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Research perspectives for the study of Neandertal subsistence strategies based on the analysis of archaeozoological assemblages
Sabine Gaudzinski-Windheuser a, b, *, Lutz Kindler a, b
a b

Palaeolithic Research Unit, Rmisch-Germanisches Zentralmuseum, Schloss Monrepos, D-56567 Neuwied, Germany Johannes Gutenberg-Universitt 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 the social behaviour of Neandertals. However, the translation of proposed subsistence strategies into predictions about Neandertal social organisation still remains a challenge. The paper discusses the current state of archaeozoological research with respect to Neandertal subsistence. It is concluded that the methodological research focus in archaeozoology has shifted from its original holistic perspective to intensied/specialised studies of particular taphonomic components. The authors argue for a return to a more holistic perspective to develop the full potential of archaeolozoology in order to obtain a comprehensive overall perspective of Neandertal social behaviour. Here, two avenues are suggested to reate the processual character of taphonomy: 1. by conducting actualistic studies, which should serve to test the homogeneity of a faunal assemblage; and 2. by concentrating on sites from ecologically welldened 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 rst time, consequences of large-mammal hunts for the social structure and behaviour of our ancestors were discussed (Lee and De Vore, 1968). Large game hunting enables hominins to exploit animals larger and heavier than themselves, permitting an invasion in ecological niches formerly exclusively occupied by large carnivores. The development of subsistence strategies is also closely connected with innovations in hunting technology, but also with radical changes in individual and social behaviour, shaping the life history of 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 game hunting is considered a key element for the social behavioural evolution of hominins. The rising awareness of the consequences of hunting behaviour soon led to new questions addressing this strategy which developed into a new and important focus for research: faunal analysis.
* Corresponding author. Palaeolithic Research Unit, Rmisch-Germanisches Zentralmuseum, Schloss Monrepos, D-56567 Neuwied, Germany. E-mail address: Gaudzinski@rgzm.de (S. Gaudzinski-Windheuser). 1040-6182/$ e see front matter 2010 Elsevier Ltd and INQUA. doi:10.1016/j.quaint.2010.11.029

New methodology was developed to explain the nature and origin of a faunal accumulation. Applied to faunal assemblages from Pleistocene sites, this new methodology led to the almost immediate invalidation of previously held conceptions of our ancestors hunting abilities (Binford, 1985). This approach sharpened the perception of the problems encountered in discussing former subsistence strategies. Further methodology soon developed in this eld (e.g. to assess scavenging opportunities (Blumenschine, 1986), population structures (Stiner, 1994), the depositional history of bones (Lyman, 1994), carcass exploitation strategies (e.g. Binford, 1981; Brain, 1981; Shipman, 1986; Domnguez-Rodrigo, 2002)) Q2 which helped to systematically view Pleistocene faunal assemblages against an ecological perspective. This methodology successfully proved its potential, as evidenced by the changing paradigm in the eld. Hunting as a way of life of our ancestors has been shown to be deeply rooted in our past, suggesting that modern forms of behaviour already have a long evolutionary history (Gaudzinski, 2004a; Domnguez-Rodrigo et al., 2007). This has far reaching consequences for our current understanding of hominin dispersal. However, suggesting that hunting almost always characterised the hominin niche is not enough. Evidence for hunting obtained from the faunal record is meaningless unless the social organisation behind this strategy is addressed. Several approaches have been made in this direction, for

