Evolution

The emergence of lithic technology by ∼2.6 million years ago (Ma) is often interpreted as a correlate of increasingly recurrent hominin acquisition and consumption of animal remains. Associated faunal evidence, however, is poorly preserved prior to ∼1.8 Ma, limiting our understanding of early archaeological (Oldowan) hominin carnivory. Here, we detail three large well-preserved zooarchaeological assemblages from Kanjera South, Kenya. The assemblages date to ∼2.0 Ma, pre-dating all previously published archaeofaunas of appreciable size. At Kanjera, there is clear evidence that Oldowan hominins acquired and processed numerous, relatively complete, small ungulate carcasses. Moreover, they had at least occasional access to the fleshed remains of larger, wildebeest-sized animals. The overall record of hominin activities is consistent through the stratified sequence – spanning hundreds to thousands of years – and provides the earliest archaeological evidence of sustained hominin involvement with fleshed animal remains (i.e., persistent carnivory), a foraging adaptation central to many models of hominin evolution.

We report here on the zooarchaeological record of bovid remains. These dominate the assemblages in terms of overall abundances (representing a minimum of 56 individuals), and are amenable to analysis using published protocols and experimental datasets [21]–[30], [56]–[63]. Analytically, we group remains by bed (e.g., ‘KS-1’, ‘KS-3’) rather than by excavation [49]. We further sort specimens by body size class [21], grouping animals into ‘small’ (e.g., Grant’s gazelle, Gazella granti) and ‘medium’ (e.g., Topi, Damaliscus lunatus) sizes. Extinct bovids of intermediate size and weight (e.g., Parmularius sp.) are treated as medium-size animals. Larger bovids (e.g., buffalo, Syncerus caffer) are poorly represented in the assemblages and are not treated in detail here. Following convention, we incorporate taxonomically-unidentifiable long bone fragments in all appropriate analyses.

In our study of bone surface modifications, three investigators (JVF, BLP, and JSO) jointly analyzed specimens, shared observations, and discussed interpretations before providing individual assessments of bone damage [17]. Analysts employed low–power magnification (10×-40×) and strong light sources to identify modifications. They attributed agency (e.g., hominin, carnivore) to modifications only after excluding all possible alternatives (including potential confounds detailed in [32], [64]–[69]).

Values for minimum numbers of skeletal elements (MNE) reflect considerations of animal size and developmental age, extensive refitting efforts, and, for long bones, element identification of shaft portions [21]. High-survival elements (HSE) include the cranium, mandible, humerus, radius, metacarpal, femur, tibia, and metatarsal [61]. Point estimates of Shannon evenness follow published methods [30], [70], whereas interval estimates are constructed using Bayesian models [71].

Bone surface modification frequencies are known to accurately reflect the timing and context of both hominin and carnivore involvement with animal remains. We use them here to assess the identity and sequence of actors and behaviors responsible for forming and modifying the assemblages.

Hominin-modified specimens (i.e., fossil bones bearing cut marks and/or hammerstone percussion damage) are present through the entire KS-1 through KS-3 sequence (Table 2 and Table S1). These specimens provide unambiguous evidence of hominin processing of bovid remains (Figure 2), and indicate a functional relationship between artifactual and faunal materials. When considering the anatomical placement of cut marks, we report bone damage consistent with both defleshing and disarticulation activities [17]. Frequencies of cut-marked limb specimens range from 1.9% to 6.3% in summed (i.e., total bed) assemblages, with similar frequencies observed irrespective of analyst, bed, or animal body size. The overall uniformity of these results suggests a relatively consistent pattern of carcass exploitation through time (within-analyst test for the homogeneity of cut mark frequencies across beds: homogeneity not falsified, all p-values >0.1).

With respect to the timing of hominin access to these smaller-sized individuals, actualistic studies in a modern East African grassland (the Serengeti) show that small bovid carcasses are, almost without exception, completely consumed by lions and/or hyenas within the first few minutes to hours following death [63]. Given the relative abundance of small bovid carcasses at KJS (Table S3), the relative dearth of carnivore tooth marks on their remains (Table S1), and the inferred rarity of such scavenging opportunities in grassland settings, our results strongly suggest that hominins acquired many of these animals very early in their resource lives (i.e., fairly close to the moment of death). At present, the summed evidence that Oldowan foragers acquired, defleshed, and demarrowed numerous, complete, small bovids throughout the formation of all three assemblages plausibly represents the earliest archaeological record of hominin hunting activities.

