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Home , Cleaver (Stone Age tool), Flake tool, Ground stone, Lithic technology, Lomekwi, Lower Paleolithic

doi 10.4436/jass.95017 JASs Invited Reviews Journal of Anthropological Sciences Vol. 95 (2017), pp. 67-108

Form and function in the Lower Palaeolithic: , progress, and continued relevance

Alastair J. M. Key1 & Stephen J. Lycett2

1) School of and Conservation, University of Kent, Canterbury, Kent, CT2 7NR, United Kingdom e-mail: [email protected] 2) Department of Anthropology (Evolutionary Anthropology Laboratory), University at Buffalo, SUNY, Amherst, NY 14261, U.S.A.

Summary - Percussively flaked stone artefacts constitute a major source of evidence relating to hominin behavioural strategies and are, essentially, a product or byproduct of a past individual’s decision to create a with respect to some broader goal. Moreover, it has long been noted that both differences and recurrent regularities exist within and between Palaeolithic stone artefact forms. Accordingly, archaeologists have frequently drawn links between form and functionality, with functional objectives and performance often being regarded consequential to a ’s morphological properties. Despite these factors, extensive reviews of the related concepts of form and function with respect to the Lower Palaeolithic remain surprisingly sparse. We attempt to redress this issue. First we stress the historical place of form–function concepts, and their role in establishing basic ideas that echo to this day. We then highlight methodological and conceptual progress in determining artefactual function in more recent . Thereafter, we evaluate four specific issues that are of direct consequence for evaluating the ongoing relevance of form–function concepts, especially with respect to their relevance for understanding evolution more generally. Our discussion highlights specifically how recent developments have been able to build on a long historical legacy, and demonstrate that direct, indirect, experimental, and evolutionary perspectives intersect in crucial ways, with each providing specific but essential insights for ongoing questions. We conclude by emphasising that our understanding of these issues and their interaction, has been, and will be, essential to accurately interpret the Lower Palaeolithic archaeological record, tool-form related behaviours of Lower Palaeolithic hominins, and their consequences for (and relationship to) wider questions of .

Keywords - Artefacts, Morphology, Flake, Biface, Handaxe, Stone-tool Function.

Functional concepts within Lower (i.e. waste), their production was nonetheless the Palaeolithic result of flaking a stone object for functional pur- poses. Consequently, functional concepts have Palaeolithic stone artefacts are the product of maintained a prominent position within studies a past individual’s intention to modify a natural of lithic artefacts (Semenov, 1964; Keeley, 1980; such that it is capable of performing a defin- Odell 1981; Torrence, 1989a; Nelson, 1991; able objective or activity. That is, they are a prod- Kuhn, 1994; Shea, 2007; Roche et al., 2009; uct of creating a tool capable of modifying an Gowlett, 2011a; Braun, 2012; Key, 2016). Most aspect of an individual’s physical or social envi- notably, archaeologists have frequently drawn ronment in respect to some broader goal. Hence, links between form and functionality, with while many artefacts excavated from Palaeolithic functional objectives and performance often sequences may have been considered a byproduct being regarded as consequential to a stone tool’s

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morphological properties. In turn, ‘function’ is their potential ability to influence the percep- frequently presented as an explanatory hypoth- tion and behaviour of other individuals or esis for the forms of stone artefacts recovered groups within hominin social systems. Hence, in archaeologically. theory, change or stability in the formal proper- In many respects, function may be consid- ties (material, mass, size, shape, etc.) of artefacts ered a principal influence on stone tool form through time can be explained, in part, by selec- during the Lower Palaeolithic. Certainly, their tive processes that increase or maintain a tool’s role as raises important questions relating to performance characteristics within a given func- the choices underlying their production and how tional context (Meltzer 1981; Schiffer & Skibo this relates to their ability to be applied to utilitar- 1987, 1997; O’Brien et al. 1994; Key & Lycett, ian tasks and the wider behavioural strategies of in press; Lycett et al. 2016). Performance charac- hominin populations (Torrence, 1989b). Given teristics may be defined as “an interaction-and- the potentially contentious of implying activity-specific capability of a person or artefact” ‘function’ to be a principal determinate of stone (Schiffer et al. 2001, p.731). In this way, a tool’s tool morphologies, it is important to stress that performance characteristics describe its ability to the term ‘principal’ need not imply that function undertake specific techno-, socio- and ideo-func- is necessarily the most frequent cause of variation tions, based on its formal properties (Schiffer & between artefactual forms, or even the variable Skibo 1997). Moreover, such various functional with the strongest potential influence. Indeed, categories are not necessarily mutually exclusive an important role of the present review is to and could be incorporated simultaneously into highlight limitations to the explanatory power artefactual forms (Sackett, 1977). Accordingly, of form–function relationships and the fact that the interaction between a tool’s formal attributes in many situations the application of functional and performance could, in principle, lead to the models should not only be done with caution, exertion of selective pressure on artefactual form but often leads directly to the need to consider (O’Brien et al., 1994). other explanatory hypotheses. Despite these factors, extensive reviews of The ‘function’ of a lithic artefact within Plio- the related concepts of form and function with contexts was potentially diverse and respect to the Lower Palaeolithic remain sparse. could accordingly be defined too narrowly. As This could be considered surprising given that noted by Schiffer & Skibo (1987), artefactual stone artefacts constitute one of the most impor- ‘function’ may be considered in either utilitar- tant data sets relating to hominin behaviour and ian (techno-function), social (socio-function) the central role that function may have played or ideological (ideo-function) terms. Functional in forming that record. Here, we attempt to discussions within an archaeological context do, redress this issue. First we stress the historical then, imply the use of an artefact for a particular place of form–function concepts, and their role purpose. Most functional studies of lithic arte- in establishing basic ideas that echo to this day. facts are concerned with the use of stone tools However, we show how little progress beyond during cutting behaviours, where the forceful listing plausible functions was made in the early application of an object (usually with a clearly phases of the discipline. We next, therefore, defined edge) to material results in fracturing highlight methodological and conceptual pro- and deformation of that material. The vari- gress in determining artefactual function in more ability of potential cutting actions undertaken recent years. Thereafter, we evaluate four specific by stone is diverse and includes issues that are of direct relevance to determining slicing, cleaving, piercing, grinding, scraping whether form–function interactions were of rel- and drilling, among others (Atkins, 2009; Key, evance to Lower Palaeolithic hominin behaviour. 2016). Their possible role as percussive tools In one sense, the notion of form and function or as projectiles has also been discussed, as has with regard to the Lower Palaeolithic may seem A. J. M. Key & S. J. Lycett 69

quaint or even outdated entirely. In conclusion, large flakes and possess a sharp, straight edge at therefore, and based directly on the issues raised, one end, which is left from the original flake sur- we attempt to untangle the extent to which face (Isaac, 1977; Clark & Kleindienst, 2001). form and function may have been entwined in Many studies have focused on documenting and Lower Palaeolithic stone technologies. Moreover, analysing size and shape variation within and we assess the continued relevance of these con- between handaxe and assemblages distrib- cepts for Palaeolithic archaeology, especially with uted across and Eurasia (e.g., Isacc, 1977; respect to their ongoing relevance for under- Roe, 1981; Wynn & Tierson, 1990; Saragusti et standing human evolution more generally. al., 1998; McPherron, 1999; Vaughan, 2001; McNabb et al., 2004; Sharon, 2007; Lycett & Gowlett, 2008; Lycett, 2008; Chauhan, 2010; Lower Palaeolithic Stone Iovita & McPherron, 2011; Shipton & Petraglia, 2011; Wang et al., 2012; Gowlett, 2011a, 2015), The Lower Palaeolithic is characterised—at identifying both similarities and differences in its most basic level—by the production of large artefactual forms. The appearance of ‘Levallois’ (>10cm) or small flake (<10cm) tools (modified prepared-core (or Mode 3) technologies at least or unmodified) and bifacially flaked core tools, 300 KYA is often regarded to signify the end produced by means of direct percussion using of the Lower Palaeolithic and beginning of the hard (lithic) or soft (bone/antler/wood) ham- Middle Palaeolithic (Moncel et al., 2011; Tryon mers and/or through bipolar percussion or anvil & Faith, 2013). (passive) techniques. The potentially oldest and While the above discussion is by no means deliberately flaked stone-tool (although a definitive account of Lower Palaeolithic tech- see Proffitt et al., 2016), are the recently described nological or morphological variability (see e.g., 3.3 million--old large flakes (and associated Semaw et al., 2009; Hovers, 2012; Gowlett, cores) from 3, West Turkana, 2015; Harmand et al., 2015 for discussion of (Harmand et al., 2015). From 2.6 million years broader and additional features), it is clear that ago, the Palaeolithic record is characterised by there is considerable variation in the size and ‘’ assemblages and becomes more abun- shape of stone tools that are being produced by dant in terms of the number of artefacts and sites hominins during the Lower Palaeolithic. As has (Hovers & Braun, 2009; Roche et al. 2009; Rogers long been recognised, an obvious question to & Semaw, 2009). Although ‘Oldowan-like’ (or address is, therefore, whether some of that vari- Mode 1) flake and core industries are produced ation might be influenced by—or have conse- throughout the Lower Palaeolithic (and beyond), quences for—functional issues. many archaeologists regard the ‘Oldowan’ to end around ~1.7–1.5 MYA coinciding with the appearance of bifaces (Semaw et al., A brief history of functional and 2009: 173; Diez-Martín et al., 2015). morphological concepts within Lower The Acheulean (or Mode 2) techno-complex Palaeolithic research: 1800–1950 is defined by the appearance of new manufac- turing methods, including the ability to shape John Frere (1800, p.204) is known for relatively large (> 10cm) flakes or nodules into providing the first suggestion for handaxe func- an elongated, but broadly ‘oval’, ‘tear-drop’, or tion, describing artefacts recovered from Hoxne ‘triangular’, bifacial form (Clark, 1970; Gowlett (UK) as “evidently weapons of war, fabricated 1988; Goren-Inbar & Saragusti, 1996; Lycett & and used by a people who had not the use of met- Gowlett, 2008; Semaw et al., 2009; Sharon, 2010; als”. While the great antiquity of these tools went Diez-Martín & Eren, 2012). Cleavers, also char- unrecognised and their form was not reported in acteristic of the Acheulean, were produced from detail, it appears that Frere (1800) viewed these

