large carnivores and earliest european hominids: changing determinants of resource availability...

Alan Turner Hominid Palaeontology Research Group, Department of Human Anatomy and Cell Biolou, University of Liverpool, P.O. Box 147, Liverpool L69 3BX, U.K.

Received 28 May 199 1 Revision received 18 October 1991 and accepted 2 1 October 199 1

Keywordsxavenging, hominid dispersion, larger carnivore guild, earliest European occupation.

Large carnivores and earliest European hominids: changing determinants of resource availability during the Lower and Middle Pleistocene

The archaeological record seems to imply sporadic occupation of Europe during the Lower Pleistocene, but it may always be difficult to reach firmer conclusions from this alone. A framework for viewing the pattern ofEuropean settlement is proposed based on the changing availability of scavengeable remains oflarger ungulates. The structure ofthe larger carnivore guild would have conditioned that availability, and is likely to have presented obstacles to successful settlements until the latter part of the Middle Pleistocene when the guild came to resemble that ofmodern-day Africa. These changes imply that initial hominid colonizations during the Lower Pleistocene simply could not be maintained, and that only in the later Middle Pleistocene did conditions become suitable for more permanent occupation.

Journal of Human Evolution (1992) 22, 109-126

Introduction

One of the most significant landmarks in the evolution of the Hominidae after their origin in Africa (Simons, 1989) is their first appearance outside that continent. At a recent meeting in Paris, most contributors concluded that the earliest colonization of Europe probably occurred in the period 1.5-1.0 Ma, although the possibility of earlier traces coming to light could not be excluded (Ackerman, 1989; Delson, 1989; E. Bonifay & Vandermeersch, in press). But it remains true that the evidence for more intensive occupation comes from the Middle Pleistocene, as Isaac (1975) pointed out nearly 20 years ago, suggesting that the Lower Pleistocene witnessed no more than initial, sporadic attempts at colonization.

If Isaac’s interpretation remains broadly correct, then Lower Pleistocene evidence for hominid presence will continue to be relatively scanty. But how are we to distinguish between sporadic presence and incomplete recovery? Investigations dependent upon archaeology alone, or the discovery of hominid remains, will simply reinforce uncertainty about the true extent of Lower Pleistocene occupation until such times as a sufficient quantity of material has accumulated to satisfy the sceptical. The need for more data will probably continue to be a prominent part of any conclusion.

In this paper, an effort is made to develop an independent framework for viewing the archaeological evidence, one based on a consideration of the pattern of resource availability for hominids in Europe over the period of interest. The approach builds on the conclusion that larger ungulates would have been the major source of food available to hominids entering Europe, and on the inference that scavenging would have been the most probable method of exploitation. Successful scavenging depends very much on the presence of both carcase producers and competitors (Kruuk, 1972), to the extent that changes in the structure of the competitor community, in this case the larger carnivore guild (Turner, 1990), may be used to infer the likely success of the strategy through time (Turner, 1988). The resultant changes in the pattern of resource availability should not only offer support or rejection of interpretations based on the existing archaeological record, but should also result in predictions for the pattern offuture discovery.

The paper commences with an outline of the environmental background and the likely importance of ungulates as a source of hominid food in Europe, and emphasizes the value of

0047-2484/92/020109 + 18 $03.00/O 0 1992 Academic Press Limited

110 A. TURNER

scavenging as a means of procurement, before going on to discuss changes in the larger carnivore guild and their probable impact. The lower boundary of the Pleistocene is taken as 1.7 Ma based on the Vrica marine section in Calabria (Berggren et al., 1985).

Plio-Pleistocene environments and food resources

Environmental background

Marine sediments provide a detailed record of global climatic change (Shackleton, 1975; Shackleton & Opdyke, 1973, 1976, 1977; Shackleton et al., 1984; Kukla, 1977; Thunell & Williams, 1983; Prentice & Denton, 1988). They show a marked shift in climate at around 3.2 Ma, seemingly a reflection of the onset of glaciation in Iceland, followed by a drop in ocean temperatures and an increase in ice-rafted debris at around 2.5 Ma with the “first major northern hemisphere glacial event” (Shackelton et al., 1984:622). A further cooling took place 1.7-1.6 Ma, but perhaps the most significant climatic change of all occurred just after the Jaramillo magnetic reversal event of 0.95-O-9 Ma, with the first of the very cold phases seen at stage 22 of the ocean core data. Events since stage 22 mark the fully glacial phase of the Pleistocene, with the intensity and frequency of changes presenting the familiar pattern of glacial and interglacial cycles.

The effects of these global changes are seen in the terrestrial records of the Netherlands where detailed palynological analyses combined with studies of lithostratigraphy and cryoturbic structures provide information back as far as the late Pliocene (Zagwijn & De Jong, 1984; Zagwijn, 1985; De Jong, 1988). C orrelations have also been made between the Netherlands and the Mediterranean region (Sue & Zagwijn, 1983; Zagwijn & Sue, 1984; Sue, 1984). The results suggest that in northern Europe, warm-temperate and moist conditions gave way to cycles of cooling at around 3.2 Ma (Sue & Zagwijn, 1983)) while in the Mediterranean a change to a climatic rhythm of summer drought has been identified (Sue, 1984). In both the Netherlands and the Mediterranean, the vegetational changes record a shift to dryer and colder conditions at 2.4 Ma, and also show the beginnings of the climatic oscillations that characterize the later Quaternary. Those oscillations increase in amplitude and frequency in the Netherlands sequence just after the Jaramillo event and parallel those visible in the deep-sea core record. During the Jaramillo event the deposition of loess began to increase in abundance and frequency (Kukla, 1977). The cooling recorded in the deep sea cores at about 1.6-1.7 Ma may parallel the transition from the Tiglian to the Eburonian pollen zones of the Netherlands, if correlations suggested by Van Montfrans (1971) are correct.