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

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instance, by the contextualisation of physiological data with results obtained from faunal analysis (Soerensen and Leonard, 2001; MacDonald et al., 2009). Faunal analysis alone, however, equipped with a methodological apparatus developed to shed light on subsistence patterns only, is unsuitable to highlight this evolutionary history of hominin social behaviour. Faunal analysis illustrates changes in diet breadth and character through time (e.g. Stiner, 2002) which can only be considered against an ecological background. These studies are founded on a synopsis of case studies of individual sites. The signicance of observed patterns, the sustainability of the deduced subsistence strategies and especially the social systems behind these strategies remain unconsidered and thus invisible. In consequence, translation of subsistence patterns into social forms of hominin organisation with questions addressing e.g. the role of hominin interactions in broader subsistence systems still remains a challenge. This is illustrated by a summary on the current knowledge about Neandertal subsistence strategies in Europe. Faunal analysis has the potential to considerably contribute to knowledge of the social evolution of Neandertals and offer perspectives 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 in ecologically well-dened locales such as interglacial deposits. Strict application of the methodological apparatus to decipher biostratinomic sequences in a site context is needed. To test individual biostratinomic sequences, actualistic studies and experimental simulations are suggested. The comprehensive contextualisation of biostratinomic data with all plausible site variables can identify patterns in the archaeological record which express hominin social behaviour. 2. Methods in archaeozoology Today, faunal analysis is embedded in a taphonomical theoretical framework. Taphonomic analysis founded on observations of modern processes is today generally considered fundamental to all zooarchaeological interpretations of deposits containing bones with certain modications (Cleghorn and Marean, 2007). Since the late 1960s, the principles of bone taphonomy have become more and more inuential for the interpretation of archaeological assemblages. In this perspective, hominins are among the various contributors shaping the composition of faunal accumulations. It is generally assumed that these various agents leave their distinct imprint on bone preservation and/or surface modication. Thus, the identication of patterns of bone preservation should enable the identication of the various agents responsible for a bone accumulation. Analogies taken from ethnoarchaeology (Binford, 1981), carnivore feeding behaviour (e.g., Haynes, 1983; Blumenschine, 1986; Marean et al., 1992; Domnguez-Rodrigo, 1999), demographic studies (e.g., Stiner, 1994), actualistic and comparative studies, such as hydrodynamic sorting (e.g., Voorhies, 1969; Behrensmeyer, 1982; Coard, 1999; Fernndez-Jalvo and Andrews, 2003), trampling (e.g., Olsen and Shipman, 1988; Nielson, 1991; Blasco et al., 2008; Gaudzinski-Windheuser et al., 2010) climatically 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, by means of various quantitative indices (e.g. using NISP, MNE, MNI see Lyman, 1994). Certain individual bone surface modications, such as e.g. cutmarks and gnawing marks, allow the identication of modifying agents in Pleistocene faunas. Quantications of the observed modications are used to assess the degree of impact of the modifying agent. These quantications very often lead to results which allow

