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Research perspectives for the study of Neandertal subsistence strategies basedon the analysis of archaeozoological assemblages

Sabine Gaudzinski-Windheuser a,b,*, Lutz Kindler a,b

a Palaeolithic Research UnitQ1 , Römisch-Germanisches Zentralmuseum, Schloss Monrepos, D-56567 Neuwied, Germanyb Johannes Gutenberg-Universität Mainz, Schloss Monrepos, 56567 Neuwied, Germany

a r t i c l e i n f o

Article history:Available online xxx

a b s t r a c t

The discipline of archaeozoology holds the potential to considerably contribute to knowledge about thesocial behaviour of Neandertals. However, the translation of proposed subsistence strategies intopredictions about Neandertal social organisation still remains a challenge. The paper discusses thecurrent state of archaeozoological research with respect to Neandertal subsistence. It is concluded thatthe methodological research focus in archaeozoology has shifted from its original holistic perspective tointensified/specialised studies of particular taphonomic components. The authors argue for a return toa more holistic perspective to develop the full potential of archaeolozoology in order to obtaina comprehensive overall perspective of Neandertal social behaviour. Here, two avenues are suggested toreflate the processual character of taphonomy: 1. by conducting actualistic studies, which should serve totest the homogeneity of a faunal assemblage; and 2. by concentrating on sites from ecologically well-defined environments with high temporal resolution, such as interglacial sites.

� 2010 Elsevier Ltd and INQUA.

1. Introduction

During the mid-1960s, with the “Man the Hunter” symposium,the hunting way of life was placed in an evolutionary perspective.For the first time, consequences of large-mammal hunts for thesocial structure and behaviour of our ancestors were discussed (Leeand De Vore, 1968).

Large game hunting enables hominins to exploit animals largerand heavier than themselves, permitting an invasion in ecologicalniches formerly exclusively occupied by large carnivores. Thedevelopment of subsistence strategies is also closely connectedwith innovations in hunting technology, but also with radicalchanges in individual and social behaviour, shaping the life historyof the genus Homo (e.g. Cartmill, 1993; Aiello, 1998; Kaplan et al.,2000; Stanford and Bunn, 2001; Stiner, 2002; Roebroeks, 2007;Hublin and Richards, 2009). Against this background, large gamehunting is considered a key element for the social behaviouralevolution of hominins.

The rising awareness of the consequences of hunting behavioursoon led to new questions addressing this strategy which devel-oped into a new and important focus for research: faunal analysis.

New methodology was developed to explain the nature and originof a faunal accumulation. Applied to faunal assemblages fromPleistocene sites, this new methodology led to the almost imme-diate invalidation of previously held conceptions of our ancestors’hunting abilities (Binford, 1985). This approach sharpened theperception of the problems encountered in discussing formersubsistence strategies. Further methodology soon developed in thisfield (e.g. to assess scavenging opportunities (Blumenschine, 1986),population structures (Stiner, 1994), the depositional history ofbones (Lyman, 1994), carcass exploitation strategies (e.g. Binford,1981; Brain, 1981; Shipman, 1986; Domínguez-Rodrigo, 2002 Q2))which helped to systematically view Pleistocene faunal assem-blages against an ecological perspective.

This methodology successfully proved its potential, as evi-denced by the changing paradigm in the field. Hunting as a way oflife of our ancestors has been shown to be deeply rooted in our past,suggesting that ‘modern’ forms of behaviour already have a longevolutionary history (Gaudzinski, 2004a; Domínguez-Rodrigoet al., 2007). This has far reaching consequences for our currentunderstanding of hominin dispersal. However, suggesting thathunting almost always characterised the hominin niche is notenough.

Evidence for hunting obtained from the faunal record is mean-ingless unless the social organisation behind this strategy isaddressed. Several approaches have beenmade in this direction, for

* Corresponding author. Palaeolithic Research Unit, Römisch-GermanischesZentralmuseum, Schloss Monrepos, D-56567 Neuwied, Germany.

E-mail address: Gaudzinski@rgzm.de (S. Gaudzinski-Windheuser).

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Quaternary International

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1040-6182/$ e see front matter � 2010 Elsevier Ltd and INQUA.doi:10.1016/j.quaint.2010.11.029

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instance, by the contextualisation of physiological data with resultsobtained from faunal analysis (Soerensen and Leonard, 2001;MacDonald et al., 2009). Faunal analysis alone, however, equip-ped with a methodological apparatus developed to shed light onsubsistence patterns only, is unsuitable to highlight this evolu-tionary history of hominin social behaviour.

