valente 2004 - humans and carnivores in the early upper paleolithic in portugal - data from pego do...

8/8/2019 Valente 2004 - Humans and Carnivores in the Early Upper Paleolithic in Portugal - Data From Pego Do Diabo Cave

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ISSN 0253-6730

1 Universidade do Algarve, Campus de Gambelas, FCHS, 8000 Faro, Portugal, Email : [email protected]

Revue de Palobiologie, Genve (dcembre 2004) 23 (2): 611-626

Humans and Carnivores in the Early Upper Paleolithic in Portugal :

Data from Pego do Diabo Cave

Maria Joo VALENTE1

AbstractDuring the Upper Pleistocene human hunters had to compete with large carnivores in their ecological setting. This paper analyses thefauna from the small cave of Pego do Diabo, with an occupation classied as Aurignacian, and considers human-carnivore relationshipsin the Portuguese Upper Paleolithic. Similar to other Early Upper Paleolithic cave deposits in Portugal (Salemas, Caldeiro, Casa daMoura and Buraca Escura), Pego do Diabo faunal assemblage includes remains of several carnivore species, such as wolf, hyena andiberian lynx.Some evidences suggest important changes in the relationship between humans and carnivores at the end of the last glacial in PortugueseEarly Upper Paleolithic sites : rst, the relative abundance of carnivore remains, associated with a high degree of carnivore-modiedbones ; second, a signicant reduction in carnivore species diversity during the Solutrean and Magdalenian. A model is developedemphasizing short and discontinuous human occupations of small cavities during the Early Upper Paleolithic, which left these sites freefor carnivore denning. These patterns changed during the Solutrean, as large carnivores started to disappear from the archaeologicalrecord and sites containing signicant numbers of carnivore-damaged fauna begin to be rare.

Key wordsUpper Paleolithic, Portugal, Carnivores, Ecology.

INTRODUCTION

In the past decades, many researchers have noted thatduring the Upper Pleistocene human hunters had to

compete with carnivores in their ecological setting,frequently overlapping in the use of space and inforaging strategies (e.g. BINFORD, 1981 ; BLASCO, 1997 ;BLUMENSHINE, 1995 ; BLUMENSHINE & MAREAN, 1993 ;BRANTINGHAM, 1998 ; BRUGAL & JAUBERT, 1991 ; BRUGALet al., 1997 ; FOSSE, 1995 ; HOCKETT, 1993 ; STINER, 1994 ;STRAUS, 1992). This paper discusses the faunal remainsfrom the cave Pego do Diabo (in English meaning deephollow of the Devil). The site contains a short humanoccupation classied as Aurignacian by J. ZILHO (1997),and affords an opportunity to analyse human-carnivorerelationships at early Upper Paleolithic Portuguese sites.

Despite not being numerous, Pego do Diabos faunalcollection shows specic taphonomic features that agreewith other Pleistocene deposits in small cave sites withEarly Upper Paleolithic assemblages (like Caldeiro,Salemas, Cova da Moura and Buraca Escura). Dataindicate that a relatively large number of carnivorespecies were in direct competition with human groupsat this time, not only for the use of space but also for thesame animal resources.

BACKGROUND, STRATIGRAPHY AND DATING

Pego do Diabo Cave (9136W, 385152N) is a smallcavity located in the Portuguese Estremadura about

20 km north of Lisbon (Fig. 1). The site stands in aTuronian limestone ridge that lies on the southern slopeof the Loures River Valley, near its conuence with theLousa River, at 250 m a.s.l. From the cave we have a vastview shed of the surrounding area.The cave has two sections : a narrow passage about13 m long and a room at the end measuring about 2 mwide (Fig. 2). Bedrock can be seen at the entrance and ismarked by a distinct depression toward the back of thecave. In this area there was a well-preserved sedimentarysequence about 1.4 m deep.Pego do Diabo was excavated in 1976 by the Grupo para

o Estudo do Paleoltico Portugus (GEPP, 1979) and inthe late 1980s by ZILHO (1988 and 1997). In total, anarea of 11 m2 has been opened, representing about half ofthe deposits presently accessible. In 1993, J.L. CARDOSOpublished a paleontological study on the PleistoceneLarge Mammals of Portugal, in which he analized asmall sample of Pego do Diabo faunal assemblage(only elements adequate for osteometric data) ; and in1995, J. ZILHO nished is Ph.D. thesis, which includeda complete study of the lithic assemblage (published in1997).

Actes du XIVe Congrs UISPP Hommes & Carnivores au Palolithique , Lige 2001,BRUGAL, J.P. & P. FOSSE(Eds).


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The stratigraphic prole (Fig. 2) shows the 6 unitsrecognized by ZILHO (1997, 2 : 94-95). The top layer(Level A) was only found at the back of the cave ; ityielded historical material and has a probable funerary

occupation. Level 1 contains a mixture of modern andPleistocene material (including fauna) that representa redeposition of level 2. Level 2 comprises UpperPaleolithic artefacts in association with bones. Level3 also has Pleistocene fauna, and displays a distinctcoloration, probably made by manganese oxide. At thebottom of the deposit, level 4 has rare archaeologicalmaterial and level 5 contains no material at all.The faunal analysis and the results presented here arefrom level 2. We also included in the study all theremains from level 1 that exhibited the same surfacecharacteristics of level 2 (colour and fossilization).

The four radiocarbon dates obtained are shown in Table 1.Based on laboratory information, the two younger dateswere rejected : in the ICEN-306 date of level 2, recentcarbon from the top layer probably contaminated thesample ; in the ICEN-491 date of level 3, there was aprobable contamination incident or a decient collagenextraction resulting in an impure sample (ZILHO, op.cit. : 98).The accepted radiocarbon dates (ICEN-490 and ICEN-732) and the lithic analysis, especially the six Dufourbladelets, led J. ZILHO (1997) to classify level 2 asAurignacian (see Note 1). The 28.000 years BP datewas interpreted as representing the age of the human

occupation at the base of level 2, and the 23.000 yearsBP date was considered the end of a slow depositionsequence (or a contaminated sample). Consequently,levels 3 and 4 are expected to be Middle Paleolithic, evenif the scarce lithic industry (n = 6) has no diagnostic tools(ZILHO, op. cit. : 95).