The skeletal remains of medium-sized bovids reflect a slightly different taphonomic history. Although specimens from all skeletal regions are represented, cranial remains predominate (Figure 5B). Within each assemblage, skeletal element abundances are positively correlated with bone densities (rs range: 0.401 to 0.666; all p-values <0.10) [59], and HSE abundances are not significantly correlated with either standardized food utility values (rs range: −0457 to −0.241; all p-values >0.20) [62] or within-bone nutrient values (rs range: 0.107 to 0.657; all p-values >0.10) [28], [29]. When considering the sum of surface modification data, Shannon evenness values (range: 0.808 to 0.944), and theoretical considerations of transport behaviors [61], [62], the record from KJS most parsimoniously indicates that Oldowan hominins introduced the partial remains of medium-sized carcasses to the site, with specific foraging behaviors varying with respect to body region (e.g., head versus postcrania) and timing of access to carcasses [63].

The overall taphonomic history of medium-sized postcrania is thus fairly equivalent to that of the smaller-sized carcasses. In both cases, remains are present at abundances that far exceed natural landscape accumulation norms (Table 1), and bone surface modification frequencies and skeletal part analyses indicate that hominins had primary access to soft tissues (Table 2, Figure 3, Figure 4). The evidence is consistent with scenarios in which hominins introduced a relative abundance of fleshed medium-sized postcrania to the site. In contrast to the record of smaller-sized bovids, however, skeletal element representation and element evenness scores suggest an increased measure of hominin selectivity in skeletal part choice and transport decisions when dealing with medium-sized remains (Table S3, Table S8). Long bone elements are fairly numerous relative to axial remains (as measured by %MAU) (Figure 5B, Table S3); and the more proximal limb elements (i.e., humerus, radio-ulna, femur, and tibia) are relatively more abundant than metapodials (Figure 5B, Table S3). This patterning differs substantially from that of the smaller-sized bovids. The latter’s remains are more evenly-distributed across the entire postcranial skeleton (HSE’s+low survival elements [LSE’s]), as well as across the six major long bones (Figure 5A, Table S3), and presumably reflects the introduction of numerous, fairly complete small bovids to the site. At issue here: what strategies did hominins follow when selecting and transporting medium-sized remains?

The record is potentially consistent with two main scenarios. In the first, hominins introduce an abundance of compete (or relatively complete) medium-sized carcasses to the site. This follows a ‘food maximizing’ strategy in which hominins face negligible-to-minor transport constraints and transfer most or all of the edible remains from death sites to KJS [75]. As a result, they treat both small and medium-sized bovids in a relatively similar manner when making carcass transport decisions. Observed differences in skeletal element records on-site (smalls vs. mediums) would then presumably reflect systematic differences in post-depositional carnivore scavenging behaviors. In the second scenario, hominins preferentially transport limb remains from medium-sized carcasses, plus some axial elements whenever possible. This follows a ‘weight minimizing’ strategy in which transport constraints (e.g., the number of available carriers, distance to destination, predation risk, etc.) limit hominins to carrying away only a subset of all edible tissues [75]. In this case, carnivore treatment of skeletal remains on-site would be relatively consistent across size groups [25], and observed differences in the skeletal element record (small vs. medium) would instead predominantly reflect systematic size-based differences in hominin transport practices.

Here, comparisons between size groups are particularly informative. For small bovids, LSE values are not grossly disproportionate to those of HSE’s (Figure 5A, Table S3). In fact, their overall skeletal record corresponds fairly well to expectations for dual-patterned hominin-first assemblages, [22], [25], [27], [29]. Note too that skeletal remains of smaller-sized individuals are usually at far greater risk of destruction than those of medium-sized animals, especially in grassland contexts [43], [63].This makes the latter’s record at KJS all the more interesting. In each of the assemblages, medium-sized bovids are fairly depauperate in postcranial axial remains relative to both head and limb elements (Figure 5B, Table S3). As the smaller-sized bovids are more evenly represented across the skeleton (both with and without considerations of cranial remains), we discount the possibility that hominins introduced a substantial amount of medium-sized postcranial axial elements to the assemblages (or, alternatively, that those remains were somehow introduced ‘naturally’; e.g., via mass death). In short, if an abundance of medium-sized axial remains were originally present on-site in substantial numbers, and they were largely deleted by scavenging carnivores, then the overall skeletal record of smaller-sized bovids should reflect a substantially more biased record (both in terms of head remains relative to postcrania, and HSE’s relative to LSE’s). The alternative, a null hypothesis in which all bovids were originally present on-site as similarly-apportioned carcasses, would require that medium-sized postcrania (LSE’s+HSE’s) were preferentially deleted by carnivores relative to all smaller-sized remains. We argue that this is unlikely (especially for the record of HSE’s), and note that tooth-mark frequencies are relatively similar across the remains of both size groups (Table S1). In turn, we argue that the KJS record provides robust evidence that hominins largely – but certainly not exclusively – followed a ‘weight-minimizing’ strategy at KJS when selecting and transporting remains from fleshed medium-sized carcasses.