Faunal analysis illustrates changes in diet breadth and characterthrough time (e.g. Stiner, 2002) which can only be consideredagainst an ecological background. These studies are founded ona synopsis of case studies of individual sites. The significance ofobserved patterns, the sustainability of the deduced subsistencestrategies and especially the social systems behind these strategiesremain unconsidered and thus invisible.

In consequence, translation of subsistence patterns into socialforms of hominin organisation with questions addressing e.g. therole of hominin interactions in broader subsistence systems stillremains a challenge. This is illustrated by a summary on the currentknowledge about Neandertal subsistence strategies in Europe.

Faunal analysis has the potential to considerably contribute toknowledge of the social evolution of Neandertals and offerperspectives how to improve the potential of faunal analysis.However, a higher temporal resolution is needed in faunal analysis.This can be achieved by the focus on high-resolution archives inecologically well-defined locales such as interglacial deposits. Strictapplication of the methodological apparatus to decipher bio-stratinomic sequences in a site context is needed. To test individualbiostratinomic sequences, actualistic studies and experimentalsimulations are suggested. The comprehensive contextualisation ofbiostratinomic data with all plausible site variables can identifypatterns in the archaeological record which express hominin socialbehaviour.

2. Methods in archaeozoology

Today, faunal analysis is embedded in a taphonomical theoreticalframework. Taphonomic analysis founded on observations ofmodern processes is today generally considered fundamental to allzooarchaeological interpretations of deposits containing bones withcertain modifications (Cleghorn and Marean, 2007). Since the late1960s, the principles of bone taphonomy have become more andmore influential for the interpretationof archaeological assemblages.

In this perspective, hominins are among the various contribu-tors shaping the composition of faunal accumulations. It is gener-ally assumed that these various agents leave their distinct imprinton bone preservation and/or surface modification. Thus, the iden-tification of patterns of bone preservation should enable theidentification of the various agents responsible for a bone accu-mulation. Analogies taken from ethnoarchaeology (Binford, 1981),carnivore feeding behaviour (e.g., Haynes, 1983; Blumenschine,1986; Marean et al., 1992; Domínguez-Rodrigo, 1999), demo-graphic studies (e.g., Stiner, 1994), actualistic and comparativestudies, such as hydrodynamic sorting (e.g., Voorhies, 1969;Behrensmeyer, 1982; Coard, 1999; Fernández-Jalvo and Andrews,2003), trampling (e.g.,Q3 Olsen and Shipman, 1988; Nielson, 1991;Blasco et al., 2008; Gaudzinski-Windheuser et al., 2010) climati-cally induced weathering (Behrensmeyer, 1978; Andrews and Cook,1985) or density studies (Lam et al., 2003; Lam and Pearson, 2004,2005) are used to explain patterns of fossil bone preservation, bymeans of various quantitative indices (e.g. using NISP, MNE, MNIsee Lyman, 1994).

Certain individual bone surface modifications, such as e.g. cut-marks and gnawing marks, allow the identification of modifyingagents in Pleistocene faunas. Quantifications of the observed modi-fications are used to assess the degree of impact of the modifyingagent. These quantifications very often lead to results which allow

only ambiguous interpretation. A density mediated faunal assem-blage could reflect diagenetic destruction, selective transport and/ordestruction by hominins and/or carnivores as well as hydrodynamicbone transport or even an amalgam of all these processes. Thisdilemma of ambiguities in data interpretation is often described asthe problem of “equifinality” (Rogers, 2000a; Lyman, 2004).

Several types of studies in archaeozoology are currently tryingto meet this challenge. Actualistic studies are currently designed toimprove and fine-tune the body of bone surface modificationsattributable to the actions of known modifying agents (e.g.Fernández-Jalvo and Andrews, 2003; Q4Dominguez-Rodrigo andBarba, 2006; Blasco et al., 2008; Q5Dominguez-Rodrigo et al., 2009).Intensive discussions centre around the units of quantificationapplied in faunal analyses (e.g. body-part profiles vs. survival sets(e.g., Cleghorn and Marean, 2004; Stiner, 2004)). In addition, anincrease in sophisticated statistical procedures is an obviousrequirement to improve the resolution of variation seen inarchaeofaunas (Rogers, 2000a, 2000b; Domínguez-Rodrigo andYravedra, 2009). Finally, advances in technology lead to improve-ments within fields that deal with particular aspects of archae-ozoology, e.g., isotopes studies or skeletochronology (Richardset al., 2008; Britton et al., 2009; Rivals et al., 2009; Rendu, 2010).The majority of these important approaches constitute an integralpart of classical faunal analyses with the superordinate aim tocharacterise the role of hominins in faunal assemblages.

