Are Lithics and Fauna a Match Made in (Prehistoric) Heaven?



Lithic artifacts and animal bones form the bulk of the material remains of the Paleolithic. This has led archeologists to interpret these two types of finds as tethered components of subsistence systems. Differences observed through time and space in the lithic repertoire were considered as functional adjustments, designed to maximize gains from a diverse faunal resource base. While we do not challenge the general notion that lithic artifacts were used (also) for exploiting faunal (and other) resources, we note that significant lithic technological breakthroughs, clearly directed towards higher efficiency of procurement of faunal resources (e.g., hafting, projectile weapons), are few and far-apart in the evolution of material culture. Based on case studies from the Levantine Middle Paleolithic (MP), we question the degree to which fauna-based subsistence determined lithic variability. Current research focus on functional relationship between lithics and fauna may preclude consideration of other causes for lithic typo-technological diversity. These may include technological traditions, differences in cultural transmission processes, or the level of within- and between-group connectivity, various combinations of which may have operated in the MP.


Middle Paleolithic Lithic diversity Faunal resources Functional paradigm 


Lithic artifacts and animal bones form the bulk of the material remains from the Paleolithic period. They are often interpreted as tethered components of subsistence systems. Accordingly, differences in the lithic repertoire, observed through time and space, are considered as functional adjustments, designed to maximize gains from diverse faunal resource bases.

Such an explicit emphasis on the functional link between stone artifacts and subsistence is a relatively new concept in Paleolithic archaeology. As part of the zeitgeist of the early days of archaeology and until the mid-twentieth century, human survival through the production and use of material culture was axiomatically perceived as the main characteristic of humans, that which made them unique in the animal kingdom and placed them apart from (and above) other species, “substantially distanced himself from his animality […] as a […] culture dependent hominid” (Tobias 1991:845). An early case in point is that of Australopithecus africanus, which Dart (1925) endowed with an “osteodontokeratic” material culture. Perhaps the most striking example of this attitude (as late as the 1960s) is the inclusion of dexterity for stone toolmaking as one of the defining characteristics of Homo habilis (Leakey et al. 1964).

By the same token, technology-assisted faunal procurement was deemed an inseparable part of the human condition since the appearance of the earliest stone tool assemblages. Prehistorians have inferred a causal relationship between stone tools, hominin carnivory, and the cognitive and cultural evolution of the hominin lineage from early days of Paleolithic research (see Ambrose 2001; Isaac 1971).

For researchers of the late nineteenth and early twentieth centuries, the functionality of tools was axiomatic rather than a questionable issue. The typological classifications of modified artifacts exemplify this attitude. Lithic morphotypes were defined by names that connoted function with no doubts expressed as to the factual validity of such designations (Dark 1995; Trigger 2006).

Major changes occurred in the mid-twentieth century with the appearance of quantification and statistical descriptions of assemblages (e.g., Bordes 1950, 1961; Roe 1969) and the “New Archaeology” paradigm (e.g., Binford 1973; Clarke 1968). Emphasis shifted from observations of the archeological record to attempt to understand the processes underlying its formation. Within this emerging worldview, all archeological phenomena had to be explained, not “merely” described (though see Gould 1989:97–100 on the importance of descriptive monographs in historical sciences). Much of the research has shifted to the construction of formal hypotheses, anchored in a plethora of disciplinary theories (e.g., anthropological, sociological, and biological) that could be tested against the archeological evidence (see also Wiessner 2016). Questions about the ubiquitous co-occurrence of stone artifacts and animal bones throughout the prehistoric record were now formally addressed from the perspective of a functional “cause-and-effect” relationship. In the context of Middle Paleolithic (MP) studies, this difference between the older approach of unquestioned assumptions and the new processual thinking is best epitomized by the Bordes-Binford debate in the 1970s (Binford 1973; Bordes and de Sonneville-Bordes, 1970; and see Wargo 2009).

