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CAN ANATOMICALLY-MODERN HUMANS BE USED AS ANALOGUES FOR NEANDERTAL FORAGING PATTERNS?

Benjamin Collins

IntroductionCurrent frameworks and models for interpreting past foragers’ subsistence patterns have been constructed through extending ethnoarchaeological and ethnographic observations of modern peoples (Binford, 1978; David and Kramer, 2001; Hudson, 1993; Kelly, 2007). A strong critique of this approach during the 1970s and 1980s (Gould, 1978, 1980; Wobst, 1978) has led to both a refinement in the extent to which modern foraging behaviour is extended to past foragers, and more accurate interpretations of past foraging behaviours (Wylie, 1985, 2002). These models have also been used to interpret the behaviours that are preserved in archaeological and zooarchaeological assemblages attributed to other hominins, perhaps uncritically so.

Neandertals present an interesting case, as they are the most closely related hominin to anatomically-modern humans (AMH). This close relationship is demonstrated by both genetic (Green, et al., 2006, 2010; Krause, et al., 2007; Noonan, et al., 2006) and morphological research (Conroy, 2005; Harvarti, et al., 2004; Trinkaus, 2007),

with the genetic data indicating that the two populations diverged roughly 400,000 years ago and were capable of interbreeding. However, it has been strongly contested that Neandertals lack the cognitive complexity associated with AMH (Barnard, et al., 2007; Klein, 2008, 2009; Mellars, 2005, 2006; Wynn and Coolidge, 2004).

Are interpretive models based on studies of AMH foragers therefore suitable for interpreting Neandertal foraging behaviours? This question will be addressed by assessing the arguments for differences and similarities between Neandertal and AMH cognition, with a focus on the archaeological record, biological differences and cognitive models. A recent study will also be discussed to demonstrate how the ethnographic record can be used to infer Neandertal behaviour.

AMH and Neandertal archaeological assemblagesThe archaeological record has been extensively used to argue for significant cognitive differences between

Within the past 25 years, ethnographic and ethnoarchaeological studies of modern foragers have been critically used to create and refine models for inferring past human foraging behaviours. These models have also been extended to interpret the foraging behaviours of other hominins. However, critical evaluations of whether ethnographic and ethnoarchaeological studies of AMH are relevant to other hominins are rare. This article assesses whether the foraging patterns of Neandertals, our closest relatives, can be accurately inferred when using models derived and refined from studies of anatomically-modern human (AMH) foragers.

It is suggested that Neandertal foraging patterns can be accurately inferred when using optimal foraging models derived from behavioural ecology and refined with ethnographic and ethnoarchaeological studies of AMH. However, potentially symbolic or cultural aspects of Neandertal foraging behaviours are contended to be beyond the scope of these models, as the differences between AMH and Neandertal cognition are yet to be understood.

2. Can anatomically-modern humans be used as analogues for Neandertal foraging patterns? 9

Neandertals and AMH, a position that stems from the ‘symbolic explosion’ associated with the Upper Palaeolithic cultural tradition (UP, approximately 40 kya to 10 kya) (Klein, 2009; Mellars, 2006). The UP is represented by the appearance of new lithic technologies and material culture (see Table 1) that are either absent or rare during the Middle Palaeolithic (MP, approximately 250 kya to 40 kya) (Mellars, 1996, 2005). Material symbolic expression and complex behaviour are considered a uniquely AMH phenomenon, as AMH are argued to be exclusively associated with the UP (Klein, 2008, 2009; Mellars, 2005, 2006). Neandertals are considered to be solely represented by MP material culture in Europe and only contentiously associated with several transitional industries that have UP characteristics, such as blades and personal ornaments (Bar-Yosef and Kuhn, 1999; d’Errico, 2003; d’Errico, et al., 1998; Mellars, et al., 2007; Zilhao, et al., 2007).