However, a holistic taphonomic perspective, in which homininsare one biostratinomic variable in the chain of taphonomicprocesses, is decreasingly addressed. Consequently, studies of site-formation processes become more and more detached from theoriginal taphonomical perspective, with bones being analysedisolated from their embedding milieu.

More than a decade ago, Pleistocene archaeology in Europe lostamajor researchparadigm, thequestionofhuntingvs scavenging,dueto discoveries of weapons made at the German site of Schöningen(Thieme,1997, 2008). Nevertheless, efforts continue atfine tuning themethodology that was developed to solve the hunting vs. scavengingdebate from an ecological perspective. Consequently, only theecological factors influencing hominin hunting strategies such ascoevolution/competition with carnivores or the ethology of wildgame (by seasonality studies) are recognised. From amethodologicalpoint of view, the social factors influencing hominin subsistence suchas learning, communication, cooperationand foodsharingstill remainmostly invisible in the archaeological record (Isaac, 1978; Lovejoy,1981; Kaplan et al., 2000; Kuhn and Stiner, 2006). This is also truefor variation in subsistence strategies, which can be translated intobroader systems of subsistence, modes of land use and social orga-nisation in a given ecological setting. The translation of taphonomicsignals suffers from uncertainties in their interpretation. Con-textualisation of these data with results obtained by other lines ofresearch such as lithic technology, isotope studies or paleoanthro-pological studies remains difficult because of differences in the reso-lution of data obtained. Therefore, the discipline of archaeozoologyremains isolated and disconnected from the anthropological andhistorical context (comp. e.g. Domínguez-Rodrigo et al., 2007).

Numerous evidence suggests that the hunting way of life isdeeply rooted in our past (Gaudzinski, 2004a; Domínguez-Rodrigoet al., 2007), suggesting thatmodern forms of behaviour have a longevolutionary history. Consequently special emphasis must be puton testing the functional context of archaeological sites and thesocial organisation behind subsistence strategies. Such a shift inresearch perspectives could be substantiated by a return to theholistic perspective of taphonomy established during the 1980s(Behrensmeyer and Hill, 1980; Binford, 1981; Gifford, 1981;Behrensmeyer, 1984; Q6Andrews and Cook, 1985; Behrensmeyerand Kidwell, 1985; Koch, 1989; Lyman, 1994).

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The need for a substantial shift in research questions in faunalanalyses andmethodological approaches can be considered againstthe background on current knowledge about Neandertal subsis-tence in Europe. In this perspective, a detailed reconstruction of thechronologyof biostratinomic and site-formationprocesses, inwhichhominins are just one variable is imperative before questions abouthominin social organisation can be answered.

3. The current European evidence for Neandertal subsistence

Major elements shaping the subsistence strategies of Neander-tals have been uncovered in recent years through contextualisationof results obtained from analyses of the archaeological record. Thediscussion becomes more and more detached from the actualarchaeozoological evidence.

This trend results from limitations of the methodologicalapparatus, and only very rarely is it possible to narrow down thetemporal depths within archaeozoological assemblages. Even theinterpretation of relatively unambiguous in-situ archaeologicalevidence uncovered at the German site of Schöningen (Thieme,2008) suffers from the difficulty of ascertaining the contempora-neity of the horse individuals scattered between the woodenweapons. Even though it can be plausibly argued that these are theleftovers of a hunting scenario, its social context remains obscure. Itis the contextualisation of archaeozoological data with dataobtained through micromorphological site analysis, dental analysisor isotope studies which helps to make predictions on this socialcontext (Hublin and Richards, 2009). This is illustrated by majoraspects of our current knowledge on Neandertal subsistence basedon the archaeozoological evidence.

Even though important aspects of the behavioural repertoire andsocial organisation of Neandertals begin to emerge in a new light(HublinandRichards,2009), anensembleviewofMiddlePalaeolithic

subsistence is still far from being clear. Evidence from stable isotopestudies of Neandertal remains (Richards et al., 2000) shows thatanimal products formed an important part of their diet. Competitionwith carnivores was apparently low (Kindler, 2007), as indicated byinterpretations of analyses of isotope signatures for sites in South-western France. Though both hyenas and Neandertals subsisted onherd animals, Neandertals additionally focused heavily on mega-fauna such as rhinoceros and mammoth (Bocherens et al., 2005).However, the regularexploitationofmegafaunaremains isnot visiblein the archaeozoological record (Bratlund, 2000).

