Quaternary Research 74 (2010) 395–404

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Quaternary Research

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Bovid mortality profiles in paleoecological context falsify hypotheses of endurance – and passive scavenging by early Pleistocene hominins

Henry T. Bunn a,⁎, Travis Rayne Pickering a,b a Department of Anthropology, University of Wisconsin-Madison, 1180 Observatory Drive, 5240 Social Science Building, Madison, Wisconsin, 53706, USA b Institute for Human Evolution, University of the Witwatersrand, Private Bag 3, WITS 2050, Johannesburg, South Africa article info abstract

Article history: The world’s first archaeological traces from 2.6 million years ago (Ma) at Gona, in Ethiopia, include sharp- Received 21 February 2010 edged cutting tools and cut-marked animal bones, which indicate consumption of skeletal muscle by early hominin butchers. From that point, evidence of hominin meat-eating becomes increasingly more common Keywords: throughout the Pleistocene archaeological record. Thus, the substantive debate about hominin meat-eating Uniformitarianism now centers on mode(s) of carcass resource acquisition. Two prominent hypotheses suggest, alternatively, Bovid mortality profiles (1) that early Homo hunted prey by running them to physiological failure and then dispatching Carcass foraging Early Homo them, or (2) that early Homo was relegated to passively scavenging carcass residues abandoned by carnivore FLK 22 Zinjanthropus predators. Various paleontologically testable predictions can be formulated for both hypotheses. Here we test four predictions concerning age-frequency distributions for bovids that contributed carcass remains to the 1.8 Ma. old FLK 22 Zinjanthropus (FLK Zinj, Olduvai Gorge, Tanzania) fauna, which zooarchaeological and taphonomic data indicate was formed predominantly by early Homo. In all but one case, the bovid mortality data from FLK Zinj violate test predictions of the endurance running-hunting and passive scavenging hypotheses. When combined with other taphonomic data, these results falsify both hypotheses, and lead to the hypothesis that early Homo operated successfully as an ambush predator. © 2010 University of Washington. Published by Elsevier Inc. All rights reserved.

Introduction Notably, the latter context is comparable to the reconstructed savanna–bush–woodland habitats in which early Homo evolved A recent, high-profile hypothesis contends that early Homo (e.g., Hay, 1976, 1990; Cerling, 1992; Cerling et al., 1988; Reed, employed long distance, endurance running (ER) to hunt and 1997; Sikes, 1994). We proposed that an understanding of predator– scavenge. More particularly, the hypothesis asserts ER was the prey dynamics, as reflected in large ungulate mortality profiles, specific mode of a general type of —persistence hunting enables the development of testable hypotheses about ER-PH in early (PH)—in which ungulate prey is pursued over a long distance until it Homo. We then noted that an actual test of ER-PH using the published fails physiologically and can then be easily dispatched by the hunter ungulate mortality profile from FLK 22 Zinjanthropus (FLK Zinj, (s) (Bramble and Lieberman, 2004). In our recent discussion of that Olduvai Gorge, Tanzania), the best available assemblage of butchered hypothesis, we emphasized paleoecological and, specifically, vegeta- fossil bones from a Pleistocene context (e.g., Bunn and Kroll, 1986, tional context as a key determinant of the viability of ER-PH in 1988; Bunn, 2007; Domínguez-Rodrigo et al., 2007a), flatly contra- tracking and hunting prey animals (Pickering and Bunn, 2007). As a dicts the test predictions of the ER-PH hypothesis, falsifying it. contribution to a uniformitarianist database relevant to paleoanthro- In a response, Lieberman et al. (2007) imagine we practice the pology, we cited two ethnographic examples from our own empirical “tyranny of ethnography,” but do not comment on the impact of research: one from the Kalahari (Botswana), in which sparse vegetational context on the evolution of tracking skills in early Homo vegetation and a sandy substrate provide an ideal setting for except to assert definitively that early Homo possessed the necessary successful hunting by Bushmen using skilled tracking and skills to track. Further, Lieberman et al. are reticent regarding our (our example) or ER (Bramble and Lieberman's example from the proposal and testing of ER-PH with mortality data on ungulate prey, literature), and one from the Lake Eyasi region of Tanzania, in which and they offer, instead, a claim about the unlikelihood of efficient much heavier, savanna–bush–woodland vegetation inhibits tracking hunting or scavenging by hominins lacking bows and arrows or stone- of wounded prey or any involvement in ER-PH by Hadza foragers. tipped spears, except by ER-PH. We continue to believe, nevertheless, that ungulate mortality data provide a potentially decisive means of testing ER-PH and other ⁎ Corresponding author. current hypotheses about hominin strategies of prey acquisition in the E-mail address: [email protected] (H.T. Bunn). early Pleistocene, and that hominin foraging adaptations are most