only ambiguous interpretation. A density mediated faunal assemblage could reect diagenetic destruction, selective transport and/or destruction by hominins and/or carnivores as well as hydrodynamic bone transport or even an amalgam of all these processes. This dilemma of ambiguities in data interpretation is often described as the problem of equinality (Rogers, 2000a; Lyman, 2004). Several types of studies in archaeozoology are currently trying to meet this challenge. Actualistic studies are currently designed to improve and ne-tune the body of bone surface modications attributable to the actions of known modifying agents (e.g. Fernndez-Jalvo and Andrews, 2003; Dominguez-Rodrigo and Q4 Barba, 2006; Blasco et al., 2008; Dominguez-Rodrigo et al., 2009). Q5 Intensive discussions centre around the units of quantication applied in faunal analyses (e.g. body-part proles vs. survival sets (e.g., Cleghorn and Marean, 2004; Stiner, 2004)). In addition, an increase in sophisticated statistical procedures is an obvious requirement to improve the resolution of variation seen in archaeofaunas (Rogers, 2000a, 2000b; Domnguez-Rodrigo and Yravedra, 2009). Finally, advances in technology lead to improvements within elds that deal with particular aspects of archaeozoology, e.g., isotopes studies or skeletochronology (Richards et al., 2008; Britton et al., 2009; Rivals et al., 2009; Rendu, 2010). The majority of these important approaches constitute an integral part of classical faunal analyses with the superordinate aim to characterise the role of hominins in faunal assemblages. However, a holistic taphonomic perspective, in which hominins are one biostratinomic variable in the chain of taphonomic processes, is decreasingly addressed. Consequently, studies of siteformation processes become more and more detached from the original taphonomical perspective, with bones being analysed isolated from their embedding milieu. More than a decade ago, Pleistocene archaeology in Europe lost a major research paradigm, the question of hunting vs scavenging, due to discoveries of weapons made at the German site of Schningen (Thieme, 1997, 2008). Nevertheless, efforts continue at ne tuning the methodology that was developed to solve the hunting vs. scavenging debate from an ecological perspective. Consequently, only the ecological factors inuencing hominin hunting strategies such as coevolution/competition with carnivores or the ethology of wild game (by seasonality studies) are recognised. From a methodological point of view, the social factors inuencing hominin subsistence such as learning, communication, cooperation and food sharing still remain mostly invisible in the archaeological record (Isaac, 1978; Lovejoy, 1981; Kaplan et al., 2000; Kuhn and Stiner, 2006). This is also true for variation in subsistence strategies, which can be translated into broader systems of subsistence, modes of land use and social organisation in a given ecological setting. The translation of taphonomic signals suffers from uncertainties in their interpretation. Contextualisation of these data with results obtained by other lines of research such as lithic technology, isotope studies or paleoanthropological studies remains difcult because of differences in the resolution of data obtained. Therefore, the discipline of archaeozoology remains isolated and disconnected from the anthropological and historical context (comp. e.g. Domnguez-Rodrigo et al., 2007). Numerous evidence suggests that the hunting way of life is deeply rooted in our past (Gaudzinski, 2004a; Domnguez-Rodrigo et al., 2007), suggesting that modern forms of behaviour have a long evolutionary history. Consequently special emphasis must be put on testing the functional context of archaeological sites and the social organisation behind subsistence strategies. Such a shift in research perspectives could be substantiated by a return to the holistic perspective of taphonomy established during the 1980s (Behrensmeyer and Hill, 1980; Binford, 1981; Gifford, 1981; Behrensmeyer, 1984; Andrews and Cook, 1985; Behrensmeyer Q6 and Kidwell, 1985; Koch, 1989; Lyman, 1994).

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The need for a substantial shift in research questions in faunal analyses and methodological approaches can be considered against the background on current knowledge about Neandertal subsistence in Europe. In this perspective, a detailed reconstruction of the chronology of biostratinomic and site-formation processes, in which hominins are just one variable is imperative before questions about hominin social organisation can be answered. 3. The current European evidence for Neandertal subsistence Major elements shaping the subsistence strategies of Neandertals have been uncovered in recent years through contextualisation of results obtained from analyses of the archaeological record. The discussion becomes more and more detached from the actual archaeozoological evidence. This trend results from limitations of the methodological apparatus, and only very rarely is it possible to narrow down the temporal depths within archaeozoological assemblages. Even the interpretation of relatively unambiguous in-situ archaeological evidence uncovered at the German site of Schningen (Thieme, 2008) suffers from the difculty of ascertaining the contemporaneity of the horse individuals scattered between the wooden weapons. Even though it can be plausibly argued that these are the leftovers of a hunting scenario, its social context remains obscure. It is the contextualisation of archaeozoological data with data obtained through micromorphological site analysis, dental analysis or isotope studies which helps to make predictions on this social context (Hublin and Richards, 2009). This is illustrated by major aspects of our current knowledge on Neandertal subsistence based on the archaeozoological evidence. Even though important aspects of the behavioural repertoire and social organisation of Neandertals begin to emerge in a new light (Hublin and Richards, 2009), an ensemble view of Middle Palaeolithic