Outside Europe, there is evidence from China of a change in vegetation at the Gauss/ Matuyama boundary (Azzaroli et al., 1988), while at around 1.0 Ma there is evidence for extensive loess deposition in the northern part of the country and in Central Asia (Dodonov, 1980, 1982; Derbyshire, 1983; Kukla, 1987). In the Himalayas there is evidence of increased tectonic activity at around 1.0 Ma (Azzaroli & Napoleone, 1982), which itselfis likely to have precipitated extensive changes in climate and vegetation. In east Africa, palynological data (Bonnefille, 1983, 1985) suggest an opening of habitats, supported by studies of small- mammal faunas (Wesselman, 1984, 1985) and Bovidae (Vrba, 1985, 1988). Apparently contemporaneous changes in the bovid faunas at South African sites (Vrba, 1988), together with alterations in the composition of sediments from deep-sea cores taken off the coast of western Africa (Janacek & Ruddiman, 1987), p oint to the continent-wide nature of these

LARGE CARNIVORES AND EARLIEST EUROPEAN HOMINIDS 111

environmental changes. A further shift towards open habitats is seen l&1*6 Ma at Omo,

Turkana and Olduvai (Bonnefille, 1983, 1984, 1985).

Implicationsforfood resources in Europe

The climatic data summarized above strongly imply that the past 3.0 million years in Europe (and probably in temperate Eurasia as a whole) have been dominated by conditions colder than those of the present day. Gamble (1986) stressed this point for the period since stage 22, arguing that over most of the last 900,000 years the climate was both colder and drier than at present in Europe. The true interglacials were of relatively short duration.

In such circumstances, as Gamble (1986:97) p ointed out, the key food resources available to hominids in Europe would have been animal. Even during the interglacials, times of greatest warmth, the vegetation could simply not have provided sufficient food on an annual basis without a well-organized means ofexploiting and storing the seasonally available items. The option of winter dormancy, a strategy successfully adopted by modern and fossil ursids (Kurten, 1958; Andrews & Turner, 1991) is simply not available. Terrestrial mammals are therefore likely to have been a key resource for hominids in Europe whatever the time of their arrival. That is not to argue for a total neglect ofvegetable resources by European hominids, simply to put the value of animals as a year-round food resource into perspective. Nor is it to ignore the potential contribution ofmarine foods, although variations in sea level are likely to have destroyed potential sources ofevidence for the extent ofsuch contributions. But even the most generous allowances for other food sources cannot disguise the fact that all claims for early hominid appearance are from localities with abundant terrestrial mammal remains.

Such implied dependence upon animals raises further questions about the proportion of protein consumed and the ability ofhominids to subsist on a meat-rich diet. Speth (1989) has recently shown that prodigious consumption of meat alone is simply not possible on a sus- tained basis for modern humans because of physiological limits to protein intake and the unbalanced nature of the diet. However, if fats are also consumed then a diet rich in meat is possible. Fats are therefore as much ofa target as meat when people place a heavy reliance on animals as food, not only implying that the condition of the animals eaten by humans highly dependent upon them is extremely important, but also underlining the point that a scavenger could find much of value on a partly-eaten carcase.

African wild ungulates, even when in prime condition, average no more than 4% carcass fat content, a figure that would only occur in “stressed or even starving ungulates in temperate and northern latitudes” where fat content of carcases tends to be higher, according to Speth (1989). It should perhaps be said that precise figures for such comparisons are rather difficult to obtain because of variations in methods of assessment, and the matter is perhaps not as clear-cut as Speth implies (C. Pond, pers. comm.). However, a comparison of the original African data provided by Ledger (1968: Appendix II), and quoted by Speth, with that available for temperate-zone species (Pond, 1978: Table 2) suggests that Speth’s interpretation is broadly correct. More intensive exploitation of ungulates as a food source would therefore have been possible for hominids in Eurasia in comparison with African if Lower Pleistocene European taxa shared the higher body fat levels of their modern counterpart.

Questions about the actual levels offood availability represented by the ungulate fauna of Europe need not detain us. Estimates of biomass for palaeoecosystems are fraught with difficulties and probably offer no more than a spurious air of precision. The long-term diversity in taxa and body size of the European PlioPleistocene ungulate community, and

112 A.TURNER

Table 1 Net primary production for some world ecosystems expressed in grams per square metre per year’

Vegetation unit Mean Range

Tropical grassland 900 200-2000 Temperate grassland 600 200-1500 Tundra and Alpine 140 1wN?O

‘After Whittaker and Likens (1975: Table 15-1).

the range of predators that exploited it, in themselves afford sufficient indication that a substantial food supply was available to a meat eater. Of course diversity, whether among ungulates or predators, if not a measure of biomass, and should not be considered a simple reflection ofit. Indeed, high diversity and high biomass among modern larger ungulates tend to be mutually exclusive, except in areas where a range ofhabitats is available (Andrews et al.,

1979). The very diversity of the European ungulate fauna points to such a range of habitats, including temperate grasslands and woodland of the kind implied by the palynological data (Sue & Zagwijn, 1983; Zagwijn & Sue, 1984), and temperate grasslands are among the most productive of modern-day ecosystems (Table 1).