As evidence from sites once perceived to reflect Neandertals asobligate scavengers has been refuted (Binford, 1988; Grayson andDelpech, 1994), and new evidence of hunting discovered (Thieme,2008), regular large-mammal hunting is now accepted as themajor source of meat acquisition by Neandertals. Numerousarchaeozoological studies have long established that our MiddlePalaeolithic predecessors were highly specialised hunters of largemammals (Gaudzinski, 1996; Gaudzinski and Roebroeks, 2000)focussing on high return rates. This behavioural strategy can befollowed in various environmental contexts (Gaudzinski, 2004b).

Monospecific faunal assemblages serve to illustrate this(Gaudzinski-Windheuser, 2006). From MIS 9 or 7 onwards andespecially in steppe environments during the dry-temperateclimates of the late Pleistocene, archaeofaunas are dominated bya single taxon (Fig. 1). These sites reflect the interception of entireherds or aggregations of many individuals at waterholes, waterstreams or along their migratory routes.

Sites with monospecific faunas are characterised by a highdominance of remains of a single species represented bya minimum number of individuals of up to over 100 animals,associated with lithic artefacts (Gaudzinski-Windheuser, 2006). InEurope these assemblages occur in open-air sites and caves fromMIS 9 onwards, although themajority of sites are recorded from the

Fig. 1. Late Pleistocene (MIS 5 or younger) Middle Palaeolithic sites in Eurasia characterised by monospecific or species dominated faunal assemblages.

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Early Weichselian towards the end of the Middle Palaeolithic. Cut-marks and hammerstone-induced impact notches attest directhominin interactionwith the fauna. It was argued (e.g., Gaudzinski-Windheuser, 2006) that subsistence tactics which led to theseaccumulations were obviously quite varied and demanded flexi-bility in hunting tactics and thus careful preparation and planning.

Faunas from Western and Central European Middle Palaeolithicsites dominated by large herbivores attest to a selective long-termexploitation of individuals in particular age classes. Exploitation oflarge bovids centered on prime age individuals, whereas formegafaunal exploitation, there is a focus on young individuals(Gaudzinski-Windheuser, 2006). Subsistence tactics were linked toparticular locales in the landscape which were repeatedly occupiedduring short visits. From the archaeozoological record, it remainsmostly ambiguous if these long-term exploitation strategies werelinked to a particular season. Prey selection and prey exploitationmay be subject to seasonal variation as suggested by (e.g.) toothanalysis of small assemblages from Middle Palaeolithic cave sites(Rendu, 2010). From the faunal evidence of the Levant, it was evenconcluded that intensive hunting by Neandertals led to a decline inthe populations of certain prey taxa (Speth and Clark, 2006).

In contrast to these long-term strategies, there are seasonallyrestricted mass hunting events accompanied by carcass exploita-tion geared towards resources with high return rates. The faunalassemblage from Salzgitter-Lebenstedt serves as a good example.The German reindeer-dominated site of Salzgitter-Lebenstedt sawshort-term mass death scenarios characterised by unselectivekilling of entire reindeer herds. At Salzgitter-Lebenstedt, the largestgroup of animals in the assemblage e adult reindeer males e diedaround September, as discerned from the stage in development ofmale antlers, as well as by means of tooth eruption stages in youngindividuals. The reindeer remains display abundant evidence ofbutchery, including filleting of meat and standardised processing ofbones for marrow. During marrow extraction, a clear selectionagainst sub-adults, as well as against bones with low marrowcontent occurred (Gaudzinski and Roebroeks, 2000, 2003).

Detailed studies of sites with species dominated faunal assem-blages give no indication that parts of the carcass exploitationsequence were associated with particular locales in the sites.Transport of particular animal parts remains largely invisible, andall stages of the butchery process can be reconstructed. Althoughdirect evidence for intra-site organisation is poor, analyses of thebone and stone assemblages suggest that other activities besidekilling and butchering must have taken place. To name a few: hideworking, production and curation of stone artefacts, preparation offood, perhaps also for storage, import of gathered goods, such ascollected bones for artefact production and - as some researcherssuggest - for architectural purposes (Gaudzinski et al., 2005).

Thus, indication is given that during the Middle Palaeolithica clear functional differentiation of the living space did not exist asis also obvious from other aspects of Neandertal life which is e.g.expressed in the lack of living structures (Kolen, 1999). It seemshighly plausible to assume that kill and butchery sites are subse-quently transferred into temporal residential camps.

