant salmon runs and densities in singlefigures are more common (Mace et al., 1994; Miller et al.,1997). Taken overall they are likely to have been mostimportant as guild members when competing locally for accessto caves or shelters. The large Ursus deningeri, the likelyforerunner of U. spelaeus, is known in some numbers fromBacton (Bishop, 1982) and has previously been identified in thematerial from the West Runton Freshwater Bed deposits,although more recent work now suggests some similarities tothe living brown bear, Ursus arctos, which if confirmed wouldmake this one of the earliest occurrences of this species (Lewiset al., 2009). The entry in Table 2 reflects this uncertainty. Ursusdeningeri is very abundant at Westbury (Bishop, 1982), wheredenning activity is clearly evident (Andrews and Turner, 1992;Turner, 1999b), and it is recorded at Boxgrove (Roberts, 1990).



In contrast, Ursus spelaeus is much rarer. It is recorded atSwanscombe (Sutcliffe, 1964), and seems mostly to haveconfined its activities to the Continent. Currant (1984)described the bear from the Intermediate Complex and LowerBreccia at Pontnewydd as having ‘spelaeoid characters’, butdid not assign it to any particular species. However, morerecently he has argued that there and at a small number of cavesites at Torbryan in Devon, including the MIS 5e ‘Bear Stratum’of Tornewton Cave, U. spelaeus may show a more drawn-outcontinuity with and eventual replacement by Ursus arctos(Gilmour et al., 2007; Currant, pers. comm.). Brown bear aloneis known from most later deposits, and is certainly present inIpswichian and Devensian age localities, and is represented atBanwell Cave (MIS 5a) in the Mendips by a particularly largeform. In Ireland it occurs in several deposits with radiocarbondates back as far as 40 ka BP (Woodman et al., 1997).

Climate changes and the prey fauna

As argued elsewhere (Turner, 1995a; Turner and Anton, 1999;Anton et al., 2005), there can be little doubt that many aspectsof the evolution of the large carnivore guild in Europe reflectchanges in the ungulate fauna and that those changes in theungulates were themselves under the ultimate influence ofchanges in climate and part of larger-scale patterns of Plio-Pleistocene dispersals, many of which originated in Asia (Sher,1992). The wide distribution in time and space of the Europeanguild clearly suggests that the main constraints on distributionwere the availability of food rather than climate per se, and thatchange in the structure of the prey community driven bychanges in vegetation (Zagwijn, 1992) may have been animportant factor. The appearance of larger-bodied cervids andbovids in Europe towards the end of the Villafranchian mayhave disadvantaged the machairodont cats in particular (Turnerand Anton, 1999), and contributed to their disappearance or, atthe very least, to a reduction in numbers.

Table 3 shows the larger mammal prey species that can beassigned to the oxygen isotope stages of the British Isles, and itshould be clearly understood that the assignments to stagesrepresent a summary of points amplified in the text. Thesespecies are grouped into four broad size classes, from small(ovicaprid or roe deer) through medium, large and very large(rhino and above) as explained in the table, since size is ofmajor concern to any potential predator. I have concentratedon species from the ‘Cromerian Complex’ onwards for reasonsmentioned above, although just as we know of a poorlyrepresented pre-‘Cromerian Complex’ carnivore guild we alsohave some generalised indication of the kind of prey speciesavailable to its members (Stuart, 1982; Gentry, 1999). Theseinclude the proboscideans Anancus arvernensis (a gom-phothere), Zygolophodon borsoni (a mastodont) and Mam-muthus meridionalis (an early form of mammoth) and theperissodactyls Hipparion sp. (three-toed horse), Equuscf. E. stenonis and E. bressanus (respectively zebrine andcaballine true horses), Tapirus arvernensis (a tapir) andDicerorhinus megarhinus (a rhino). While the horses and tapirwould have been obvious prey, the others were surely too largeto have figured frequently in the diet other than as carrion.More probable contenders included Leptobos sp. (a lightly builtlarge bovid), Gazella sp. (a gazelle) and one or more speciesof Eucladoceros (a medium-sized deer). Gentry (1999) alsoidentifies a smaller cervid from the Siliceous Member atWestbury, although allocation to species is in his viewuncertain. While details thus remain unclear, it is evident that


the carnivores of the later Pliocene and Early Pleistocene wouldhave had little difficulty in finding food.