subsistence is still far from being clear. Evidence from stable isotope studies of Neandertal remains (Richards et al., 2000) shows that animal products formed an important part of their diet. Competition with carnivores was apparently low (Kindler, 2007), as indicated by interpretations of analyses of isotope signatures for sites in Southwestern France. Though both hyenas and Neandertals subsisted on herd animals, Neandertals additionally focused heavily on megafauna such as rhinoceros and mammoth (Bocherens et al., 2005). However, the regular exploitation of megafauna remains is not visible in the archaeozoological record (Bratlund, 2000). As evidence from sites once perceived to reect Neandertals as obligate scavengers has been refuted (Binford, 1988; Grayson and Delpech, 1994), and new evidence of hunting discovered (Thieme, 2008), regular large-mammal hunting is now accepted as the major source of meat acquisition by Neandertals. Numerous archaeozoological studies have long established that our Middle Palaeolithic predecessors were highly specialised hunters of large mammals (Gaudzinski, 1996; Gaudzinski and Roebroeks, 2000) focussing on high return rates. This behavioural strategy can be followed in various environmental contexts (Gaudzinski, 2004b). Monospecic faunal assemblages serve to illustrate this (Gaudzinski-Windheuser, 2006). From MIS 9 or 7 onwards and especially in steppe environments during the dry-temperate climates of the late Pleistocene, archaeofaunas are dominated by a single taxon (Fig. 1). These sites reect the interception of entire herds or aggregations of many individuals at waterholes, water streams or along their migratory routes. Sites with monospecic faunas are characterised by a high dominance of remains of a single species represented by a minimum number of individuals of up to over 100 animals, associated with lithic artefacts (Gaudzinski-Windheuser, 2006). In Europe these assemblages occur in open-air sites and caves from MIS 9 onwards, although the majority of sites are recorded from the

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

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Early Weichselian towards the end of the Middle Palaeolithic. Cutmarks and hammerstone-induced impact notches attest direct hominin interaction with the fauna. It was argued (e.g., GaudzinskiWindheuser, 2006) that subsistence tactics which led to these accumulations were obviously quite varied and demanded exibility in hunting tactics and thus careful preparation and planning. Faunas from Western and Central European Middle Palaeolithic sites dominated by large herbivores attest to a selective long-term exploitation of individuals in particular age classes. Exploitation of large bovids centered on prime age individuals, whereas for megafaunal exploitation, there is a focus on young individuals (Gaudzinski-Windheuser, 2006). Subsistence tactics were linked to particular locales in the landscape which were repeatedly occupied during short visits. From the archaeozoological record, it remains mostly ambiguous if these long-term exploitation strategies were linked to a particular season. Prey selection and prey exploitation may be subject to seasonal variation as suggested by (e.g.) tooth analysis of small assemblages from Middle Palaeolithic cave sites (Rendu, 2010). From the faunal evidence of the Levant, it was even concluded that intensive hunting by Neandertals led to a decline in the populations of certain prey taxa (Speth and Clark, 2006). In contrast to these long-term strategies, there are seasonally restricted mass hunting events accompanied by carcass exploitation geared towards resources with high return rates. The faunal assemblage from Salzgitter-Lebenstedt serves as a good example. The German reindeer-dominated site of Salzgitter-Lebenstedt saw short-term mass death scenarios characterised by unselective killing of entire reindeer herds. At Salzgitter-Lebenstedt, the largest group of animals in the assemblage e adult reindeer males e died around September, as discerned from the stage in development of male antlers, as well as by means of tooth eruption stages in young individuals. The reindeer remains display abundant evidence of butchery, including lleting of meat and standardised processing of bones for marrow. During marrow extraction, a clear selection against sub-adults, as well as against bones with low marrow content occurred (Gaudzinski and Roebroeks, 2000, 2003). Detailed studies of sites with species dominated faunal assemblages give no indication that parts of the carcass exploitation sequence were associated with particular locales in the sites. Transport of particular animal parts remains largely invisible, and all stages of the butchery process can be reconstructed. Although direct evidence for intra-site organisation is poor, analyses of the bone and stone assemblages suggest that other activities beside killing and butchering must have taken place. To name a few: hide working, production and curation of stone artefacts, preparation of food, perhaps also for storage, import of gathered goods, such as collected bones for artefact production and - as some researchers suggest - for architectural purposes (Gaudzinski et al., 2005). Thus, indication is given that during the Middle Palaeolithic a clear functional differentiation of the living space did not exist as is also obvious from other aspects of Neandertal life which is e.g. expressed in the lack of living structures (Kolen, 1999). It seems highly plausible to assume that kill and butchery sites are subsequently transferred into temporal residential camps. These monotonous patterns in the Middle Palaeolithic record of central Europe which only reects Neandertal behaviour with respect to larger herbivores prevent a more comprehensive understanding of Neandertal subsistence and ecology, in particular the inuence of sympatric competitors. However, what becomes obvious from these single-taxon sites is that Neandertal land-use in the northern part of Europe seems to be highly dependent on seasonal aggregation prey, which is not obvious from bone analysis alone. Isotope studies also emphasize a specialisation of Neandertal subsistence to large migrating ungulates and megaherbivores (Bocherens and Drucker, 2003; Bocherens et al., 2005).