These monotonous patterns in the Middle Palaeolithic record ofcentral Europe which only reflects Neandertal behaviour withrespect to larger herbivores prevent a more comprehensiveunderstanding of Neandertal subsistence and ecology, in particularthe influence of sympatric competitors. However, what becomesobvious from these single-taxon sites is that Neandertal land-use inthe northern part of Europe seems to be highly dependent onseasonal aggregation prey, which is not obvious from bone analysisalone. Isotope studies also emphasize a specialisation of Neandertalsubsistence to large migrating ungulates and megaherbivores(Bocherens and Drucker, 2003; Bocherens et al., 2005).

That seasonal exploitation must have been a driving force is alsounderlined by the unique evidence from the German site of BalveCave. The cave was frequently occupied because of its logisticalvalue, as could be demonstrated by zooarchaeological studies. Theanalysis of the numerous faunal remains attests to a seasonal“harvesting” of hibernating bears (Kindler, 2007).

The evidence from sites with faunas dominated by a single taxonmight also indicate seasonal aggregations of Neandertal groupsthat conduct hunting focused on ungulate herds. These groupsmust have split up into smaller bands of Neandertals, retuning tologistically attractive locales in their territories, which wereexploited in an opportunistic way, as seen in the case of Balve Caveand other cave sites in Central Europe (Jöris, 2005; Kindler, 2007).

The evidence for differences in subsistence tactics during theMiddle Palaeolithic (i.e. mass hunting vs. long-term exploitation ofindividual age classes within a taxon) was interpreted in terms offlexible resource utilization against a background of unstableenvironmental conditions, which is prone to equivocal interpreta-tions in terms of highly efficient hominin adaptations, mereopportunism or a combination of both. However, the overallconsolidating evidence leaves little doubt that Neandertal subsis-tence strategies were driven by obtaining high energy return rates,which in turn called for a high degree of flexibility in the tacticsemployed (Gaudzinski-Windheuser and Niven, 2009).

In more southern latitudes, fluctuations in return rates mighthave been compensated by a broadening of the dietary spectrum.For more southern latitudes, the exploitation of fast-moving smallprey can be demonstrated (Blasco and Férnandez Peris, 2009), thesmall prey spectrum being generally shown to encompass onlyslow moving or static prey such as tortoises, marine molluscs oreggs (Stiner, 2002; Blasco, 2008).

For north-central Europe, the inclusion of small mammals andbirds in hominin diet remains largely undocumented. In most caveand open-air sites, there is an underrepresentation of smallmammals and birds in the archaeofauna, possibly relating to taph-onomic and excavation bias. Small mammal and bird exploitationhas largely remained unstudied from a taphonomic perspective dueto the overall scarcity of anthropic marks on the bones. This rendersit difficult to attribute singular skeletal elements of small mammalsto the hominin dietary spectrum rather than to the natural back-ground fauna (Gaudzinski-Windheuser and Niven, 2009). However,the number of sites with unambiguous evidence of utilization ofsmall game and especially birds by Neandertals, consisting of thepresence of cut-marked bones has increased in recent years due tothe recognition of researchers for this important topic (e.g. Fioreet al., 2004; Dibble et al., 2009).

This brief summary illustrates the major focus of Neandertalsubsistence, i.e. to obtain high-energetic food, coupled with theincreasing tendency for a broadening of the dietary niche. Thisperspective is in accordance with the potential of our methodo-logical apparatus as was already argued. Consequently the socialorganisation behind these subsistence tactics is indeed addressedin most interpretations of archaeozoological data but cannot betested. Thus the interpretation of subsistence strategies solelyremains embedded in an environmental frame of reference.

4. Results: perspectives for faunal analyses

The limitation in narrowing down the temporal resolution of anarchaeozoological context represents a major obstacle for the inter-pretation of obtained data. The temporal frame of reference refers toa geological perspective which prevents us from corroboratingaspects of social behaviour. As a consequence, a higher resolution ofthe sequence of taphonomic processes and of formationprocesses ofarchaeological sites is needed. It is this line of researchwhich shaped

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the taphonomic discussion beginning in the 1980s (Behrensmeyer,1975, 1982; Isaac, 1978, 1983, 1984; Behrensmeyer and Hill, 1980;Binford, 1981, 1985, 1987, 1989; Badgley, 1982; Behrensmeyer andSchindel, 1983; Allison and Briggs, 1991; Bunn, 1991), and whichreceded into the background in recent years. The present authorsstrongly support the return to this holistic perception of taphonomy.Study of faunal assemblages from high-resolution archives and thedetailed high-resolution reconstruction of the individual bio-stratinomic sequencing of a site provide perspectives for futureresearch. Contextualisation and combination of results obtained byhigh-resolution analysis will reveal patterns in the archaeologicalrecordwhichmight enable further analysis (e.g., digital modelling ofsite-formation processes) and can be addressed in the context of theevolution of hominin social behaviour.