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objects as having been intentionally produced. late 1850s, particularly following excavations at Frere’s discovery was, however, largely ignored at Brixham (UK) (Grayson, 1990). A nota- the time (Wymer, 1968; Sackett, 2000) and it ble moment in associating Palaeolithic stone was not until the second half of the 19th century tools and faunal remains came with the visit of and developments in other fields, most notably Joseph Prestwich, Charles Lyell, and John Evans geology, that the deep antiquity of human origins to Amiens and in the Somme Valley, and recognition of an ‘Old Stone Age’ came to be who having observed the terraces for themselves, (Sackett, 2000). The emergence of Palaeolithic “all came away convinced” of the lithic-faunal archaeology as an academic field during the 19th associations that had earlier been proposed by and early 20th centuries has been described in Boucher de Perthes & Rigollot (Grayson, 1990, detail elsewhere (Dennell, 1990; Sackett, 2000; p.10). Prestwich’s (1860, pp.295-296) descrip- Davis, 2009; Gowlett, 2009a; Pettitt & White, tion of the implements from these sites is, 2011; Henke, 2015). Here, to set contemporary however, particularly noteworthy: work and concepts in broader historical context, we are concerned specifically with the history “In these implements we find three principal types: and development of hypotheses and evidence one, lance-shaped; a second, almond shaped; and surrounding potential functions of different a third smaller form, a flattened ovoid … These Lower Palaeolithic stone tool forms. forms are constant, and each type presents a nearly The association between Palaeolithic imple- constant relation between the length and breadth ments and the remains of extinct animals was of the specimens … [T]he cutting edge extends first reported in France and Belgium (Pettitt & all round … it is to be observed that these cutting White, 2011; Henke, 2015). Of particular note edges are always on one plane, and that they were is Philippe-Charles Schmerling (1833) who pub- produced by blows applied at the edge …and that lished a detailed monograph on 40 Palaeolithic the sides are equilateral.” sites in Belgium documenting associations between stone artefacts and faunal remains. Prestwich (1860, p.296) was, of course, Boucher de Perthes (1847) similarly recognised describing the variable forms among what would an association between stone implements and later be referred to as ‘Acheulean handaxes’ extinct animals in the gravel terraces of Abbeville (Fig. 1). However, he went further, stating that: in the Somme Valley. Somewhat surprisingly, “one object is apparent throughout, that of giving however, he characterised the handaxes recovered to a hard durable substance a shape either sharp- not as utilitarian tools, but instead as Sackett pointed or cutting.” It is the consideration given (2000, p.45) notes “symbolic items used for pur- to the functional or potential ‘design’ elements poses of ritual, exchange, and trade”. This is, as that is notable for the time – certainly, the reader far as we know, the earliest suggestion for Lower can be left in little doubt what Prestwich thought Palaeolithic tools having a social function. de regarding the intent to produce specific forms, Perthes’ work was followed by Jérôme Rigollot and that he considered their functional capacities (1854) who supported the contemporaneity of to centre on their pointed shape and sharp edges. stone tools and extinct animals in the Somme Evans (1859; reproduced in Prestwich Valley, most notably at Saint-Acheul, which [1860]) had, however, slightly earlier alluded would eventually become the eponymous site for to functional implications when describing the the ‘Acheulean’ techno-complex. In all instances, Abbeville and Saint Acheul artefacts, noting the however, the claims associating stone tools and more “pointed weapons” (handaxes) to be analo- faunal remains were largely dismissed by their gous to “lance or heads”, while the “oval contemporaries (Sackett, 2000; Gowlett, 2009a). or almond-shaped” handaxes were characterised Associations between faunal remains and by the presence of a cutting edge all around. No stone tools began to be debated seriously by the precise purpose for handaxes is given, with Evans A. J. M. Key & S. J. Lycett 71

Fig. 1 - Selection of ‘flint implements’ described by Prestwich (1860) from Saint Acheul (a, b, d) and Abbeville (c). These artefacts clearly show what would later be commonly described as ‘handaxes’ and are representative of the variable forms often found in northern . Length a = 7.8cm; length b = 19.4cm; length c = 9.1cm; length d = 12.9cm. himself noting “it is impossible to say for what with flint flakes on fresh bones. Contemporaries purpose they were intended” (reproduced in such as Desnoyers (1863) and Lyell (1863) also Prestwich, 1860, p.311), merely speculating that speculated that cut marks on fossil bones from more pointed forms may be spear tips or for split- northern France were produced through the use ting wood or “grubbing in the ground”, while of flint tools (for discussion see Grayson, 1986). the oval and almond-shaped forms may have A little later, Evans (1864) published evi- been or “sling-stones”. Evans also noted, dence of “Reindeer-horn” having been cut and however, that “flakes or splinters arising from carved by “flint instruments” that were mixed the chipping of the flint, will of necessity pre- with bones of other animals. While many have sent sharp cutting edges, and are certain in con- stressed (e.g. Clark, 1962; Henke, 2015) the sequence to be utilized”, potentially as “- importance of this as early evidence of the con- heads or ” (Prestwich, 1860, p.310). At temporaneity of and extinct animals, the same time Lartet (1860) suggested cut marks the modified antler is also among the first-docu- on Pleistocene bones discovered north of Paris mented examples of direct Palaeolithic stone tool were caused by stone tools, and in a remarkably use (albeit within an Upper Palaeolithic context). prescient example of experimentation, indicated It is, however, a volume of combined works by that he had been able to produce similar marks Lartet & Christy (1865-75) that provides one of

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Fig. 2 - Reproduction of Lartet & Christy’s (1865-75, p.17) figure detailing large flake cutting tools, with the larger tool (a) being described as “being one-edged cutting-instrument, or chop- per…chipped on both faces along one margin, so as to produce a sharp cutting-edge”. Length a = 12.7cm; length b = 9.5cm. the clearest insights into emergent ideas concern- the three implement forms that he described in ing Palaeolithic tool function. In their discussion 1859 (see above), but adds a number of additional of handaxes (albeit Middle Palaeolithic) from Le forms, due in part to the fact that he frequently Moustier they describe them as “cutting or chop- focuses on individual artefacts. Evans (1872, p.565) ping, hatchet-like implements”, including one does, however, amend his earlier functional propo- where “the but-end is blunt, so that the ‘hatchet’ sitions and now considered it unlikely that pointed can stand on end” (Lartet & Christy, 1865-1875, forms (handaxes) were tips, although he still p. 78). Perhaps the most intriguing of items to regarded them as “weapons of offense [rather] than be described by Lartet & Christy (at least given mere tools or implements”. Evans (1872) also modern Acheulean literature) are two artefacts helpfully provides a review of potential handaxe that would best be described today as large flake functions proposed by his contemporaries, includ- cutting tools (Fig. 2). One of which is described ing their use as ice chisels, wedges for splitting as being a “one-edged cutting-instrument, or wood, hatchets, axes to cut down trees, prepare …finely chipped on both faces along firewood or carve out , as well as their use one margin, so as to produce a sharp cutting- as throwing implements, food-cutting tools and edge in the form of a segment of a circle. The implements. He even recited Lubbock’s other margin is left with the natural crust of the (1862, p.250) opinion that “almost as well might flint, and can be conveniently held in the hand” we ask to what would they not be applied”. (Lartet & Christy, 1865-1875, p.17). Clearly, Notably, Evans did not repeat the social func- the potential functional applications of varying tions for handaxes proposed by de Perthes (1847). forms of handaxes and other artefacts in both the Regarding Acheulean flake tools, Evans (1872, Lower and Middle Palaeolithic were not lost on p.562) suggests “their use appears to have been for these early pioneers of Palaeolithic archaeology. cutting and scraping whatever required to be cut Evans’ (1872) volume The Ancient Stone or scraped”. In sum, there appears to be few cut- Implements, Weapons, and Ornaments, of Great ting activities that Lower Palaeolithic stone tools Britain not only reconfirms distinctions between were considered not to have been applied. A. J. M. Key & S. J. Lycett 73

The association between extinct fauna and useful example. These tools were among the first Palaeolithic artefacts continued to be noted from Lower Palaeolithic artefacts to be reported from the 1870s onwards (e.g. Spurrell, 1880; Hughes, sub-Saharan Africa (Seton-Karr, 1896), and as a 1897), including by French prehistorian Gabriel result, it is the establishment of human prehis- de Mortillet (1867), who is perhaps better known tory in a new geographical context that is of for using tool typologies to differentiate between upmost concern to the author, not their func- Palaeolithic industries in the late 1860s. Spurrell tion. Certainly, Seton-Karr (1909, p.182) gives (1884) provides a perceptive account concern- little consideration to their function beyond that ing the production of Palaeolithic stone tools; they were “probably for barter and exchange”. however, it is his distinction between the func- The period between 1910 and ~1930 is tional capabilities of modified and unmodified notable for publications by French prehistorian flake tools that is of particular interest. Indeed, Abbé Henri Breuil (Breuil, 1912, 1926; Breuil & Spurrell (1884, p.117) suggests “trimmed” flake Koslowski, 1934). Accordingly, Dennell (1990, edges with rounded sides would be less likely to p.553) contends that Breuil produced “argu- pierce skin during skinning relative to an unmod- ably the most important Lower Palaeolithic ified flake. The functional value Spurrell (1884) study of the 1930s” with his work in the Lower assigns flake tools, despite discussing other tech- Palaeolithic sequences in the Somme valley nologies such as handaxes, demonstrate that the (Breuil & Koslowski, 1934). However, such relative simplicity of these artefacts did not result works, although important in defining techno- in him overlooking their functional potential. logical aspects of sequentially progressive periods Reports of Palaeolithic artefacts became (that would not be superseded until the 1960s), more frequent and geographically widespread did little to further understanding of the use of between 1870–1910 (Laing, 1892; Wymer, Palaeolithic artefacts. For example, while dis- 1968). However, it would not go too far to cussing large flake assemblages (‘’) in state that few advances were made in terms of Britain, little comment is made regarding their either broadening ideas about the potential potential function beyond “l’association à une uses of stone tools or providing more rigorous faune chaude à Eléphant antique est certaine” investigations of whether certain forms were (Breuil, 1926, p.225). The three chapters dedi- preferentially suited to particular tasks. Davies cated to the ‘Chellean’ and Acheulean periods (2009, p.130) describes this theoretical situation in Arthur Keith’s (1915) The Antiquity of Man as a consequence of the typological distinctions are similarly sparse. Certainly, it is the geological constructed by de Mortillet and others proving context of artefacts, the way they were produced, a “straightjacket” to developments. The eolithic and their relationship to the antiquity of ‘man’ debate likely also distracted from functional that arguably continues to be of chief concern considerations as it kept discussion focused on among prehistorians. whether artefacts were of anthropogenic or geo- As is apparent, functional concepts during logical origin, even though actualistic research the 19th and early 20th century were predomi- sometimes formed part of this debate (Grayson, nantly informed by analogy with modern tools, 1986). However, the relative lack of attention considerations of intent regarding the produc- given to behavioural inferences is due also, in tion of sharp edges, and contextual associations part, to the struggle to establish chronological with faunal remains. The influence of early and geographical frameworks for the artefacts ethnographic accounts of modern stone tool being discovered. In other words, prehistorians using populations must, however, also be noted. were concerned with other fundamental ques- Certainly, indigenous peoples encountered in the tions that were deemed of greater importance , and other areas of the world at the time. Seton-Karr’s (1896, 1909) descrip- came to influence ideas on the potential uses of tions of handaxes from Somaliland provides a Palaeolithic artefacts (Gosden, 1999; Pettitt &

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White, 2011) and, on occasion, led early prehis- greatest value, noting that he was able to “cut up torians to actively seek the opinion of (European) a large in less than two hours, using individuals who had spent time among such only a cleaver” (Leakey, 1950, p.73). Indeed, communities (e.g., Lartet & Christy, 1865-75). after readily admitting failure to cut animal skin Despite these accounts being a product of their “with his hands, nails or even his teeth”, Leakey time (i.e. subjective, biased and subscribing to (1950, p.71) appears convinced of the adaptive notions of ‘primitive’ populations), and in many benefit of stone tools and their importance in instances being offensive to the populations animal food processing. described (e.g., Tylor, 1894), these basic eth- At this time more explicit statements con- nographic descriptions provided one of the few cerning tool (form) attributes and function pieces of evidence relating to plausible uses for appear, with Barnes (1939, p.109), for instance, stone tools. Accordingly, ethnographic accounts highlighting the need for “acute edges (less than formed a prominent contributor toward early 90°) for cutting and scraping” and so providing ideas concerning Palaeolithic stone tool function early commentary on edge angle, which would (e.g., Lubbock, 1865; Tylor, 1894; Aiston, 1928). become increasingly discussed in later years. In Between World Wars I and II, Palaeolithic 1948, Praus provides an early comparative exam- studies continued to be dominated by Breuil’s ination of Palaeolithic and modern metal cutting evolutionary ideas regarding the (progressive) tools and also suggested that modern engineer- technological development of Lower Palaeolithic ing/mechanical theory might be used to shed sequences (Dennell, 1990). For the first time, light on prehistoric tools. Moreover, he indirectly however, there is a substantial shift in focus confirms the novelty of Leakey’s experiments by towards Palaeolithic industries in Africa and noting that functional inferences had previously Asia (e.g., Wayland, 1934; de Terra & Teilhard been derived only through ethnographic analogy de Chardin, 1936; Movius 1944, 1948), includ- or general appeals based on shape. It is with Praus ing their associations with cut-marked faunal (1948, p.158) too, that there is early assertion remains (Pei, 1933). Famously, this includes that handaxes “cannot be considered a true ax[e]”. ’s first visits to However, evidence of the continued importance (Leakey, 1931, 1951) and his assessment of of ethnography in related debates also appear in functional hypotheses through experimentation. works of the time (Mitchell, 1949). In Adam’s Ancestors Leakey (1953, p.58) notes Movius (1944, 1948) is widely known for how “experiments suggest that the hand- his work around this time that would later lead was a kind of general utility tool rather than a to establishment of the famous ‘Movius Line’. weapon…with a hand-axe it is possible to dig up However, Movius (1948, p.349) also noted how wild edible roots, to dig holes to serve as pit-traps, the terminology he used in describing Asian to dig along the burrows of rodents until the nest assemblages was based entirely on artefactual chamber is reached, to chop the smaller bones of forms and production techniques “rather than animals when cutting up a beast, and, of course, any hypothetical functions”. Movius (1948) to cut up meat”. Leakey also describes the more would not, however, be alone in highlighting the limited ability of handaxes to sharpen wooden functional connotations inherent within many stakes or skin an animal relative to flake tools, typological systems and the potential problems and the potential functions of cleavers, scrapers this can pose (Wymer, 1968; Clark, 1970; Odell and ‘Clactonian’ tools. A published lecture by 1981; Toth, 1985). Leakey in 1949 (Leakey, 1950) reaffirms these During the second half of the 20th century, functional statements, but adds that handaxes however, functional concepts began to receive “were certainly not missiles” but may have been more earnest attention. Importantly, distinct ana- used as “clubs or choppers”. Regarding butch- lytical techniques focusing on the use of stone tools ery behaviours, it is cleavers that Leakey assigns emerged, and empirical analyses of tool forms A. J. M. Key & S. J. Lycett 75