Any successful exploitation of animals is likely to have been dependent on the availability of tools as a means of getting through the thick skin of larger species, for detaching body parts and for breaking bones. The use of stone tools in Africa from at least 2-O Ma, perhaps even 2.4 Ma (Harris, 1983), suggests that there was no inherent technological impediment to dispersion into temperate areas from that time so far as food acquisition was concerned. However, the requirements for clothing and shelter, as well perhaps as the necessity for fire, may have acted as tighter constraints on movements into such regions.

Scavenging

Previous rejections of the likely importance of scavenging to hominids, based on studies of carcases in East Africa (Schaller & Lowther, 1969), employed the argument that it was inevitably an erratic source of food and therefore likely to have been of little overall significance. This conclusion was seriously undermined in the studies undertaken by Kruuk (1972) on the ecology of the spotted hyaena, Crocuta crocuta, where it became clear that scavenging could provide a substantial proportion of the diet to a suitably organized carnivore. Blumenschine (1986, 1987a,b, 1989) has now provided much more detailed information about the potential of scavenging from an anthropological perspective. He underlines the importance of seasonality, habitat, carcase size and predator behaviour in the formation of a scavangeable resource, and stresses that the structure of the larger carnivore community, especially the ratio of flesh-eaters to carcase-destroyers, is one of the most significant determinants of availability of carcases on the ground. Scavenging, according to Blumenschine, may indeed be an erratic source of food, but the circumstances in which it is feasible are predictable. We may deduce that the availability of carcases in Pleistocene Europe would have been conditioned in large part by the structure of the carnivore community, and changes in that structure would have therefore have affected the scale of scavenging

. . opportumtles. If colonizing hominids were largely dependent on scavenging, then resource


availability would have impinged directly on the success of that colonization and therefore on the pattern of occupation.

Scavenging as a potential source of food is one thing: scavenging as a demonstrable strategy is another, and there has been much archaeological concern in recent years with the question ofwhether early hominids hunted or scavenged. The view that hunting was a major feature of early hominid lifestyle has a long tradition, but it is an assumption which has largely been based on the association of stone tools and animal bones rather than on any clearly articulated criteria. So far as the function of those tools is concerned, Higgs & Jarman (1972:9) pointed out several years ago that “many industries which (by archaeological hypothesis) relate to hunter-gatherer economies [are] without unequivocal hunting equip- ment”, while “industries which are most prominent in possession ofsuch items are frequently chronologically late and apparently connected with farming economies”. Binford (1981, 1984, 1985, 1988) has employed comparative studies ofskeletal part frequency and damage patterning to mount a forceful argumsent against regular hunting prior to the later stages of human evolution, and although some of his claims for the ubiquity of scavenging may be questioned (Bunn & Kroll, 1988; Turner, 1989) it is clear that the image of”man the hunter” has been permanently tarnished. In so far as any consensus can be identified among those who have examined early hominid-associated assemblages, it would seem that scavenging is considered to have been a significant factor in their formation, based on distributions ofbody parts, carnivore toothmarks and hammerstone percussion, and that most disagreement centres on whether scavenging was active and confrontational or passive (Shipman, 1986; Bunn, 1986; Bunn & Kroll, 1986; Blulmenschine, 1991). In contrast, there seems to be little direct evidence to support the notion ofhunting.

Changing patterns in the mammalian fauna of Europe

The term “Villafranchian”, commonly used for the later Pliocene and earlier Pleistocene mammalian fauna of Europe, derives from the Italian fossil-bearing locality of Villafranca d’Asti. Although it is not strictly a chronostratigraphic term, its use may be taken to cover the period 3.2-0.9 Ma (Kurten, 1968; Azzaroli, 1977, 1983a; Azzaroli et al., 1988). Post- Villafranchian faunas of the earlier Middle Pleistocene are often referred to as Galerian, a name first proposed by Ambroseti et (21. (1972). These terms will be employed as a useful shorthand in the discussion.

Fauna1 turnover

The larger mammal fauna of Europe for the last 3.0 Ma is shown in Tables 2-5. A major turnover in species has long been recognized at the end of the Villafranchian in Europe (Kurten, 1963, 1968; Kahlke, 1975; Maglio, 1975; Azzaroli, 1977, 1983a; Azzaroli et al.,

1988; Aguirre & Morales, 1990), although the absolute chronology has been unclear until relatively recently. Azzaroli (1983a) placed the transition to the Galerian fauna between 1.0-0.9 Ma, based on the sequence of mammalian faunas in Italy, and referred to it as “a turning point in the history of Eurasia”.