That seasonal exploitation must have been a driving force is also underlined by the unique evidence from the German site of Balve Cave. The cave was frequently occupied because of its logistical value, as could be demonstrated by zooarchaeological studies. The analysis 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 taxon might also indicate seasonal aggregations of Neandertal groups that conduct hunting focused on ungulate herds. These groups must have split up into smaller bands of Neandertals, retuning to logistically attractive locales in their territories, which were exploited in an opportunistic way, as seen in the case of Balve Cave and other cave sites in Central Europe (Jris, 2005; Kindler, 2007). The evidence for differences in subsistence tactics during the Middle Palaeolithic (i.e. mass hunting vs. long-term exploitation of individual age classes within a taxon) was interpreted in terms of exible resource utilization against a background of unstable environmental conditions, which is prone to equivocal interpretations in terms of highly efcient hominin adaptations, mere opportunism or a combination of both. However, the overall consolidating evidence leaves little doubt that Neandertal subsistence strategies were driven by obtaining high energy return rates, which in turn called for a high degree of exibility in the tactics employed (Gaudzinski-Windheuser and Niven, 2009). In more southern latitudes, uctuations in return rates might have been compensated by a broadening of the dietary spectrum. For more southern latitudes, the exploitation of fast-moving small prey can be demonstrated (Blasco and Frnandez Peris, 2009), the small prey spectrum being generally shown to encompass only slow moving or static prey such as tortoises, marine molluscs or eggs (Stiner, 2002; Blasco, 2008). For north-central Europe, the inclusion of small mammals and birds in hominin diet remains largely undocumented. In most cave and open-air sites, there is an underrepresentation of small mammals and birds in the archaeofauna, possibly relating to taphonomic and excavation bias. Small mammal and bird exploitation has largely remained unstudied from a taphonomic perspective due to the overall scarcity of anthropic marks on the bones. This renders it difcult to attribute singular skeletal elements of small mammals to the hominin dietary spectrum rather than to the natural background fauna (Gaudzinski-Windheuser and Niven, 2009). However, the number of sites with unambiguous evidence of utilization of small game and especially birds by Neandertals, consisting of the presence of cut-marked bones has increased in recent years due to the recognition of researchers for this important topic (e.g. Fiore et al., 2004; Dibble et al., 2009). This brief summary illustrates the major focus of Neandertal subsistence, i.e. to obtain high-energetic food, coupled with the increasing tendency for a broadening of the dietary niche. This perspective is in accordance with the potential of our methodological apparatus as was already argued. Consequently the social organisation behind these subsistence tactics is indeed addressed in most interpretations of archaeozoological data but cannot be tested. Thus the interpretation of subsistence strategies solely remains embedded in an environmental frame of reference. 4. Results: perspectives for faunal analyses The limitation in narrowing down the temporal resolution of an archaeozoological context represents a major obstacle for the interpretation of obtained data. The temporal frame of reference refers to a geological perspective which prevents us from corroborating aspects of social behaviour. As a consequence, a higher resolution of the sequence of taphonomic processes and of formation processes of archaeological sites is needed. It is this line of research which 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 and Schindel, 1983; Allison and Briggs, 1991; Bunn, 1991), and which receded into the background in recent years. The present authors strongly support the return to this holistic perception of taphonomy. Study of faunal assemblages from high-resolution archives and the detailed high-resolution reconstruction of the individual biostratinomic sequencing of a site provide perspectives for future research. Contextualisation and combination of results obtained by high-resolution analysis will reveal patterns in the archaeological record which might enable further analysis (e.g., digital modelling of site-formation processes) and can be addressed in the context of the evolution of hominin social behaviour. Studies in the Late Upper Palaeolithic illustrate this point. Some faunal assemblages from Late Upper Palaeolithic open-air sites, e.g. Gnnersdorf (Germany) (Street and Turner, in press), Oelknitz (Germany) (Gaudzinski-Windheuser, in press), Pincevent (France) (Enloe, 2003) display very high temporal resolution compared to Middle Palaeolithic sites. Detailed retting studies and correlation of retting data with the spatial record reveal individual site specic patterns interpreted in terms of human economic and social interaction (Enloe, 2003). 4.1. The study of high-resolution archives It is particularly worthwhile to study Neandertal presence in ecologically well-dened environments, at sites with high temporal resolution. Interglacial deposits represent such high-resolution archives for studying hominin behaviour. Interglacial deposits are