The paucity of symbolic expression recovered during the MP has led to Neandertals typically being considered cognitively inferior to AMH. However, d’Errico and colleagues (2003, 2009), amongst others (Speth, 2004; Wolpoff, et al., 2004), have noted that there are sporadic and increasing indications of Neandertal symbolic behaviour during the MP. For example, over 40 MP sites yield evidence of pigment use, with over 500 pieces of ochre being recovered from Pech de l’Azé I and IV in France (Soressi, et al., 2007). Incised bone fragments are also becoming more common, as are personal adornments, such as shell beads and worked animal teeth (Burke and d’Errico, 2008; Davies and Underdown, 2006; d’Errico, et al., 2003, 2009; Zilhao, et al., 2010). In fact blades, once used as an indicator of UP AMH archaeological assemblages, are now frequently found in both MP and African Middle Stone Age (MSA) contexts (Bar-Yosef and Kuhn, 1999; d’Errico and Stringer, 2011; McBrearty and Brooks, 2000).

Comparisons between MP and UP faunal assemblages also provide an interesting perspective. UP AMH assemblages are argued to contain a diverse range of fauna, indicating broad spectrum foraging patterns that incorporate all aspects of the environment, including small and hard to catch prey, aquatic resources, seasonal prey and large, dangerous prey

(Klein, 2001; Mellars, 1996, 2005; Stiner and Munro, 2002). This has been contrasted with the MP, during which Neandertal faunal assemblages appear to be composed predominately of large herbivores Hockett and Haws, 2005; Patou-Mathis, 2000; Richards and Trinkaus, 2009).

Isotopic studies comparing Neandertals and UP AMH have frequently distinguished between the two groups, with Neandertals displaying specialised, terrestrial carnivore diets and UP AMH broader diets that included aquatic resources (Richards, et al., 2001; Richards and Trinkaus, 2009). Furthermore, a recent study involving a Neandertal skeleton recovered from an underwater coastal context did not give any indication of an aquatic dietary component, furthering the specialised, terrestrial carnivore hypothesis (Hublin, et al., 2009).

Other zooarchaeological studies contest this hypothesis and instead suggest that some Neandertal faunal assemblages indicate a broad spectrum diet, similar to UP AMH (Bar-Yosef, 2004; Stringer, et al., 2008). These studies have more closely related the past environmental conditions to the faunal assemblages and found that Neandertals living in warmer, non-steppe environments were acquiring aquatic and small prey, and using plant resources (Adler, et al., 2006; Blasco, 2008; Blasco and Fernandez Peris, 2009; Henry, et al., 2011; Lev, et al., 2005; Madella, et al., 2002; Stringer, et al., 2008). Furthermore, several zooarchaeological studies have demonstrated continuity in subsistence strategies across the terminal MP and early UP in steppe environments (Adler, et al., 2006; Grayson and Delpech, 2003; Morin, 2008).

Neandertal zooarchaeological assemblages can be argued to indicate that Neandertals were able to forage in a similar manner to AMH. However, there are still some important differences regarding the social and cultural aspects of MP and UP foraging. As noted above, the UP is characterised by symbolic expression, which is also a strong feature of modern hunter-gatherer subsistence (Bird-David, 1999; Brightman, 2002; Ingold, 2000; Willerslev, 2007). This is indicated by UP cave art, zoomorphic figures and modified faunal remains, such as pierced animal tooth pendants. Evidence for symbolic aspects of Neandertal foraging is sparse and represented by a small number of perforated animal teeth and some modified bone in late MP or controversial transitional assemblages (Burke and d’Errico, 2008; d’Errico, et al., 2009; Mellars, et al., 2007; Zilhao, et al., 2007).

The similarities and differences between Neandertal and AMH foraging patterns necessitates careful consideration, as the zooarchaeological assemblages suggest that Neandertals and AMH both potentially foraged in a similar fashion. However, the absence of a substantial symbolic component associated with the Neandertal assemblages suggests some cognitive differences may have been present.

Table 2.1. The major archaeological differences between the UP and MP (Mellars, 1996, 2005; Klein, 2009). Note that symbolic artefacts include person, portable and parietal art

Upper Palaeolithic Middle Palaeolithic Blade-focused industries Flake-focused industries Complex spatial patterning Simple spatial patterning Symbolic artefacts common Symbolic artefacts rare Broad spectrum subsistence strategies

Narrowly-focused subsistence strategies (focus on medium to large herbivores)

Elaborate burials Simple burials

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AMH and Neandertal biologyThere are several physiological characteristics that are argued to differ between AMH and Neandertals. Most important for this discussion are developmental growth rates and skull morphology.