The most obvious result of the end-Villafranchian event is the eventual Middle Pleistocene appearance of familiar species such as Rangayer tarandus, Cervus elaphus, Megaceros giganteus,

Bison priscus, Bos primigenius, Elephas antiquus, Mammuthus primigenius, Coelodonta antiquitatis,

Canis lupus, Crocuta crocuta and Panthera Leo. More recent work has drawn attention to other

114 A. TURNER

Table 2 European VJllafranchian-Galerian fauna. Proboscidea, Perissodactyla, Hippopotami&e and Suidae 6

3.0 2.5 2.0 1.5 1.0 0.5 0 AGE Ha 1....0....1....0....1....0....1

MAMWJTIDAE Mammut borsoni GOMPHOTHERIIDAE Anancus arvernensis ELEPNANTIDAN Mammuthus meridionalis Mammuthus m. tanamensis Mammuthus armeniacus Mammuthus primigenius Elephas antiquus TAPIRIDAE Tapirus arvernensis RNINOCEROTIDAe Dicerorhinus jeanvireti Dicerorhinus etruscus Dicerorhinus hemitoechus

? ? ? ? ---- -----

-__---_____-______

7

Dicerorhinus kirchbergensis Coelodonta antiouitatis EPDIDAe Hipparion sp. Equus livenzovensis ___~ Equus stenonis Equus bressanus Equus stehlini Equus sussenbornensis Equus altidens Equus hydruntinus Equus ferus HIPPOPOTAMIDAN Hippopotamus antiquus SUIDAE Sus minor --- ? ? ?

Sus strozii sus scrofa

? ? -----____-_

AGE Ma 1 3.0' . .205

7

???????????

? ? --------- ? ? ? ? ---------

----? ? ? ? ? ? ? -____________

. . 1....0....1....0....1 2.0 1.0 1.5 0.5 0

Sources: Alberdi & Bonadonna (Eds) (1989); Azzaroli (1977, 1983a, 19836); Azzaroli et al. (1988); Aguirre (in lit.); Bruning (1980); Forsten (1988); Giuli (1986); Giuli & Masini (1986); Giuli et al. (1986); Heintz (1970); Heintz et al. (1974); Kahlke (1975); Lister (1986, 1988); L umley, et al. (1988); Masini & Lovari (1988); Terre (in lit).

turnover events in the Villafranchian, again based in large part on the Italian evidence but with details from other areas also brought into the discussion (Torre, 1987; Azzaroli et al.,

1988; Aguirre & Morales, 1990; Masini & Torre, 1990; Turner, in press). At around 1.7 Ma the “wolf event (Azzaroli et al., 1988; Masini & Torre, 1990)) named after the appearance of Cunis etruscus, is marked by the appearance of the Olivola fauna of Italy (Azzaroli, 1977, 1983a; Azzaroli et al., 1988), and appears to correlate with the beginning of the Pleistocene as defined at the Vrica section. An earlier change at 2.5 Ma, termed the Elephant-Equus event, is marked by the disappearance of several archaic elements and by the appearance of a range of species that includes the first European members of the genus Equus. In Italy, this new fauna is recorded in the Montopoli fauna1 unit (Azzaroli, 1977, 1983a). All of these fauna1 events appear closely correlated with the major climatic changes of the Plio-Pleistocene outlined above (Azzaroli et al., 1988).

The mammalian fauna of Europe therefore changed markedly over time. The fauna represented in Tables 2-5 is, of course, an amalgamation from a large area, but then it seems that most of the larger species were widely distributed and provincial faunas within the continent are hard to identify. This is shown by the Lower and Middle Pleistocene

LARGE CARNIVORESANDEARLIESTEUROPEANHOMINIDS 115

Table 3 European ViUafranchian-Chlerian fauna. Cetvidae. Sources as for Table 2

3.0 2.5 2.0 1.5 1.0 0.5 0 AGE Ma 1....0....1....0....1....0....1

Cervus pardinensis Croizetoceros ramosus 7 Cervus perrieri Arvernoceros ardei Cervus rhenanus ? ?-__-_? 7 ? Dama nestii Dama dama Eucladoceros falconeri Eucladocercs tegulensis Eucladoceros dicranios Eucladoceros tetraceros ? ? ? ? ? ? ? ? ?_-___--__--__- Cervus acoronatus Cervus elaphus Meqaceros verticornis Megacer0s solilhacus ? -----~_ Megaceros savini Meqaceros gigantsus Cervalces qallicus Cervalces latifrons Alces alces capreo1us caprec1us Ranqifer tarandus

AGE Ma 1.. ..o.... l....o.... l....O....l 3.0 2.5 2.0 1.5 1.0 0.5 0

Table 4 European Villafrancbian-Calerian fauna. Bovidae. Sources as for Table 2

AGE Ma

Pliotraqus ardeus Hesperoceras merlae Gazella borbonica Galloooral meneahini Gazeliospira to;ticornis Leptobos stenometopon Leptobos meslai Leptobos etruscus LerXobos vallisarni Edbison sp. Bison schoetensacki Bison priscus Bos trocheros Bos primigenius Bubalus murrensis Praeovlbos priscus Ovibos mozchatus Meqalovis latifrons Ovis musimon Capra ibex Procamptoceras brivatense Rupicapra rupicapra Salga tatarica Hemitragus bonali Soerqelia elizabethae