THE PLEISTOCENE FAUNA AT PEGO DO DIABO

CAVE

A very diversied Pleistocene fauna was found in the

cave, including small animals (such as birds, smallrodents, bats and amphibians), medium mammals

Fig. 1 : Approximate locations of the main archaeological sitesmentioned in the text. (1) Pego do Diabo Cave, (2)Salemas Cave, (3) Casa da Moura Cave, (4) Lapa doPicareiro, (5) Caldeiro Cave, (6) Buraca Escura.

Table 1 : Radiocarbon dates for Pego do Diabo Cave (ZILHO 1997, 2: 94-95).

Provenience Level Sample Lab No. Date B.P.

M13 2 Charcoal ICEN-306 2.410 +/- 80M11 + L11 2a Macrofauna ICEN-490 23.080 +/- 490

M11 + L11 2b Macrofauna ICEN-732 28.120 - 780 + 860M13 + M14 3 Rabbit and macrofauna ICEN-491 18.630 +/- 640

612 M. J. VALENTE


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(mustelids and leporids), and large mammals. Excludingmicrofauna and birds, fourteen taxa were identied(Table 2). There are ve species of herbivores : wildboar (Sus scrofa), red deer (Cervus elaphus), ibex(Capra pyrenaica), chamois (Rupicapra rupicapra) and

horse (Equus caballus). Seven species of carnivores arepresent : fox (Vulpes vulpes), wolf (Canis lupus), brownbear (Ursus arctos), hyena (cf. Crocuta crocuta spelaea),iberian lynx ( Lynx pardina), badger (Meles meles) andanother species of mustelid. Also two species of leporidsare present : rabbit (Oryctolagus cuniculus) and hare(Lepus sp.). This faunal association does not differ fromother Portuguese collections from Initial Pleniglacial(transition OIS 2/3, from 32 to 22 000 year BP).

Large mammals

The mammal assemblage has a total of 1472 largemammal remains (Table 3). Of these, 450 (about 31 %)

can be identied to the species level. Whenever possible,besides taxonomical and anatomical determination,size and age classication were recorded. Number of

identied specimens (NISP), Minimum Number ofIndividuals (MNI) and Minimum Number of Elements(MNE) were calculated for each taxon.Herbivore remains (NISP = 375 ; MNI = 21) aredominated by red deer and horse, followed by caprids.

For all these species there is a signicant amount of juvenile specimens, some of them representing fetal orneonatal individuals (see below).An important quantity of carnivores was revealed, bothin number of individuals (NISP = 74 ; MNI = 12) and invariety of taxa (seven different species). The carnivorecollection is dominated by iberian lynx, and followed bywolf and fox (Table 4).Data on animal behaviour and results on the age prolesfor each carnivore species suggest differences in thenature of the cave occupation. Juvenile brown bearremains are present, and it is probable that bears used thecave to hibernate. Juvenile wolf remains are also present,

and in all probability they used the site to reproduceand protect offspring. No juvenile hyena remains werefound, but this species is also known to den in small

Fig. 2 : [A] Plan view of Pego do Diabo Cave. Diagonal lines show excavated squares. [B] Stratigraphy of Pego doDiabo Cave. (Both adapted from ZILHO 1997).

Humans and Carnivores in the Early Upper Paleolithic in Portugal 613


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cavities (e.g. BARTRAM & VILLA, 1998 ; BRUGAL et al.,1997 ; FOSSE, 1995 ; FOSSEet al., 1998). In addition, wolfand hyena may bring herbivore bones into caves to beconsumed. Small carnivores such as the iberian lynx, fox

and badger are known to prey on smaller animals andto take shelter in cavities either to protect themselvesor to eat. The carnivore remains in general can be dueto starvation (thus rare, this can happen to bear cubsduring hibernation ; cf. ROGERS, 1987 and 1992), tocarnivore predatory behaviour or to competition betweencarnivores (interspecic) or between carnivores andhumans for the use of the cave.Wolf, hyena and humans are the most obvious predatorsof the herbivores found in the cave, and they are likelythe main contributors to the bone assemblage. But wecan also associate the iberian lynx to this predator group,considering that under special circumstances this speciesis known to prey on small herbivores (juvenile deer andcaprids ; cf. AYMERICH, 1982 ; BLTRANet al., 1985 ; BRO,1993 ; and DELIBES, 1980) and occasionally carryingthem into shelters (FERNANDEZ & PALOMARES, 2000). Thisfact was recently documented at Buraca Escuras UpperPaleolithic levels by J.-Ph. BRUGAL (AUBRYet al., 2001).At Pego do Diabo, almost 52 % of the herbivore MNI arerepresented by juveniles remains (40 % of the herbivoreNISP). It is possible that most of the medium herbivores(caprids) and one third of the larger herbivores (the juvenile individuals) could have been preyed on by

iberian lynx.Brown bear was considered an unlikely candidate asa predator since it rarely feeds on animals (even ifoccasionally scavenges) and rarely transports carrion orbones to their hibernation den. In fact, the reduction ofsmells is one of its priorities to avoid attracting wolvesand humans during hibernation when a bear is oftenvulnerable given its reduced metabolic state (LINNELetal., 2000 ; ROGERS, 1987 ; STINER, 1998 ; TIETJE & RUFF,1980 ; also see ARGANT & PHILIPPE, 1997 on the utilityof bear fur). It is interesting to note that, contrary to thecave bear (Ursus spelaeus), brown bears are quite rarein strictly paleontological sites and relatively frequent inarchaeological sites, which may conrm the competitionfor the use of caves between brown bears and humans(ARGANT & PHILIPPE, 1997).

Table 2 : Pego do Diabo identied mammal taxa (microfauna excluded).

Order Artiodactyla Family Suidae Sus scrofa, L. 1758Family Cervidae Cervus elaphus, L. 1758Family Bovidae Capra pyrenaica, SCHINZ 1838

Rupicapra rupicapra, L. 1758Order Perissodactyla Family Equidae Equus caballus, L. 1758Order Carnivora Family Canidae Canis lupus, L. 1758 Vulpes vulpes, L. 1758

Family Ursidae Ursus arctos, L. 1758 Family Mustelidae Meles meles, L. 1758

Family Felidae Lynx pardina, TEMMINCK 1824 Family Hyaenidae Cf. Crocuta crocuta spelaea, GOLDFUSS 1810Order Lagomorpha Family Leporidae Lepus sp. Oryctolagus cuniculus, GRAY 1874

Note: Another species of carnivore, probably a mustelid (notMeles meles), was identied.