Developmental growth rates are thought to have differed substantially between Neandertals and AMH and potentially reflect different life cycles, social structures and group mobilities (Bird and O’Connell, 2006; Conroy, 2005; O’Connell, et al., 1999; Pettit, 2000). However, Ponce de Leon and colleagues (2008) have recently argued that despite Neandertal brains being larger than those of AMH, both finish growing at around the same age. The authors suggest that Neandertals had similarly slow or possibly even slower life histories. An interesting caveat to this study is the contention that the smaller AMH brain was re-organised to become more efficient, possibly resulting in a competitive advantage over the larger and more costly Neandertal brain (Ponce de Leon, et al., 2008, p. 13767).

Neandertals and AMH exhibit significant differences in the external shapes of their skulls, with Neandertals generally demonstrating greater robusticity, especially in the brow ridge and occipital regions. This results in a platycephalic, or long and low-shaped Neandertal cranial vault, in contrast to the globular-shaped vault characteristic of AMH. AMH also generally display more gracile cranial features and a pronounced chin (Conroy, 2005; Stringer and Gamble, 1993; Schwartz and Tattersall, 2000; Trinkaus, 2007).

Differences in the external shape of the brain case have been used to infer variation in internal brain morphology, particularly in the frontal cortex and occipital regions (Wynn and Coolidge, 2004). The frontal lobes and frontal portion of the brain are argued to play a significant role in speech and language production, which is an important condition for AMH cognition (Amanti and Shallice, 2007; Davies and Underdown, 2006; d’Errico, et al., 2009). Similarly, the temporal-parietal area differs in both hominins and is also argued to play a major role in the ability to symbolise and for abstract thought (Henshilwood and Dubreuil, 2011; Wynn et al., 2009). If different brain morphologies existed and had functional consequences, Neandertal communication and cognitive abilities may have been, at the very least, qualitatively different from AMH.

Weaver and colleagues (2007) have reached an interesting conclusion in their comparison of Neandertal and AMH cranial differences. The authors sought to determine whether the external morphological variation present in AMH and Neandertal crania was the result of genetic drift or natural selection. Weaver and colleagues (2007) concluded that they could not reject their null hypothesis: morphological differences between AMH and Neandertal crania were the result of genetic drift and not natural selection. However,

there are two caveats to this study: it tested for natural selection’s role in producing variation and not maintaining the same general shape and functional morphology; and it focused on the external morphology only, and not the internal morphology, which has a greater bearing on cognition.

Roseman and colleagues (2011) have also compared human and Neandertal cranial morphology. In this study, the authors compared the patterns of cranial integration, or the covariance of specific cranial traits, between W.W. Howell’s collection of AMH cranial data and data from a set of 20 Neandertal skulls. Using these data, the authors demonstrated that AMH and Neandertals share a similar pattern of cranial integration for roughly three-quarters of the 37 traits studied and differed significantly for several others, specifically those related to the frontal and occipital bones. The authors conclude that their results indicate that the two hominins reacted differently to evolutionary forces (Roseman, et al., 2011).

Davies and Underdown (2006, p. 149), however, have noted issues with ‘paleo-phrenological’ approaches, as external differences in cranial morphology have not been shown to elicit significant cognitive differences amongst AMH. Two studies directly support this argument, with Bookstein and colleagues (1999) and Semendeferi and Damasio (2000) both demonstrating that despite significant differences in external cranial morphology amongst several hominoid species, frontal lobe morphology has remained relatively stable over time. This conclusion has been supported by other palaeo-neurological research that has also failed to detect significant differences between the neural development of Neandertals and AMH (Holloway, 1995; Semendeferi, 2001).

However, recent research by Bruner (2010), has demon-strated that the parietal portions of the skull and the parietal lobes of the brain in AMH are distinct from other hominins. AMH generally have larger parietals enabling parietal lobes that that are larger and more integrated with other parts of the brain. Bruner (2010) notes that the parietal lobes are involved with the interpretation of external spatial environments and its integration with inner perception, for example internal mental images, imagined worlds, thought experiments and abstract representations. These are all associated with enhanced working memory (Wynn and Coolidge, 2004; and see below) and modern cognition (Henshilwood and Dubreuil, 2011), and may represent different modes of cognition between AMH and Neandertals.