From the ‘Cromerian Complex’ onwards we have a clearerpicture of structure and changes in the herbivore faunas, at leastin so far as presence and absence are concerned. Of these, thevery large taxa – the elephants, rhinos and hippos – are againlikely to have been relatively immune to predation as adults,although juveniles may have been taken and dead animals fromnatural mortality are always likely to have formed a scavenge-able resource. Hunted prey are more likely to have come fromamong the equids, cervids, bovids and suids, with the largelitters of the latter likely making an attractive source of easilycaught food for all the larger predators (Anton et al., 2005).However, the importance of some members of these families ishard to judge because the records are scarce – the bovidsOvibos, Soergelia and various Caprini would certainly fall intothis category. The few examples that are known can hardlyhave been isolated individuals, but we have insufficientmaterial and therefore no basis on which to infer anythingelse about populations, distributions or predator activities andpredator–prey relationships. Broadly speaking, the best infor-mation for such matters comes from concentrated deposits incaves where predator accumulation can be reasonablyinferred, and these are best known from Ipswichian interglacial(MIS 5e) and Devensian glaciation (MIS 5d–2) periods, with theaddition of earlier Mendip localities such as Westbury (MIS 13)and Bleadon and Hutton (MIS 7). The MIS 5e and 3 assemblagesin particular have a wider importance, since they give us largesamples of remains and represent two very different sets ofclimatic and vegetational conditions. In both periods, Crocutaseems to been largely responsible for much of the accumulationof assemblages, but the respective population structures of thehyaena seem in turn to have been affected by the conditions ineach period (Turner, 1995a, and next section).

In Ireland the prey fauna of the last cold stage was impoveri-shed relative to that of Britain, but nonetheless consisted ofmammoth, reindeer, red deer, Megaloceros and horse, with thenotable absence of woolly rhino and bison (Woodman et al.,1997). Stuart (1995) notes an interesting addition to the list inthe form of musk-ox, Ovibos, identified by Lister.

A number of points in the ungulate fauna distributionssummarised in Table 3 require emphasis. First is the absence ofhorses in the relatively diverse MIS 5e assemblages incomparison with their ubiquity and abundance in MIS 3assemblages. After extensive previous debate this Stage 5eabsence is now widely accepted as real. Secondly, in MIS 3assemblages, and perhaps as early as MIS 4 (Gilmour et al.,2007; see below for discussion of this allocation), horsesreappear in quantity and are joined by often large assemblagesof remains of Rangifer tarandus, the reindeer, and in many casesby extremely large quantities of bones and teeth of Coelodontaantiquitatis, woolly rhinoceros. At many Devensian sites theyare accompanied by bison (Bison priscus), giant deer(Megaloceros giganteus) and red deer (Cervus elaphus) andconsistent although fragmentary evidence of Mammuthusprimigenius, the mammoth. There is little evidence of regionalvariation in these distributions, although red deer appears tohave been absent from the assemblages of Robin Hood’s,Church Hole and Pin Hole Caves at Creswell Crags inNottinghamshire (Turner, unpublished data). The third pointto make is the impoverished but widely recognised ungulatefauna of bison and reindeer, identified as a consistent feature atperhaps a dozen sites across England, accompanied by arestricted carnivore guild of Canis lupus and an at times verylarge Ursus arctos, and previously allocated by Currant andJacobi (2001) to MIS 4. More recent work involving dating ofstalagmites at a number of key localities now leads them to



allocate the assemblage to MIS 5a (Gilmour et al., 2007). I shallreturn to these and other points in the next section.