become more explicitly integrated with functional The presence of sharp edges on flakes, considerations. Advances in radiometric dating in handaxes, cleavers and other core forms has the 1950s, the subsequent freeing from a purely ensured that recent discussion has continued to typological–chronological perspective, and the focus primarily on their potential role as cutting increasing focus of archaeologists to specialise by implements (e.g., Posnansky, 1959; Toth, 1985; technological period, also guided research away Shea, 2007; Roche et al., 2009; Barsky et al., from generalised discussions of ‘Palaeolithic’ tech- 2011; Braun, 2012; Hovers, 2012; Key, 2016). nologies. Ultimately, this shift also contributed It is, however, impossible to discuss the poten- toward the famous Bordes‒Binford debate, where- tial functions of Lower Palaeolithic technologies, upon it was argued that variability as inferred from aspects of their form, without might better be explained by functional param- mentioning their potential utility within homi- eters than culture-historic alternatives (Binford & nin social systems. Most prominently, Kohn & Binford, 1966). However, these broader shifts also Mithen (1999, p.519) argued handaxes may contributed toward greater focus on the Lower have been “integral to the process of mate choice Palaeolithic as a distinct period, with greater within socially complex and competitive [homi- awareness of the evolutionary context that inevi- nin] groups”. Kohn & Mithen’s (1999) argument tably contributes to debate on form and function hinges on the assumption that handaxes are over- during this timeframe. Consequently, a strictly engineered for functional purposes alone and, chronological review, as has been undertaken thus given that they have potential to be costly (time far, may become unclear in terms of describing and energy) to produce, may, therefore, have how different approaches have combined to form acted as viable indicators of mate quality. That current perspectives. Moreover, current research is, “fine symmetrical handaxe[s]” may have acted builds directly on works from the 1960s-80s, and as an indicator of “good genes”, demonstrating their inclusion in a historical review could mis- their producers had knowledge of resource dis- represent their continued importance. Hence, tributions, an ability to complete complex tasks discussion continues in the following section first and social awareness (Kohn & Mithen, 1999, by means of current evidence surrounding the p.521). While the sexual selection hypothesis is possible functions of Lower Palaeolithic artefacts, an intriguing one (Currie, 2014), it has received and then by means of current knowledge regard- credible criticism (Machin, 2008; Nowell & ing relationships between stone tool forms and Chang, 2009). Similarly, Spikins (2012) and functional application. Hiscock (2014) have proposed that because handaxes may go beyond the immediate need to produce a viable tool this might indicate roles The present state of research extending beyond strictly techno-functional concerning the form and function of applications. Pope et al. (2006) meanwhile con- Lower Palaeolithic stone tools tended that discarded handaxes may have acted as signals and behavioural cues to other hominin What were Lower Palaeolithic stone tools used for? individuals or groups. As intriguing and inven- Current ideas surrounding the possible func- tive as they are, few of these hypotheses, how- tions of Lower Palaeolithic implements build on ever, are currently supported by direct evidence ~60 years of increasingly detailed and steadily or through formal analyses of data independent cumulative research. This is not to say, however, of that from which they are built. that the number of suggested functions for these Shea (2006) has also highlighted that a con- tools has changed considerably since the 19th cen- siderable portion of the stone artefacts recov- tury. Rather, the amount of evidence supporting ered from Palaeolithic contexts may have been or refuting the possible functional applications the product of imitative behaviours by juve- has increased. niles or bouts of practice by unskilled

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individuals. If this supposition is correct, the 1960s and 70s (Ingersoll et al., 1977; Coles, function of these ‘training pieces’ would strictly 1979; Outram, 2008) and from this point the be for increasing knapping proficiency (be this functional utility of Lower Palaeolithic artefacts motor-processes, technological capabilities, etc.). began to be addressed more earnestly through Theoretically, artefacts resulting from such learn- experimental means. These early functional ing bouts might be discernible through a greater experiments were described by some of their number of mistakes (step fractures, breakages, practitioners as largely “freewheeling” (informal mauled platforms, etc.); however, as Shea (2006) and qualitative) in nature, often conceived with notes, there are many challenges to identifying the objective of increasing “understanding, from these activities reliably. an archaeologist’s point of view, of the service- Ethnographic accounts have continued to ability of various items of a diversified lithic tool provide important information relating to the kit” (Crabtree & Davis, 1968, p.426). Although potential functions of Lower Palaeolithic tools few experiments were quite so openly explora- over the past 60 years. Accounts from a variety tive as those of Crabtree & Davis (1968), similar of populations have detailed the use of unmodi- experiments utilised broad, investigative meth- fied flakes when producing spears, digging odologies to assess the suitability of specific sticks, , bone needles, arrow shafts, tool forms to undertake functional tasks (e.g., handles, and more generally during numerous Sonnenfeld, 1962; Swauger & Wallace, 1964; scraping and cutting actions on wooden, shell Huckell, 1979; Jones, 1980). and bone objects, as well as butchery (White & Since the 1980s, Thomas, 1972; Binford, 1986; Hampton, 1999; has flourished to the extent that it contributes Sillitoe & Hardy, 2003; Shott & Sillitoe, 2005; a substantial proportion of new information Holdaway & Douglass, 2012; Hayden, 2015). As concerning Palaeolithic technologies (Eren et noted by Shott and Sillitoe (2005, p.654) when al., 2016). The effective capabilities of unmodi- describing tool use in New Guinea, flakes “were fied flakes for butchery has now been confirmed ubiquitous and mundane. They served in all by numerous experimental studies (Toth, 1985; tasks for which stone was required and ground- Jones, 1994; McCall, 2005; Dewbury & Russell, stone axes could not be or were not used” (see 2007; Braun et al., 2008a; Merritt, 2012, 2015). also, Holdaway & Douglas, 2012). Ethnographic Similarly, handaxes and cleavers have demon- accounts describing the use of bifacial/core tools strated efficacy during butchery activities (Jones, are notably rarer than for flakes (Kelly, 1988). 1980; Toth, 1985; Mitchell, 1995; Machin et al., However, Gould (1980) recorded use of hand- 2007; McCall, 2005; Bello et al., 2009; Toth & held choppers (cobbles with few flakes removed) Schick, 2009; Galán & Domínguez-Rodrigo, for tasks, including the cutting of 2014). Such studies have even demonstrated the spear-throwers and spear shafts. Hayden (2015) ability of both flakes and bifaces to butcher mega- similarly noted the use of handaxe-like imple- faunal pachyderms such as elephants (Frison, ments in Papunya (Australia) for the manufac- 1989; Jones, 1994; Toth & Schick, 2009, p.333; ture of wooden winnowing troughs. As has long Gingerich & Stanford, in press). It is notable that been noted, however, while ethnography can experiments examining the suitability of Lower demonstrate a range of plausible tool applica- Palaeolithic technologies for plant processing tions (Skibo, 2009; McCall, 2012), it does not tasks are relatively rare. Those that have been mean that Palaeolithic populations necessarily undertaken, however, indicate that flakes and used them in these ways, nor that there are not bifaces are capable of numerous woodworking additional uses that have no modern analogue activities, including amongst others, the pro- (Wobst, 1978). duction of spears, bamboo knives, and digging Experimental archaeology developed as a sticks (Toth, 1985; McNabb, 1989; Jones, 1994; distinct methodological approach during the Hardy & Garufi, 1998; Toth & Schick, 2009; A. J. M. Key & S. J. Lycett 77

Bar-Yosef et al., 2012). Experiments examining In subsequent years, evidence for stone-tool the suitability of handaxes for digging appear to related bone modification has become more have been largely overlooked (although see, Toth chronologically and geographically widespread. & Schick, 2009). The suitability of handaxes Currently, the earliest examples may be as old as as projectile tools/weapons has also been tested 3.4 MYRs (McPherron et al., 2010), although, experimentally. While the conclusions of these notably, these are contentious (Domínguez- experiments have at times been inconsistent, Rodrigo et al., 2012). In sites dating from 2.5 generally, they have been shown to make unre- MYRs through to the Middle Palaeolithic tran- liable projectiles (O’Brien, 1981; Whittaker sition such evidence is, however, more regu- & McCall, 2001; Samson, 2006; McCall & larly recorded (e.g., de Heinzelin et al., 1999; Whittaker, 2007). Domínguez-Rodrigo et al., 2005; Choi & One novel source of evidence emerged in the Driwantoro, 2007; Bello et al., 2009; Stiner et 1960s from studies of tool use by wild great apes, al., 2009; Ferraro et al., 2013; Diez-Martín et al., most notably (Pan troglodytes) (e.g., 2015; Starkovich & Conard, 2015; Landeck & Lawick-Goodall, 1968; Struhsaker & Hunkeler, Garriga, 2016). The number of species butchered 1971). Through phylogenetic parsimony, these by Lower Palaeolithic hominins has also increased studies provided evidence that early hominin since the 1980s and now includes catfish, perch, stone-tool behaviours were likely to have emerged turtle, crocodile (Braun et al., 2010; Archer et during subsistence-related strategies (McGrew, al., 2014), beaver (Lebreton et al. 2017), bear 1992; Panger et al., 2002). Indeed, following (Gaudzinski, 2004), monkeys (Pobiner et al., decades provided widespread evidence that other 2008), bison (Landeck & Garriga, 2016), rhi- nonhuman , including bonobos (Pan noceros (Landeck & Garriga, 2016), elephants paniscus), capuchins (Cebus sp.) and macaques (Yravedra et al., 2010), deer (Rabinovich et al., (Macaca sp.) are capable of using tools for food 2008; Stiner et al., 2009), felids (Kolfschoten acquisition and processing (Gumert et al., 2009; et al., 2015) and additional hominin species Visalberghi et al., 2009; Shumaker et al., 2011). (Saladié et al., 2012). The age, context and form Such evidence further bolsters suggestions that of cut-marks found has sometimes been used to food-related tool use in hominins has deep roots. suggest that either flakes or bifaces were specifi- Studies of faunal remains and cut-marks by cally used (e.g., Bunn, 1994; de Heinzelin et al., Shipman, Bunn and Potts in the 1980s were 1999; Domínguez-Rodrigo et al., 2005; Pobiner instrumental in providing further evidence of et al., 2008; Bello et al., 2009; Yravedra et al., stone-tool-assisted butchery in Lower Palaeolithic 2010), indicating that in at least some instances contexts. In particular, their work at Olduvai both tools were used for butchery. Cut marks Gorge provided some of the first unequivocal present on the ~300 KYA Schöningen wooden (direct) evidence that early stone tools were used spears from Germany also indicate that flint tools for butchery (Bunn, 1981; Potts & Shipman, were used during their production (Schoch et al., 1981; Shipman & Rose, 1984; Bunn & Kroll, 2015). Similarly, the wooden Clacton (spear) 1986; Potts, 1988). Although their work more point (~400 Kya) displays striations consistent widely across the should also be noted with having been worked by flint tools (Oakley et (e.g., Shipman et al., 1981; Shipman & Rose, al., 1977). Hominin-imposed modifications on 1983a,b; Bunn, 1986, 1994). The species identi- faunal remains and wooden artefacts, therefore, fied as having been butchered included numer- unequivocally indicate that Lower Palaeolithic ous bovid, giraffid, hippopotamus, antelope and stone tools were used to butcher animal carcasses equid species through to primates such as geladas and make wooden spears. and even mid-to- (Bunn, Additional direct methods of identify- 1981; Potts & Shipman, 1981; Shipman et al., ing Palaeolithic tool functions were also being 1981; White, 1986; Potts, 1988; Bunn, 1994). attempted by the early 1980s, namely lithic