Modelling Neandertal and AMH cognitionWithin the past decade, Enhanced Working Memory (EWM) has been a very influential concept for describing the evolution of AMH cognition (Wynn and Coolidge, 2004). Wynn and Coolidge (2004) in particular have developed a

2. Can anatomically-modern humans be used as analogues for Neandertal foraging patterns? 11

model that uses both archaeological research and cognitive neuropsychology to posit an understanding of AMH and past hominin cognitions (Davidson, 2010).

Wynn and Coolidge (2004; Coolidge and Wynn, 2005) use the concept of working memory to propose a model that differentiates between AMH and Neandertal cognitions (Baddeley, 2001; Baddeley and Logie, 1999). Working memory is described as a tripartite cognitive system consisting of a phonological storage system, related to speech information; a visual-spatial sketchpad, relating to visual and spatial information; and a central executive that is responsible for maintaining attention and decision-making, and is located in the frontal lobes (Wynn and Coolidge, 2004, p. 469).

The authors propose that differences in working memory can be used to distinguish between Neandertal and AMH cognitions and that these differences are reflected in the archaeological record. To briefly summarise their model, Wynn and Coolidge (2004) suggest that a genetic mutation in AMH approximately 50 kya created an ‘enhanced’ working memory capacity and modern cognitive abilities, a position also advanced by Klein (1995, 2001, 2008, 2009). Enhanced working memory (EWM) is reflected by an increase in the capacities of the central executive and an expanded phonological storage system. Together, these enhancements are argued to have resulted in greater native and fluid intelligences, better attention spans and the development of the subjunctive, or ‘what if?’ mode of speech. Wynn and Coolidge (2004; Coolidge and Wynn, 2005) further argue that this would have facilitated thought experiments, increased innovative potential, and created introspection, all of which are strongly related to increased cultural complexity and symbolic expression (see Rossano (2009) for a discussion of EWM’s role in symbolic ritual).

Introspection and self-reflection are argued to be particularly important and uniquely human, as in conjunction with the subjunctive mode of speech, they establish the conditions for reflecting on past experiences and extending them to the future. Wynn and Coolidge (2004) argue that AMH cognition is thereby reflected in the establishment of highly abstract, symbolic culture, illustrated by the UP in Europe and Later Stone Age in Africa. And that Neandertals, while demonstrating complex and very similar cognitions to AMH, lacked EWM and were therefore limited in their innovative and symbolic capacities.

Much of the EWM model has been constructed using studies of individuals with frontal lobe damage, as this is the suggested site of the central executive portion of working memory. Wynn and Coolidge (2004; Coolidge and Wynn, 2007) are careful to note criticisms that brain-damaged AMH are not the best models for Neandertals (Beaman, 2007). However, they contend that the research on impaired frontal lobe neural structures, in combination with the archaeology, provides a heuristic window into the Neandertal mind.

Other recent models of Neandertal-AMH cognition largely draw upon the EWM concept as discussed above (Amati and Shallice, 2007; Barnard, et al., 2007; Cole, 2009; Davidson, 2010; Henshilwood and Dubreuil, 2011; Rossano, 2009). These models all form the same conclusion: AMH display more complex cognitive abilities, reflected by a fluid intelligence that is ascribed to an internal dialogue. This dialogue facilitates complex reflection on the past and future, and provides the foundation for an enhanced innovative capacity (see also Mithen, 1996).

The archaeology, zooarchaeology and cognitive models have all pointed to both broad similarities and particular differences in Neandertal and AMH cognition. These differences are posited to be in the AMH capacity for abstract thought and internal reflection manifested through symbolic expression. However this position has been contested with the presence of some symbolic artefacts associated with Neandertal occupations (d’Errico, 2003; d’Errico and Vanhaeren, 2007; Zilhao, 2006; Zilhao et al., 2010), as well as the genetic closeness of Neandertals to AMH (Green, et al., 2010).