3.0 2.5 2.0 1.5 1.0 0.5 0 l....o.... l....O....l....O....l

? ? -----

~_________? ? ? -_____________

AGE bia 1....0....1....0....1....0....1 3.0 2.5 2.0 1.5 1.0 0.5 0

assemblage lists drawn up by Kahlke (1975). A zzazoli et al. (1988) mention only the absence of the cervid genus Cervalces from Spain (while pointing out that it has only recently been recorded in Italy) and the apparent absence of the genus Bos from higher latitudes. The Carnivora, as might be expected from an order ofsuch generalists, were ubiquitous. Kurten (1968) observed that the dirk-toothed cat Megantereon cultridens appeared to be broadly

116 A.TURNER

Table 5 European Villaframhian-Galerian fawta. Larger Carnivora

3.0 2.5 2.0 1.5 1.0 0.5 0 AGE Ma 1....0....1....0....1....0....1

FELIDAE Acinonyx pardinensis Panthera gombaszoegensis Panthera lea Panthera pardus ? ? ?_---_-----------_--- Homotherium latidens Megantereon cultridens EYAJZNIDAg Pachycrocuta perrieri Pachycrocuta brevirostris Chasmaporthetes lunensis crocuta crocuta CANIDAF, Canis etruscus Canis arnensis 3 Canis falconeri 7 Canis mosbachensis Canis lupus ?---- Nyctereutes megamastoides --~---~------~~---~---~---~ ? Xyenocyon lycaonoldes Cuon alpinus ? ? __________~__

URSIDAE Ursus minimus ursus etruscus __~-__--__~-__-__--__~--------- Ursus deningeri ursus spelaeus --____ ursus arctos Ursus thibetanus

AGE Ma l....o.... l....O....l....O....l 3.0 2.5 2.0 1.5 1.0 0.5 0

Sources as in Table 2 plus Kurten & Poulianos (1977, 1981); Masini & Tome (1990); Pens-Moya (1987) and direct observation.

circum-Mediterranean in its European range, but it is worth stressing that the range includes sites in the Massive Central such as Seneze and Les Etouaires, as well as central and eastern Europe. Indeed, its overall range was from southern Africa to northern America (Turner, 1987).

Only a very specific requirement, as in the case of hippos and water, seems to have produced clear-cut geographical restrictions, but even then the pattern is essentially intra- regional (R. D. Kahlke, pers. comm.). However, some local partitioning of fauna1 elements must have taken place; the most obvious would have been between more open and more closed habitats. Seasonal movements would also have occurred, and the climatic cycles that intensified during the Middle Pleistocene would themselves have produced a degree of segregation between faunas of colder and warmer periods. But the cold-tolerant fauna of the glacial periods that one inevitably associates with human activity in Europe is a phenomenon of the later-Middle and Upper Pleistocene (and even then reindeer are recorded from Spain). The Villafranchian fauna of Europe is less differentiated in this way, and its changes are more chronological. Overall, it would seem that for much of the tim.e many of the larger ungulates were present in any given region.

Hominids equipped with a lithic technology would therefore have encountered a significantly different range of ungulate species depending on the time of their arrival rather than on the region of settlement. Those differences alone might have affected the success of colonization attempts to some extent, although we have no direct way of assessing the impact of change in the ungulate fauna on resource exploitation. Hominids would also have encountered a variety of large predators equally interested in the food resources represented


Table 6 Larger Camivora of eastern Africa from later Pliocene. Data from Turner (1990)

Felidae Hyaenidae Canidae

At 2.0 Ma Homotherium latidens Megantereon cultridens Dinofelispiueteaui Panthera lea Panthera perdus AcinonyxJubatus

After 1.5 Ma Panthera lea Panthera pardm Acinonyxjubatus

crocuta crocuta Hyaena hyaena

Crocuta crocuta Hyaena hyaena

Canis mesomelas Lycaon pi&s

Cams mesomelas Lycann pi&s

by the ungulate species. Changes in the ungulate fauna over time would in turn have affected the larger carnivores, producing a complex, shifting pattern of resource availability.

The guild of larger carnivores

Olle of the most notable features of the Pleistocene fauna of Europe lies in the size and structure of the larger carnivore guild. The guild, which is shown in Table 5, includes the Ursidae for completeness, but these are not considered further in the following discussion since their role in carcase production and destruction is likely to have been minimal in comparison with the other species present.

The nature of the European guild is highlighted if we compare it with that of eastern Africa, the likely point of origin of the European hominids and the scene of much of the modern comparative work on predator-prey behaviour. By ca. 1.5 Ma in eastern Africa the guild consisted of the extant species (Turner, 1990) (Table 6), whereas in Europe during the same period it was composed almost entirely of archaic taxa (Table 5). Only by ca. O-4 Ma did the structure of the European carnivore guild come to resemble that of Africa over the past 1.5 Ma. In the intervening period the European carnivore community expanded markedly (Figure 1) as species invaded, probably from Africa in the case of the lion (Punthera

Leo), leopard (Panthera pardus) and spotted hyaena (Crocuta crocuta) (Turner, 1990, in press) and from Asia in the case of the large European hunting dog (Xenocyon lycaonoides) and smaller dhol, Cuon alpinus (Sher, 1986). At ca. 0.7 Ma the situation was further complicated by the reappearance in Europe of the large hyaena, Pachycrocutaperrieri, not seen in the area since the end of the Pliocene (Table 5).