Table 3 : Quantifying data on the fauna present in Pego doDiabo (level 2).

NISP MNIHERBIVORESSus scrofa 12 2Cervus elaphus 197 7Capra pyrenaica 29 6Rupicapra rupicapra 22 2Cf. Caprid 42Equus caballus 73 4Sub-Total Herbivores 375 21

CARNIVORESCanis lupus 16 2Vulpes vulpes 15 2Ursus arctos 3 2Meles meles 6 2Cf. Mustelid* 1 1Lynx pardina 28 2Crucuta c. spelaea 6 1Sub-Total Carnivores 74 12Sub-Total Large Mammals 450 33Unidentied Large Mammals 1022TOTAL Large Mammals 1472

LEPORIDSLepus sp. 10 2Oryctolagus cuniculus 539 27Sub-Total Leporids 549 39Unidentied Leporid 158TOTAL Leporid 707

TOTAL NISP Herbivores + Leporids 924TOTAL Animal 2179

Note: (*) NotMeles meles.

614 M. J. VALENTE


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Researchers have used several criteria to identify themain agents of herbivore bone accumulation in caves,such as relative abundance between herbivore andcarnivore remains, presence of coprolites, presence of

artefacts, artefact abundance compared to carnivorebones, herbivore mortality proles, representation ofanatomical parts, and studies of bone modications suchas gastric dissolution, gnawing damage, cut marks orpercussion marks.The Carnivore Index is based on the ratio of herbivoreand carnivore remains (for a recent review of thisindex, see BLASCO-SANCHO, 1996). In this ratio, onlythe carnivores that are known to collect herbivorebones and contribute actively to bone accumulationswere considered (for reasons explained above and byBRUGALet al., 1994) : wolf, hyena and lynx. This ratiois 0,13 when using NISP [carnivore NISP (wolf, hyenaand lynx) = 49/herbivore NISP = 375]. Following STINER(1994 :91, table 4.14) in her study of Upper Pleistocenefaunal accumulations in Italian caves, it is believable thatthis result most probably corresponds to a den with faunacollected by carnivores.The use of the Carnivore Index, however, is somewhatproblematic, given that most carnivore remains foundin caves are probably due to in situ deaths, either bynatural causes or by carnivores preying on one another ;therefore, under these circumstances, it is plausible tothink that the full carcass will be present in the deposit.In contrast, herbivores are not cave users and are hunted

outside the cavity, so perhaps only a part of the carcasswill be deposited in the cave. As a result of these differentprocesses, we are likely to have an ination of carnivoreremains in the faunal accumulation. Using the MNIinstead of the NISP can help to alleviate this problemsince that unit is unaffected by differential transportation(cf. BLASCO-SANCHO, 1995). The index ratio of 0,24[carnivore MNI (wolf, hyena and lynx) = 5/herbivoreNISP = 21] increases the probability that carnivoreswere the most important agent in the accumulation ofherbivore bones at Pego do Diabo.Another line of evidence to take into consideration is thepresence of coprolites (BARTRAM & VILLA, 1998 ; DAVID,

1994 ; FOSSE et al., 1998 ; STINER, 1994). Though few,seven fragmented coprolites (hyena ?) were found atPego do Diabo Cave.Comprehensive research of the predatory agents involvedin the deposition of bones to Pego do Diabo Cave mustinclude a critical evaluation of direct evidence forhuman use of the cave, such as artefacts and replaces,in comparison to the presence of carnivore remains.For instance, a large quantity of lithic material coupledwith small numbers of carnivore bones normally reectsintense human activity in the fossil assemblage (cf. DAVID,1994 ; STINER, 1994). At Pego do Diabo, only 32 lithics

(11 tools and 21 akes or chips) and one bone pointwere recovered (ZILHO, 1997, 2 : 98-99). The relativelylow ratio of artefacts to carnivore NISP (33/52, or 0,63)compares favourably to STINERs (1994) ratio obtained

Table 4 : Quantifying NISP and MNI for herbivores andcarnivores at Pego do Diabo.

Adult

Immat. Total

% Herb

or Carniv

HERBIVORES

NISP

Sus scrofa 5 7 12 3,2

Cervus elaphus 114 83 197 52,5

Capra pyrenaica 14 15 29 7,7

Rupicapra r. 22 22 5,9

Cf. Caprid 28 14 42 11,2

Equus caballus 45 28 73 19,5

TOTAL 228 147 375

MNI

Sus scrofa 1 1 2 9,5

Cervus elaphus 3 4 7 33,3

Capra pyrenaica 2 4 6 28,6

Rupicapra r. 2 2 9,5

Equus caballus 2 2 4 19,1

TOTAL 10 11 21

CARNIVORESNISP

Canis lupus 13 2 15 20,0

Vulpes vulpes 12 3 15 20,0

Ursus arctos 1 2 3 4,0

Meles meles 7 7 9,3

Cf. Mustelid 1 1 1,3

Lynx pardina 28 28 37,3

Crocuta c. s. 6 6 8,0

TOTAL 67 7 74

MNI

Canis lupus 1 1 2 16,7

Vulpes vulpes 1 1 2 16,7

Ursus arctos 1 1 2 16,7

Meles meles 2 2 16,7

Cf. Mustelid* 1 1 8,3

Lynx pardina 2 2 16,7

Crocuta c. s. 1 1 8,3

TOTAL 9 3 12

Note: (*) NotMeles meles



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on Italian Pleistocene carnivore den assemblages, andthis reinforces the idea that carnivores used Pego doDiabo cave more frequently than did humans. As to thepresence of hearths, none was found, but a few burned

bones were recovered (see below).Herbivore mortality proles also assist in theidentication of the agents of faunal accumulation incaves. Most of the literature reports that wolf, hyenaand iberian lynx have a predilection for weak or younganimals, especially in large game such as red deer andhorse (on wolf see BEAUFORT, 1987 ; also see DOMINGUEZ-RODRIGO, 1993 for an oppositing view ; for the hyenasee DOMINGUEZ-RODRIGO, 1993 ; FOSSE, 1995 ; UADELLI,1989 ; HILL, 1989 and KRUUK, 1972 ; for the iberianlynx see BLTRANet al., 1985 and BRO, 1993). On theother hand, some modern hunter-gatherer groups favour

prime-age adults that are valuable in terms of caloriesand secondary products ; however, several exceptions areknown, and thus this strategy is highly variable amonghuman foragers (LYMAN, 1987, 1994 ; STINER, 1990).