We do not challenge the general notion that lithic artifacts were made for functional purposes, or that stone tools were used—already in deep prehistoric time—for the procurement and exploitation of faunal as well as other resources. This, however, is not the same as saying that changes in techno-morphological properties of lithic artifacts, or of lithic assemblage structure, were driven by the need to maximize gains from faunal exploitation, a way of thinking that characterized much of the twentieth century archeological discourse.

From a diachronic perspective, there are but few and widely separated significant lithic technological breakthroughs that are unambiguously linked with increasing efficiency of faunal resource procurement and exploitation (e.g., hafting, stone-tipped weapons, and projectiles). On the other hand, one can observe many shifts in typological diversity, and technological variation in lithic assemblages throughout the prehistoric record. We contend that emphasis on the functional relationship between lithics and the exploitation of fauna as the default explanation for changes in lithic variation precludes consideration of other potential causes of the observed phenomena. The diversity of material culture among extant hunting and gathering groups—so often unrelated to functional considerations—serves as a cautionary tale (see below).

In this paper, we bear on the MP Levantine record to illustrate the difficulties arising from a still-prevalent emphasis on a strictly functional relationship between lithics and fauna, considered as a main driver of the prehistoric record in general, and of stone tools variability in particular. Our review suggests that on a number of different analytical levels, the inter-relationship between lithic variability and the exploitation of faunal resources is far from being as straightforward as often posited in the literature. We then suggest potential alternative avenues for understanding the variability of the MP lithic record.

The Archeological Record

A Note About Analytical Approaches

Hypotheses pertaining to relationships between stone tools and human subsistence are based on two main traits of the archeological record—artifact form and context.

Preliminary perceptions about the ways stone tools were used stem from artifact morphologies. Regardless of formal typological classifications, different configurations of lithic artifacts have different kinetic potentials, more conducive to particular motions (i.e., linear cutting motions, angled motions in scraping; e.g., Hovers 1997; Kuhn 1995). Binford and Binford’s (1966; Binford 1973) arguments about the functional nature of Mousterian variability were established on such premises.

A second line of relevant archeological evidence is that of context. Stratigraphic and spatial associations between stone tools and animal or plant remains trigger inferences about their functional relationship.

Such preliminary inferences are further evaluated through four additional and independent lines of enquiry—ethnographic analogy, microwear, residue analysis, and experimentation. Notably, microwear and residue analyses constitute formal tests of functional hypotheses, whereas the other approaches serve to narrow down the range of processes that potentially could have led to the formation of the observed phenomena. Thus, a number of analytical tools were developed to address function-form relationship. To evaluate the effectiveness of armature pieces1 and to understand how diachronic changes in their designs may be linked to increasing effectiveness in meat procurement, a number of analytical parameters were constructed. These focused on diagnostic impact fractures (DIF) on the tips of stone points as the result of their use as hunting armature (Shea 1988, 1993, 1997). In addition, Welner lines, identified microscopically on stone points, could be used to assess impact velocity of the hunting weapon (Hutchings and Bruchert 1997). Experimental and ethnographic data suggested that lower velocity indicated the use of stone points as tips of thrusting spears or javelins, whereas higher velocity indicated their use as darts and projectiles (Hutchings 2011; Hutchings and Bruchert 1997).

Another analytical approach focuses on armature delivery systems. The notion that points are elements of weapons led to the construction of quantitative measures of their penetration potential and ballistic properties. Hughes (1998) showed that the higher the value of its tip cross-sectional area (TCSA), the lower the projectile’s flight velocity, reducing its effectiveness, whereas a lower TCSA value improves projectile penetration. Another metric measure, tip cross-sectional perimeter (TCSP), was more recently argued to be a better proxy of the force needed to lethally penetrate a target (Sisk and Shea 2011). Furthermore, an extensive ethnographic survey conducted by Hughes (1998) showed that variation in TCSA values is linked to different classes of lithic armatures. Shea (2006) and Sisk and Shea (2011) subsequently argued that TCSA values could be cautiously used to assign a delivery system of projectiles.