0033-5894/$ – see front matter © 2010 University of Washington. Published by Elsevier Inc. All rights reserved. doi:10.1016/j.yqres.2010.07.012 396 H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404 appropriately viewed in an ecological context. In this paper, we 1998). Bunn and Kroll (1986) reported a catastrophic mortality profile develop further the application of ungulate mortality data and for the larger from FLK Zinj, but its meaning was left in doubt vegetational context to the ER-PH hypothesis. in deference to Klein's (1982: 153, 1986: 446–447, 1999: 461) In addition, our data provide a relevant test of the well-known preference for a very young life-history threshold for defining the passive scavenging hypothesis (e.g., Binford, 1981, 1985, 1988; onset of old age (i.e., 40%–50% of potential ecological longevity [PEL] in Blumenschine, 1986, 1987, 1988, 1991, 1995; Blumenschine et al., African buffalo [Syncerus caffer] and, he argued, other large ungulates as 1994), which contends Pleistocene hominins enjoyed very limited well (Klein, 1978, 1982:153;Klein and Cruz-Uribe, 1984: 50, 56, 1996: access to fleshed carcasses. According to this view, even those carcass 322)). parts that hominins infrequently secured were already picked-over by Here, bolstered by a new, behaviorally relevant consideration of carnivores, leaving no appreciable “surplus” resources for hominin bovid age classes (summarized here and fully explicated in an scavengers to share. Binford's argument for passive scavenging was accompanying paper; Bunn and Pickering, 2010), we use the FLK Zinj basedsimplyonarejectionoftheresultsofcomprehensive bovid mortality data to test four discrete predictions that emanate taphonomic analysis of skeletal element representation and cut- from the ER-PH and passive scavenging hypotheses. One prediction mark abundance that belied his beliefs on the bone assemblage from for each hypothesis concerns large, size group 3 bovids and one for FLK Zinj. Blumenschine, in contrast, acknowledged the taphonomic each hypothesis concerns smaller, size group 1–2 bovids (see Brain evidence, including the abundance of cut marks on once-meaty long (1974, 1981) or Klein (1976) for definitions of bovid size classes, and limb bone shafts, but questioned its interpretation by arguing that a Bunn (1986; Bunn et al, 1980) for definitions of more inclusive perceived abundance of carnivore gnawing on long limb elements mammal size groups developed from them). indicated that most meat had already been consumed by primary carnivores prior to acquisition of the carcasses by hominins. According Test prediction 1 (large bovids) to Blumenschine, passive scavenging from defleshed and abandoned lion kills of large bovids and leopard (Panthera pardus) kills of small If early Homo used ER-PH to obtain the size 3 bovids at FLK Zinj, bovids would have yielded marrow bones but marginal remnants of then the mortality data should match what predators of meat, without requiring either hunting or aggressive scavenging by large bovids are known to kill (which, preferentially, is the relatively hominins. vulnerable young and old, more than prime adults). Additional actualistic and experimental research, however, contra- dicts the evidence and reasoning underlying the passive scavenging Test prediction 2 (large bovids) hypothesis in several ways. First, observations of lion (P. leo)killsby Domínguez-Rodrigo (1999) document complete defleshing of midshaft If early Homo passively scavenged from large felids to obtain the portions of upper long limb elements (i.e., humeri and femora), leaving size 3 bovids at FLK Zinj, then the FLK Zinj mortality data should match no meat to remove in the precise anatomical locations where defleshing what lions (as the best available modern proxy for large felid cut marks are abundant in the FLK Zinj fauna. Second, strong similarities predators of size 3 bovids) are known to kill (i.e., a relatively higher in the anatomical patterning of cut marks on long limb bone shafts from percentage of prime adults than cursorial predators such as spotted thorough defleshing of fully-fleshed limbs by Hadza foragers and from hyenas [Crocuta crocuta]). hominin butchery at FLK Zinj, indicate that hominins at FLK Zinj Even though ER-PH and passive scavenging are generally consid- similarly possessed fully-fleshed limbs and butchered them for ered