Mortality proles are generally based on the analysis oftooth wear, and we have distinguished four age classesfor the herbivores : infant, juvenile, adult and old (seeBLASCO-SANCHO, 1995). Although Pego do Diabos

collection is relatively small (see Tables 4, 5), severalinteresting trends can be noted. These results, displayedin Figure 3, show herbivore mortality curves with abimodal distribution dominated by very young (infant)and prime-age adults for deer, horse and ibex. These dataalso inform on seasonality considerations. The presenceof infants with only deciduous dentition (a few with dPsstill erupting and others with dPs already erupted butwithout wear) suggest fetal or neonatal animals. Reddeer remains include two left dP

2completely unworn and

one right dP2

with intermediate stage wear, representingthree infant individuals (fetal to a few weeks old) and a

season of death between late spring and summer (LEGGE

& ROWLEY-CONWY, 1988 ; MARIEZKURRENA, 1983). Horsedeciduous remains include two dP2 with intermediateuse wear (infant), also matching late spring or summer

Fig. 3 : Mortality proles for wild boar, red deer, ibex, chamois and horse. Age classes: infant, juvenile, adult and old.

Table 5: Teeth NISP for each herbivore and carnivore taxon at Pego do Diabo.

Sus scrofa Cervus elaphus Capra pyrenaica Rupicapra r. Cf. Caprid Equus caballusDeciduous teeth 5 51 15 4 19Incisors/Canines 1 3 2 2 2? 2Pms 11 2 5Ms 33 7 5 2

Pms / Ms 3 21 3 1 6 32TOTAL 9 119 29 13 12 55

Canis lupus Vulpes vulpes Ursus arctos Lynx pardina CrocutaDeciduous teeth 1 4 2Incisors/Canines 1 1 3 1Pms 1 2 5 1Ms 1TOTAL 4 7 2 8 2

616 M. J. VALENTE


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killing. Ibex remains include two left unworn dP2 andtwo left intermediately worn dP

3, suggesting late spring

to summer kills (PEREZ RIPPOL, 1988). Finally, wild boardisplays one infant individual with unworn dPs, but since

this species has a reproduction period extending betweenFebruary and June, its season of death can correspond to aperiod that goes from late winter/early spring to summer(BULL & PAYNE, 1982). In sum, the abundance of isolatedteeth does not allow us a detailed study of the herbivorepopulation age, but the use wear features on deciduousteeth (unworn or intermediately worn) suggests that 40 %of the deer, 50 % of the horse and 66 % of the ibex werekilled in late spring or summer. This interpretation doesnot apply to adult prey, which could have been hunted atany other season, but implies preference for temperateseasons at least for the predation of immature animals.

Dening presence and absence of some anatomical partsmay help identify the agents of bone deposition. Mostpredator lairs are known to contain a large number ofcomplete or nearly complete herbivore long bones,whereas human sites present a different pattern withcranial elements predominating and higher long bonebreakage [for hyenas see DOMINGUEZ-RODRIGO, 1994 ;FOSSE, 1995 ; HILL, 1989 ; for wolves see DOMINGUEZ-

RODRIGO, 1994 ; HAYNES, 1980 ; STINER, 1994 ; for theiberian lynx there are no adequate data, but for otherfelids see NASTI, 1996 (puma) and RUITER & BERGER,2000 (leopard)].

At Pego do Diabo, the distal elements of the limbsare particularly prevalent, in contrast to the axialand cranial bones (see Table 6). Red deer exhibits anoverrepresentation of the appendicular extremities, suchas metapodials and phalanges. Horse is representedprimarily by limb extremities and other forelimbelements (radii, ulnae and carpals). Wild boar displaysfew remains, but again long bone ends predominate.Finally, caprids are represented mainly by humeri,astragali, metapodials and phalanges.Aware of the possibility that these results could havebeen caused by post depositional factors that affectfragmentation, particular attention was devoted tounidentiable bone fragments ; again, considerablenumbers of long bones were present.The evidence shows an overrepresentation of someanatomical parts, especially appendicular extremities,together with a few forelimb bones. These skeletalsegments do not represent the highest nutritional valueof the carcass, but they may correspond to carnivorespreying on leftovers discarded in the cave.

Table 6 : Bone NISP for each herbivore and carnivore taxon at Pego do Diabo.

Sus Cervus Rupicap Cf. Caprid Equus Canis Vulpes Ursus MelesCf. Mustelid Lynx Crocuta

Antler/horn 1 2Maxilla (2) (1) 1Mandible 2+(1) 1 1Vertebra 3+(1) 2Innominate 1Scapula 1Humerus 1+(1) 2+(2) 1Radius (1) 1 4 1Ulna 1 1 2Carpal 4+(3) (4) 4 1Metacarpal (1) 2 (2) 2+(3) 1

FemurTibia 1 (1) 1 1Calcaneus (1) 1 1 1Astragalus 3+(1) 3 (1) 1 1Tarsal 1 2 1Metatarsal 5 1 2 3 2Sesamoid 1 1Metapode 6+(2) 1+(3) (1) 2 4 11 Phalanx (1) 5+(11) 3 2+(3) (1) 1 2 6 12 Phalanx 8+(6) 4 1 1 1 1 13 Phalanx (1) 1+(3) 2 2 (1)Sub-total (3) 45+(33) 9 18+(12) 9+(10) 10+(1) 8 1 7 1 20 4TOTAL 3 78 9 30 19 11 8 1 7 1 20 4

Note: (x) immature

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Bone modications are the best diagnostic tool to identifythe accumulator or modier of the faunal assemblage(HAYNES, 1983). Table 7 summarizes bone NISPdisplaying gastric dissolution, gnawing damage, cut

marks and burning. There is direct evidence for carnivoreactivity, such as remains with gastric dissolution (mostlyin red deer but also in chamois ; see Fig. 4), scoring,punctures and a shortage of long bone ends. In contrast,there are a few clear cut-marks, normally on long boneshafts (Fig. 5 and 6). The only identied species that havecut-marks are the chamois, all on shafts, and red deer, ona distal antler branch.Although no hearths were found in the cave, somecarbonised remains were found. A total of 14 bones wascharred, but only two were identied taxonomically andto element, and both were carnivores : a wolf maxilla

fragment and a proximal radius of lynx.