MP Points

Form and Function Arguments

“Points” (and more specifically, projectile points) is a category of artifacts that has long captivated the attention of prehistorians, ethnographers, and ethnologists (e.g., Iovita and Sano 2016; Knecht 1997; Pétrequin and Pétrequin 1990). Armature items have drawn such attention not the least because their use is associated with high cognitive abilities and complex behaviors (see below). In the early days of archaeology, the association of armature with hunting as well as warfare was based on the morphological similarities with armatures known from ethnographic contexts. This analogy held sway across the twentieth century paradigm changes, and more recently was paraphrased as a rule of thumb, invoking the principle of uniformitarianism (Shea 2006).

The appearance of pointed morphotypes is a hallmark of the MP/Middle Stone Age (MSA), distinguishing it from the preceding period in the Levant and elsewhere (Hovers and Belfer-Cohen 2013; Malinsky-Buller 2016a, b; McBrearty and Brooks 2000; Tryon and Faith 2013).2 It has also been a common a priori assumption that production and use of points from the early MP/ MSA onwards co-occurred with the systematic use of hafting techniques.3 Combined, these two technological novelties have been repeatedly viewed as portraying true weaponry and as a major improvement in the efficiency of hunting technology, contributing to hominins’ cognitive evolution and the broadening of their ecological niche (Coolidge et al. 2016; Lombard 2005). As a direct result of the importance assigned to pointed morphotypes, archeologists have engaged in attempts to find analytical evidence to support such perceptions.

Publications related to the early MP site of Misliya Cave serve to illustrate the notion about the link between the shapes of stone points and faunal resource acquisition. The faunal assemblage of Misliya Cave exhibits clear evidence for hunting (Yeshurun et al. 2007). Zaidner and Weinstein-Evron (2014) divided the points in the associated lithic assemblages into seven typological groups, based on their shapes and dimensions. This typological diversity was attributed to blank selection and purposeful design in the process of making each type rather than to a reduction continuum. Ensuing microwear studies and TCSP calculations of four typological point groups from the site (“Levallois,” “Abu Sif,” “Hummal,” “Misliya”; Yaroshevich et al. 2016) suggested that the variability in point forms is related to their use as components of different weapons: “Thrusting or throwing spears, as well as darts may have been in use, presumably for different game or biotopes. Faunal evidence supports this possibility.” (ibid. 130). Referring to this study, Zaidner and Weinstein-Evron (2014:20) stated: “… the small points (Misliya points) could be used as tips of hafted projectiles, while larger points (e.g., Levallois points, Abu Sif points) as tips of thrusting spears.” In contrast, Groman-Yaroslavski et al. (2016:318) claimed, based on residue analyses, that a “… comprehensive use-wear analysis of Abu Sif points [from the same assemblages] found almost no indication of their having been used as spear points, or even as tools somehow related to hunting (for example, butchering at the kill site).” Rather, these points were related to the acquisition of vegetal foods and to craft-related activities (ibid.). This conclusion is congruent with experimental results by Shea et al. (2001:814), following which they suggested that Levantine early MP Levallois points “… were probably designed for use as knives”.

Zaidner and Weinstein-Evron (2014) noted that the morphological richness of points in later Levantine MP assemblages is much lower, as armatures found in these later assemblages (e.g., Umm el-Tlel, Qafzeh XV, Kebara, Amud, Quneitra; Hovers 2009: Table 8.3 and references therein; Pagli 2010) are nearly exclusively Levallois points. Although Shea (1991, 1995) identified those items (in Qafzeh, Kebara, Tor Faraj, dating to MIS 5–3) as multi-functional tools based on microwear analysis, it is their role as elements of armatures that has been emphasized in subsequent publications (e.g., Shea 1993, 1997, 1998; Shea et al. 2001). Levallois points in the Levantine later MP were suggested to have been used as spearheads, based on the similarity of their calculated TCSA to those of experimental thrusting spears. Indeed, their TCSA values are high (see Shea 2006: Fig. 7), unlike those of ethnographic dart tips or arrowheads.