less productive and less likely as strategies for acquiring small significant amounts of meat (Bunn, 2001, 2007). Third, Domínguez- bovids, test predictions are similar to those for large bovids. Rapid Rodrigo and Barba (2006; Domínguez-Rodrigo et al., 2007a) established destruction by feeding carnivores (i.e., large felids or hyenas) of entire that the incidence of carnivore gnaw marks on long limb elements at FLK small carcasses, bones and all, renders passive scavenging for small Zinj is actually far lower than what Blumenschine reported, because bovids unlikely, although the possibility that early Homo, as a non- biochemical processes damaged bone surfaces and left many marks hunter, could have scavenged from abandoned, tree-stored leopard mistakenly identified as carnivore gnawing by Blumenschine. For kills, has been proposed (Cavallo and Blumenschine, 1989). For expanded discussion of carcass acquisition by early Pleistocene present purposes, test predictions using mortality data as the test can hominins, see reviews by Domínguez-Rodrigo (2002) and by Pickering be tailored directly to these scenarios for small bovids. and Dominguez-Rodrigo (2006). Test prediction 3 (small bovids) Predictions and tests of the ER-PH and passive scavenging hypotheses If early Homo used ER-PH to acquire smaller, size group 1–2 bovids We test the ER-PH and passive scavenging hypotheses using the at FLK Zinj, then the FLK Zinj mortality data should match what bovid mortality data from the ca. 1.8 million year old (Ma) FLK Zinj cursorial predators are known to kill (again, the relatively vulnerable zooarchaeological assemblage excavated by Mary Leakey (1971) in young and old). Bed I at Olduvai Gorge. As discussed above, the FLK Zinj assemblage exhibits the strongest signal of butchery by hominins of any Bed I site, Test prediction 4 (small bovids) and indeed, any early Pleistocene site, which implicates early Homo as the dominant taphonomic agent in the transport and systematic If early Homo passively scavenged from tree-stored leopard kills to butchery of large prey animals recovered there (Bunn, 1981, 1982, obtain small bovids at FLK Zinj, then the FLK Zinj mortality data should 1986; Bunn and Kroll, 1986, 1987, 1988; Potts, 1988). From match what leopards are known to kill (i.e., a non-age-selective, taphonomic analysis of the FLK Zinj assemblage, Bunn reported more living-structure sample). than 200 bones with cut marks, abundant hammerstone percussion damage, and an abundance of limbs and mandibles of an estimated 48 Materials and methods large mammals (minimum number of individuals, MNI) represented. More recent taphonomic analyses have refined the evidence and Paleontological samples debated its meaning but without changing the fundamental conclusion that early Homo had the dominant role in forming the bone assemblage In addition to testing the ER-PH and the passive scavenging (Blumenschine, 1995; Bunn, 2001, 2007; Capaldo, 1997; Domínguez- hypotheses with the bovid mortality data from the FLK Zinj Rodrigo and Barba, 2006; Domínguez-Rodrigo et al., 2007a; Selvaggio, assemblage, we also present comparative paleontological data from H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404 397 eight other excavated early Pleistocene levels in Tanzania and South From the same perspective, we also analyze fossil bovid mortality Africa. As with FLK Zinj, three of these levels are in Bed I at Olduvai data from three South African cave sites (five stratigraphic levels), Gorge. But, in contrast to FLK Zinj, FLK North level 1–2 (FLK N 1–2), which are inferred predominantly as products of large carnivore FLK North North level 2 (FLK NN 2) and FLK North level 6 (FLK N 6) predation, although recent taphonomic analyses have established that show little or no evidence of anthropogenic input. Leakey (1971) hominin butchery activities also contributed to site formation in some interpreted FLK N 1–2 as a hominin living floor based on her recovery cases (Brain, 1981, 1993; Watson, 1993a; Pickering et al., 2004a,b, of ~1205 stone artifacts and 3294 taxonomically varied large mammal 2005, 2007, 2008; Adams, 2006; Thackeray, 2007). Swartkrans, bones (plus abundant microfauna) from the ~65 cm-thick level at the Kromdraai and Gondolin all formed as phreatic maze-caves in the site. From preliminary taphonomic analysis, Bunn (1982, 1986) dolomites of the Chunniespoort Group, probably sometime in the tentatively agreed, based on the finding of a few cut-marked Miocene and only opened to the land surface at various times specimens but a notable lack of cut marks in anatomical locations throughout the Pliocene and Pleistocene, when they were then able to where they are common at FLK Zinj (e.g., distal humerus shafts) and a capture and incorporate terrestrial debris into their