The evolving guild of larger Carnivora

The larger European picture

Across the continent of Europe as a whole there is a pattern ofmajor change in both the structural and the taxonomic makeupof the larger carnivore guild as we move from the later Plioceneinto the Middle Pleistocene (Turner, 1995a; Turner and Anton,1999). One of the most obvious features of the later Plioceneguild is the almost complete absence of the canids, andcertainly of larger taxa, and thus the apparent absence of acursorial, pack-hunting species. However, the two hyaenas ofthe time, Pliocrocuta and Chasmaporthetes, were undoubtedlyable to operate in such a manner. Pliocrocuta, while less robustthan the later Pachycrocuta and Crocuta, was a large hyaena,one clearly also able to act as a scavenger and no doubt capableof operating in a group, consuming bone and destroyingcarcasses. Although I have previously suggested (Turner,1995a) that Chasmaporthetes should be classed with thespecialist flesh eaters on the basis of its dentition and gracilebuild, this may give rather the wrong impression of itscapabilities. Clearly it lacked the robust and even hyper-robustdentition of the bone-cracking hyaenas, but at the same time itwas certainly better equipped to deal with bones than even alarge canid. We may therefore see in Chasmaporthetes the finalstage in the long chain of events by which the early hyaenas ofthe late Miocene and Pliocene played the ecological role latertaken over by the canids (Werdelin and Turner, 1996a,b; Turneret al., 2008).

By the Early Pleistocene the relatively long-lived andseemingly fairly stable guild of the European later Pliocenewas changing, with Pachycrocuta brevirostris replacingPliocrocuta and medium-sized and larger canids replacingChasmaporthetes. Among the cats, the machairodonts Homo-therium and Megantereon continued, as did Acinonyx andseemingly Puma, but the appearance of Panthera gombaszoe-gensis added another, fairly large cat to the mix of species.Megantereon was the first of these to disappear after its lastknown appearance at Untermassfeld in Thuringia at aroundthe time of the Jaramillo event, perhaps the most vulnerable ofthe species to climatic changes and their effects in opening thevegetation somewhat at around that time (Turner and Anton,1999; Anton et al., 2005). But an even more impressivetransition occurred in the Middle Pleistocene with an increasein taxon numbers, leading to the co-occurrence of the modernPanthera leo, P. pardus and Crocuta crocuta with the remnantsof the older, Villafranchian guild and its Early Pleistoceneadditions of Pachycrocuta, Panthera gombaszoegensis and thecanids. The Middle Pleistocene reappearance, at least in someparts of Europe, of Pliocrocuta perrieri swelled the guildnumbers even more, and if the smaller cat at Vallonnet isindeed a leopard then the felid component in the later stages ofthe Early Pleistocene and the earlier part of the MiddlePleistocene was particularly large. However, that situation wasshort lived and appears to have come to an end by around 0.5Ma with the demise of the archaic members of the guild,leaving a European community broadly similar in membershipand structure to that of East Africa today (Turner, 1990; Turnerand Anton, 1999). In fact comparison with Africa is ratherinstructive, because it can show us that such a large andseemingly crowded guild as we appear to have had during the


earlier part of the European Middle Pleistocene is not such anoddity. A rather similar pattern of overlap and eventualreplacement had already taken place in Africa, but with a verydifferent timescale. Most of the modern large carnivore specieswere present in Africa by 3.0–3.5 Ma, whereas many of thearchaic elements of the guild such as Homotherium, Mega-ntereon and Chasmaporthetes only became extinct therearound 1.5 Ma (Turner, 1990; Turner and Anton, 1999,2004). However, despite having a Miocene guild that includedthe ursid tribe Ursavini and the bear-dog family Amphicyo-nidae, Africa was never clearly occupied by members of theursid tribe Ursini, the bears as we think of them in Eurasia andthe Americas and which have been a constant presence in thoselatter continents (Turner and Anton, 2004).

The British Isles

For the British Isles, some components of that whole latePliocene–Middle Pleistocene transition, a period in excess of1.0 Ma, are clearly missing. The most notable absentees areMegantereon and Chasmaporthetes, but it is possible to seeothers in place for at least some of the time. Pachycrocuta andPanthera gombaszoegensis, Homotherium and (albeit scarcelyrepresented) Acinonyx appear to have been establishedmembers of a guild that may have had the generalisedEuropean Early Pleistocene structure of large and medium-sized felids (including a machairodont cat) together with bone-crushing hyaenas and a cursorial component made up by thecheetah and possibly by the hyaena Pliocrocuta. Bears of somedescription may have been present, but we lack any detail.What is notably missing, however, is a pack-hunting cursorialelement among the Canidae; there are no known EarlyPleistocene dogs here, and it is hard to judge whether this isa real absence or simply a taphonomic artefact. With theappearance in the Middle Pleistocene of the lion, spottedhyaena, two kinds and sizes of dog (and perhaps even theleopard), both Homotherium and Panthera gombaszoegensismanaged to continue to coexist for some time in a guildcontaining several large cats, a large bear, pack-hunting canidsand a very capable cursorial pack-hunting and scavenginghyaena.