An Organizational System?

Many researchers have looked at the connections between behaviors of resource search, procurement, and transport, linking them with the manner by which hominins coped with ecological constraints and possibilities. The production, use, and discard of stone tools are regarded as outcomes of complex considerations, including economic decision-making and optimization aimed at making tools efficiently available when and where they are needed. Stone tools are perceived as having been designed and used to minimize risk and effort and to maximize gains in resource exploitation. In most reconstructions of MP subsistence and mobility, modes of game procurement are (explicitly or implicitly) implicated in the timing and occurrence of various technological behaviors (e.g., Chase 1986; Kuhn 1995; Speth 2004; Speth and Clark 2006; Stiner and Kuhn 1992). Strategies of tool design (“maintainable” and “reliable” tools and toolkits) and of raw material economy (provisioning of locales vs. provisioning of individuals, curation, expediency) are regarded as having been adjusted to the environmental conditions, resource use, and mobility patterns (e.g., Bamforth and Becker 2000; Binford 1979, 1980; Bleed 1986, 2002; Kelly 2013; Nelson 1991; Parry and Kelly 1987).

Within the framework of this burgeoning literature, researchers dealing with the Eurasian MP record have tried to link modes of mobility and meat acquisition with changing technological procedures of tool production. For example, in their diachronic study of lithic and faunal assemblages from four Italian cave sites, Stiner and Kuhn (1992; Kuhn 1995; Stiner 1994) linked flaking from platform cores with reduced mobility and more emphasis on hunting, whereas use of centripetal cores was linked to higher mobility and greater reliance on scavenging. A somewhat similar approach was taken by Delagnes and Rendu (2011), who argued that various lithic flaking systems (e.g., Levallois and laminar; Quina; discoidal-denticulate; and the Mousterian of Acheulian Tradition) reflected distinct mobility strategies as a response to different hunting practices. This case study, from a single region and from a restricted time window, is based on associations of faunal and lithic evidence but the causal relationship is not well elucidated, as noted by the authors themselves.

In the context of Levantine MP studies, organizational models focusing on relationship between faunal resources and stone tools have led to some additional, far-reaching claims, beyond economic strategies. Lieberman and Shea (1994) looked at differences in armature frequencies, the occurrence of DIF, and the results of seasonality studies, attempting to tie these data with behaviors of the local MP human populations (anatomically modern humans or Neanderthals). They concluded that local Neanderthals hunted more frequently and were more residentially mobile compared to modern humans. Moreover, on the basis of an assumed relationship between hunting methods and lithic repertoire,4 they had suggested that in the Levant, the adaptation of Neanderthals was possibly more predatory than that of early modern humans (ibid.).

Interestingly, the European Neanderthals, who have been promoted in the literature as top carnivores (based on isotopic studies, tooth wear studies, faunal remains, and a plethora of optimization and ecological reasoning; e.g., Stiner 1994, 2002, 2006; Wißing et al., 2016), do not exhibit a similar organizational behavior. The frequencies of armatures in European lithic assemblages are generally low (e.g., Shea 2006, 2009 and references therein; Villa et al. 2009) and their identification as such often relies on unsupported assumptions (Rots and Plisson 2013; and see Rots et al. 2016). In an attempt to explicate this conundrum, Shea (2006) turned to broad demographic and ecological perspectives. He reasoned that in the small groups of the European Neanderthals, the effects of personnel losses/injuries when interacting with dangerous animals were magnified and highly disadvantageous compared to those effects on larger groups, and therefore avoided more strictly. Also, due to low population densities, coalitionary killing in inter-group conflicts was not regularly adopted. Hence, the minimal use of armatures.

This explanation begs the following:
  • If armatures were not essential for hunting activities, why should we hold on to the notion that they are?

  • Why should one accept that economic organizational systems were structured for maximizing gains from the acquisition of faunal resources?