Leporids : rabbits and hares

Approximately seven hundred leporid bones have been

recovered from the Early Upper Paleolithic level. Thesebones represent more than four hundred elements, or27 rabbits and two hares (Tables 3 and 8).Both small carnivores such as lynx, wildcat, fox orbadger, or larger ones like wolves, are known to huntrabbits and accumulate their bones in shelters (e.g.DELIBES & HIRALDO, 1981 ; GIL-SANCHEZ et al., 1999 ;HENRYet al., 1988 ; JAKSIC & SORIGUER, 1981 ; MARTINetal., 1995 ; ALOMARESet al., 2001 ; REVILLA & PALOMARES,2002). As HOCKETT (1991, 1993, 1995 ; also HOCKETT &HAWS, 2002) pointed out, assemblages created by smallercarnivores may be characterized by the accumulation of

entire rabbit carcasses, with large numbers of completeor nearly complete limb elements usually with one ofthe epiphysis attached to long bones. On the other hand,

Fig. 4 : Carnivore damage: digested red deer bones (hyenascats?).

Table 7 : Bone NISP damage in Pego do Diabo Cave.

Dissol. Gnawed Cutmarks BurnedHERBIVORESSus scrofa 1?Cervus el. 9 7 1Rupicapra r. 1 2 2Cf. Caprid 1 6Equus c. 2 1?

CARNIVORESCanis lupus 1? 1Vulpes v.Ursus arctosMeles m.Lynx pardina 1Crocuta 1?

Unidentied 71 10 4+1? 12TOTAL 82 27+3? 7+2? 14

Fig. 5: Anthropical modications : indeterminate long bonefragment with cut-marks.

Fig. 6: Anthropical modications: indeterminate long bonefragment with cut-marks.

M. J. VALENTE618


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wolves tend to bite and chew rabbit bones leaving lesscomplete elements, and humans usually create limbshafts, breaking the proximal and distal ends of longbones (e.g. humeri, femora and tibiae) to extract bonemarrow. Sometimes humans also leave cut-marks onrabbit bones, predominantly on mandibles and limbs,as has been shown for the Magdalenian levels at Lapado Suo (about 15 % of the collection shows cut-marks,many of them seen only under a microscope ; HAWS &VALENTE, 2001 ; VALENTE, 2000 ; also see PREZ-RIPOLL,1992 for higher percentages).

In the Pego do Diabo assemblage, all skeletal bones areequally present, showing no differential transportation ofany part of the rabbit carcasses. There are no cut-marksand most of the limb bones have at least one of theirends (Fig. 7), reinforcing the absence of anthropic andlarge carnivore activity. There are also two limb bonesthat show opposite tooth punctures on the distal end (inparticular the femora ; see Fig. 8), which is typical ofsmall carnivore activity, such as lynx and fox (HOCKETT,1993, 1999). The potential human inuence is limited tofour burned bones. Five remains display the characteristicsingular puncture of raptor activity.

As mentioned before, there is a small number of harebones present. None of these specimens represents anentire element, and all long bones have both or one of its

epiphyses. One proximal tibia exhibits a single puncture,typical of raptor damage.

CONCLUSIONS ON PEGO DO DIABOS

ASSEMBLAGE

Gathering all the evidence, Pego do Diabos Early UpperPaleolithic faunal assemblage can be characterizedas a palimpsest of different depositional agents, mostpossibly affected by bioturbation and sedimentary

or hydrological transportation. This interpretation isconrmed by the radiocarbon dates and reinforced by thenoted differences in mortality age proles (bimodal), andin particular by the probable multiple numbers of agentsthat accumulated and modied bones, such as carnivores,raptors and humans.In 1997, ZILHO said that Pego do Diabo Cave wassporadically used by small human groups, who organisedand maintained their hunting equipment (1997, 2 : 99).According to him, the large carnivores found in the caveaccumulated most of the faunal remains. This studydoes not contradict his interpretations, but the processes

that produced the Pego do Diabo faunal assemblageseem to be more complex ; besides wolf and hyena, weshould associate humans, iberian lynx and other small

Table 8 : Leporid bone NISP and NME, and NISP damage at Pego do Diabo.

Oryctolagus cuniculus Lepus sp. DamageNISP MNE NISP MNE Punct BurnedAdult Juve Infant Adult Juve Infant Adult Juve Adult Juve

Head 1 1Maxilla 6 3Mandible 21 11Scapula 9 1 5 1 1 1Humerus 15 4 12 2 1 1 2Radius 21 1 1 13 1 1 1 1 1 1 1Ulna 22 1 17 1 1Metacarpal 23 1 22 1Vertebra 21 17 1Rib 5 3Sacrum 2 2

Innominate 43 17 1 1 1Femur 19 1 1 9 1 1 2 1 1Tibia 28 7 3 16 6 1 1 1 1Calcaneus 36 3 1 36 3 1 1 1Astragalus 9 9Metatarsal 124 14 83 14 2 1 2 1Metapode 2Phalanx 95 96 1Unidentied 1