For the ‘Cromerian Complex’ and MIS 13 a reasonablywell-established guild of Crocuta, lion, Homotherium,P. gombaszoegensis and at least one species each of dogand bear seems to have been present, although numbers ofspecimens at any one locality are never great and parameterssuch as population size, body size, sex ratios, sexualdimorphism and of course typical age at death are thus difficultto discern. But such a relatively large guild at least points tostructure and thus complexity, with solitary and group hunters,ambush and cursorial predators represented. Although thearrival of lion and P. gombaszoegensis may have impinged onHomotherium, all three seem to have coexisted until the time ofthe Westbury deposits in MIS 13. The one species that is betterknown is the large bear, Ursus deningeri, for which the largesample from Westbury has permitted some insights intodenning activities and the clear presence of females withyoung in one part of the cave at one period and of largely malebears in another part during another period (Andrews andTurner, 1992; Turner, 1999b). The quantities of bear remainsthere imply a long period of occupation with presumably asingle adult denning each year, and other predators thereforepresent at different times. The Westbury guild assemblage is infact one of the most diverse, with – as has been pointed out – theonly recorded presence of the large Canis lycaonoides in theseislands, but it is hard to discern the nature of relationships



between the guild members in view of the scarce quantities ofmaterial, and I have previously suggested that some combi-nation of the locality and the form of the cave may have meantthat only bears used it consistently (Turner, 1999b). Canislycaonoides was presumably a pack-hunting predator, butquite why it proved so unsuccessful in the longer term here andelsewhere in Europe is less clear, unless some combination ofthe presence of Crocuta and Canis mosbachensis proved toomuch in the way of competition for resources, since it does notappear to have overlapped in time with the larger Canis lupus(see below).

For warm stages MIS 11 and 9 the evidence of change in theguild structure and composition is equivocal (Table 2). DuringMIS 11, the Hoxnian interglacial, lion and P. gombaszoegensisare present along with the smaller C. mosbachensis but Crocutais unrecorded. The cave bear, U. spelaeus, is also recorded forthe first time from this stage, but there is now no evidence forHomotherium. However, the fate of Homotherium, not only inthe British Isles but also in northern Europe in general, hasrecently come under renewed scrutiny. The North Seamandible reported by Reumer et al. (2003), an undoubtedHomotherium with a determination of 28 100� 220 14C a BP,remains the only directly dated specimen, but a presence inLate Pleistocene deposits from Kent’s Cavern in Devon wassuggested by Kurten (1968) and has more recently beenrepeated by Proctor et al. (2005). However, as the latter authorsconcede, Cook and Jacobi (1998) have argued that thespecimens – isolated teeth – could as well have beenintroduced into those deposits by Late Pleistocene humans,and indeed could even have been derived from the earlier,Middle Pleistocene deposits that are also present in the cave.Whatever the true origin of the Kent’s Cavern finds, if the datesare correct then the animal must by implication have led a veryquiet existence that kept it out of most deposits and the fossilrecord, as Anton et al. (2005) have argued. In comparison, eventhe solitary and secretive leopard managed to leave traces at acouple of sites here in MIS 7 and a scattering of sites on theContinent. Anton et al. (2009) concluded that this lack of otherconfirmed records for the later Middle Pleistocene and LatePleistocene strongly suggests that the lion was the dominantlarge felid in most if not all parts of Europe. Of course, ifHomotherium were still present then it would have playedsome part in guild structure and operation, although how we goabout assessing that impact is unclear. When it was present,Homotherium clearly operated as a cursorial group hunter andflesh eater, as pointed out by Anton et al., although as they alsowent on to argue, it is unlikely, given its dental characteristicsand likely social structure, that it left significantly more in theway of scavengeable resources for competitors than any otherfelid. I shall return to the role of the lion below, but it is clearthat the North Sea Homotherium warrants further examination,including redating using the ultrafiltration methods describedby Jacobi et al. (2006), a point reiterated more recently byAnton et al. (2009).