By the Middle Pleistocene the ratio of carcase destroyers to flesh eaters in Europe had settled at unity, the same as that of eastern Africa since the beginning of the Pleistocene [Figure 1 (C)l. However, at the same time the flesh-eating component of the community in Europe continued to include the machairodont cats Homotherium latidens and, at least until ca. 1.0 IMa, Megantereon cultridens, species formerly present in eastern Africa but extinct by 1.5 Ma (Turner, 1986, 1987, 1990). The continued presence of the archaic cats is extremely significant, since although the number of carcase destroying species in Europe grew in comparison with eastern Africa, the importance of the purely flesh-eating carnivores as sources of partly-consumed carcases was enhanced by the activities of the machairodonts.

118 A. TURNER

Figure 1. Numbers oflarger carnivore guild species in Europe over the past 2.0 Ma. A, Hyaenidae; B, larger Canidae; C, Machairodontinae; D, larger Felinae; E, flesh eaters (machairodontine and feline cats); F, carcase destroyers (hyaenas and larger canids); G, ratios of carcase destroying species: flesh eaters for Europe (--) and eastern Africa (- - -).

Ewer (1954) was one of the first authors to point out that the specialized dentition of the machairodont cats would have prevented them from eating all but the most easily accessible portions of flesh on a carcase, and she stressed the likely quantities of food that might thus be made available to a scavenger (Ewer, 1967). This point was made in a recent discussion of scavenging potential for eastern and southern African hominids (Turner, 1988), and has been repeated recently at length by Marean (1989). It is likely that between 1.5-l .O Ma in Europe, a significant proportion of the killing of larger ungulates would have been undertaken by H. latidens and M. cultridens, the former the size of a modern lion and the latter the size of a large leopard. Both were powerful animals (Ballesio, 1963; Schaub, 1925)) and were probably able to take larger prey than the cheetah, Acinonyx pardinensis, and the jaguar, Panthera gombaszoegensis. The latter was larger than the extant jaguar of the New World (Simpson, 1941; Kurten, 1965, 1973; Table 7), but although it was evidently a stocky, powerfully built animal, it had a typically pantherine dentition lacking any specialized bone- destroying abilities. The European cheetah was also a considerably bigger animal than the extant African variety (Ficcarelli, 1984), Table 7, but it had the somewhat specialized, sectorial dentition ofits modern counterpart and was equally unable to consume bone of any but the smaller ungulates. Indeed, the cheetah is likely to have been restricted to smaller,


Table 7 Sizes of fossil and modem cheetahs and jaguars based 0x1 length of lower camassial

n Mean+S.E.M. Range C.V.

Acinonyx pardinensis (various European localities) Acinonyx_iubatus (southern Africa) Panthera gombaszoegensis (various European localities) Panthera onca’ (South America)

10 21.1 kO.45 19.2f23.8 6.7

51 18.5_fO.11 17.0-21.3 4.3

17 24.2 k 0.45 21.0-27.7 7.6

8 21.2f0.35 20.0-24.0 5.9

‘From Simpson (1941:12). S.E.M., standard error of the mean; C.V., coefficient ofvariation.

although perhaps faster moving prey, than the jaguar or either of the machairodont species. It is therefore clear that during much ofthe Lowerand Middle Pleistocene the structure ofthe European carnivore guild would have provided high quality carcases to any scavenger species. The possible appearance of the leopard as early as 1.2 Ma at sites such as Venta Micena in Spain (Pons-Moya, 1987) and Ceyssaguet in the Massive Central (A. Turner, unpublished data) would have added to that potential resource.

After 1.0 Ma the number of flesh-eating, and therefore carcase-producing, species rose with the incursion of the lion added to the already present leopard, seemingly balancing the loss of M. cultridens. European lions were large (Kurten, 1985; Turner, 1984), and the size of prey that can be taken by lions acting in concert (Schaller, 1972) may have increased the average size of carcases, thereby augmenting the potential food available to a scavenger. But any such increase in the food supply was met by a consistently high ratio of bone destroying dogs and hyaenas (Table 5, Figure 1)) and the parallel incursion of the spotted hyaena alone is likely to have had a significant effect on carcase availability. Although long unjustly vilified as a mere scavenger unable to hunt for itself, the spotted hyaena is indeed a prodigious consumer of bone and a pack is capable of consuming an entire adult zebra (Kruuk, 1972).

As Table 5 and Figure 1 show, the size and nature of the European larger carnivore guild changed dramatically at around 0.5 Ma, leaving only the leopard, lion, spotted hyaena and the wolf as the major large predators. Although the ratio of carcase destroyers to flesh eaters remained the same at unity, the details ofthe structure had in fact changed far more dramatically than this figure alone suggests. The changes involved the loss of three large cats, including most significantly H. latidens, as carcase providers, and the disappearance of the hunting dog and two large hyaenas. The reasons for these losses are undoubtedly complex and interlinked, but are also likely to have been casually related to the changes in the ungulate fauna. As Azzaroli (1983a) has pointed out, events towards the end of the Villafranchian include the major appearance of heavier bodied bovids of the genera Bison

and Bos, which replace the more lightly-built members of the genus Leptobos, and the appearance oflarge-bodied cervids of the genus Megaceros. Such increases in average size are unlikely to have been to the advantage of the larger predators, since prey size is a major determinant of hunting success and efficiency. Large prey are more difficult to kill, while smaller animals

120 A, TURNER

hardly repay the effort. The best size is at or about the body size of the predator, unless group huntingisadopted (Pienaar, 1969;Mech, 1970; Schaller, 1967,1972;Kruuk, 1972; Sankhala, 1978). However, large ungulates do mean the eventual availability of large carcases.