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microwear (use-wear) and residue analyses () was butchery, alongside occasional gath- (Semenov, 1964; Stemp et al., 2015). Keeley’s ering of herbaceous or woody plants. Others pioneering work in the early 1980s specifically have since interpreted microwear on Lower attempted to apply microwear analysis to Lower Palaeolithic handaxes and flakes to indicate Palaeolithic artefacts (Keeley, 1980; Keeley & their use for plant/animal processing (e.g., Ollé Toth, 1981). Indeed, Keeley & Toth (1981) iden- et al., 2012; Sahnouni et al., 2013; Lemorini et tified wear traces on nine Oldowan artefacts from al., 2014; Viallet, 2016). It is notable that there (dating to 1.5 MYRs) and determined is still (Shea, 2011) a lack of Lower Palaeolithic that these tools had been used to cut both ani- microwear analysts employing the use of high- mal and plant tissue. Similarly, Keeley’s (1980) powered 3D microscopy techniques widely analysis of British Lower Palaeolithic artefacts advocated in other areas of lithic microwear resulted in diverse functional assignments for research (e.g., Stevens et al., 2010; MacDonald, flake and core tools, including their use during 2014; Stemp et al., 2015). Nonetheless, taken at plant processing and butchery behaviours. These face value, current literature indicates that micro- publications had a substantial impact on Lower wear traces have potential to yield important evi- Palaeolithic research, and in combination with the dence directly relating to the functions of Lower early analyses of cut-marked bones by Shipman, Palaeolithic tools, and according to existing stud- Bunn & Potts, purportedly provided the first ies, both meat and plant-processing tasks may direct evidence relating to the function of Lower have been undertaken with flakes and handaxes. Palaeolithic stone tools. Isaac (1983, p.18) noted Other analysts have combined microwear the impact of these works at the time, stating that and residue analyses, which has potential to prior to this point “we could and did speculate provide greater security of conclusion. However [about the function of early stone tools], but we to date, it has only been utilised on three occa- had no direct evidence”. sions within Lower Palaeolithic contexts. Rots In the ~20 years immediately following and colleagues’ (2015) analysis of flakes from Keeley’s publications there were only a small Schöningen (~300Kya, Germany) identified number of microwear studies examining Lower residues and wear indicating woodworking and Palaeolithic stone tools, most of which are meat/hide processing. More recently, flint arte- limited in artefact numbers and methodologi- facts dating to ~300 Kya from cal detail (Roberts et al., 1986; Beyries, 1990; (Israel) revealed collagen and bone residue in Mitchell, 1998; White et al., 1998). In part, this association with microwear indicating longitu- has been assigned to problems such as the natu- dinal cutting motions (Zupancich et al., 2016). rally rough surfaces of volcanic rocks commonly Combined with the recovery of a deer bone used, taphonomic damage (Beyries, 1990) and showing signs of having been worked by a flint questions over the replicability and precision of tool, this indicates that at least some of these Keeley’s methods (Unger-Hamilton, 1984; Levi- artefacts were used to deliberately modify bone. Sala, 1996; Newcomer et al., 1986; Shea, 1987). Solodenko et al. (2015) also combined microwear In 1999 Carbonell et al. published the first and residue analysis of a biface, flake and substantial analysis of Lower Palaeolithic micro- from Revadim Quarry (Israel), finding fat residue wear traces since the early 80s. Their work on (likely elephant) on the biface and scraper. the ~780 Kya lithic artefacts from Gran Dolina, There have been additional studies of resi- Atapuerca (Spain), resulted in the identification dues on tools from Lower Palaeolithic contexts. of 24 flake tools displaying wear traces suggested Domínguez-Rodrigo et al. (2001) is perhaps to indicate they were primarily used to cut wood the best known, having identified wood phyto- and meat. Lemorini et al. (2006) similarly sug- liths on three handaxes dating to ~1.7–1.5 Mya gested the primary function of Lower–Middle at Peninj, . Use of Lower Palaeolithic Palaeolithic (382-152 Kya) flakes at Qesem Cave tools for woodworking has also been supported A. J. M. Key & S. J. Lycett 79

Fig. 3 - Illustrations depicting two hypothesised stone tool functions that have been a mainstay of Lower Palaeolithic archaeology for over 150 years. Depicted here is the butchery of a large mam- mal (left) and the production of a wooden implement (right), both of which are supported by direct evidence. However, these specific actions potentially form only part of the functional repertoire of Lower Palaeolithic stone tools. through residue analyses by Loy (1998). The rel- As Desmond Clark (1970, pp.99-100) stated ative importance of Loy’s (1998) discovery was, at the time: “unfortunately, we have, as yet, no however, in the purported identification of ani- direct means of knowing what many of these mal proteins on Oldowan tools dating to 2 Mya tools were used for”. Following the introduc- from , . More recently tion of microwear, residue and cut-mark analy- Nowell et al. (2016) identified 17 artefacts (15 ses from the 1980s onwards, workers have been flake artefacts and two handaxes) from the late exploring potential direct links between artefacts Acheulean site of Azraq (Jordan) displaying and their use, providing evidence that at least traces of horse, rhinoceros, camel and duck anti- some Lower Palaeolithic artefacts were used serum. This study is particularly important in for plant and animal processing. Despite these substantiating the use of tools on a diverse range advances, however, general consensus regard- of species, including the processing of water- ing the primary functions of Lower Palaeolithic fowl. Moreover, it is worth noting that the two stone tools has not changed considerably in the handaxes were identified as having been used to past 60 years (Wymer, 1968; Clark, 1970; Isaac process horse and rhinoceros, both larger animals 1971, 1981; Roe, 1981; Villa, 2001; Shea, 2007; among those represented. Roche et al., 2009; Diez-Martín & Eren, 2012; In sum, prior to the 1980s functional infer- Plummer & Bishop, 2016). Certainly, there has ence continued to rely heavily on interpretations long been consensus that Lower Palaeolithic of tool forms, analogy with modern or ethno- stone tools were not only likely to have been graphic tools, and contextual associations with multifunctional tools, but were used to butcher faunal remains. The expansion of experimental a variety of sized animals and cut and modify archaeology and primatological studies in the plant materials (Fig. 3). Discussion regarding 1960s and 70s provided new and important the adaptive significance of these behaviours and sources of evidence, however, it is nonetheless how they may have come to influence the tool the case that evidence continued to be indirect. forms found in the artefact record is, however,

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not only more contentious, but less frequently As noted previously, ethnographic accounts addressed. It is to these issues which we turn in have the potential to provide important func- the following section. tional information. Across a wide range of studies it has been observed that the form of a flake’s cut- ting edge is an important morphological Potential relationships between for individuals when choosing potential tools stone-tool forms, functional utility (White & Thomas, 1972; Gould, 1980; Sillitoe and hominin behaviour & Hardy, 2003; Hiscock, 2004; Holdaway & Douglass, 2012). Most notably, edge angle in Suggestions of functionally related inten- flakes has been described as consequential to the tionality and selectivity (either cultural or nat- type of task undertaken. For example, Gould ural) during the production and use of Lower (1980, p.119) noted that among Western Desert Palaeolithic stone tools raises four important Aborigines “flakes with edge angles of 40-89 considerations. The first is whether variation degrees … were called purpunpa and were con- in artefact form has the potential to influence sistently used for wood scraping and chopping their effectiveness in functional tasks. Secondly, tasks in procuring and shaping … hard woods”, whether there was an adaptive benefit to the while those with more acute angles of 15–59 preferential use or production of particular tool degrees “were called tjimari and were used as forms. Thirdly, in the case of more cognizant knives for cutting flesh, sinew, and fibrous mate- selection, the extent to which hominins were rials”. Such accounts alone raise the question of capable of preferentially ‘selecting’ or recognizing whether Plio-Pleistocene hominins made similar more effective tool forms. Finally, there is a need functional distinctions, although of course, can- to identify whether Lower Palaeolithic artefacts not prove it. provide evidence that hominins were producing It is from experimental research, however, or using certain tool forms dependent on their that arguably the greatest detail regarding the functional performance characteristics. potential influence of morphological variation on stone-tool performance has been provided. Here, Would the form of a Lower Palaeolithic stone tool we deliberately focus on three sets of experimen- have influenced its functional abilities? tal research that are specifically relevant to the As we have noted, the primary function utilitarian impact of form in Lower Palaeolithic of Lower Palaeolithic stone tools would likely artefacts. The first is concerned with the com- have been to cut plant and animal materials. parative utility of differing flake forms, the sec- Accordingly, it is logical to question whether ond examines differing biface forms, while the the form of a stone tool could impact its cut- third investigates the comparative use of flakes ting capabilities. Certainly, mechanical and ergo- and bifaces directly. nomic investigations of modern hand-held tools often detail how the form of cutting edges and Flakes manual contact areas (e.g., handles) are impor- Three studies have examined the impact of tant in determining their effectiveness (e.g., Hall, flake size on their cutting abilities. Jobson (1986) 1997; McGorry et al., 2005; Atkins, 2009; Rossi identified a significant relationship between et al., 2014; Schuldt et al., 2016), and the value increased flake length and greater efficiency (cut- of this literature to Palaeolithic archaeology has stroke count) during the butchery of rabbits recently been reviewed elsewhere (Key, 2016). (n = 24 flakes and rabbits), most likely due to Here, we are concerned specifically with evi- their longer cutting edges, greater penetration, dence from lithic-centric sources that detail what and manipulability. When flake size was exam- impact, if any, a Lower Palaeolithic stone tool’s ined in terms of mass, however, this appeared form may have on its functional performance. to have limited impact on performance. In A. J. M. Key & S. J. Lycett 81

turn, Jobson (1986, p.15) suggested that “flake range of variability examined was limited (10°– geometry supersedes sheer mass with respect to 47°) (Jobson, 1986). Key & Lycett (2015) also how well a tool performs”. Prasciunas (2007) examined the influence of edge angle, but inves- similarly investigated the impact of flake size (n tigated its influence during a more standardised = 30 flakes) on cutting but used a standardised cutting task, using a greater range of edge angles material (denim) during the experiment. Her (12°–67°) and specifically recorded loading levels results suggested a relationship between increas- applied across flakes of varying size. Their results ing flake size and the amount of material cut in highlight the potentially complex relationships 10 minutes, with a drop in efficiency once tools that can exist between different aspects of a tool’s dropped below 5g or had a surface area below form and its performance. Specifically, although 7cm2 (Prasciunas, 2007). In each of these stud- there is a relationship between edge angle and ies, however, the size variation examined was lim- increased cutting efficiency in smaller flakes, as ited, with only one and two flakes (respectively) tool size increases, the potential for greater work- in each assemblage weighing over 50g. Key & ing load also increases. Consequently, larger Lycett (2014) recently addressed this issue by flakes with more obtuse edge angles can cut as examining cutting efficiency in 342 flakes rang- effectively as smaller ones with acute angles (Key ing between 27–105 mm length and 3–408 g in & Lycett, 2015). mass. Their results indicated that there is not an absolute relationship between increased flake size Bifaces and efficiency; rather, results indicated that once The number of studies that have systemati- flake sizes reach a specific ‘threshold’, increased cally examined the functional performance char- size has limited influence on effectiveness. Below acteristics of bifaces varying in form is more lim- this threshold, however, the use of smaller flakes ited than for flakes. Diez-Martín & Eren (2012, reduces performance. p.341) highlighted this, noting that experimen- The remaining flake-specific studies have tal use of handaxes has principally been limited investigated the impact of edge form on cutting to demonstrating “what a hand ax[e] could have efficiency. Walker (1978) investigated the com- been used for and the experimental falsification parative effectiveness of 20 flakes with unmodi- of those potential uses”. To our knowledge, only fied edges and 25 flakes with bifacial edges. Machin et al. (2005, 2007) and Key, Lycett and Through butchery of a variety of large mam- colleagues (Key & Lycett, 2017a; Key et al., mals, Walker (1978) concluded that both edge 2016) have specifically investigated how the forms were effective, but bifacially flaked edges cutting performance of handaxes might co-vary were less effective in most tasks, except skinning. with variation in form. However, it should be Collins (2008) undertook a more controlled noted that Whittaker & McCall (2001) experi- experiment, examining the impact of edge curva- mentally examined whether handaxes can be ture and angle on flake effectiveness during wood used as projectiles, comparing both a large and scraping. Collins (2008) identified that convex small handaxe in terms of ease of use and dis- edges resulted in significantly greater amounts of tance thrown. Distinctions between tool forms material being removed relative to straight edges. were noted, with the smaller tool capable of Edge angle was also determined to have a statisti- being thrown further, but less stable in flight. cally significant impact on scraping, with more Machin and colleagues (2005, 2007) looked acute-edged tools removing material at greater at the potential influence of handaxe symmetry rates (Collins, 2008). Jobson (1986) also exam- on efficiency during the experimental butchery ined the impact of flake edge angles on butchery of deer. Their first study was limited in scale efficiency. Contrary to Collins (2008), his results (eight deer), but led to an initial conclusion that suggested no relationship between these two “there is not a strong correlation between either variables, however, it should be noted that the frontal or side symmetry and the effectiveness of