This guild structure of MIS 11 seemingly continues in the nextwarm stage MIS 9, with Crocuta identified at Purfleet only onthe basis of a coprolite (Schreve, 2001b, 2004) although there isalso a small amount of skeletal material from Grays Thurrock inthe NHM collections, and lion is scarce. A small number ofsmall canid remains from Grays Thurrock are assigned to thisstage, and the presence of the living brown bear, U. arctos, maybe noted. In MIS 7 we get the first evidence for the larger Canislupus, the extant wolf, in a guild that now clearly includesCrocuta, lion, brown bear and the only known records ofleopard. If nothing else, the presence here of larger, pack-hunting canid in competition with Crocuta is likely to havealtered the balance of predator-prey systems and to have


affected the quantities of scavengeable material available in thelandscape. On the face of things it may appear odd that such acursorial predator should have appeared together with anambush hunter like the leopard, but wolves can operateefficiently in wooded terrain and the overall presence of lion,leopard, wolf and spotted hyaena points to a mosaic of habitats,different in details but similar in structure to that seen inmodern-day eastern Africa. This guild structure, perhaps minusthe leopard, seems to have been reasonably well suited toconditions during MIS 5e and 3, the Ipswichian interglacial andthe Middle Devensian glaciation, judging from the number ofsites at which it occurs in those stages, but as Table 2 indicateschange still took place.

The presence of both Crocuta and lion in MIS 5e and 3assemblages (and the hyaena also in MIS 5c), but their absencefrom MIS 5a, has a potential bearing on likely contacts betweenthe British Isles and the European mainland. Age structure andtooth size data for Crocuta of Ipswichian and Devensian agehave already suggested a disjunction between the twopopulations (Turner, 1995a, Table 2), and I have previouslyargued for the extinction of the British population at the end ofthe interglacial with a likely recolonisation during the lastglaciation in MIS 3. Even a small presence in MIS 5c at BaconHole would not contradict this general interpretation. Thisdisjunction is now supported by evidence of intermediate-ageassemblages such as Banwell, Windy Knoll, Pickens Hole andoverall some dozen localities where wolves together with anoften particularly large form of brown bear seem to have beenthe sole predators before Crocuta reappeared (Currant, 2004a).This is the Banwell Bone Cave MAZ assemblage previouslyassigned by Currant and Jacobi (1997) and Currant (2004a) toMIS 4 but more recently reassigned to MIS 5a on the basis ofrefinements in dating methods (Gilmour et al., 2007). However,comparisons between the admittedly smaller samples of lionsof Ipswichian age and Devensian age show no such diffe-rentiation in age structure or metrical data (Turner, unpublisheddata), but again their absence from MIS 5a assemblages ofBanwell type seems clear and strongly implies a similardisjunction.

Of course the precise meaning of this disjunction is anothermatter. Although I have previously argued for a recolonisationfrom the Continent, at least so far as the Devensian populationof Crocuta is concerned, one should also perhaps consider thepossibility of relict populations within the rest of the British Islesproviding the Devensian populations of lion and hyaena. Bisonand wolf continue from MIS 5e through into 3 without suchobvious breaks in occupation, and one could argue that lionsand hyaenas were simply restricted to other, perhaps morenortherly areas. It is even possible that the appearance in smallnumbers at Bacon Hole in MIS 5c deposits represents a relictpopulation in southern Wales, although the small sample size(perhaps two individuals) at Bacon Hole precludes aligning itwith either the MIS 5e or MIS 3 populations. However, we mayalso note the MIS 5a presence in some numbers of reindeer; it ishard to envisage likely refugia within the British Isles for thatspecies during stage 5e and much more probable that they atleast represent a true recolonisation. Gilmour et al. (2007) seethe MIS 5a fauna as an island assemblage of high-latitudespecies that remained here after moving into the British Islesduring a cooling event in MIS 5b. If so, that might make the laterparallel recolonisation of at least Crocuta, and perhaps also oflion, more probable; if they were present in northern areas ofBritain during MIS 5a then it is hard to see what could haveexcluded them from more southerly parts. This patterningtherefore suggests that the absences of lions and hyaenas in theBritish Isles after MIS 5e–c were real, and should perhaps beconsidered in conjunction with the later late Quaternary



megafaunal extinctions across Europe as a whole. I shall returnto this point in the final section.