Studies in the Late Upper Palaeolithic illustrate this point. Somefaunal assemblages from Late Upper Palaeolithic open-air sites, e.g.Gönnersdorf (Germany) (Street and Turner, in press), Oelknitz(Germany) (Gaudzinski-Windheuser, in press), Pincevent (France)(Enloe, 2003) display very high temporal resolution compared toMiddle Palaeolithic sites. Detailed refitting studies and correlationof refitting data with the spatial record reveal individual sitespecific patterns interpreted in terms of human economic andsocial interaction (Enloe, 2003).

4.1. The study of high-resolution archives

It is particularly worthwhile to study Neandertal presence inecologically well-defined environments, at sites with high temporalresolution. Interglacial deposits represent such high-resolutionarchives for studying hominin behaviour. Interglacial deposits are

comparatively rare during the Middle and Upper Pleistocene interms of percentages of the overall time of the periods (ca. 8%). Therehas been considerable debate about the capacities of Neandertals tosurvive in such forested environments (Gaudzinski-Windheuserand Roebroeks, in press).

Many interglacial sites are known from Europe (Gaudzinski-Windheuser and Roebroeks, in press) (Fig. 2). Many of these sitesshare major characteristics as their finds are embedded in a fine-grained sedimentary matrix and the excavated and investigatedareas are large. The amount of finds is extremely high, theircondition tends to be excellent, in that plant remains are preservedand bone surfaces do not show signs of (post-) depositional alter-ation. Many still exhibit the finest cut- and scrape-marks. The sitesare spatially structured by archaeological material. Most impor-tantly, the correlation of the palynological succession of an Inter-glacial with the sedimentary envelopes of the sites allowsa subdivision into defined and restricted time slices, representingnarrow time windows during which the accumulation of thearchaeological material occurred. This is a unique situation in theMiddle Palaeolithic record which distinguishes interglacial sitesfrom other well preserved sites with finely stratified andwell datedsequences allowing vertical and horizontal control when analysingfaunal assemblages.

The Eemian interglacial provides an especially interesting casestudy of Neandertal exploitation of forested habitats. This periodhas been subject to detailed studies, as it was hoped to gaininformation that could be relevant for current discussions on globalclimate change (Kukla et al., 2002; Shackleton et al., 2003; Breweret al., 2008; Tzedakis et al., 2009). The Eemian started around125,000 years ago and lasted approximately 10,000e12,000 years

Fig. 2. Eemian archaeological sites (MIS 5e) in Europe, which combine extraordinary environmental resolution and rich and well preserved faunal accumulations in one locale.A remarkable study area is located directly along the fringes of the Drenthe glacier dating to MIS 6 (indicted in a light grey; for comparison the extent of the Weichselian glacificationis indicated in dark grey). With retreat of the Drenthe glacier numerous lake basins were formed in the underground and filled during the succeeding Eemian Interglacial.

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in the northern part of Europe (Müller, 1974; Brewer et al., 2008). InPleistocene terms, this is a short period of time and a ratherhomogeneous environment.

Excavations undertaken at the Eemian site of Neumark-Nord2 illustrate the potential of interglacial deposits for disentanglingthe temporal resolution of Middle Palaeolithic archives. Neumark-Nord 2 represents one of the rare examples among Eemian inter-glacial sites that were recently excavated with modern techniqueswhere excavations were not hampered by a rescue situation. At thissite, part of a shallow lake basin with shore area was excavated infine-grained sediments. Over 120,000 well preserved bone remainsrepresenting taxa of a typical interglacial fauna were uncoveredtogether with almost 20,000 artefacts made from Baltic flint. Thebones show distinct cut-marks and hammerstone-induced conicalimpacts that clearly indicate hominin interaction with the fauna.Results of geomorphological and palynological studies show thatthe accumulation of the sediments that yielded these archaeolog-ical remains could be reconstructed to have lasted less than 500years (Kindler et al., 2010; Sier et al., in press). At Neumark-Nord 2 itwill be possible to reconstruct the formation of the palimpsestrepresented here and it will be possible to evaluate the contem-poraneity of elements of the faunal assemblage.

4.2. Testing the homogeneity of faunal assemblages

An additional avenue which could be approached in order tonarrow down the temporal resolution of an archaeozoologicalassemblage and to test whether a faunal assemblage is homoge-neous in character refers to actualistic studies. These shouldattempt to reconstruct the individual chronology of biostratinomicfactors and processes at a particular site. Such studies should try toobtain insight into the additive sequencing of a taphonomicprocess. These studies must pay attention to individual site-formation histories that can be recognised in archaeological sites,as well as incorporate already known contexts, linking the actual-istic study to the taphonomic history of a given faunal assemblage.Thus, the specific mode of operation of a taphonomic process ata given site rather than the general recognition of patterns oftaphonomic signals must be central to these studies.