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a handaxe for butchery” (Machin et al., 2005, (butt) portion of handaxes showed the strongest p.28). The second study was more extensive, relationship to efficiency. These results are con- comparing 60 handaxes varying in frontal and sistent with suggestions that more obtuse angles side symmetry during the butchery of 60 deer in the butt of the handaxe facilitate greater appli- carcasses (Machin et al. 2007). Despite the cation of force during use and, in turn, increase larger sample, a relationship between symme- cutting efficiency. try and butchery efficiency was still not firmly established, with results suggesting “at the very Flakes vs. Bifaces most, frontal symmetry has a small role to play” Direct comparison of flakes and bifaces has (Machin et al., 2007, p.891). Indeed, the two received greater attention than the influence of participants undertaking the butchery provided variation within a single tool type. Jones (1980, contrasting results, and any positive relation- 1994) undertook a series of well-known experi- ships were dependent on subjective scorings or ments at Olduvai Gorge functionally comparing the removal of an outlier. flakes with handaxes. One of his main conclu- More recently, Key & Lycett (2017a) com- sions was that relatively larger tools (ideally with pared cutting efficiency (time) across 500 a bifacial edge) were preferable butchery tools handaxes in a standardised experimental task. compared to smaller tools. Jones (1980) attrib- The replica tools displayed considerable vari- uted this to their increased mass, edge length, ation in size (4–30 cm in length and 8–4484 ease of gripping and applying force, and capacity g) and morphometric shape properties, which for prolonged use. Serrated edges were also sug- deliberately exceeded those typically found gested to be the “sharpest and longest lasting”, archaeologically in order to push the range of although retouched edges were considered less variability examined. Results showed that varia- efficient than fresh edges (Jones, 1994, p.293). tion in handaxe shape had little effect on cutting For woodworking, Jones (1994) suggested that efficiency. Moreover, comparison of efficiency unmodified edges, such as those found on flakes, and mass across all 500 handaxes also revealed were unsuitable and that more robust, modified a statistically insignificant effect (Key & Lycett, edges with thicker profiles were preferable due to 2017a). However, their results did demonstrate reduced fracturing. evidence for a ‘threshold’ effect at the lower end A series of experiments comparing flakes and of size variation, beyond which, as handaxes bifaces was summarised recently by Toth & Schick reduced in size, their functional performance (2009). This work stresses the advantages pro- decreased accordingly (Key & Lycett, 2017a). vided by larger tool forms for chopping or shap- This suggests ‘minimal size’ may be a pertinent ing wood, although lighter woodworking can be consideration in handaxe cutting effectiveness. undertaken using flakes. Based on experiments In a follow-up study, Key et al. (2016) exam- comparing efficiency during disarticulation of ined edge angles in these replica handaxes and pork ribs, Toth & Schick (2009) also suggested investigated their effect on cutting effectiveness. that although “refined and crude handaxes had Variation was investigated at a whole handaxe similar cutting efficiency”, flakes dulled quicker level (mean angles) and more localised levels than handaxes. They concluded that “Early (individual points around the edge). Somewhat Stone Age groups that habitually butchered counterintuitively, results at a whole-tool medium to large mammals would likely develop level showed that more obtuse edges actually either large bifacial handaxe/cleaver technolo- increased cutting efficiency. However, this effect gies like the Acheulean or flake denticulate tech- was observed only up to a threshold of around nologies” (Toth & Schick, 2009, p.277). Based 70°, beyond which, increased angles resulted in on other experiments, McCall (2005), however, decreased efficiency. Key et al. (2016) were fur- argued that flakes could be more efficient than ther able to show that angles in the proximal core tools due to their acute angles. Galán & A. J. M. Key & S. J. Lycett 83

Domínguez-Rodrigo (2014), meanwhile, com- that the scalloped edges would likely increase pared the efficiency of eight basic (unmodi- their efficiency compared to straight-edged tools fied) flakes, seven retouched flakes, two small when blunt (Key & Lycett, 2017b). handaxes and one large handaxe during butchery. They concluded that “small-sized handaxes can Would there potentially have been an adaptive be more efficiently used … than flakes” and that benefit in preferring specific tool forms? “large handaxes typical of the ESA Acheulian do Flaked stone technology is generally con- not seem to be as efficient for butchery as the sidered to have conveyed an adaptive benefit to small handaxes” (Galán & Domínguez-Rodrigo, hominins, in turn, facilitating their widespread 2014, p.1071). Their for this suggestion production among populations across the Old include the larger handaxe being more difficult World (Ambrose, 2001; Shea, 2007; Roche et to handle and its more obtuse edges. Somewhat al., 2009; Plummer & Bishop, 2016). Indeed, conversely, Merritt’s (2012, 2016) experiments as Braun (2012, p.223) notes, “it seems unlikely using flake and core tools led him to suggest that hominins invested time and energy in the these might be of equal value during butchery development of a complex series of behaviors tasks. A butcher involved in one of these tasks that had no impact on their eventual genetic fit- did, however, describe personal preference for ness”. Typically, the purported adaptive benefits the longer, straighter edge of flakes compared of stone technology centre on the ability to effec- with bifaces (contra Jones, 1980, 1994). Merritt tively process animal and plant foods. However, (2016) also examined the influence of tool mass the use of stone tools to procure other items (e.g., and edge angle on butchery time, with neither shelter, firewood, spears) might also have been displaying significant relationships. important. The extent to which it would have Key & Lycett (2017b) have recently high- been beneficial for Lower Palaeolithic hominins lighted that the form–function disparities to modify or produce specific tool forms to opti- observed across these studies may be due to the mise their utilitarian value, however, has received differing tasks undertaken in each case. Indeed, less attention. when Key and Lycett (2017b) compared the effi- The underlying assumption of specifically ciency of flakes, handaxes, and flakes of equal adaptive arguments in behavioural models is size to handaxes (n = 60 in each instance), they that energetic efficiency would have been of observed that the relative utility of each tool was concern to individuals, and selective pressures dependent on the type of cutting activity under- contributed toward optimising this (Codding taken. ‘Basic’ flakes were significantly more effec- & Bird, 2015). Although the ecological niche of tive than handaxes during tasks requiring greater Lower Palaeolithic hominins would have been precision and where the amount of material was highly variable due to the broad geographical relatively limited. Conversely, handaxes were and chronological boundaries of the period, the significantly more efficient than ‘basic’ flakes optimisation of energetic expenditure relative when cutting more substantial portions of resist- to gain would have been a prevailing selective ant material. This suggests the adoption and use force across populations. Indeed, optimal forag- of each tool type by Palaeolithic hominins were ing modelling (or more broadly, human behav- likely linked to the material contexts in which ioural ecology) demonstrates that in situations they were used. Interestingly, the comparative where energy resources are constrained and/or efficiencies of handaxes and flakes of equal size costly to procure, there are adaptive benefits to were less pronounced. This prompted these increasing the efficiency of energy procurement authors to reiterate the value of considering addi- behaviours (Smith & Winterhalder, 1992; Bird tional factors motivating handaxe production, & O’Connell, 2006; Codding & Bird, 2015). such as their potential use as cores, capacity for Hence, given that stone-tool use is hypothesised resharpening, greater length of cutting edge, and to have been an essential component in Lower

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Palaeolithic subsistence strategies, the efficiency tool-use activity, but also by how regularly a spe- with which these technologies undertook cutting cific task is likely to be undertaken (Stevens & activities is likely to be of evolutionary conse- McElreath, 2015). Indeed, “if a multifunctional quence. Accordingly, it is frequently stated that tool is cheaper [less costly] to make, and has a energetic efficiency is a driving force of tool pro- decent return rate across multiple tasks, it can duction and selection decisions undertaken by outcompete more specialized tools”, however, hunter-gatherer populations (Bamforth, 1986; “specialized tools, even if they are more expensive Bleed & Bleed, 1987; Fitzhugh, 2001; Bird & to manufacture, can outcompete multifunctional O’Connell, 2006; Kelly, 2013; Plummer & tools, but only when the tasks they are designed Bishop, 2016). At a broad level, then, it would for are performed often enough, or occur with have been adaptive for hominins to have pref- enough certainty, to make it worthwhile” (Stevens erentially produced and selected tool forms that & McElreath, 2015, p.108). More broadly, how- reduced costs and were functionally efficient. ever, this implies that if a hominin cannot reliably The extent to which more energetically opti- predict a tool’s functional context at the point of mised stone tool forms would always have been production, or there is a likelihood that it will produced by Lower Palaeolithic hominins is, how- be used in multiple functional contexts, then a ever, more complex than this generalised model. multifunctional tool would be optimal. Indeed, the costs and benefits associated with Finally, it is important to emphasise that using stone technology cannot be solely meas- these studies specifically refer to energetic con- ured through the direct utilitarian application of siderations in anatomically modern humans. the tool, but must take into account raw materi- While such models might reasonably be applied als costs, investment of time, body stress, mobility to hominins associated with Acheulean technol- patterns, risk, tool reliability and maintainability, ogies (e.g., s.l.) (Bird & O’Connell, among other considerations (Bamforth, 1986; 2006), greater caution is necessary for the pro- Bleed, 1986; Torrence, 1989a,b; Jeske, 1992; ducers of Lomekwian and Oldowan technolo- Kuhn, 1994; Bamforth & Bleed, 1997; Beck et gies. Certainly, differences in anatomy likely pre- al., 2002; Brantingham, 2003, 2010; Bousman, clude direct energy-consumption comparisons 2005; Bettinger et al., 2006). As Ugan et al. for some behaviours (e.g., raw-material transpor- (2003, p.1323) put it, “the costs of technology are tation costs), and thus may convey additional/ every bit as important as their benefits when try- reduced energy requirements relative to modern ing to assess investment decisions”. Accordingly, humans. This does not alter underlying require- stone-tool production and selection choices could ments to optimise energetic expenditure relative represent solutions shaped by multiple selective to gains and the adaptive benefits of using more pressures (Bleed & Bleed, 1987). efficient tool forms. Indeed, this emphasizes that Moreover, Lower Palaeolithic stone tools were understanding the relative efficiencies of varying likely utilised in a variety of situations, suggest- tools forms might provide an important ‘anchor’ ing a further need to consider trade-offs between variable within a wider range of complex and using a greater number of specialised tools ver- potentially unknowable considerations. sus more generalised tools (Bleed, 1986; Nelson, 1991). Stevens & McElreath (2015) have recently Were Lower Palaeolithic hominins capable of modelled such considerations in intentionally selecting or producing functionally tools, which have attendant impli- preferable tool forms? cations for Palaeolithic technologies. Their results The tool selection and modification behav- again highlight that tool form production and iours of nonhuman primates provide important selection decisions are not only underpinned insights into the potential capabilities that were by the relative efficiency with which a tool may likely displayed by Lower Palaeolithic homi- complete a task, or the relative duration of a nins. Indeed, parsimony suggests that if extant A. J. M. Key & S. J. Lycett 85