The record of medium-sized cranial elements requires a bit more explanation. Specifically, these remains are disproportionately abundant within each of the assemblages (Figure 5B, Table S3). If hominins largely followed a ‘weight-minimizing’ strategy, and solely had access to complete medium-sized carcasses, they would not have preferentially transported crania and mandibles to KJS. The reason is clear: head remains are quite heavy given their tissue yields, and will often be ignored at death sites in favor of a slew of higher-ranked remains [75]. These same arguments hold when discussing medium-sized limb HSE’s. The preponderance of head remains on-site (as well as the paucity of long bone remains) is thus unlikely to reflect either simple utility or density-related phenomena. Instead, the record strongly suggests the purposeful introduction of a fair number of isolated heads to the site by Oldowan foragers.

But why acquire, transport, and process an abundance of medium-sized heads? In living animals, these remains contain a wealth of fatty, calorie-packed, nutrient-rich tissues: a rare and valuable food resource in a grassland setting where alternate high-value foodstuffs (fruits, nuts, etc.) are often unavailable [2], [3], [29], [49], [52], [63], [76]–[78]. Medium-sized heads are also relatively dense and durable elements, and their internal contents are generally inaccessible to all but hyenas and tool-wielding hominins [63], [79], [80]. As a result, they are often seasonally-available as scavengable resources in East African grasslands [63], [76], [79]–[83]. Additionally, bone surface modification studies at KJS clearly demonstrate that hominins accessed internal head contents: several cranial vault and mandibular fragments bear evidence of percussion striae. Considered in sum, the presumed availability of these isolated remains across the landscape, the relative abundance of these remains in the KJS assemblages, and unambiguous material evidence that hominins exploited their contents on-site is most parsimoniously interpreted as reflecting very early archaeological evidence of a distinct hominin scavenging strategy – one that included a strong focus on acquiring and exploiting fatty, nutrient-rich, energy-dense within-head food resources (e.g., brain matter, mandibular nerve and marrow, etc.) [e.g., 24,63,76,82,84–86].

The total abundance of remains on site, (Table 1), the number of animals represented (Table 1), the high taxonomic diversity present [17], [50], [52], the relatively low frequency of tooth-marked specimens (Figure 3, Figure 4, Table S1), and a sedimentological record wholly inconsistent with a fluvial accumulation of remains [49], [52] also combine to suggest that the KJS assemblages are unlikely to represent in situ death or ‘background scatter’ accumulations formed by non-hominin agencies. Similarly, the skeletal element record of medium-sized bovids suggests that they were unlikely to have been present on-site as complete carcasses, an expectation of most ‘kill-site’ and/or landscape accumulation models. When limiting discussion to medium-sized postcrania, the high abundance of limb remains (including many isolated epiphyses) relative to axial elements is also the inverse expectation for landscape assemblages (Figure 5B) [63].

Finally, as with many zooarchaeological assemblages, the KJS skeletal inventories are dominated by numerous unidentifiable long bone shaft fragments. At issue: who or what created these fragments from whole bones? The relative rarity of ‘dry bone’ fractures, coupled with abundant evidence of ‘green bone’ breakage, strongly suggests the involvement of behavioral agents of modification, especially given the inferred low-energy depositional setting at KJS [17], [49]–[52]. Bone surface modifications (e.g., percussion marks and notches; tooth marks and notches) indicative of access to within-bone resources, however, are found at relatively low frequencies in each of the assemblages (Figure 3; Figure 4; Table 2; Table S1; Table S2) [17]. This result is surprising as it is inconsistent with known outcomes of both hominin and carnivore bone breakage practices, where surface modification frequencies are, on average, substantially higher [e.g., 22,23,25,57,58]. A similar pattern of an abundance of shattered but largely unmodified long bone specimens is observed in many other Paleolithic assemblages [31,45,72,73; Table S2], suggesting to us that current bone breakage models may not fully account for all relevant variables. Notably, at KJS there is no evidence that post-depositional sediment compaction and/or bone weathering influenced the bone breakage record [17]. Further experimental research may be required to fully explain these observations.

3.2. Middle Palaeolithic Hunting: Sites such as Zwolen (Gautier, 1989) and Mauran (Farrzy & David, in press; Girard-Farrzy & Leclerc,1981) preserve clear evidence of active hunting. 