In this context, the results of actualistic studies undertaken forthe EarlyMiddle Pleistocene Israeli site of Gesher Benot Ya’aqov (fora compilation of the current state of analysis, see Alperson-Afilet al., 2009) have successfully demonstrated the homogeneity ofthe faunal assemblage, with important consequences for the overallinterpretation of the site (Gaudzinski-Windheuser et al., 2010;Rabinovich et al., 2008, in press).

Tumbling and trampling experiments were initiated to mimicthe individual environmental conditions at the site, to gain quali-tative insights into processes of bone modifications and to assessthe timing of the biostratinomic chronology. It was assumed thatboth mechanisms were responsible for the formation of striationsdocumented on the bone surfaces documented for Layers V-5 andV-6.

Efforts succeeded in replicating the chronology of bio-stratinomic factors and processes at the site and, for the first time,were able to obtain insight into the additive sequencing of a taph-onomic process (Gaudzinski-Windheuser et al., 2010; Rabinovichet al., in press) (Fig. 3).

These observations have major implications for taphonomicstudies as they suggest that interpretations based on separatedisolated taphonomic processes may be misleading. Against a bio-stratinomic background, bone surface modifications result froma process in which bone surfaces are continuously altered. This isimplying that interpretations of biotically- and abiotically-inducedmodifications according to diagnostic templates but without theirintegration into their complete individual biostratinomic sequenceare highly ambiguous.

Recent studies on isolated taphonomic features, e.g. thefrequency of cut-marks (Domínguez-Rodrigo and Yravedra, 2009)and percussion-marks (Galàn et al., 2009) on bones, illustrate thehigh variability of these traces which makes their verification andquantification more difficult. These studies acknowledge themodifying qualities of other additional subsequent and super-imposing biostratinomic processes. They attempt to replicateindividual site specific taphonomic signals with the aim to makegeneral prediction on their origin from a superordinate perspective.It is not attempted to analyse these signals in the individual bio-stratinomic context. As a result interpretations of significantcorrelations between the various variables involved in these

Fig. 3. Model of the biostratinomic process replicated in the trampling experiments simulating environmental conditions at the Middle Pleistocene site of Gesher Benot Ya’aqov,Israel. Observable bone alterations are given on the ordinate. On the abscissa the progress of the biostratinomic process is given on an ordinal scale as taphonomic time (afterLyman, 1994, 358).

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Fig. 4. Trampling experiment 5 simulating environmental conditions at the Middle Pleistocene site of Gesher Benot Ya’aqov, Israel. Hyena ribs, a) before trampling, b) after 5 h oftrampling. Reduction of the bones into a fibrous consistency and bone disintegration occurred after trampling, resembling bone alteration due to climatic induced weathering.Arrows indicate the locations of the bones which are shown in detail.

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studies which are expressed in clearly discriminable patterns stillremain confronted with equifinality.

A consequent mandatory problem concerns the question ofwhether certain taphonomic patterns are diagnostic or not withregard to specific processes and agents. Due to their individualstructure andmorphology, bones that pass the same biostratinomicsequence can display completely different patterns of bone pres-ervation and bone modification, regarded as significant for tapho-nomical interpretations.

Studies of climatically induced weathering and abrasion usuallyoutline that the degree of bone destruction correlates with the timebetween the death of an individual and its final burial. As a conse-quence, homogeneity in bone surface preservation is considered anindicator for the assumption of homogeneous environmentalconditional and/or homogeneity of an entire bone assemblage. Inour experiments at Gesher Benot Ya’aqov a very short abrasiveprocess produced certain diagnostic bone surface modificationswhich are indistinguishable from bone surface modifications due tolong-term climatically inducedweathering (Fig. 4). Moreover it wasdemonstrated that the process of trampling is characterised byformation, modification and erosion of traces sharing the samemorphology as hominin cut- and carnivore tooth-marks. In addi-tion, cut-marks and most likely also tooth-marks, can be altered bythe abrasional process to the degree that they finally mimicstriations.

As a consequence, strict determination of the time of cessationof the biostratinomic process is mandatory for diagnosis andinterpretation of bone surface modifications. Against this back-ground generalised interpretations of fossil assemblages deprives

Q7 them of their individual history and leaves us without theirpotential scientific value (Gaudzinski-Windheuser et al., 2010;Rabinovich et al., in press).