nonhuman primates can preferentially select and that chimpanzees are displaying an appreciation produce different tool forms, then early humans of tool raw-material quality prior to undertaking may similarly have been capable of doing so. a task, selectively transporting certain tool types to Accordingly, it is often argued that a food resource, modifying tools prior to use, and tool use is an effective model for understanding repeatedly producing tools of specific proportions the technological and behavioural capabilities and morphologies, an inference further supported from which Lower Palaeolithic tools emerged experimentally (Manrique et al., 2010). (McGrew, 1992; Ambrose, 2001; Wynn et al., Wild chimpanzees have also been observed 2011; Carvalho & McGrew, 2012). Moreover, preferentially selecting nut-cracking hammers if such capabilities are not only displayed in our depending on their material and form proper- closest relations within the genus Pan, but more ties (Sirianni et al., 2015), which is a behaviour widely across the order, then such claims only previously having been noted in captive are strengthened. populations (Schrauf et al., 2012). Specifically, In some chimpanzee communities, termite Sirianni et al. (2015) observed chimpanzees of fishing involves the forceful opening of a mound’s the Taï National Park (Côte d’lvoire) prefer- exterior with a sturdy ‘perforating twig’ (Sanz et ring stone hammers over wooden alternatives, al., 2004), followed by use of more slender stems heavier stone hammers relative to light stone to extract termites (Sanz et al., 2009). Sanz et ones, increasingly heavier hammer stones the al. (2009) examined the design of termite-fish- closer they are to an anvil, and the preferential ing tools from chimpanzees in the Goualougo selection of lighter hammer stones when crack- Triangle (Republic of Congo) and identified spe- ing nuts on a wooden anvil. This demonstrates cific tool modifications by chimpanzees, includ- a complex, multidimensional tool-selection ing the fraying of probe ends, splitting the probe, strategy that may be considered representative separating plant fibres, and modifying the length of foraging optimisation through appropriate (Sanz et al., 2009). Experimental data indicated tool-form selection (Sirianni et al., 2015) simi- that those with modifications were “significantly lar to that discussed above for humans. Schrauf more effective in gathering termites” (Sanz et al., et al. (2012) further demonstrated that captive 2009, p.295). Intentional tool-modification pro- chimpanzees are not only capable of selecting the cesses are further noted in chimpanzees that hunt most effective for nut cracking, for prosimians (Galago senegalensis) with ‘spear but that they can identify tool mass as central to like’ probes. Up to five modification behaviours the effectiveness of the tool. Schrauf et al. (2012, are sometimes undertaken prior to use of these p.12) conclude that “encoding the requirements tools (Pruetz & Bertolani, 2007). In ~50% of that a nut-cracking tool should meet (in terms cases, they are used before all steps of modification of weight) to be effective therefore lies within have been undertaken, implying that individuals chimpanzees’ capabilities”. Chimpanzees are, are assessing the effectiveness of partially modified then, not only capable of preferentially altering tools and only continue with further modification the form of a tool so that it may better undertake if necessary (Pruetz & Bertolani, 2007). a functional task, but are able to assess the rela- Boesch et al. (2009) have observed chimpan- tive performance attributes of varying tool forms zees in Loango National Park, Gabon, undertak- and, accordingly, make informed choices. ing sequential tool choice and modification pro- Insights into the stone-tool modification cedures, producing several different tools used and selection capabilities of Pan also come from together for honey extraction. These ‘tool-sets’ experiments with captive apes. Famously, the consistently contain of a number of branches bonobo was shown how to knap sharp of distinct forms and raw materials that are stone flakes (Toth et al., 1993; Schick et al., employed in different stages of the honey extrac- 1999). Kanzi was then encouraged to use these tion process. Hence, Boesch et al. (2009) note flakes to cut pieces of cordage and gain access

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to a food reward. Notably, Kanzi preferentially forms dependent on their lack of cortex, edge selected the largest flakes and then tested their durability, size and completeness (Stiles, 1991; sharpness using his tongue (Schick et al., 1999). Reti, 2016). The demonstrable understanding of Although this is only a single individual, it is stone fracturing and knapping skills by Oldowan indicative of a capability to assess both the ergo- hominins (Roche et al. 1999; Delagnes & Roche, nomic benefits of a larger flake and the necessity 2005; Stout et al., 2010), which exceeds even of a sharp edge when undertaking cutting tasks. those of Kanzi (Toth et al., 2006), and their ability Old World monkeys also preferentially to display differential tool-production strategies select tools based on their functional capabili- depending on the relative costs of raw materials ties. Gumert et al. (2009) presented evidence that (Blumenshine et al., 2008; Gurtov & Eren, 2014; wild long-tailed macaques (Macaca fascicularis) Reti, 2016) may further underline the selective select and use different stone tools depending on capabilities of Oldowan hominins. A conscious the type of food-processing activity undertaken. awareness of the feedback mechanisms relating Specifically, macaques used smaller stones and to tool performance (ease of use, effectiveness, their pointed ends when detaching oysters, whereas etc.) that underpin tool production decisions and tools found at anvils (for processing larger mol- inform individuals of the consequences of tech- luscs and nuts) tended to be larger and displayed nological choices (Schiffer & Skibo, 1997) may, evidence of broader surfaces having been utilised then, have been within the cognitive capabilities (Gumert et al., 2009; Gumert & Malaivijitnond, of these Lower Palaeolithic hominins. 2013). Some New World monkeys have similarly After ~1.8 MYA, some hominin species begin displayed such abilities. Fragaszy et al. (2010), for to display substantially increased cranial capaci- example, experimentally demonstrated that wild ties relative to Pan and on the order of twice bearded capuchins (Sapajus libidinosus) can dis- those (upwards of ~800cc) of earlier hominins criminate between the differing resistance of nut (Antón et al., 2014). Consequently, the sophis- species and the mass of different hammerstones, in ticated tool selection and modification processes turn selecting combinations of tools and nuts that displayed by chimpanzees are arguably going to minimise the number of strikes. It have been exceeded by Acheulean hominins. This has also been noted that both visual and manual is potentially further supported by the tool pro- cues are used by capuchins when assessing the duction strategies displayed by Acheulean homi- functional suitability of hammerstones, prompting nins (Wynn, 2002; Goren-Inbar, 2011; Stout et Visalberghi et al. (2009, p.215) to conclude that al., 2014). The extent to which Acheulean homi- they “consistently and immediately selected func- nins could exceed the tool selectivity and func- tional tools, regardless of the condition intricacies”. tionally related modification processes displayed Evidence from the fossil record indicates by Pan is, however, a point that requires further that the encephalization quotients of likely attention in the . Although, as the follow- Lomekwian and Oldowan stone-tool producers ing discussion highlights, work with relevance to were only marginally larger—if at all— compared this discussion is beginning to be undertaken. with Pan (Antón et al., 2014). It may, therefore, be suggested that the earliest stone-tool produc- Do Lower Palaeolithic artefacts indicate that ing hominins would likely have been as capable relatively more effective tool forms were being of the tool-choice proficiencies displayed by the intentionally produced or selected by hominins? living nonhuman primates discussed. Evidence To some extent, the evidence for hominin tool of raw material selectivity based on cutting-edge selectivity and their likely capabilities to do so will durability further suggests that Oldowan homi- have changed over the course of time engulfed by nins were capable of relevant tool selection deci- the ‘Lower Palaeolithic’ (i.e. c.3.3 MYRs to ~300 sions (Harmand, 2008; Braun et al. 2009), as does KYRs). Accordingly, we consider these issues on a their ability to selectively transport specific flake chronological and technological basis. A. J. M. Key & S. J. Lycett 87

The Lomekwian performance and minimising material costs asso- The 3.3 million-year-old stone tools found ciated with producing larger (but fewer) flakes at Lomekwi 3 (West Turkana, Kenya) have only per unit of raw material (Brantingham, 2010; recently been described, although evidence to Surovell, 2009). Given the controlled reduction date suggests that relatively large flake tools were strategies exhibited in many of these assemblages being intentionally produced (Harmand et al., (e.g., Semaw, 2000; de la Torre, 2003; Delagnes 2015). Indeed, the mean length and mass of & Roche, 2005), there is to predict that published Lomekwian flakes (n = 26) is 120mm Oldowan hominins were preferentially produc- and 842g (Harmand et al., 2015). Key & Lycett ing flakes above experimentally described func- (2014) demonstrated that flakes just under tional thresholds to produce flake tools that this size (103mm in mean length) are not only could be efficiently used as a hand- or finger-held effective cutting tools, but are as effective flakes cutting tools. Of course, the form of Oldowan 88–58mm in length and are more efficient than flakes may depend on multiple factors (Kimura, flakes below ~43mm in length. Moreover, flakes 1999; Braun et al., 2008b; Blumenshine et al., of similar size (mean length = 145mm, mean 2008; Roche et al., 2009; Stout et al., 2010; Reti, mass = 677g) to the Lomekwian material have 2016), many flakes within excavated assemblages been shown experimentally to be effective cut- may not been used (Roche et al., 1999; Lemorini ting tools during a variety of different tasks (Key et al., 2014) and hominins may have displayed & Lycett, 2017b). On the basis of such results, only limited selectivity once flakes met mini- there appears little to reason to suggest that the mum thresholds (Hiscock, 2004; Holdaway & large ‘Lomekwian’ flakes were necessarily ineffec- Douglass, 2012). Nonetheless, flake data does tive if ever used as cutting tools. Key & Lycett indicate that—on average—Oldowan homi- (2014) also demonstrated that the loading capa- nins were producing functionally efficient flake bilities of flake tools increases with size. Whether sizes. Some Oldowan sites, however, exhibit dis- this indicates that loading potential was impor- proportionally high numbers of small flakes, at tant to the hominins at Lomekwi is currently times with mean lengths of ≤25mm (de la Torre, unclear. However, if the makers of these flakes 2004; Zaidner, 2013; Gallotti & Mussi, 2015). were restricted in their physical ability to exert Experimental evidence would, then, seem to downwards momentum during use (Schick et al., indicate that these flakes displayed somewhat 1999; Domalain et al., in press), the greater load restricted functional potential (Jobson, 1986; potential afforded by large flakes may have been Key & Lycett, 2014), perhaps being particularly of value (Key & Lycett, 2017b). suited to tasks requiring limited force, restricted use times, or involving relatively small amounts The Oldowan of material. However, while additional factors Oldowan flake artefacts are highly variable require consideration (e.g., the availability of dif- in shape and size. Across broad chronological ferently-sized raw materials [de le Torre, 2004]), and geographic ranges, however, mean flake the presence of such assemblages suggests some lengths of individual assemblages are typi- uncertainty regarding the extent to which homi- cally between 35–50mm (Semaw, 2000; de la nins were always capable of optimising flake size, Torre, 2003; Delagnes & Roche, 2005; Stout at least on the basis of experimental data. et al., 2010; Harmand et al., 2015). Broadly, Edge morphology data are rarely reported this coincides with the more efficient flakes uti- from Oldowan assemblages, with discussion lised by Jobson (1986) and is above the mini- tending to centre on the value and intentional mum size-efficiency thresholds identified by production of ‘sharp’ cutting edges (e.g., Toth, Prasciunas (2007) and Key & Lycett (2014). 1985; Semaw, 2000; Barsky et al., 2011). Arguably, flakes of this size may correspond to Lemorini et al. (2014) provide one of the few an optimisation between maximising functional studies of edge angles for Oldowan flake tools