While Neandertal expressions of symbolism are not as ubiquitous as those associated with UP AMH, some similarities arise when they are compared to contemporaneous AMH from the African MSA. The MSA and MP display similarities in lithic technology, as well as with the paucity of symbolic materials. On this basis, Neandertals and MSA AMH were thought to be cognitive ‘equals’, with AMH cognition and behaviour only appearing with the transition to the Later Stone Age around 40 kya to 50 kya (Klein, 2001; Mellars, 2006; Willoughby, 2007).

However, recent excavations focusing on the African MSA have provided substantial archaeological material that can be related to symbolism and complex cognition. McBrearty and Brooks (2000) provided a summary of some of these materials over a decade ago and demonstrated how the African MSA’s material culture argued for the presence of modern cognition. More recently, excavations from Blombos Cave (Henshilwood, et al., 2002, 2009), Pinnacle Point (Marean, 2010; Marean, et al., 2007) and Sibudu Cave (Wadley, 2007, 2010a, 2010b; Wadley and Plug, 2006; Wadley, et al., 2009) have all demonstrated archaeological remains that are used to argue for complex symbolic (modern) behaviour (see Table 2). Furthermore, the Still Bay (c.73 kya to c.70 kya) and Howieson’s Poort (c. 65 kya to c. 59 kya) lithic industries are argued to bear strong resemblance to those from the LSA and UP (Jacobs, et al., 2008a, 2008b). These similarities include styles that span large geographical regions and are temporally limited; and the frequent occurrence of microliths and backed tools in the Howieson’s Poort (Wadley, 2007; Lombard, 2011; Villa, et al., 2009; Wurz, 1999). Both industries are associated with symbolic expressions and are preceded and followed by hiatuses.

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The saltational occurrences of symbolic behaviour during the MSA have recently been characterised as demographic pulses, with larger populations consisting of well-connected sub-populations capable of information sharing displaying more complex behaviour, followed by smaller, less well-connected populations displaying less complex behaviour. Both the greater population size and the ability to share information are argued to facilitate novel and more complex cultural innovations to be acquired, spread and maintained (Powell, et al., 2009; Richerson, et al., 2009). The much used example of the decrease in complexity of the lithic technology used by the Tasmanian Aborigines fits well with this argument (Henrich, 2004).

Taking into account the hypothesised low population densities for Neandertals during the MP, may explain why symbolic expression is rare and patchy during this period and increases substantially in the archaeological record with the transition to the UP, which is associated with the arrival of AMH and increased demographic pressure (Adler, et al., 2006; d’Errico, 2003; d’Errico, et al., 2003; Hockett and Haws, 2005; Zilhao, 2006). d’Errico and Stringer (2011) have recently contested that the patchy appearance of complex, ‘modern’, behaviours during the MP may be linked to climatic and environmental variation. In this regard, symbolic or ‘modern’ behaviours appear as adaptations to eco-cultural niches, which contract and expand through time based on demographic and environmental change.

A complicated picture arises in trying to understand Neandertal cognition and behaviour. What can be said is that despite differences reflected in the archaeological record, Neandertals are not, to put it bluntly, stupid and must have been quite similar to AMH in order for interbreeding to occur (Green, et al., 2010). Potential differences in cognition should then be considered to reflect cultural variation, rather than different levels of cognitive complexity. This conclusion merits further consideration for the extent to which frameworks and models used for understanding AMH foraging behaviours can be extended to Neandertals.

Interpreting Neandertal behaviourAnalogical reasoning is fundamental for the construction of archaeological models and interpretive frameworks, whether explicitly acknowledged or not (Wylie, 1985, 2002). Ethnoarchaeology contributes insight into past behaviours by studying modern peoples and the use of analogical extension, either directly (formal analogic reasoning), or more commonly indirectly (relational analogic reasoning) (David and Kramer, 2001). Through combining ethnoarchaeological studies with explanatory frameworks from evolutionary theory, behavioural ecology can be used to construct models for interpreting past subsistence practices, specifically those that have arisen from optimal foraging theory (Bird and O’Connell, 2006; Lupo, 2007; Kelly, 2007; Winterhalder and Smith, 2000). These models were developed in the biological sciences, and have been borrowed to interpret human behaviour.