Discussion

From Table 5 and Figure 1 it might seem that it would have been advantageous for scavenging hominids to have attempted to colonize Europe l-5-I.0 Ma. With two machairodont cats plus the cheetah, the jaguar and perhaps the leopard killing prey and producing carcases, and only one hyaena species and two dogs in contention for the remains, the scope for a scavenging hominid to enter the scene appears to have been considerable.

Against this must be set the reaction of the incumbent scavengers. The populations of these species are likely to have expanded as a result of the availability of food, an effect seen by Kruuk (1972, 1975) in his studies ofspotted hyaenas and lions in Tanzania. Ifthis happened, then intense competition for food may have followed as the population numbers increased, leading to few carcases remaining uneaten for very long, as Kruuk found in the Ngorongoro Crater. The largest and strongest of the predators, or those able to amass the greatest numbers at carcase sites, are then likely to have fared best. Two other events recorded in Table 5 and Figure 1 lend weight to this interpretation in the fossil context. The first is the extinction of the cursorial hyaenid species Chasmaporthetes lunensis, unknown in Europe later than the Olivola fauna in which Pachycrocuta brevirostris first appears. With its relatively gracile dentition, C. lunensis is likely to have been less effective as a scavenger than most other hyaenas, but it is widely recorded in Eurasia and was an apparently successful predator (Kurten & Werdelin, 1988). However, it was unable to survive the end of the Pliocene. The second, and perhaps the more significant event, is the local disappearance of the other large hyaenid Pachycrocuta perrieri, also last seen at Olivola. This species re-immigrated during the earlier part of the Middle Pleistocene, but its absence for ca. 1-O Ma came after it had become well-established in Europe is noteworthy. Pachycrocutaperrieri, which may have been no more than a larger variant of the extant brown hyaena, Hyaena brunnea (Turner, 1990), was dentally equipped for bone destruction on a considerable scale, as may be seen in the excel- lent illustrations of the specimens from St Vallier provided by Viret (1954). The even larger P. brevirostris is aptly referred to as a gigantic species of lion size (Boule, 1983; Kurten, 1968)) as attested at the later Lower Pleistocene site of Untermassfield, Germany, where its bone consuming abilities are clearly evident in the quantity ofcoprolites and damaged bones (Turner, in prep.). The basal skull length of the type specimen of P. brevirostris, from Sainzelles in the Auvergne, is 322 mm (Boule, 1893) and may be compared with a mean of 294 mm (range 242-328 mm) for a sample of 19 southern African male lions (Turner, unpublished data). The Sainzelles specimen is clearly not an outlier in the size range, as comparison with dental measurements from other specimens clearly shows (Turner, in prep.). A clear parallel for the possible exclusion of P. perrieri by P. brevirostris may be seen today in the distributions of the spotted hyaena, which occupies much of the sub-Saharan area of Africa, the entirely southern African brown hyaena and the eastern and northern African striped hyaena (Haltenorth & Diller, 1980). According to Mills (1990) the large spotted hyaena generally manages to displace the smaller brown species in areas where they overlap, and has a similar effect upon the even smaller striped hyaena in eastern Africa (Kruuk, 1972, 1975). If the disappearance of C. lunensis and P. perrieri do indeed point to the difficulties encountered at the beginning of the Pleistocene by two hyaena species, at least one of which was a well-


adapted scavenger, then it is difficult to see why things should have been easier for incoming hominids.

Despite the subsequent incursion of the lion and the certain presence of the leopard, the structure of the larger carnivore guild after 1.0 Ma does not offer any clear improvements from the point of view of potential hominid colonists. Although the cheetah and the sabre- toothed cat H. latidens were still active in Europe, and the lion and the leopard would have provided scavengeable carcases, the loss ofMegantereon is likely to have reduced the frequency with which carcases could be found with only their more fleshy portions consumed. More- over, the appearance of the spotted hyaena and the subsequent reappearance of P. perrieri

may have more than offset the effect ofcarcase provision by the cats, even with the addition of the lion. It is true that the spotted hyaena and P. breuirostris do not frequently occur in the same deposits, while P. brevirostris and P. perrieri, when the latter re-appears, seem to have been mutually exclusive (Kurten & Poulianos, 1977, 1981). P.p errieri and the spotted hyaena are recorded together at a number of sites, such as Mosbach (Bruning, 1980), Petralona (Kurten & Poulianos, 198 1) and L’Escale (Bonifay, 197 1; Turner, 1990), pointing to the overall complexity of the relationships between the hyaenas of Pleistocene Europe. Even if they all operated allopatrically, and setting the presence of large canids on one side, the existence of two, and later three, such large carcase-destroying hyaenas in the earlier Middle Pleistocene implies stiff competition for ungulate remains until around 0.5 Ma when P.

brevirostris and P. perrieri, together with A. pardinensis, P. gombaszoegensis and H. latidens,

became extinct. In contrast, the dramatic change at 0.5 Ma left a large carnivore guild in Europe

essentially similar in overall structure to that of eastern Africa since the beginning of the Pleistocene. Stressed here must be the fact that there is similarity rather than precise identity between the two faunas, since the cheetah and the striped hyaena continue to exist in Africa and the canids differ. The similarity in structure to the eastern African guild is important because Blumenschine (1986,1987a,b, 1989) has shown that carcase availability to a hominid there today can be considerable when there is a seasonal glut through die-off in the dry season, or in woodlands where hyaenas scavenge less (Kruuk, 1972). In Blumenschine’s scheme, confrontational scavenging employed at other times of the year would be necessary to ensure access to carcases, but if this were adopted then virtual dependence on scavenging would be possible in present conditions in the region. On this basis, and considered purely in terms of the structure of the carnivore guild, only after 0.5 Ma do opportunities for scavenging by hominids appear to have been at their best in Europe.