Whether this patterning, whatever our interpretation, holdsany implications for the timetabling of appearance by lions andhyaenas during previous warm–cold stage transitions isunclear, but it serves as a warning against any naiveassumptions of population continuity for these and perhapsother Middle and Late Pleistocene predator species at points inthe sequence where resolution is less. In more general terms,the absence of such a major and evidently capable predator asCrocuta during MIS 5a may suggest that the absence of the otherlarge hyaenid, Pliocrocuta perrieri, from the British Isles, whenit reappeared in central and southern Europe during the MiddlePleistocene, is not an artefact of recovery. However, this shedslittle light on the question of whether the apparent absences ofthe cursorial hyaenid Chasmaporthetes and dogs from latePliocene and Early Pleistocene deposits here is real or simplyanother effect of the poorly represented fauna of these periods.

It is from MIS 3 assemblages that the Irish fauna is known, andseems to reflect the operation of a reduced guild of Crocuta andwolf, albeit with the addition of brown bear, taking acomparatively reduced but still fairly diverse prey faunawithout potential interference from lions. That guild structureis probably unique, although some aspects of the MIS 5a guildwithin Britain mentioned above seem to parallel it.

Guild interactions

The absence of lions and hyaenas from MIS 5a assemblagescorrelates interestingly with the large size of the brown bear ofthe period, as seen in particular at Banwell. Brown bears can befast moving and are clearly opportunistic predators. They willdrive wolves from kills, and can themselves take adult elk(Alces alces – moose in North American terminology), reindeerand red deer (elk in North American terminology), and canbecome adept at seasonal kills of calves of all three species(Boertje et al., 1988; Gunther and Renkin, 1990; Young andMcCabe, 1997). Alaskan brown bears tend to be the largestliving form of the species, and the size plasticity of the speciesduring the Pleistocene was extensively catalogued by Kurten(1958, 1959). Meiri et al. (2004) have confirmed previousinterpretations of size variation in living and fossil brown bearsas conforming to Bergmann’s rule, at least in North America,and studies by Hilderbrand et al. (1999) point to dietary meat,and in particular the availability of salmon, as a major factordetermining body size. Banwell Cave, situated as it is at thewestern end of the Mendips, would have been within strikingdistance of the coastline and any rivers of the time and thus ofany salmon runs, the type of area incidentally where populationdensities of living brown bears also tend to be greatest (Milleret al., 1997). This, combined with the absence of competinglarge predators other than the wolf, may go far to explain thesize of the MIS 5a bears from the site.

So far as size variation in other guild members is concerned, itis clear that British populations of Crocuta were composedof animals as large as, or larger than, any living members ofthe species (Kurten, 1958), although estimating the size ofPleistocene lions is less straightforward than it might appear.Strong sexual dimorphism in pantherine cats makes simpleinferences based on often scanty material surprisingly difficult,but we can reasonably infer that European lions in general werecomparatively large throughout the Pleistocene (Turner andO’Regan, 2002; Lewis et al., 2009). This implies that wemight revise upwards any ideas about the prey size limitations


imposed on European Pleistocene lions based on modern-daycomparisons, and may suggest that juvenile mammoths andrhinos may have been an attractive source of food. As discussedby Turner and Anton (1999), lions can survive in open habitatslargely because of their social structure, but retain very strongforelimbs and the highly propulsive hindquarters of an ambushhunter and can readily operate in closed conditions, much like atiger, as shown by their presence in virtually all phases of theMiddle and Late Pleistocene of Europe. The latter authors alsopointed out that lions readily turn to hunting smaller prey in timesof shortage, something likely to have given them a significantadvantage over the machairodont cats like Homotherium.