Optimality models are, however, fundamentally reductionist and, as such, necessarily limit the number of variables that are taken into account. This approach often yields false hypotheses (see Sih and Christensen (2001) for a review), which can act as heuristics for further research. Archaeologists have therefore used ethnographic and ethnoarchaeological research to refine these optimality models and make them more applicable to understanding both present and past hunter-gatherers’ foraging decisions (for example: Broughton, 2002; Faith, 2008; Grayson and Delpech, 1998, 2003; Morin, 2008; Nagaoka, 2002, 2005). In this regard, can optimal foraging models refined using ethnoarchaeological studies of AMH be used to infer Neandertal subsistence strategies?

This question depends strongly on two factors: the extent to which AMH and Neandertals are cognitively similar and the extent to which the subsistence behaviours being studied can be related to differences in cognition. The above comparison of Neandertal and AMH cognitions, based on the biological and archaeological information, as well as the proposed cognitive models has elicited a complicated picture.

Table 2.2. Recent Middle Stone Age finds that demonstrate symbolism and complex cognition

Artefact How it demonstrates symbolic thought References Engraved ochre and ostrich eggshell

Intentionally engraved abstract design that has no functional purpose. Possibly art.

Henshilwood et al., 2002, 2009; MacKay and Welz, 2008; Texier, et al., 2010

Shell beads No functional purpose. Used as pendants for displaying personal and group identity.

Henshilwood, et al., 2004; d'Errico, et al., 2005, 2008

Compound tools Multi-component, compound tools require complex mental preparation and planning.

Wadley, 2010b; Haidle, 2010; Reuland, 2010

Trapping and snaring Requires ability to make traps and snares, and long-term planning and scheduling to place them correctly and remember to check them

Wadley, 2010a; Clark and Plug, 2008

Ochre use Ochre could be used as adhesives for multi-component tools, and for symbolic painting of body and objects

Watts, 2010; Wadley, et al., 2009;

Marine resource use Requires specialised technology and long-term scheduling Marean, 2010; Marean, et al., 2007; Henshilwood, et al., 2002

2. Can anatomically-modern humans be used as analogues for Neandertal foraging patterns? 13

Genetics, parts of the endocranial skull morphology and the faunal assemblages indicate a substantial degree of cognitive similarity between the two hominins. However, substantial differences are found in material symbolic expressions, ubiquitously associated with UP AMH, but only sparsely so with Neandertals. While this difference may be related to differences in demographic intensity (d’Errico and Stringer, 2011), it has also led to models proposing that Neandertals lacked complex, abstract thoughts and internal dialogues (Barnard, et al., 2007; Wynn and Coolidge, 2004). Cognitive models and the archaeology therefore necessitate a consideration of the validity for extending AMH foraging behaviours to Neandertals.

Taking the above into account requires that any attempt to infer more cultural aspects of Neandertal subsistence must necessarily be met with caution. If Neandertals are characterised by a different cognitive system (which appears likely, although these differences may be more akin to cultural variation than cognitive complexity), then they have no modern analogue (Barnard, et al., 2007). This weakens any attempt to infer the social and cultural contexts surrounding Neandertal foraging behaviours, especially since such contexts have been strongly linked ethnoarchaeologically to AMH forager decision making (Egeland and Byerly, 2005; Lupo, 2006; 2007). However, behavioural ecology and optimal foraging models can be used to infer the less cultural aspects of Neandertal foraging behaviours.

Behavioural ecology and optimal foraging models employ an evolutionary framework that focuses on fundamental behavioural patterning conditioned by natural selection (Bird and O’Connell, 2006; Winterhalder and Smith, 2000). Optimal foraging models, such as the diet-breadth model, patch-choice model and central-place foraging model (see Kelly, 2007, pp. 73–110), used in a reflexive manner can be used to establish a link between AMH and Neandertal behaviour, especially considering these models’ reductionist tendencies. This reductionism is what gives optimal foraging models their analytical power (Bettinger, 2009), as they do not incorporate the symbolic aspects of foraging, which differ immensely between different cultures and more than likely between Neandertals and AMH (Kelly, 2007; Ingold, 2000).

Dusseldorp (2012) has recently provided a detailed study of several Neandertal faunal assemblages from Europe. This study used optimal foraging models, specifically the diet-breadth model to assess the relationship between climate change and changes in Neandertal foraging strategies in northern European steppe and forest environments. Through analysing the species that were hunted and their relative abundances, Dusseldorp was able to demonstrate that solitary species were focused upon during the warmer periods, while gregarious species were focused on during colder periods.