Summary and conclusions

The whole of the Pleistocene period in Europe, as well as the latter part of the Pliocene, has been characterized by conditions that were on average colder and probably drier than today. In such circumstances, and without the technology to permit storage of vegetable products throughout the year, animals are likely to have been the most important single food resource for hominids for most of the time provided that fats were included in the diet.

The composition of the European mammalian fauna between 1.5-0.5 Ma was notable for the continued existence of machairodont cats. Such animals were incapable of consuming more than the most fleshy portions oftheir prey because of the highly derived, sectorial nature oftheir cheek teeth, and would haveleft aconsiderable quantity ofscavengeable food at a kill.

122 A. TURNER

The presence of the large hyaena P. brevirostris, the extinction of C. lunensis and the parallel disappearance until the Middle Pleistocene of the other large hyaena P. perrieri, suggest that competition for carcases was marked and that the scale of scavenging opportunities for incoming hominids may have been small. Between 1 .O and perhaps 0.5 Ma the situation may have deteriorated further, from the hominid perspective, as M. cultridens became extinct and the carcase-destroying component of the carnivore fauna expanded with the immigration of the spotted hyaena and the re-appearance of P. perrieri. Only after 0.5 Ma do we see a shift towards the structure of the modern-day carnivore guild of eastern Africa and only then did the pattern of resource availability in Europe, judged in terms of the composition of the predator community, improve for the hominid occupants.

The structure of the mammalian community in the period 1.5-1.0 Ma may have encouraged colonization attempts that simply could not be maintained, leading to a patchy record that will remain frustratingly difficult to interpret, if considered in isolation from events affecting the rest of the terrestrial biota. The structure of the carnivore guild therefore implies that the existing interpretation of the archaeological record may be broadly correct. It also offers the interesting conclusion that conditions prior to 1.5 Ma, when the flesh eating component of the carnivore guild was dominated by machairodonts and before P. brevirostris

appeared in Europe, may have been even better for a scavenger (Table 5). This would imply that there is at least no evident palaeoecological barrier to claims for later Pliocene hominid presence.

The framework provided by the fossil evidence permits us to go beyond merely supporting an existing interpretation of the archaeological record and to test that interpretation by predicting something of the course of future discovery. First, it implies that O-5 Ma will probably be the time at which an increasing density of more permanent hominid occupation will be seen to have occurred. Before that time, occupation is likely to have been much more sporadic, at least back as far as 1.5 Ma. Secondly, it implies that any discernible patterning in the evidence for hominid occupation prior to 0.5 Ma should relate to patterning in the availability of resources, perhaps in the form of congruent distributions of archaeological evidence and major carcase producers. The suggestion made by Kurten (1968) for a circum- Mediterranean distribution of M. cultridens might therefore warrant closer scrutiny, and claims for earliest hominid appearances in Spain, southern France and Italy (Bonifay & Vendermeersch, in press) might prove to make palaeoecological sense, for example. It is also clear that the distribution of vegetational cover during the Pleistocene would have favoured southern areas (Zagwijn & Sue, 1984), and the increased availability of unconsumed carcases in such areas might have been a further factor in the distribution of hominid activities.

Thirdly, the emphasis placed by Blumenschine (1989) on the importance of confrontational scavenging raises a significant point. The discussion presented here has dealt with changing resource availability in the context of essentially passive scavenging by hominids. Active, confrontational scavenging would obviously have the potential to change the scale of availability at any given time. However, it should also result in a different pattern of body part frequency and damage in the fossil record, as Blumenschine (1987a,b, 1989) has stressed, since the timing of access to a carcase, and its consequent condition, would be altered. In principle, it should therefore be possible to distinguish the two types of scavenging in assemblages, to observe changes in the frequency of each type, and in consequence to monitor any developments that may have taken place. This raises a clear challenge for future work on appropriate fossil assemblages to integrate the study of damage and body part frequency


with a fuller understanding of the opportunities for resource exploitation reflected in the structure of the larger carnivore guild.

Acknowledgements

An earlier version of this paper was presented at the LOBRA ‘87 meeting on earlier European hominid occupation, held in Milan in 1990. I thank G. Cadeo for making attendance at the meeting possible. I am grateful to the following people for access to material and/or for discussions on the topics mentioned here: E. Aguirre, R. Ballesio, E. Bonifay, M.-F. Bonifay, G. Ficcarelli, J. Franzen, A. Currant, H.-D. Kahlke, R.-D. Kahlke, A. Lister, F. Massini, J. Morales, M. Phillipe, G. Petter, D. Torre. I also thank P. Andrews, R. Blumenschine, J. D. Speth and an anonymous referee for constructive comments on an earlier draft of the current paper. The research was supported by a grant from The Leverhulme Trust.

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