A Functional Impasse

That prehistorians focus on stone tools and bones as the target of their research is a default of the archeological archive, given that typically there is little else for us to work with (Hovers 1998), especially in deep prehistory. This should not be confused with a presumption that stone tools were made primarily for obtaining meat or for exploiting faunal resources by butchering, filleting, etc. And yet, despite an expanding body of theoretical discussions, nutritional and ethnographic data (e.g., Speth 2010 and references therein; and see below) arguing for a lesser importance of meat in human evolution, a notion persists that the exploitation of faunal resources was a prime mover in human biological, cognitive, and material culture evolution (e.g., Braun 2010; McPherron et al. 2010; Plummer 2004; Plummer and Bishop 2016; Schick and Toth 2009; Stanford and Bunn 2001; and see also Aiello and Wheeler 1995).

While “[w]e know [our emphasis] that Early Paleolithic stone tools were used from about 2.7 Myr onward to aid early hominins in butchering animal carcasses” (Shea 2007:224), this knowledge does not derive from morphological or other properties of the tools but from stone tool-induced marks on bones (ibid.) and does not allow for a more specific identification of the tools themselves. From periods as remote as the African Early Stone Age, microwear and residue analyses repeatedly suggest that stone tools were used for diverse purposes, more directly related to maintenance than to subsistence (Hayden 2008; Shea 2007). The sparse archeological data speaking to the early use of lithics for obtaining meat (at Dikika some 3.4 myr ago; McPherron et al. 2010) are controversial (Domínguez-Rodrigo and Alcalá, 2016; Njau 2012; Sahle et al. 2017), and the earliest stone tools recognized to date (at the Lomekwi site 3.3 myr ago; Harmand et al. 2015; Lewis and Harmand 2016) comprise mainly heavy anvils and hammers suitable for the processing of hard vegetal materials.5 During the first 600 kyr of the Oldowan, conspicuously few modified bones attest to any form of treatment of animal tissue using stone tools (Domínguez-Rodrigo and Martínez-Navarro 2012; Hovers 2012). This is inconsistent with the pivotal role often assigned to faunal exploitation in early human evolution (e.g., Braun 2010; Plummer and Bishop 2016; McPherron et al. 2010; Schick and Toth 2009).

With a diachronically growing repertoire of stone tool standardized morphologies, explicit predictions were formulated about patterned co-variations between bones and stone tools due to causal, functional relationship. In the context of the MP, many researchers argued that Levallois points present a relatively standardized form, intentionally shaped by a particular reduction process for triangular blanks (e.g., Boëda et al. 1990; Goval et al. 2015, to name but a few). These artifacts are repeatedly regarded as stone tips of hunting weapons based on their morphology, in tandem with experimentation, DIF patterns, and TCSA/TCSP calculations. Notably, these analytical approaches, with which such form-function assumptions could be tested, provided contradictory results due to methodological problems but also because the items were used for more than a single function. In their review of the methods of use wear studies, Rots and Plisson (2013) highlight a series of problems in the very same methods that are designed to expand on the functional links between armatures and faunal resources. They caution that the underlying assumptions of such claims are rather unfounded. Namely, researchers have looked at only a few types of damage patterns (focusing mainly on “tip damage” burination as DIF); or applied inadequate experimentation procedures (e.g., small sample sizes used to replicate and explain variability in damage patterns); or too often neglected to consider and test the damage patterns of alternative activities, which could result in similar patterns.

Given such analytical uncertainties, it is small wonder that controversies exist as to the causes of observed impact fractures, as to whether they should be considered as diagnostic of hunting (e.g., Douze 2014; Plisson and Beyries 1998 and debate within; Sahle et al. 2013), or even as to the identification of the materials on which they were used. Similarly, recent studies of ethnographic material and experimental results suggest that TCSA/TCSP statistics are not robust measures of projectile performance, or reliable proxies for inferring delivery systems (Clarkson 2013). For example, a study of hafted stone projectile points and knives demonstrated that Australian aborigines did not construct their tools with the aim of optimizing TCSA values, thus challenging TCSA-based narratives about projectile weaponry (Newman and Moore 2013).