TOTAL 500 28 11 372 27 7 10 2 9 2 8 4

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carnivores, as well as raptors as probable contributors ofbones to the assemblage.Inter-species competition for the use of the cave waspronounced. It is clear that during the Upper Pleistoceneseveral carnivore species (at least hyenas, wolves, bears,

lynxes, foxes and mustelids) used Pego do Diabo asshelter and/or den, often transporting herbivore preyremains inside the cave. With the exception of wild boar,

all the herbivore species represented contain remainsdisplaying carnivore gnawing. Hyenas are surely one ofthe carnivore agents that modied the herbivore remains,as gastric action is one of their signatures on chamois and

red deer remains. Wolves and lynxes are a possibility aswell ; the latter may have preyed on immature red deerand horse, and possibly caprids. In addition, raptors,foxes, badgers, and above all lynxes, are potentialaccumulators of the leporid bone assemblage.It is also plausible to think that encounters amongcarnivores occurred ; perhaps a few of them had mortalconsequences, thereby producing at least part of thecarnivore bone assemblage found inside the cave (twocarnivore bones show possible gnawing marks, onebelonging to wolf, the other to hyena).The same kind of competition for the use of the cavity

was established between humans and carnivores, sincewe found some artefacts and a few herbivore remainswith unmistakable anthropic modications. Of all theherbivores, chamois seems to have the highest number ofevidence for human action : prime age prey (the literaturesuggests that hyenas, wolves and lynxes prey primarilyon immature herbivores), cut-marks, and a shortageof typical carnivore modications. Plus, even if mostcaprid bones were not identied to the species level,some of them may be chamois, including some elementswith high economic utility (e.g. pelvis and humerus ; cf.BINBFORD, 1978 ; METCALFE & JONES, 1988). The evidencealso suggests that humans modied red deer remains.

It is difcult to detail human intervention in Pego doDiabos assemblage and numerous hypothesis could beoffered. Humans could have transported animal bonesinto the cave as a result of hunting activities or secondaryexploitation of carcasses found in the vicinity ; or,alternatively, humans could have recovered some partsof carcasses brought to the cave by carnivores, either touse them as food (BRUGAL & JAUBERT, 1991) or for otherpurposes, such as manufacturing bone tools.

THE NATURE OF HUMAN-CARNIVORERELATIONSHIP DURING THE EARLY UPPER

PALEOLITHIC IN PORTUGAL

The consecutive use of Pego do Diabo Cave both byhuman communities and carnivores is something wealso can perceive in other settlements dated to the EarlyUpper Paleolithic in Portugal. Here, we do not havemany archaeological contexts dated from this period,and those with preserved and studied fauna are evenfewer (see Note 2). Besides Pego do Diabo, we presentlyhave ve cave sites that are early Upper Paleolithic in

age, all in Central Portugal (see Fig. 1 for locations) :Salemas, Caldeiro, Casa da Moura, Buraca Escura andLapa do Picareiro. In the absence of quantitative data, a

Fig. 7 : Carnivore damage : rabbit femura (upper row) andtibiae (lower row).

Fig. 8 : Carnivore damage : rabbit femur with oppositepunctures by small carnivore.

M. J. VALENTE620


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taxonomical list for these sites (and others) is presentedin Appendix 1. Radiometric dates are documented inAppendix 2.Salemas Cave is a thin gallery (1 m in width and 30 m

in length) located on a limestone cornice situated notfar from Pego do Diabo. Level III was classied asPerigordian by ZBYSZEWSKIand ROCHEin the 1960s and1970s (ZBYSZEWSKI, 1963), but recently ZILHO (1997,2 : 91) re-evaluated the lithic industry, classifying it asAurignacian and Early Gravettian. The faunal collectionwas never completely studied, but CARDOSO (1993)analysed the specimens that allowed paleontological andosteometrical observations (n = 142). Species presentand their relative frequencies are : Felis sylvestris -14,8 %, Lynx pardina - 21,8 %, Canis lupus - 14,3 %,Vulpes vulpes - 2,1 %, Ursus arctos - 1,4 %, Sus scrofa- 4,9 %, Cervus elaphus - 31 %, Rupicapra rupicapra- 7,7 %, andBos primigenius - 1,4 % (p. 544). No exactinformation on the age of the animals is available, butthe existing data show an assemblage similar to Pego doDiabos level 2.Caldeiro Cave is a narrow meandering gallery (3 min maximum width and 20 m in length) located onthe slope of a small valley at the bottom of which astream ows into the Nabo River. The site has EarlyUpper Paleolithic occupations : levels Jb and Ja wererespectively classied as Early and Final Gravettian byZILHO (1997, 2 : 118, 121). He also concluded that thehuman role in the bone accumulation was rather modest.

Recent studies by S. DAVIS (2002) showed that largecarnivores, especially hyena, but also cave lion, werecommon in these levels. Other carnivores are bear, lynx,and fox. Some indicators suggest that the cave, during theearly periods of occupation (Middle Paleolithic and EarlyUpper Paleolithic), was used as a hyena den : presenceofCrocuta remains, hyena coprolites and gastric etchedbones associated to a relatively high number of juvenileremains of red deer and equids, scarce burned remainsand a small density of lithic materials (opposed to a highnumber of faunal remains). Based on the increasing ratioof rabbit remains to herbivore remains from the olderMousterian and Early Upper Paleolithic levels to the

Magdalenian ones, it is also suggested that there was agradual increase of hunting pressure on the environment,maybe due to human population increasing.At Cesareda plateau, on its north edge, is Casa da MouraCave, with a 50 m sinuous gallery. Its outer room (theone with cultural deposits, measuring about 10 x 20 m)has a disturbed layer (1b) with materials that accordingto ZILHO (1997, 2) correspond to three occupations,Early and Final Gravettian (corresponding to the baseof layer 1b) and Solutrean (top of layer 1b). Excavatedby Delgado in the 19th century, level 1b delivered scantevidence for human occupation and a faunal assemblage

dominated by rabbit and a wide variety of carnivores(iberian lynx, leopard, wild cat, hyena, european polecat,fox, wolf, bear) along with horse, red deer, ibex, chamoisand wild boar (ROCHE, 1951). More recent studies by