Of course, it is hardly surprising that interpretations of thestructure, evolution and interaction of the British largercarnivore guild prove difficult. In addition to the basic problemsof dealing with an incomplete record and an imperfectly knownseries of past ecosystems, we have the fact that nowhere todaydo animals such as lions and spotted hyaenas compete withwolves and bears (much less sabretoothed cats) for prey such asreindeer, bison, horses and mammoths. In other words, wehave no direct comparisons for much of what went on, and wemust therefore generalise from modern observations of largepredators. Take the question of what ate what among thespecies listed in Tables 2 and 3. It may seem that in comparisonwith ecosystems and ungulate faunas such as those found inAfrica now and in the past (Turner and Anton, 2004), the preyfauna of the British Isles shown in Table 3 is somewhat reduceddespite a reasonable range of large predators. But in fact thelarge carnivores in an ecosystem tend to prey on the mostabundant suitably sized ungulate species, however great thetotal range may be, as pointed out by Anton et al. (2005). Thusdespite the range of potential prey the lions, leopards, cheetahs,spotted hyaenas and wild dogs of Kruger National Park in SouthAfrica take the abundant impala as their main prey, although itis a little small for the lions to bother with and a little big for thecheetah to handle (Mills and Biggs, 1993); there is no simpleniche separation in the manner beloved of ecologists. In thesame way, the tigers, leopards and dholes of the BandipurReserve of India mainly take the small chital deer, despite thepresence of larger sambar deer, which might seem a moreobvious target for the tigers (Johnsingh, 1992). On this basis wewould expect that the large carnivores of the British Isles wouldhave tended to select from among the more abundant preyspecies, and that herding animals such as reindeer, othermedium-sized deer and perhaps horses would have formed thebulk of the kills, with obvious variations depending upon theclimate and vegetation of particular periods. Young, old andweak of other species close to the preferred size range mightalso be taken, but those of other, larger species are likely tohave been tackled much less frequently. Thus the kind ofdetailed ecological hierarchy of predation proposed byHemmer (2001) based on the assemblage from the EarlyPleistocene locality of Untermassfeld in Thuringia may beoffering too neat, too precise and possibly even too complex animage of prey apportionment.

In many ways perhaps the most interesting but so farunstudied interaction between guild members is that of Crocutaand the large wolf, Canis lupus. Both Crocuta and wolf aretoday efficient, adaptable, cursorial hunters, with strong socialstructures that allow them to live and hunt in groups (Mech,1970; Kruuk, 1972; Cooper et al., 1999; Mech et al., 2001;Honer et al., 2002). Nowhere do their ranges overlap today,and in Africa Crocuta is clearly dominant over the much smallerwild dog, Lycaon pictus, which anyway tends to take smallerprey (Kruuk, 1972). Both also scavenge, although the hyaena isthe supreme exponent of this strategy with its ability to consumebone and thus derive nourishment from a carcass that has no



useable food value to another predator, including wolves.Hyaena remains vastly outnumber those of wolves in MIS 5eand 3 assemblages (Turner, 1995a), and the greater proportionof all bone specimens in these deposits shows evidence ofgnawing and consumption by hyaenas, implying largepopulations. But wolves are also frequently represented andwere clearly contemporaries, and it remains difficult to discernthe actual living patterns of interaction from these assemblages.Both species would seem well suited to hunting reindeer andother cervids, including perhaps the largest in view of theevident success of wolves with elk. Horses would clearly be agood target for hyaenas, in view of the success of Crocuta intaking equids, but the wolves may not have been so suited. Datafrom Kruuk (1972) suggest that hyaena groups hunting zebrastend to be larger than those hunting other species. This is clearlya reaction to the anti-predator tactics of the zebras, wherestallions defend their females and young aggressively, and it isthe vast size of hyaena clans, the result of rather fluid socialsystem, that enables them to form a large enough hunting groupto tackle zebras successfully. Wolves, in contrast, tend to formsmaller packs that centre on and may be entirely composed ofrelated individuals (Mech, 1970), and although several wolvesmay be needed to hunt elk successfully the solitary nature of theprey requires nothing like the number needed by hyaenas totake zebra. An obvious distinction between prey preferences ofthe two predators in Pleistocene Europe may thus be inferred.

The obvious place to look at interaction between the twopredators, apparently stripped of interference from otherspecies except the bear, would be Ireland, where the guildseems to consist of only those three species. However, despitethe improvements in our knowledge of the fauna there outlinedin the discussion presented by Woodman et al. (1997), it is farfrom clear that we have sufficient evidence on which to base adetailed investigation, certainly until the dating of specimenscan benefit from the advances in methodology outlined byJacobi et al. (2006) and clarify the true extent of co-occurrence.