The Misliya Cave case study, where the same items are argued to have been either a hunting tool stone tip (based on microwear and TCSA) or a tool for procuring vegetal materials (on the basis of residue analysis), exemplifies such research shortcomings. Similarly, a series of stone tools (Levallois points included) from the later MP site of Amud Cave was subjected between the years 2001–2004 to independent microwear analyses of three specialists, using comparative as well as different methods. These analyses resulted in three different use-wear identifications suggested for each artifact (Hovers, unpublished report).

In the particular case of Misliya, a broader archeological context can be used to weigh on the issue. Should the assumed correlation or co-variation exist between the various armature types and faunal resources, one would expect the Misliya faunal record to show high variability compared to the faunal assemblages of the later Levantine MP, when point morphological variability is reduced compared to Misliya (and the early MP in general). However, the faunal studies do not lend support to the hypothesis of causal relationship between faunal assemblage structure and point morphological diversity—the Misliya bone assemblage resembles later MP ones in species composition, age structure, and the distribution of anatomical parts (Yeshurun et al. 2007). Additionally, change to arguably more efficient hunting technologies after 40 thousand years does not seem to be associated with changes in the structure of faunal resources. Recognizable variations in Levantine MP faunal assemblages correlate with geographic areas and site functions (e.g., Hovers and Belfer-Cohen 2013; Sharon et al. 2014), but not directly with point abundances, let alone their morphological variability, in the assemblages (Hovers 2009:214–218).

As mentioned above, in the Eurasian MP sphere beyond the Levant, there is weak evidence for projectile weaponry among Neanderthals. In a diachronic survey of points in MP assemblages in northwestern Europe, Goval et al. (2015:229-230) consider assemblages with up to 3% points as presenting a “ubiquitous” occurrence of pointed types; and according to their microwear studies, these are more related to butchery than to hunting.

If one accepts that MP pointed forms and faunal resources are linked functionally, the scarcity of the former in the European record is puzzling. Neanderthals in Europe have been repeatedly argued to be highly carnivorous and to have relied on various modes of meat acquisition as a main way of making a living. In such scenarios, the use of stone armature would have been beneficial. For example, reducing the need for close confrontations with dangerous animals would result in lower risks to the hunting individuals.

In sum, lithic items for cutting and pounding purposes likely existed throughout the archeological record, with faunal tissues being but one of the processed materials. Highly specialized tools emerged post-MP times (Shea 2006; Shea and Sisk 2010; Stiner 2013). As indicated by the (imperfect) analytical tools at our disposal, the functions of these tools are not necessarily linked to their morphologies (e.g., Lemorini et al. 2006, 2014; Shea 1991) beyond the general kinetic properties of artifacts. Bleed’s (1986) distinction between reliable and maintainable tools clarifies that the optimal design for armatures is that of the reliable elements—highly specialized items involving complex preparatory procedures including the preparation of hafts, adhesives, etc. In Eurasia and Africa, complex technological processes were applied in order to haft MP points (Lombard et al. 2005; Shea 1993; Wadley et al. 2009). However, flakes and retouched items, not necessarily used in fauna-related activities, were hafted in similar manners (e.g., Boëda et al. 2008; Friedman et al. 1994-5; Rots 2013; Rots et al. 2011). MP stone tools, armatures among them, likely had multiple uses (e.g., Holdaway 1989; Plisson and Beyries 1988; Shea 1988, 1991; and see reviews in Dibble et al. 2017; Nelson 1991). The cumulative evidence does not support an important role for faunal exploitation as a trigger of MP lithic morphological or assemblage variability.

Ecological and Organizational Perspectives

If a form-function relationship does not provide sufficient explanation for the variability of lithics, it is still possible that systemic behavior (i.e., organizational systems) was driven by costs and benefits involved in the acquisition and exploitation of faunal resources. Series of influential publications from the 1990s onwards (see above) promoted this view by causally linking mobility, meat procurement, and lithic production as elements of optimizing economic strategies.