CARDOSO (1993) and Altuna (on a small sample froma test excavation made in 1987 by STRAUS; STRAUS etal. 1988) again emphasized the high frequency of wolfremains. The available data strongly suggest that during

the Pleistocene Casa da Moura was essentially a wolf denintermingled with some human occupation.Buraca Escura is a small karstic cavity situated on thesouthern slope of the Poio Novo valley. It has fourlevels with Upper Pleistocene occupations, dated fromthe Gravettian, Final Gravettian and Proto-Solutrean,all with rare artefacts amongst an abundant faunalassemblage (AUBRY et al., 2001). The faunal remainswere fully analysed by J.-Ph. BRUGAL and recentlypublished (op. cit.), displaying a relatively highpercentage of carnivores. In the top layer (level 2a)horse dominates (37,2 % of the NISP), followed byibex (31,4 %) and red deer (14 %). Carnivore bones arepresent in very small quantities, most of them belongingto wolf (2,5 %, 3 specimens). All of the herbivore speciesare represented both by adults and immature (sometimesfetal or neonatal) individuals. Nearly half of the post-cranial horse remains exhibit cut-marks, and a signicantquantity of caprid bones display evidence of carnivoreaction. In layer 2b more than half the herbivore remainsare from ibex (56,1 %). Auroch (14,3 %), red deer (9,7 %)and horse (8,1 %) are also present and lynx dominatesthe carnivore assemblage (3,4 %). There is a largepercentage of immature herbivore individuals, mostlyfetal or neonatal ibex. With the exception of auroch, all

species display typical carnivore modications and a fewibex and auroch remains show cut-marks. Two speciesare well represented in level 2e : ibex (72,9 %) and lynx(11,2 %). Small quantities of horse (3,8 %), red deer (2 %)and wild cat (0,8 %) were also found. Most of the ibexbones belong to infant (foetal or neonatal) or juvenileindividuals, and carnivores modied an important numberof ibex remains. Finally, layer 2f had a small amount ofibex remains, totalling two individuals, one adult andthe other a juvenile. Three specimens show carnivoremodications. Based on these data (zooarchaeological,few artefacts and burned bones present), AUBRY et al.(2001 : 41) concluded that in Buraca Escura Cave, Upper

Paleolithic human occupations were short and intermixedwith carnivore lairs.The Lapa do Picareiro assemblage displays differentfeatures. J. HAWS (2000) summarized the evidence forcarnivore activity in all the Pleistocene cave depositsas two small marten-sized teeth, a few bones that showlimited punctures and a possible auroch bone that appearsto be acid-etched. None of these features, however, werefound on bones from the supposed Gravettian level (J)(see Note 2). This layer contained only two identiableremains belonging to red deer and caprid. In contrast,leporid remains dominate all the assemblage, including

the Gravettian occupation. HOCKETT

& BICHO

(2000)and BICHO et al. (2000 ; in preparation) point out thatthe leporid collection shows no clear evidence for thenatural accumulation of rabbit bones (no puncture marks,



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corrosion from gastric uids, thinning or polishing) ; inaddition, limb elements and cranial bones are abundantlyrepresented which is typical of human accumulations.In sum, the researchers concluded that intense rabbit

hunting occurred and rabbit constituted a principalsubsistence item near Picareiro Cave during much of theUpper Palaeolithic.

CONCLUDING REMARKS

During the Initial Pleniglacial (before Last GlacialMaximum), Central Portugal had a large number of cavesites with archaeological deposits dating from the EarlyUpper Paleolithic. These occupations are characterizedby a small amount of direct human occupation (notablyartefacts and bone modications), and by a relatively

important presence of carnivore remains (in number,but especially in diversity of species) and carnivorebone modications. The overall pattern provides uswith a scenario consistent with short and discontinuous

human occupations intermixed with carnivore dens.Furthermore, the evidence suggests that hyena, wolf andother large carnivores, lynx and other small carnivores,and humans, were competitors for the use of the caves asshelters during this time.The pattern in human-carnivore relationships seems tochange during the Final Gravettian/Proto-Solutrean,when sites with signicant evidence for carnivoreoccupation are much less common and the carnivores donot seem to play a big role in the bone accumulations. InCentral Portugal, Lapa do Picareiro (level J and I, if weaccept their Gravettian attribution ; BICHO et al., 2000

and in preparation) and Anecrial Cave [level 2 ; ZILHO

,

Appendix 1 : Portuguese Upper Pleistocene Fauna present in Early and Middle Upper Palaeolithic sites, listed by ungulates, carnivoresand small mammals (microfauna excluded).

P

egodoDiabo

S

alemas

C

aldeiro

C

asadaMoura

B

uracaEscura

L

apadoPicareiro

B

uracaEscura

B

uracaGrande

C

aldeiro

B

uracaEscura

A

necrial

L

apadoPicareiro

B

uracaGrande

Level 2 III Jb/Ja 1b 2f J 2e 9c I/H/Fc/Fb/Fa 2b/2a 2 I 9b/9a/8

Techno-complex A A

+

EG

EG/FG

(EUP)*

EG G MG? FG FG? PS/ES/

/MS/US

(Solutrean)*

PS PS PS FG/US?

HERBIVORES

Sus scrofaCervus elaphus ?Capreolus capreolusBos primigeniusCapra pyrenaicaRupicapra rupicapraEquus caballus

CARNIVORES

Canis lupusVulpes vulpes ?Ursus arctosMeles melesPanthera spelaea ?

Panthera pardusLynx pardina ?Felis sylvestris ? ?Crocuta c.spelaea ?

SMALLMAMMALS

Lepus sp.Oryctolagus cuniculus

Note: A Aurignacian, EG Early Gravettian, FG Final Gravettian, EUP Early Upper Paleolithic, G Gravettian, MG MiddleGravettian, PS Proto Solutrean, ES Early Solutrean, MS Middle Solutrean, US Upper Solutrean, ? Unsure presence of thespecies, (---)* Caldeiro Cave cultural units according to DAVIS2002.