5. Summary and discussion

In recent years numerous disciplines, among them biochemicaland/or paleoanthropological studies, as well as the contextualisa-tion of the overall archaeological record have vastly improvedknowledge of food acquisition and exploitation and thus subsis-tence strategies by Neandertals. The results of the different disci-plines are complementary, as becomes obvious when comparingresults obtained from isotope- and archaeozoological studiesemphasising the ecological specialisation on medium to large sizedherbivores. However, only archaeozoology has the potential totranslate variation in subsistence into a comprehensive overallperspective of Neandertal behaviour. Bones as the major source ofarchaeozoological analysis represent part of the archaeologicalrecord and provide direct access for the recognition of homininactions and behaviour at Pleistocene sites. Thus, archaeozoologystill remains the critical discipline for the study of subsistencebehaviour.

However, in spite of a steadily growing methodological appa-ratus and innovations in analytical techniques, archaeozoology stillfails to provide a comprehensive picture of the social organisationof Neandertal subsistence, based on the different results obtainedat individual sites. In consequence, the Neandertals’ way of liferemains ambiguous and bloodless.

Part of the dilemma results from the loss of a holistic approachin taphonomical research. This holistic perspective resulted fromresearch attempts to answer questions concerning the functionalcontext of sites representative for particular forms of socialbehaviour (compare e.g.Washburn and Lancaster,1968; Isaac,1978,1983, 1984; Behrensmeyer and Hill, 1980; Binford, 1985, 1987, 1989;Bunn, 1991). This approach to research in archaeozoology was soonreplaced by the “hunting/scavenging research paradigm” that has

in the meantime exhausted its potential as a motive for faunalanalysis. As long as the research focus is not directed towards themotives and the concomitant aspects of any subsistence behaviourentangled in the archaeological record, archaeozoological researchwill not profit frommethodological improvements in a non-holisticperspective and will only continue to recapitulate the current stateof knowledge.

The other aspect of the dilemma described above results fromthe fact that archaeozoological methodologies have become moreand more disconnected from the reconstruction of homininbehaviour. Research over the last decades has resulted in a largebody of actualistic and comparative data describing the characterand timing of individual taphonomical processes. These data helpto identify the individual processes and agents that shaped thecharacter of a given faunal assemblage. This line of research doesnot attempt to identify and/or arrange coherent and self containedchronological sequences of taphonomical processes involved in theformation of fossil accumulations. Without their integration intoa complete biostratinomic sequence, interpretations of biotically-and abiotically-induced modifications on bones according todiagnostic templates remain highly ambiguous. Thus actualistictaphonomic research must acknowledge the individual character-istics of Pleistocene sites, which form the determining variablesthat shape the individual taphonomic sequencing of an assemblage.This research must become integrated into a comprehensivepalaeoecological analysis of the embedding milieu. The quality ofthis approach can be tested by investigating sites embedded inspatial high-resolution milieus, as can be found in interglacialdeposits. As taphonomical research is not able to clearly define thetemporal resolution of a given faunal assemblage, high-resolutionmilieus provide a temporal frame of reference within which bio-stratinomic processes can operate. High-resolutionmilieus bear thepotential for the identification of entire biostratinomic sequences.This is a point of crucial importance as taphonomic processes actadditively. Within the biostratinomic chain, individual signalscaused by particular actions can be overprinted by succeedingactions, the signals of which will consequently not be identified asa taphonomically-relevant pattern. It is therefore imperative to testreconstructions of biostratinomic processes at a given archaeolog-ical site by means of actualistic studies simulating the depositionalhistory of this site.

Only if taphonomic research appreciates the immense potentialof the character of the embedding milieu at any given site asa reference frame for analysis, can taphonomical chains be identi-fied, and thus the role of hominins more precisely evaluated.

Acknowledgements

We like to thank Eudald Carbonell, Maria Gema Chacón and theIPHES working group for inviting us to the workshop “The Nean-derthal Home: Spatial and Social Behaviours”. Daniela Holst andRadu Iovita provided valuable feedback. We would like to thankRivka Rabinovich and Naama Goren-Inbar for inviting us toparticipate in the research at Gesher Benot Ya’aqov. Finally ourthanks go to all members of our Neumark-Nord working group.

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Please cite this article in press as: Gaudzinski-Windheuser, S., Kindler, L., Research perspectives for the study of Neandertal subsistencestrategies based on the analysis of archaeozoological assemblages, Quaternary International (2011), doi:10.1016/j.quaint.2010.11.029