We do not question the notion that organization of the circulation of lithic raw material and of blanks within a settlement system was structured to obtain economic gains (Perlès 1992; Soressi and Geneste 2011). At question is the underlying assumption that such gains (or constraints) were defined by the presence, predictability, or abundance of faunal resources within a land-use system. It is therefore important to ask whether this was indeed the case, as otherwise much of the organizational argumentation for the links between lithic and fauna is undermined.

As mentioned above, the focus on faunal remains in explanatory scenarios of the prehistoric record may have more to do with the limitations of the archeological record than with behavior, namely the much better preservation of faunal remains as opposed to other types of resources. More specifically for MP research, a Eurocentric research tradition associated the MP time frame with Neanderthals, who arguably subsisted within a dietary niche focused on carnivory and big-game hunting (e.g., Kuhn and Stiner 2006; Stiner 1994, 2002, 2013, and references therein).

This notion does not hold for the Levantine MP. The Levant’s location in the low middle latitudes suggests that food-procurement systems of hunter-gatherers in the region would focus on plants more than on meat resources (e.g., Binford 2001; Cordain et al. 2000). The structure of the faunal communities available to MP groups (typically territorial species) coupled with the region’s complex interplay between climate and topography all rendered the spatiotemporal distribution of plant resources less predictable than that of the main prey animals (Goring-Morris et al. 2009). Therefore, the location and availability of plant resources would be more influential than those of animal resources in determining the timing and extent of group mobility (Hovers 2009 and references therein). Also, the technological organization would be influenced by plant resource structure and distribution more than by the faunal ones (Hovers 1997, 2009:201–204).

The status of the Neanderthals as specialized high ranking carnivores has recently come into question also in regions outside the Levant. It has been argued that their diet was broader and included whatever resources were available in the local environment on a seasonal basis and according to climatic conditions (Bar-Yosef 2004; El Zaatari et al., 2016; Krueger et al. 2017; Salazar-García et al. 2013). A high-protein, meat-based diet would have severe effects on pregnant Neanderthal females and thus be unfavorable to the population’s reproductive success (Hockett 2012), which is inconsistent with Neanderthal prolonged survival in Europe. Furthermore, current evidence from a number of case studies suggests that Neanderthals enjoyed a relatively variable diet, incorporating plant and possibly aquatic resources (e.g., El Zaatari et al. 2011; Henry et al. 2014; Krueger et al. 2017; Lev et al. 2005; Madella et al. 2002; Naito et al. 2016; Weyrich et al. 2017).

Given the insights emerging from these new research directions, arguments favoring a rigid organizational link between MP stone tools and the exploitation of faunal resources across Eurasia require reconsideration. “The evidence forces us to decouple suppositions about … hunting capabilities and complex hunting weapons” (Stiner 2013: S295).

So What Are the Alternatives?

In ethnographic case studies, variability in material culture (inclusive of stone tools) is not linked exclusively to functional considerations. Newman and Moore (2013:2618) concluded that “persistence of apparently suboptimal (our emphasis) TCSA values in Aboriginal stone spear armatures is probably due to the conservative nature of technological practices”. Tool designs should be considered in the context of the reduction sequences that produced them and the dynamics of transmission of those reduction sequences across generations. Moreover, functional traits often bear symbolic meanings that are as important as the function itself.6

“Technological recipes” may be persistently favored in inter-generational transmission due to their functional benefits, but also because of conservatism, anti-novelty bias, or biases related to the status of the model from which the variant is learned rather than to its functional properties (see reviews in Mesoudi 2011, 2015; Mesoudi and O’Brien 2008; O’Dwyer and Kandler 2017). The non-reversible, reductive nature of lithic production makes it susceptible to knapper errors, such that it required learning through imitation as a means of faithful copying (Schillinger et al. 2015, 2016).