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at Kanjera South, Kenya. For the 39 artefacts to subsequently make appropriate selective deci- described, edge angles ranged between 29–90°, sions. Blumenshine (2008) similarly determined although little other morphological or meth- that tool transportation distances at Olduvai may odological information is provided making it have been related to durability of cutting edges. difficult to definitively relate such variation to Braun (2008c), Harmand (2008) and Kuman & functional issues. Zaidner’s (2013) description Field (2009) also demonstrated that raw-material of small flakes from the site of selection at Kanjera South and Lokalaei in Kenya, Bizat Ruhama (Israel), however, reported a mean and Stekfontein in South Africa (respectively), angle of 37° across the assemblage. Experimental was likely dependent on durability. These latter and ethnographic evidence indicates that flakes four studies do not directly relate to the selec- with acute edges typically display increased effi- tion of specific tool forms, but they highlight the ciency (e.g., White & Thomas, 1972; McCall, importance of maintaining functionally efficient 2005; Collins, 2008) but that small flake tools, tool forms (sharp edges) over extended periods in in particular, require more acute edges due to hominin tool production-related decisions. their reduced loading capabilities (Key & Lycett, 2015). Hence, this is consistent with hominins The Acheulean at Bizat Ruhama preferentially producing small Acheulean assemblages may contain flakes, flakes with relatively acute edges in order to bifaces, or a combination of the two. Among facilitate their more effective use as cutting tools. other hypotheses, this has sometimes been taken Retouched flakes are often rare in Oldowan to indicate that hominins may have been pro- assemblages (e.g., Roche et al., 1999, 2009). It ducing each tool in response to varying utilitar- may, therefore, be the case that Oldowan homi- ian (behavioural) demands (Domínguez-Rodrigo nins were often not modifying edges to exploit et al., 2014; Diez-Martín et al., 2015; Moncel et any potential benefits provided by increased al., 2015). Certainly, it has been repeatedly dem- scalloping/serration or sinuosity (Walker, 1978; onstrated that these two tool types can (depend- Atkins, 2009; Key, 2016). ing on the task being undertaken) display diver- There have been a few suggestions of gent performance capabilities (Jones, 1980; Toth Oldowan hominins preferentially selecting & Schick, 2009; Galán & Domínguez-Rodrigo, flake tools based on their form. Stiles (1991, 2014; Key & Lycett, 2017b), and may accordingly pp.10-13) suggested a bias in the selection and have underlain the decision-making processes that transportation of “whole flakes” from Bed II at resulted in either tool form being produced. Olduvai Gorge and that there was a “distinct Limited research has been published regard- selection for medium-sized, thin flakes”. Kuman ing the comparative forms of Acheulean flake & Field (2009) similarly suggest the preferen- tools, particularly in relation to their possible tial transportation of medium-sized Oldowan functions (Gowlett, 2015). The presence of large flakes at Sterkfontein (South Africa), ‘Clactonian’ flakes in Britain has often been in turn suggesting the preferential transportation noted (Pettitt & White, 2012) and, at times, of functionally efficient tool sizes (Jobson, 1986; has been linked to their use in specific contexts Key & Lycett, 2014). More recently, Reti (2016) (Ashton et al., 1994), such as associations with undertook a replication experiment indicating large mammal butchery sites (Wenban-Smith et Oldowan hominins at Olduvai Gorge selectively al., 2006), which may suggest heavy-duty cut- transported quartzite flakes dependent on their ting tasks (Jones, 1980, 1994; Key & Lycett, form, most likely due to levels of cortex reten- 2014, 2017b). However, it is at Iberian sites tion, more durable edges and, in turn, increased that the most detailed functional analyses of functional potential. This is arguably demon- Acheulean flake-tool forms have been attempted. strative of an ability by hominins to appreciate Terradillos-Bernal & Rodríguez (2012, 2017) the increased utility of certain flake forms and examined the mass, edge form, raw-material A. J. M. Key & S. J. Lycett 89

properties and functional capabilities of flakes handaxe mass was likely controlled for func- from several sites. They proposed three distinct tional reasons, including links between recurrent categories of flake tools were being produced average tool mass (~0.5kg) and biomechanical based on edge form (particularly edge angle) limitations. Although Key & Lycett (2017a) and tool mass (Terradillos-Bernal & Rodríguez, failed to find a significant drop in biface effi- 2012). Furthermore, they contended that use ciency levels in tools greater than 0.5kg in mass, of different raw materials affected flake-edge their experiment did not examine tool use over form and utility and, in turn, influenced the extended (i.e. >10 minutes) periods, where mass forms produced and raw materials selected may have greater influence (Jones, 1980, 1994). (Terradillos-Bernal & Rodríguez-Álvarez, 2017). These authors did, however, find an effectiveness These works are important as two of the first threshold at the lower end of handaxe variation Lower Palaeolithic artefact studies to incorpo- (Key & Lycett, 2017a). Given that handaxes rate mechanical principles when considering tool frequently appear to have been abandoned with form choices and form–function relationships. further reductive potential remaining (Pitts & However, and as noted by Key & Lycett (2015), Roberts, 1997; Goren-Inbar & Sharon, 2006; the complex interdependent relationships Petraglia & Shipton, 2008; Shipton & Clarkson, between flake size, loading, edge angle and the 2015; Gowlett, 2015; Moncel et al., 2015) it is amount of work (energy) required to cut materi- plausible that hominins managed reduction and als, need also to be considered. Further research discard behaviours with regard for such a thresh- is, therefore, needed to clarify the extent that old (Key & Lycett, 2017a). Acheulean hominins were selectively producing Examinations of handaxe edge form have different flake tool forms with respect to use. generally been limited to Middle Palaeolithic Handaxes and cleavers have received consid- bifaces, where edge-angle variation has been erable attention regarding variation and stand- linked with use in differing functional contexts ardisation in their form. However, functional (Soressi & Hays, 2003) and a possible need to considerations have not always been considered maintain effective angles following resharpen- as a leading explanatory hypothesis for such vari- ing (Iovita, 2014). However, Key et al. (2016) ation. Wynn & Tierson (1990, p.81), for exam- showed that Acheulean handaxes tend to display ple, surmised that “no one has ever made a con- increasingly obtuse edge-angles from tip to base vincing argument that mechanical function is and that mean angles in the base rarely exceeded responsible for the general shape of the biface, let 70°. Both trends were suggested to be function- along the differences in shape”. Conversely, oth- ally derived, with the former resulting in an effec- ers have proposed that biface form might have tive cutting edge toward the tip, while simultane- particular functional relevance (e.g., Roe, 1981; ously facilitating the exertion of high loading by Grosman et al., 2011; Beyene et al., 2013; Diez- the hand at its base. The latter meanwhile may Martín et al., 2014; Gowlett, 2011a, 2013, 2015; represent a need to, at least occasionally, utilise Moncel et al., 2015). Two studies by Gowlett & the base as a functional (cutting) edge (Key et Crompton (1994; Crompton & Gowlett, 1993), al., 2016). Recurrent thickness measurements in for example, demonstrated that size attributes the proximal portion of handaxes at Nahal Zihor in handaxe assemblages may display allometric (Israel) further suggest artefact forms may be scaling, meaning that an increase in the absolute linked to functional and ergonomic constraints length of a handaxe does not necessarily lead to (Grosman et al., 2011). Such factors can also be a linearly scaled increase in its thickness. They connected to previous suggestions linking the interpret these scaling relationships in terms of presence of a ‘globular butt’ and dimensional the need to maintain functionally effective tool- properties of handaxes to an increased ability to mass ranges in larger artefacts. Gowlett (2009b, stabilise, grip and balance a handaxe during use 2015) has since provided further evidence that (Gowlett, 2006, 2011b, 2013).

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Cultural evolutionary approaches to handaxe parameters, and in this sense, even ‘stylistic’ variability have also stressed the role that selective parameters can be subject to selection. Indeed, factors may have played. Simão (2002, p.419), Key & Lycett’s (2017a) study suggests there for example, proposed that symmetry may have would have been little direct functional benefit been influenced by functional pressures, or as he to specific handaxe shapes, which is in line with put it, “the selective recreation of tools depending recent studies stressing the role of cultural drift on how well they fitted their target function(s)”. in creating shape differences within and among He specifically argued that symmetry potentially different handaxe assemblages, even if there were reduces torque and maximises power by putting some selected limits to overall form (Lycett 2008; a handaxe’s centre-of-mass in line with motion Lycett et al., 2016). during use (Simão, 2002). Subsequently, using principles drawn from population genetics, Lycett (2008) argued that symmetry variation Conclusions: untangling form and in handaxes does not conform to a null model function in Lower Palaeolithic of neutral variation but, rather, displays evi- technologies and their continued dence of having been subject to selective forces, analytical relevance with techno-functional, socio-functional and aesthetic selection all being possible candidate A range of nonhuman primates have now mechanisms. More recently, Kempe et al. (2012) been observed using unmodified stone tools explicitly modelled size variation resulting from for percussion activities in the wild (Gumert copying error and compared this to variation in et al., 2009; Fragaszy et al., 2010; Carvalho & 2061 archaeological handaxes from 21 different McGrew, 2012). However, even though such sites. Their results demonstrated that artefacts behaviour sometimes results in the breakage of displayed limited size variation compared to the lithic matter (e.g., Mercader et al., 2007; Proffitt model, indicating functionally-related cultural et al., 2016), crucially, no living nonhuman pri- selection was likely operating, particularly the mate has ever been observed using a stone cutting need to grip a handaxe comfortably in the hand tool in the wild, nor deliberately flaking lithic (Kempe et al., 2012, p.6). This conclusion has material for subsequent use as a tool. This con- further been supported by experimental work trasts with evidence that occurs alongside some examining the impact of tool-user handsize on of the earliest appearances of knapped material the effectiveness of handaxes during cutting in the archaeological record, showing hominins (Key & Lycett, in press). Vaughan (2001) simi- were using stone tools for cutting activities (e.g., larly used evolutionary theory to examine varia- Semaw, 2006); a focussed tool-use pattern that tion in 251 handaxes covering some 1.5 MYRs. continued for millennia thereafter. Accordingly, Following Dunnell (1978), Vaughan reasoned as others have noted, the appearance of stone cut- that traits displaying a relatively high level of ting technology seems to be a uniquely hominin variation were likely more ‘stylistic’ and subject invention, motivated by particular needs (Roux to random evolutionary changes (drift, transmis- & Bril, 2005). Consequently, considerations of sion error), while ‘functional’ traits would be less whether stone-tool form relates to functional variable due to selection. Accordingly, Vaughan matters has deep historical roots in the discipline. (2001) argued that handaxe length may have Current perspectives clearly draw on and directly been under less selection than other attributes build upon this historical legacy. However, it has such as breadth. However, as noted by Lycett long been recognised that distinctions between (2013, p.118), Vaughan’s (2001) characterisa- tool forms, including those used to create cat- tion of ‘stylistic’ versus ‘functional’ traits does egorical types, are not necessarily tantamount to not adequately account for the idea that cultural differences in the activities to which they can be selection can operate outside of purely utilitarian applied (Odell, 1981). In turn, contemporary A. J. M. Key & S. J. Lycett 91