comparatively rare during the Middle and Upper Pleistocene in terms of percentages of the overall time of the periods (ca. 8%). There has been considerable debate about the capacities of Neandertals to survive in such forested environments (Gaudzinski-Windheuser and Roebroeks, in press). Many interglacial sites are known from Europe (GaudzinskiWindheuser and Roebroeks, in press) (Fig. 2). Many of these sites share major characteristics as their nds are embedded in a negrained sedimentary matrix and the excavated and investigated areas are large. The amount of nds is extremely high, their condition tends to be excellent, in that plant remains are preserved and bone surfaces do not show signs of (post-) depositional alteration. Many still exhibit the nest cut- and scrape-marks. The sites are spatially structured by archaeological material. Most importantly, the correlation of the palynological succession of an Interglacial with the sedimentary envelopes of the sites allows a subdivision into dened and restricted time slices, representing narrow time windows during which the accumulation of the archaeological material occurred. This is a unique situation in the Middle Palaeolithic record which distinguishes interglacial sites from other well preserved sites with nely stratied and well dated sequences allowing vertical and horizontal control when analysing faunal assemblages. The Eemian interglacial provides an especially interesting case study of Neandertal exploitation of forested habitats. This period has been subject to detailed studies, as it was hoped to gain information that could be relevant for current discussions on global climate change (Kukla et al., 2002; Shackleton et al., 2003; Brewer et al., 2008; Tzedakis et al., 2009). The Eemian started around 125,000 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 glacication is indicated in dark grey). With retreat of the Drenthe glacier numerous lake basins were formed in the underground and lled during the succeeding Eemian Interglacial.

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in the northern part of Europe (Mller, 1974; Brewer et al., 2008). In Pleistocene terms, this is a short period of time and a rather homogeneous environment. Excavations undertaken at the Eemian site of Neumark-Nord 2 illustrate the potential of interglacial deposits for disentangling the temporal resolution of Middle Palaeolithic archives. NeumarkNord 2 represents one of the rare examples among Eemian interglacial sites that were recently excavated with modern techniques where excavations were not hampered by a rescue situation. At this site, part of a shallow lake basin with shore area was excavated in ne-grained sediments. Over 120,000 well preserved bone remains representing taxa of a typical interglacial fauna were uncovered together with almost 20,000 artefacts made from Baltic int. The bones show distinct cut-marks and hammerstone-induced conical impacts that clearly indicate hominin interaction with the fauna. Results of geomorphological and palynological studies show that the accumulation of the sediments that yielded these archaeological remains could be reconstructed to have lasted less than 500 years (Kindler et al., 2010; Sier et al., in press). At Neumark-Nord 2 it will be possible to reconstruct the formation of the palimpsest represented here and it will be possible to evaluate the contemporaneity of elements of the faunal assemblage.