622 M. J. VALENTE


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1997 and BRUGAL (pers. comm.)] seem to be the oldest

contexts representing this new reality, soon to be astandard during the Solutrean (see Appendix 1).This situation during the Early Upper Paleolithic(Aurignacian and Early Gravettian) can be attributedeither to low density of human population at that time(STRAUSet al., 2000 ; BICHO, 2000), to geomorphologicalfactors affecting site preservation and/or to changes inhuman use of the landscape (ZILHO, 1997, 2001 ; seealso VILLAVERDEet al., 1996 for Middle-Upper Paleolithictransition in central Spanish Mediterranean zone).Finally, we cannot overlook that by the Solutrean, largecarnivores such as leopard (Panthera pardus), cavelion (Panthera spelaea) and hyena (Crocuta crocuta),decrease or disappear from Portuguese archaeologicalcontexts.In Europe, leopard was a frequent presence after thelast interglacial (IS 5e) and appears to become extinctduring the beginning of the Upper Paleolithic (CASTAOS,1990 ; GURIN & PATOU-MATHIS, 1996). In contrast, inIberia, this species was common in the northern area(most especially during the Aurignacian ; cf. ALTUNA &MARIEZKURRENA, 1988), where it seems to survive untilMagdalenian times (CASTAOS, 1990). CARDOSO (1993)identied most of the known Portuguese specimens[Lorga de Dine and Furninha dated from the Early

Glacial ; Casa da Moura, Fontainhas (probably Solutreanlevels ; see CARDOSO, op. cit. : 446), Pedreira das Salemas,Figueira Brava and Escoural dated from the Late Wrm]

and, recently, DAVIS(2002) also documented the presence

of leopard specimens in Cardeiro solutrean levels.Cave lion was widespread in European assemblagesduring the last glaciation. In Portugal it was found in vecave sites [Lorga de Dine (Early Wrm), and Pedreiradas Salemas, Figueira Brava, and Escoural (Late Wrm),according to CARDOSO 1993 ; and in Caldeiro EarlyUpper Paleolithic levels, see DAVIS, 2002] and seems todisappear after the Initial Pleniglacial.Crocuta crocuta is one of the most regular taxa duringthe European Middle and Upper Pleistocene. In themeridional regions of Iberia, the presence of this speciesdoes not seem to be as regular as in the northern areas ;still, in Portugal, the hyena was found in some cavesites (most data in CARDOSO, 1993) : Crocuta crocutaintermedia in two Early Wrm sites (Lorga de Dine andFurninha) ; and Crocuta crocuta spelaea in nine sitesdated from the Late Wrm (Caldeiro, Columbeira,Fontainhas, Lapa da Rainha, Porto Covo, Algar deCascais, Figueira Brava, Escoural and Pego do Diabo).The lack of detailed information does not allownal conclusions on the causes for the reduction ordisappearance of large carnivores from Portuguesearchaeological contexts. Nonetheless, factors like humancompetition and/or changes in ecological conditionssuch as a loss of herbivore biomass and the reduction

of herbivore prey availability may help to explain theobserved record (VALENTE & BRUGAL, 2002 ; BRUGAL &VALENTE, in press).

Appendix 2 : Radiometric Dates (14C) for the mentioned sites (Initial and Middle Upper Paleolithic sites with fauna).

Site Level Tecno-complex Material Lab. No. Age B.P EvaluationBuraca Escura 2a Proto-Solutream Bone OxA-5524 21,820 200 ABuraca Escura 2e Final Gravettian Bone OxA-5523 22,700 240 ABuraca Escura 2f Gravettian Bone GifA-97258 26,560 450 ABuraca Grande 9a Upper Solutrean? Charcoal Gif-9502 17,850 200 ABuraca Grande 9b Final Gravettian Charcoal GifA-93048 23,920 300 ACaldeiro Fa topo Upper Solutrean Bone OxA-1938 20,400 270 ACaldeiro Fa topo Upper Solutrean Charcoal ICEN-295 21,200 + 1,800 2,300 ACaldeiro Fc Upper Solutrean Bone OxA-2510 18,840 200 ACaldeiro H Middle Solutrean Bone OxA-2511 20,530 270 ACaldeiro H Middle Solutrean Bone OxA-1939 19,900 260 ACaldeiro I Proto-Solutrean Bone OxA-1940 22,900 380 ACaldeiro Jb Early Gravettian Bone OxA-5542 26,020 320 ACasa da Moura 1b Early and Final Gravettian Bone TO-1102 25,090 220 ?

Pego do Diabo 2a Aurignacian Bones ICEN-490 23,080 490 ?Pego do Diabo 2b Aurignacian Charcoal ICEN-306 2,400 80 RPego do Diabo 2b Aurignacian Bones ICEN-732 28,120 + 860 - 780 ASalemas VS (II) Proto-Solutrean Bones ICEN-376 20,250 320 ?Salemas VS (II) Proto-Solutrean Bones ICEN-385 19,220 300 ?Salemas VS (II) Proto-Solutrean Bones ICEN-367 17,770 420 ?

Note: A Accepted result; R Rejected result; ? Unsure result.



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Notes :

1. Some researchers maintain doubts on the existenceof an Aurignacian techno-complex in Portugal (seeBICHO, 1999, 2000 ; RAPOSO, 2000).

2. Only sites with occupations classied as Aurignacianand Early or Middle Gravettian, thus dating until circa22.000 BP (i.e. until the end of Initial Pleniglacial andbefore the Last Glacial Maximum). Level J of Lapado Picareiro was also included. This layer has noradiometric dates, but according to N. BICHO (pers.comm.) its material displays some technologicaldifferences from level I, attributed to Final Gravettian.Therefore, Level J can be preliminarily classied asMiddle Gravettian.

ACKNOWLEDGEMENTS

This paper is a revised summary of my Masters thesispresented to the Faculdade de Letras da Universidadede Lisboa, funded by Praxis XXI - Junta Nacional deInvestigao Cientca e Tecnolgica. The study alsobeneciated from two scholarships : one from CentreNational de la Recherche Scientique and Institutode Cooperao Cientca e Tecnolgica Internacional(Proc. 423/Frana ; dir. of J. ZILHO and J.-Ph. BRUGAL) ;the other from Ambassade de France au Portugaland Instituto de Cooperao Cientca e TecnolgicaInternacional (Proc. 001 A0, Dossier n 97/001 ; dir. ofL. RAPOSO and J.-Ph. BRUGAL).I am indebted to Museu Nacional de Arqueologia andto Joo ZILHO for making the Pego do Diabo collectionavailable for study. I would like to thank Jean-PhilipBRUGAL for inviting me to participate in the UISPP 2001symposium at Lige. I am particularly grateful to Jean-Philip BRUGAL, Nuno BICHO and Bryan HOCKETT for theircomments on the manuscript.

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