Planning: killings are less often occasional. Neanderthal man returns periodically (or seasonally) to special places rich in game and with a natural topography propitious to hunting activities. This testifies to an intentional and calculated choice, as at the sites already mentioned. 

Specialisation: sometimes man specialises in the capture of a particular animal species: big bovids at Mauran (Farizy & David, in press), horses at Zwolen (Gautier, 1989), wild goats at the Grotte de l'Hortus (de Lumley, 1971). 

Hunting techniques: probably some kind of game driving was practised at Mauran (Farizy & David, in press), Zwolen (Gautieq, 1989), La Quina (Jelinek, Debenath & Dibble, 7989) and La Cotte de Saint-Brelade (Scott, 1e80). 

Seasonal killings: many killings are probably seasonal, animals fall in discrete age groups at Zwolen (Gautieq, 1989) and La Quina (]elinek, Debenath & Dibble, 1989). 

Food transport: the lightest and most meaty bones (hind limbs, pulni, ribs, vertebrae) may be carried away. In kill sites man leaves big and useless parts of animal skeletons (skulls, jaws etc.). Transport of meaty skeletal parts may be exemplified at Mauran (Farizy & David, in press). 

Butchering activities: at Maurary Farizy and David (Fafizy & David, in press) notice many phases in the butchering process: dismemberment, removal of muscular masses and bone breakage for marrow extraction.

Upper Palaeolithic and Mesolithic Hunting: 

the archaeological record leaves us some direct evidence of man's hunting activities. At Meiendorf (Rust 1937) and Stellmoor (Rusf 1937), some bones of reindeer and birds still conserve weapon marks and a few pieces of silex have remained thrusted in mammalian bones; man kills reindeer with harpoons and sticks (fractured skulls), birds with bows and maybe slings. Three fractured skulls of red deer in Abri Pataud (Bouchud, 1975), and one bovid skull with a circular orifice in Saint Marcel (Allain, 1952) suggest the practice of the so called " co'up de merlin": man has delivered a blow similar to the one used today to butcher cattle. Probably the animal already immobilized (wounded or entrapped) was hit on the frontal with a big stone. At Kokorevo I (Siberia), a large scapula of bison is pierced by the upper end of a point made of bone (Boriskowksi, 1965). At High Furlong (Mesolithic), an elk was discovered with the marks of L7 wounds made by barbed points, of which two were found in the site, and by other arms. The animal had apparently been attacked at two distinct occasions: during the first one, hunters aimed at the legs to lame the animal (fig. 6), later hunters hit the thoracic region and the lungs to kill it. However the elk died in a little lake, perhaps imprisoned in the ice, and man had no access to the meat. The animal represents in fact a hunting loss (Hallam et a1.,1973). 

Planning: very good. Many sites belong to Wpe e, were occupied periodically or seasonally and specialised in the capture of a particular game (e.g., horse, reindeeq, ibex). Game drive towards cliffs have been claimed and Solutre (Combier & Thevenot,1976) has long figured as an example, but the evidence is far from conclusive. 

Scavenging: no doubt H. sapiens still killed or exploited animals in the occasional and opportunistic way of Lower Palaeolithic times. According to Lindner (Lindner,1941), hunters at Predmost utilised the carcasses of hundreds of mammoths that probably succumbed as a result of natural catastrophes, as food. 

Food transport: selective transport of the most useful animal parts is claimed for many sites. 

Specialised activities: sometimes the material is dislocated in distinct clusters that could reflect specialised activity areas as for example at Solutre (Combier & Thevenot, 1976). Site topography: some hunting sites were located in valleys enclosed by steep slopes as at Rascano (Gonziilez-Echegaray, 1979), Stellmoor (Rust, 1937), Meiendorf (Rust 1937), or at the foot of rocky cliffs at Solutr6 (Combier & Th6venot, 1,976). 

4. Conclusions 

Most of the Lower Palaeolithic sites analysed here belong to category a (butchering sites); other kind of concentrations are rare and difficult to ascertain. A number of hunting stations (category e) and a hunting stop (category f) form my sample for the age of Neanderthal man and related people. The Upper Palaeolithic is characterised by many hunting stations, while in Mesolithic times a hunting loss (category d ) was found as well as several sighting sites (category g). The foregoing distribution seems to reflect in a vague way an evolution from scavenging and haphazard opportunistic hunting to well organised, selective hunting activities. However, this reflection results no doubt in part from a priori assumptions concerning the evolution of hominid meat procurement often colouring the interpretations offered for the osseous "hard" data; these are frequently equivocal.