4.2. Testing the homogeneity of faunal assemblages An additional avenue which could be approached in order to narrow down the temporal resolution of an archaeozoological assemblage and to test whether a faunal assemblage is homogeneous in character refers to actualistic studies. These should attempt to reconstruct the individual chronology of biostratinomic factors and processes at a particular site. Such studies should try to obtain insight into the additive sequencing of a taphonomic process. These studies must pay attention to individual siteformation histories that can be recognised in archaeological sites, as well as incorporate already known contexts, linking the actualistic study to the taphonomic history of a given faunal assemblage. Thus, the specic mode of operation of a taphonomic process at a given site rather than the general recognition of patterns of taphonomic signals must be central to these studies.

In this context, the results of actualistic studies undertaken for the Early Middle Pleistocene Israeli site of Gesher Benot Yaaqov (for a compilation of the current state of analysis, see Alperson-Al et al., 2009) have successfully demonstrated the homogeneity of the faunal assemblage, with important consequences for the overall interpretation of the site (Gaudzinski-Windheuser et al., 2010; Rabinovich et al., 2008, in press). Tumbling and trampling experiments were initiated to mimic the individual environmental conditions at the site, to gain qualitative insights into processes of bone modications and to assess the timing of the biostratinomic chronology. It was assumed that both mechanisms were responsible for the formation of striations documented on the bone surfaces documented for Layers V-5 and V-6. Efforts succeeded in replicating the chronology of biostratinomic factors and processes at the site and, for the rst time, were able to obtain insight into the additive sequencing of a taphonomic process (Gaudzinski-Windheuser et al., 2010; Rabinovich et al., in press) (Fig. 3). These observations have major implications for taphonomic studies as they suggest that interpretations based on separated isolated taphonomic processes may be misleading. Against a biostratinomic background, bone surface modications result from a process in which bone surfaces are continuously altered. This is implying that interpretations of biotically- and abiotically-induced modications according to diagnostic templates but without their integration into their complete individual biostratinomic sequence are highly ambiguous. Recent studies on isolated taphonomic features, e.g. the frequency of cut-marks (Domnguez-Rodrigo and Yravedra, 2009) and percussion-marks (Galn et al., 2009) on bones, illustrate the high variability of these traces which makes their verication and quantication more difcult. These studies acknowledge the modifying qualities of other additional subsequent and superimposing biostratinomic processes. They attempt to replicate individual site specic taphonomic signals with the aim to make general prediction on their origin from a superordinate perspective. It is not attempted to analyse these signals in the individual biostratinomic context. As a result interpretations of signicant correlations 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 Yaaqov, 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 (after Lyman, 1994, 358).

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Fig. 4. Trampling experiment 5 simulating environmental conditions at the Middle Pleistocene site of Gesher Benot Yaaqov, Israel. Hyena ribs, a) before trampling, b) after 5 h of trampling. Reduction of the bones into a brous 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 still remain confronted with equinality. A consequent mandatory problem concerns the question of whether certain taphonomic patterns are diagnostic or not with regard to specic processes and agents. Due to their individual structure and morphology, bones that pass the same biostratinomic sequence can display completely different patterns of bone preservation and bone modication, regarded as signicant for taphonomical interpretations. Studies of climatically induced weathering and abrasion usually outline that the degree of bone destruction correlates with the time between the death of an individual and its nal burial. As a consequence, homogeneity in bone surface preservation is considered an indicator for the assumption of homogeneous environmental conditional and/or homogeneity of an entire bone assemblage. In our experiments at Gesher Benot Yaaqov a very short abrasive process produced certain diagnostic bone surface modications which are indistinguishable from bone surface modications due to long-term climatically induced weathering (Fig. 4). Moreover it was demonstrated that the process of trampling is characterised by formation, modication and erosion of traces sharing the same morphology as hominin cut- and carnivore tooth-marks. In addition, cut-marks and most likely also tooth-marks, can be altered by the abrasional process to the degree that they nally mimic striations. As a consequence, strict determination of the time of cessation of the biostratinomic process is mandatory for diagnosis and interpretation of bone surface modications. Against this background generalised interpretations of fossil assemblages deprives them of their individual history and leaves us without their potential scientic value (Gaudzinski-Windheuser et al., 2010; Rabinovich et al., in press). 5. Summary and discussion In recent years numerous disciplines, among them biochemical and/or paleoanthropological studies, as well as the contextualisation of the overall archaeological record have vastly improved knowledge of food acquisition and exploitation and thus subsistence strategies by Neandertals. The results of the different disciplines are complementary, as becomes obvious when comparing results obtained from isotope- and archaeozoological studies emphasising the ecological specialisation on medium to large sized herbivores. However, only archaeozoology has the potential to translate variation in subsistence into a comprehensive overall perspective of Neandertal behaviour. Bones as the major source of archaeozoological analysis represent part of the archaeological record and provide direct access for the recognition of hominin actions and behaviour at Pleistocene sites. Thus, archaeozoology still remains the critical discipline for the study of subsistence behaviour. However, in spite of a steadily growing methodological apparatus and innovations in analytical techniques, archaeozoology still fails to provide a comprehensive picture of the social organisation of Neandertal subsistence, based on the different results obtained at individual sites. In consequence, the Neandertals way of life remains ambiguous and bloodless. Part of the dilemma results from the loss of a holistic approach in taphonomical research. This holistic perspective resulted from research attempts to answer questions concerning the functional context of sites representative for particular forms of social behaviour (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 soon replaced by the hunting/scavenging research paradigm that has