Dusseldorp (2012) uses ethnographic and ethnoarch-aeological information to explain these foraging strategies. Modern hunter-gatherers will tend to aggregate and increase group size when large herds of animals are present, and disperse and decrease group size when they are not (Kelly, 2007). These strategies are related to the amount of animal biomass present in the landscape with the presence of large gregarious animals increasing the animal biomass and therefore being able to provide more food for more people.

Using the ethnographic information to interpret the archaeological record, it can be inferred that Neandertals were engaging in similar foraging strategies as AMH. Neandertals specifically exploited large herd animals when they were abundant during colder and more open conditions. To efficiently exploit these species requires large group sizes, which is reflected in the archaeology from these sites.

When the climate and environment changed to warmer and more forested conditions, herd animals were not ranked as highly and solitary species were more intensively exploited. This change in foraging strategy is linked to a decrease in group size as the available animal biomass decreased and could not support large groups.

Dusseldorp’s (2012) study demonstrates how aspects of the ethnographic and ethnoarchaeological records can be used to infer some Neandertal behaviours, such as the relationship between foraging strategies and group size. However, these records cannot be extended to infer the social and cultural changes that accompanied these behavioural shifts and further reinforces the limits to which AMH behaviours can be used to understand those of other hominins.

ConclusionWylie (1985, 2002) and others contend that strong relational analogies are constructed using multiple cables or arguments that all point to the same conclusion (Dominguez-Rodrigo, 2008; Gifford-Gonzalez, 1991; Stahl, 1993). An interpretive framework using behavioural ecology and optimal foraging models supports this contention, as it is based on fundamental behavioural similarities patterned by natural selection (Bettinger, 2009; Bird and O’Connell, 2006). This framework is strengthened by the addition of ethnographic and ethnoarchaeological studies of AMH foraging patterns, and environmental and other contextual information.

The crucial issue considered by this paper is whether these models, refined with ethnoarchaeological data from AMH and used for AMH, are applicable to Neandertals, who are considered to be equipped with a unique cognitive structure. As optimal foraging models largely focus on basic, evolutionary-patterned foraging behaviours, and Neandertal zooarchaeological assemblages closely resemble those of AMH (Adler, et al., 2006; Grayson and Delpech, 2003; Morin, 2008), it is argued that they can be reliably extended

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to interpret Neandertal faunal assemblages. This conclusion was furthered by discussing Dusseldorp’s (2012) research on the relationship between Neandertal group size, climate and environmental change and changes in foraging strategies.

The strength of the behavioural ecology interpretive framework is argued to weaken when ‘higher-order’ (Hawkes, 1954) cultural and social interpretations are attempted, stemming from possible differences between AMH and Neandertal cognitions. AMH demonstrate internal, reflexive dialogues that facilitate abstract, symbolic thoughts and behaviours. These complex cognitive abilities have been argued by some to be absent in Neandertals, thereby impeding the interpretative scope of their social and cultural behaviour (Barnard, et al., 2007; Klein, 2008; Wynn and Coolidge, 2004). This position however, is strongly contested (d’Errico, 2003; d’Errico and Vanhaeren, 2007; Zilhao, 2006).

Analogical reasoning, ethnoarchaeology, behavioural ecology and cognitive modelling have all been considered to establish the basis for and limits of an approach to inferring Neandertal foraging behaviours from faunal assemblages. The above do not consist of the only aspects contributing to a sound interpretation, with taphonomy, palaeoecology, the palaeoenvironment and the archaeology all being extremely important. These data and other potential sources of information should be continually considered, as it will facilitate a better understanding of Neandertal behaviour and the development of hominin cognition.

AcknowledgementsI would like to thank Lee Broderick, Dr Steven Churchill, Dr Lyn Wadley, Dr Manuel Dominguez-Rodrigo and an anonymous reviewer for their constructive comments on an earlier version of this manuscript, as well as comments from the Archaeology Graduate Students Writing Group from McGill University. I would also like to acknowledge financial support from a Social Sciences and Humanities Research (SSHRC) Doctoral Fellowship and the Hominid Dispersals Research Cluster.

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