concerns exhibit a focus on more precisely iden- increasingly clear that complex relationships tifying how indirect and direct evidence might exist between a stone tool’s form, the ma- shed light on form–function relationships. terial being worked and the individual tool Debate concerning the comparative performance user ‒ all of which interact to influence ef- characteristics of different tool forms, and the ficiency. Moreover, multiple performance extent to which lithic artefacts may have been characteristics (e.g., loading, manipulability, optimized with respect to these characteristics, cutting stress created) may interact to influ- has also grown in consequence. Here, we have ence the final form of a tool (Schiffer & Ski- reviewed evidence regarding these issues in Lower bo, 1987). This does not necessarily detract Palaeolithic flaked-stone-technology against this from our ability to identify important im- historical backdrop. We have proposed that four plications regarding form–function relation- broad questions may be used to address whether ships and the decisions underlying hominin Lower Palaeolithic hominins were producing tool-production choices, but rather, suggests and/or selecting different stone tool forms with that anyone modelling these relationships specific regard to their performance capabilities. needs to be measured in their approach. Current evidence in response to these four ques- • In general terms, the potentially adaptive tions may be summarised as follows: value of stone technology for Lower Palaeo- lithic hominins can be assumed with some • Experimental evidence has repeatedly shown security. Questions relating to the adaptive that morphological attributes in flakes and significance of specific stone tool forms has, bifaces have the potential to significantly however, rarely been discussed (although influence their cutting performance in a va- for examples see: Bamforth, 1986; Bleed, riety of material contexts. This includes evi- 1986; Jeske, 1992; Torrence, 1989a), par- dence suggesting that determining whether ticularly within a Lower Palaeolithic con- a flake or handaxe will be the more effective text. As discussed, it is reasonable to infer tool, depends on the physical properties of that optimising energetic expenditure rela- the task. In turn, there is potential for func- tive to gain would have been consequential tionally derived selective pressures to have to hominins. Certainly, within Lower Pal- influenced the form of stone tools during aeolithic contexts there would likely have the Lower Palaeolithic. This may be in the been strong selective mechanisms in place form of directional, disruptive or stabilising favouring not only the use of stone tools to selection. As significant as these findings are, aid in the procurement of energy resources, research indicating that artefactual varia- but also minimizing the costs associated tion may occur independently of functional with manufacturing and using these tools. performance is just as important. Indeed, in This indicates that more detailed examina- several instances it has been identified that tions of these issues in Lower Palaeolithic once tool forms are above or below form- contexts is desirable. dependent performance thresholds, there is • Studies of extant nonhuman primates dem- potential for a substantial area of ‘free play’ onstrate relatively complex tool production where tools may vary without functional and selection decisions by species with rela- consequence (Crompton & Gowlett, 1993; tive brain sizes equal to, or less than, those Lycett et al., 2016). A logical consequence of Lower Palaeolithic hominins. Impor- of this is that it is often necessary to test a tantly, such decisions are often based on a wide range of tool forms (that often go be- tool’s form or material properties and often yond those observed in the artefact record) increase performance within functional to adequately test their performance limits. contexts. Parsimony, then, allows us to in- Furthermore, in recent years it has become fer that Lower Palaeolithic hominins were

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likely capable of intentionally selecting dif- respect to their functional performance. ferent stone tool forms depending on their However, better understanding how evolv- basic attributes and performance feedback ing cognitive, anatomical and social capaci- during use, as speculated half a century ago ties to manipulate these variables relate to by David Clarke (1968, p.182, pp.217- archaeological patterns (within and beyond 218). Further, there is evidence that some the Lower Palaeolithic) clearly emphasizes Acheulean hominins may have exhibited the role form–function considerations must functionally related tool selection and pro- play in further investigation of these broad- duction capabilities in excess of those exhib- er components of human evolution. ited by extant nonhuman primates (Wynn, 2002; Key et al., 2016; Stout et al., 2014). Based on current evidence we can, therefore, In sum, there is evidence to suggest that contend that the comparative performance capa- hominins would have been able to recognise bilities of differing tool forms could theoreti- basic relative performance capabilities of cally have influenced stone tool production and different Lower Palaeolithic tool forms and selection decisions during the Lower Palaeolithic appropriately modify related behaviours in period. Moreover, hominins likely had both response. Indeed, based on data from chim- the cognitive capacity and adaptive stimulus to panzees suggesting they manufacture ‘tool respond to such pressures and, additionally, the sets’ comprised of tools with differing per- artefact record has the potential to inform us formance attributes, there seems reason to about how functionally related production and suspect Lower Palaeolithic hominins may selection behaviours came to shape the stone also have combined stone tools with differ- tool forms recovered. Being explicit about this is ent functional benefits. Most notably, given important, if stone tools are to live up to their recent data highlighting the functional ben- potential to inform us about important aspects efits of flakes and handaxes are potentially of human evolution (Gowlett, 2010; Shea, 2011; specific, the beneficial combination of both Lycett, 2013). tools as a ‘tool set’, is set in new light. ‘Function’ should not, of course, be consid- • Multiple variables interact to influence a ered a lone influence on Palaeolithic tool forms tool’s efficiency (Fig. 4). Discussed here (Roe, 2003; Gowlett, 2011a, 2011b). There are a range of artefact analyses indicating is near consensus that no single behaviour or that Lower Palaeolithic hominins were po- variable satisfactorily accounts for the variation tentially producing stone tool forms with observed within or between artefact types during respect to their functional capabilities. In the Lower Palaeolithic (Noll & Petraglia, 2003; some instances, evidence is as straightfor- McNabb et al., 2004; Machin, 2009; Gowlett, ward as the identification that flakes and 2009b; Archer & Braun, 2010; Diez-Martín bifaces were recurrently produced within et al., 2014; Eren et al., 2014; Lycett & von functionally efficient size ranges and meet- Cramon-Taubadel, 2015; Moncel et al., 2015). ing minimum thresholds. In other cases, The discussion presented here reflects this and and often with regards to bifaces, there are emphasises the importance of considering mul- specific aspects of assemblage variability tiple explanatory hypotheses when interpreting that are suggestive of tool forms being pro- variation and standardisation in artefact forms. duced that increase their functional perfor- Recent experiments identifying areas of ‘free play’ mance. When combined, there is convinc- in handaxe shape and cutting performance high- ing evidence to suggest that at least in re- light this point (Key & Lycett, 2017a) and stress gards to a number of key morphological at- the role that additional factors play in influenc- tributes, Lower Palaeolithic hominins were ing aspects of assemblage variation (McPherron, producing specific stone tool forms with 1999; Archer & Braun, 2010; Lycett & von A. J. M. Key & S. J. Lycett 93

Fig. 4 - Schematic illustrating how several variables intrinsic to the artefact (e.g., a and b) as well as external to the artefact (e.g., c and d) may conspire to determine an artefact’s efficiency with respect to a task such as cutting. A single variable such as mass (a) may have a notable effect on cutting effi- ciency, but then efficiency may be independently influenced by a second morphometric property such as edge angle (b). However, in some attributes, considerable morphological variation (‘free play’ zone in a) may be permissible with little discernible implications for efficiency except when specific morphological ‘thresholds’ are reached. The same artefact may also perform differently depending on the biometric attributes of the tool user (c). The same artefact may perform differently in two similar types of task with variable environmental properties (d), whereupon it is close to optimal in respect to one task (e.g., butchering large fauna) but relatively poorer in a similar task (e.g., butch- ering smaller fauna). The colour version of this figure is available at the JASs website.

Cramon-Taubadel, 2015; Schillinger et al., turn, functional concerns interacting with other 2017). Certainly, if experiments cannot identify factors are likely reflected in some of the strong a strong relationship between aspects of a tool’s and deep-rooted regularities observed within form and its performance characteristics across and between Lower Palaeolithic artefact assem- broad ranges of variation (i.e. the attribute is blages (Goren-Inbar & Saragusti, 1996; Lycett & influence-neutral to functional performance) or Gowlett, 2008; Hovers, 2012; Gowlett, 2015). within task-dependent ‘thresholds’, then there Moreover, the ability to experimentally discern must be alternative mechanisms influencing the compromises in tool performance, as exhibited by forms of tool produced. artefactual deviations away from more ‘optimised’ It could, however, equally be stressed that tool forms, facilitates investigation into other stone tools are functional objects and must have causal factors underlying variability (Schiffer & met basic morphological criteria enabling their Skibo, 1997). Hence, further experiments exam- use to at least minimum performance thresh- ining our relative ability to identify mechanically olds (Meltzer, 1981; Schiffer & Skibo, 1997). and ergonomically optimised tool forms are vital Specifically, stone tools must have facilitated the to not only understanding any influence of func- provisioning of returns that exceed their respective tionally selective mechanisms on artefact forms, accumulated costs, especially in the case of recur- but may profitably be used to identify the relative ring tool forms. As demonstrated in experimental influence of other causal factors. Moreover, the studies (Prasciunas, 2007; Key & Lycett, 2014, importance of direct evidence to more securely 2015, 2017a), these minimum performance levels infer the context of tool applications in archaeo- may be modelled as task-dependent thresholds. In logical settings is reemphasized.

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A functional perspective when interpreting Archer W., Braun D.R., Harris, J.W.K., McCoy artefacts forms is, therefore, vital to our ability to J.T. & Richmond B.G. 2014. Early Pleistocene accurately interpret the tool-form related behav- aquatic resource use in the Turkana Basin. J. iours of Lower Palaeolithic hominins, and their Hum. Evol., 77: 74-87. consequences for (and relationship to) wider fac- Ashton N., McNabb J., Irving B., Lewis S. & tors of human evolution. Moreover, direct, indirect, Parfitt S. 1994. Contemporaneity of Clactonian experimental, and evolutionary perspectives inter- and Acheulian flint industries at Barnham, sect in crucial ways. Given that many basic state- Suffolk. Antiquity, 68: 585-589. ments about the functions of Lower Palaeolithic Atkins T. 2009. The Science and Engineering of stone tools have changed little over more than a Cutting. Butterworth-Heinemann, Oxford century, the recognition of this interaction, and Bamforth D.B. 1986. Technological efficiency methodological improvements to investigate it, are and tool curation. Am. Ant., 51: 38-50. arguably the most outstanding contribution that Bamforth D.B. & Bleed P. 1997. Technology, developments in more recent decades have made. flaked stone technology, and risk. Archaeol. Moreover, they hint that methodological innova- Papers Am. Anthropol. Ass., 7: 109-139. tions that more rigorously allow each of these fac- Barnes A.S. 1939. The differences between natu- tors to be investigated, will also be where the great- ral and human flaking on prehistoric flint im- est contributions can be made in the future. plements. Am. Anthropol., 41: 99-112. Barsky D., Chapon-Sao C., Bahain J.-J., Beyene Y., Cauche D., Celiberti V., Desclaux E., de Acknowledgements Lumley H., de Lumley M.-A., Marchal F., Moullé P.-E. & Pleurdeau D. 2011. The early AK’s research is funded by a British Academy Post- Oldowan stone-tool assemblage from Fejej FJ- doctoral Fellowship. SJL’s work is supported by the 1a, . J. Afr. Archaeol., 9: 207-224. Research Foundation for the State University of Bar-Yosef O., Eren M.I., Yuan J., Cohen D.J. & New York. We are grateful to the editorial board Li Y. 2012. Were bamboo tools made in pre- for their invitation to undertake this review and historic ? An experimental view the anonymous reviewers for their constructive and form South China. Quat. Int., 269: 9-21. detailed comments. We are indebted to Morag Mas- Beck C., Taylor A.K., Jones G.T., Fadem C.M., Cook ters for the illustrations provided in Figure 3. C.R. & Millward S.A. 2002. Rocks are heavy: trans- port costs and Paleoarchaic quarry behaviour in the Great Basin. J. Anthropol. Archaeol., 21: 481-507 References Bello S.M., Parfitt S.A. & Stringer C. 2009. Quantitative micromorphological analysis of Aiston G. 1928. Chipped stone tools of the cut marks produced by ancient and modern Aboriginal tribes east and north-east of Lake handaxes. J. Archaeol. Sci., 36: 1869-1880. Eyre, South Australia. Papers and Proceedings of Bettinger R.L., Winterhalder B., McElreath R., the Royal Society of Tasmania, pp. 123-131. 2006. A simple model of technological intensi- Ambrose S.H. 2001. technology and fication. J. Archaeol. Sci., 33: 538-545. human evolution. Science, 291: 1748-1753 Beyene Y., Katoh S., WoldeGabriel G., Hart Antón S.C., Potts R. & Aiello L.C. 2014. W.K., Uto K., Sudo M., Kondo M., Hyodo Evolution of early Homo: An integrated biologi- M., Renne P.R., Suwa G. & Asfaw B. 2013. cal perspective. Science, 345: 1236828. The characteristics and chronology of the ear- Archer W. & Braun D.R. 2010. Variability in liest Acheulean at Konso, Ethiopia. Proc. Natl. bifacial technology at Elandsfontein, Western Acad. Sci. U.S.A., 110: 1584-1591. cape, South Africa: a geometric morphometric Beyries S. 1990. Problems of interpreting the approach. J. Archaeol. Sci., 37: 201-209. functional results for ancient periods. In K. A. J. M. Key & S. J. Lycett 95

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