in the meantime exhausted its potential as a motive for faunal analysis. As long as the research focus is not directed towards the motives and the concomitant aspects of any subsistence behaviour entangled in the archaeological record, archaeozoological research will not prot from methodological improvements in a non-holistic perspective and will only continue to recapitulate the current state of knowledge. The other aspect of the dilemma described above results from the fact that archaeozoological methodologies have become more and more disconnected from the reconstruction of hominin behaviour. Research over the last decades has resulted in a large body of actualistic and comparative data describing the character and timing of individual taphonomical processes. These data help to identify the individual processes and agents that shaped the character of a given faunal assemblage. This line of research does not attempt to identify and/or arrange coherent and self contained chronological sequences of taphonomical processes involved in the formation of fossil accumulations. Without their integration into a complete biostratinomic sequence, interpretations of bioticallyand abiotically-induced modications on bones according to diagnostic templates remain highly ambiguous. Thus actualistic taphonomic research must acknowledge the individual characteristics of Pleistocene sites, which form the determining variables that shape the individual taphonomic sequencing of an assemblage. This research must become integrated into a comprehensive palaeoecological analysis of the embedding milieu. The quality of this approach can be tested by investigating sites embedded in spatial high-resolution milieus, as can be found in interglacial deposits. As taphonomical research is not able to clearly dene the temporal resolution of a given faunal assemblage, high-resolution milieus provide a temporal frame of reference within which biostratinomic processes can operate. High-resolution milieus bear the potential for the identication of entire biostratinomic sequences. This is a point of crucial importance as taphonomic processes act additively. Within the biostratinomic chain, individual signals caused by particular actions can be overprinted by succeeding actions, the signals of which will consequently not be identied as a taphonomically-relevant pattern. It is therefore imperative to test reconstructions of biostratinomic processes at a given archaeological site by means of actualistic studies simulating the depositional history of this site. Only if taphonomic research appreciates the immense potential of the character of the embedding milieu at any given site as a reference frame for analysis, can taphonomical chains be identied, and thus the role of hominins more precisely evaluated. Acknowledgements We like to thank Eudald Carbonell, Maria Gema Chacn and the IPHES working group for inviting us to the workshop The Neanderthal Home: Spatial and Social Behaviours. Daniela Holst and Radu Iovita provided valuable feedback. We would like to thank Rivka Rabinovich and Naama Goren-Inbar for inviting us to participate in the research at Gesher Benot Yaaqov. Finally our thanks go to all members of our Neumark-Nord working group. References
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Please cite this article in press as: Gaudzinski-Windheuser, S., Kindler, L., Research perspectives for the study of Neandertal subsistence strategies based on the analysis of archaeozoological assemblages, Quaternary International (2011), doi:10.1016/j.quaint.2010.11.029