As a rule, MP group sizes were rather small (Bocquet-Appel and Degioanni 2013 and they were dispersed over relatively large territories. Hence, the interactions and “connectivity” between groups, and opportunities for acquiring novel technological “recipes”, were relatively limited. As a result, MP stone tools show variations through time in the recombination of a small number of characteristic technological traits (Hovers 2009:245; Hovers and Belfer-Cohen 2013; Kuhn 2013; Stiner 2013; Stiner and Kuhn 2006). In general, long-term directional material culture changes in the Levantine MP are rare and are not linked to variations observed in the structure of the faunal assemblages. This pattern is consistent with the concept of “rugged fitness landscape” (Palmer 1991), which explains non-directional changes as local adjustments to maintain fitness equilibria following disruptions (environmental and/or demographic), without increases in the level of fitness (Hovers 2009:245; Kuhn 2006). At times, however, particular technological “recipes” can be recognized as local technological styles (e.g., Krakovsky 2017 for the late MP in the Levant; Mackay 2011 for the Howeisons Poort in South Africa). Such observations suggest that we should consider that lithic variability is influenced also by social dynamics and cultural transmission (for a detailed discussion of this see Hovers and Belfer-Cohen 2013).

To sum up, prehistoric stone tools were made for functional purposes, and “forms of some tool types do hint at some kind of functional specificity” (Kuhn 1995:17). Our review suggests that in the case of the MP, methods developed to address this question more precisely were based on an assumption that exploitation of faunal resources and formal or organizational dimensions of lithic variability were inherently related. At close scrutiny, this does not stand the weight of the available reliable evidence. The relationship that emerges from this review between aspects of stone tools and the exploitation of faunal resources is generalized, in line with the MP lack of directional technological change. Other, non-functional causes for lithic typo-technological diversity may include the local occurrences of technological traditions formed through variable modes of cultural transmission, or the level of within- and between-group connectivity.


  1. 1.

    Here, we follow Rots and Plisson (2013) and use the term “armature” as a general designation of pointed morphotypes.

  2. 2.

    The dates (ca. 500 ka) assigned to the pointed artifacts from layer 4a, Kathu Pan 1 (Wilkins et al. 2012) are exceptionally old. This Fauresmith assemblage may represent an early MSA occurrence (ibid.), albeit based on current data it is a rather isolated one.

  3. 3.

    Evidence for earlier hafting was reported from the early Middle Pleistocene Acheulian in the Levant and from the late Middle Pleistocene in Italy (Alperson-Afil and Goren-Inbar 2016; Mazza et al. 2006), but to date, these are sporadic occurrences.

  4. 4.

    When the preferred method of obtaining meat was intercept hunting, investment in the making and using (i.e., transporting, hafting) of stone armatures was deemed more advantageous. Among less mobile groups, where encounter hunting was more common, such investment was disadvantageous (Shea 1998).

  5. 5.

    Whether stone tool use by extant non-human primates simulates a plausible precursor for hominin technological behavior (as some would argue about chimpanzee stone-assisted nut-cracking; e.g., Panger et al. 2002) or an evolutionary convergence (as is the case with capuchin monkeys stone-toolmaking; e.g., Falótico and Ottoni 2016; Proffitt et al., 2016) is not relevant for the current discussion.

  6. 6.

    Archeologically, this has been recognized only in the MSA record (e.g., Lombard 2007).



We thank Tamara Dogandžić, Geoff Smith, Karen Ruebens, and Teresa Steele for inviting us to participate in the session “Connecting Middle Paleolithic datasets: the interplay of zooarchaeological and lithic data for unravelling Neanderthal behavior” in the 82 SAA annual meeting, and for their invitation to contribute to this special issue. We are grateful to Yehuda Cohen for his critical reading of the manuscript and his helpful suggestions.

Compliance with Ethical Standards

Conflict of Interest

The authors declare that they have no conflict of interest.


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Copyright information

© Springer International Publishing AG, part of Springer Nature 2018

Authors and Affiliations

  1. 1.The Institute of ArchaeologyThe Hebrew University of JerusalemJerusalemIsrael
  2. 2.International Research Affiliate, Instittue of Human OriginsArizona State UniversityTempeUSA

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