Final extinctions

As previous discussions have shown, changes in the largecarnivore guild have been a constant feature over the longertimespan afforded us by the fossil record, and some speciesseem to have been absent for a period only to reappear later. AsI pointed out when discussing individual species, computersimulations suggest that solitary cats may have gone extincthere relatively quickly and easily (O’Regan et al., 2002), andcould imply that this was a recurrent feature of their appearancegiven the variations in climate that have taken place. However,all of the large predators discussed are now completely extinctin the British Isles, and those that are found elsewhere nowcould only come back through deliberate reintroduction. Butwhile the timetable of final disappearance is relatively clear forsome taxa, it is less obvious for others. Yalden (1999) argues forsurvival of brown bear until Roman times (perhaps a little laterin Scotland) and for the wolf into the second millennium ADbut again later in Scotland (perhaps into the 1600s). Althoughthey do not specifically refer to lion, Currant and Jacobi (2001)distinguish between their MIS 3 Pin Hole MAZ, with lion andabundant Crocuta, and their Lateglacial period Gough’s CaveMAZ, in which both lion and Crocuta are absent in a fauna thatalso lacks woolly rhinoceros, reindeer and bison. The clearimplication is that the Last Glacial Maximum has acted as someform of watershed in the pattern of distributions, with Crocutadisappearing by perhaps 30 ka. Against this is a small amount of


dating evidence from Paviland Cave in southern Wales, whereAMS determinations of 17 670� 140 14C a BP for reindeer andof 17 880� 180 14C a BP for Crocuta were originally obtained.Jacobi et al. (2006) report on new AMS radiocarbon dates usingultrafiltration techniques for several taxa, including the Pavi-land hyaena, and report a revised date of 23 120� 130 14C a BP(OxA-13659) but consider even this ‘unrealistically youngwhen compared with the results from dating spotted hyaenas atother British sites’ (p. 570), although they make no mention oflion. I have some reservations about their argument for rejectinga date, based as it must be on the assumption that all periodshave been adequately sampled, but in fairness Jacobi (pers.comm.) points out that the Paviland hyaena specimen is heavilyconserved in a way that may throw any radiocarbon deter-mination into question. Stuart and Lister (2007) largely agreewith Jacobi et al. (2006) so far as the pattern of extinction ofCrocuta is concerned, but argue for much later presence of lioninto the Lateglacial in continental Europe, although they makeno mention of the latter from the British Isles.

One might suspect that the presence of both lion and Crocutain any given area would be closely dependent on an adequatesupply of suitably sized ungulates, as indeed it seems to be forany members of the large carnivore guild (Turner and Anton,1999; Anton et al., 2005), and it is not readily apparent that thiswould have posed particular difficulties here, especially sincewolves and bears seem to have been able to survive into theHolocene. However, we also have the apparent local absenceof lion and hyaena after MIS 5e–c until MIS 3 to consider as partof a larger pattern. This too was a period of time during whichwolves and bears clearly flourished, suggesting that there wassomething about life here for lions and hyaenas that could attimes cause populations to die out. While the simulation studiescited above show that it is solitary cats that are likely to suffercatastrophic population crashes quite readily, it is at leastpossible that conditions here forced lions to be less social thanthey are today in Africa. I have also previously suggested thatthe age structure of MIS 5e hyaenas in the British Isles may pointto a heightened mortality of younger adults sufficient to cause alocal extinction during a time when the population was isolatedfrom that on the Continent, but it is hard to relate thatinterpretation to the conditions at the end of MIS 3 when theyseem to disappear finally from Europe.

Although the broad pattern of guild evolution presented hereseems well established, and despite major advances in know-ledge, it is thus clear that we are still some way short ofunderstanding the detailed patterns of megafaunal extinctions,a point stressed by Stuart and Lister (2007).

Acknowledgements I am grateful to Andrew Currant, Roger Jacobiand Anthony Stuart for information about and discussions of some of thematters discussed here, and for the comments made by two anonymousreferees. In memory of Gill, for her love, help and support during thework represented here.

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the evolution of the guild of large carnivora of the british isles during the middle and late...

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