paleoanthropo- paucity of hammerstone damage or refitting long limb bone shafts logically productive breccias. Swartkrans is the best understood of the (which are also common at FLK Zinj). A more recent taphonomic sites in terms of stratigraphy and has yielded the richest traces of early analysis reported butchery damage on these bones and interpreted hominin behavior—including Developed Oldowan/Early Acheulean the fauna as anthropogenic in origin, even more so than FLK Zinj stone tools, butchered animal bones and bone digging tools—in its (Capaldo, 1998; Blumenschine et al., 2007). In a thorough reanalysis three Pleistocene units, Members 1–3 of the Swartkrans Formation; using a more inclusive, current understanding of taphonomic and Member 3 also produced bones that were possibly burned in diagenetic processes, Domínguez-Rodrigo et al. (2007a) demonstrate hominin-tended fires (Brain, 1993). While it is important to stress convincingly that the anthropogenic damage alleged by Capaldo and that, overall, hominins contributed only minimally to the ca. 1.0– Blumenschine is, in fact, a product of trampling of the bones by large 1.8 Ma faunas from Swartkrans Member 1–3, it is equally important to animals, sediment abrasion, and large carnivore gnawing. While emphasize that those fossils that were modified by hominins are acknowledging the presence of a few bones with actual stone-tool cut informative behaviorally: the total butchered sample from each marks, Domínguez-Rodrigo et al. (2007a) attribute the fauna member is comprised of high proportions of meat-bearing upper predominantly to predation and feeding by large felids. and intermediate (i.e., radioulnae and tibiae) long limb elements, At FLK NN 2, Leakey (1971) reported a low-density scatter of 481 indicating early access to carcasses by hominin foragers (Pickering bones, but no stone artifacts, from a ~30 cm-thick layer of tuff with et al. 2007, 2008) (see preceding discussion). clayey patches, which she attributed to hyena activity. Bunn (1982, No hominin fossils have been recovered from Kromdraai A. 1986) reported abundant carnivore damage on the bones, plus one However, Thackeray et al. (2005) employed X-ray diffraction analysis specimen with stone-tool cut marks, and generally concurred with to detect bone apatite on one of the polyhedral core artifacts Leakey's interpretation, as did Potts (1988). Reanalysis by Domínguez- discovered at the site (Kuman et al., 1997). Subsequently, Thackeray Rodrigo et al. (2007a) indicated large felid, rather than hyena, activity as (2007: 67) elaborated that this finding, combined with the site's the source of the bones. carnivore:ungulate ratios and long limb bone shaft fragment size At FLK N 6, Leakey (1971) recovered an essentially entire skeleton distribution, indicates that hominins used the Kromdraai A artifacts to of an elephant (Elephas recki), 123 stone artifacts, and ~400 bones of “opportunistically…obtain bone marrow from [the site, which was other large mammals, from a ~45 cm-thick layer of silty clay. Leakey also] used at least temporarily by large carnivores such as saber-tooth interpreted the site as an elephant butchery by hominins in a swampy cats, which preyed primarily on juvenile alcelaphines.” Large area, a notable conclusion that was at least partially supported by carnivores have long been implicated as primary bone collectors at subsequent reports of rare cut marks on a few of the elephant bones Kromdraai A (e.g., Brain, 1981). Thackeray's specific accusation of (Bunn, 1982, 1986; Potts, 1988). From taphonomic analysis of the Dinofelis is based, in part, on its presence in the Kromdraai A fauna non-elephant (mostly bovid) bones, Bunn (1982, 1986) concluded (Brain, 1981). Thackeray and his co-workers (Thackeray and Van that the fauna was predominantly a non-anthropogenic, natural Leuvan Smith, 2001; Lesnik and Thackeray, 2006) argue that the background, with a bovid MNI of 35, many complete bovid long limb mortality profiles (based on molar crown height measurements) of bones (likely biased during excavation against recovery of fragmen- alcelaphines and Equus show a pattern similar to that created by tary long limb bone shafts), many specimens striated from trampling, modern lions: with the old of smaller species and juveniles of larger and several specimens (bovid long limb elements and a hippopota- species preferentially represented. Thackeray and Van Leuvan Smith mus metacarpal that had been misidentified as a rhinoceros) (2001: 11) then cite that “Lewis (1997) has previously suggested that exhibiting probable stone-tool cut marks. Domínguez-Rodrigo et al. Dinofelis displayed behavior similar to that of modern lions.” (2007a: 110) conclude that all of the linear grooves on elephant and A fossil assemblage excavated by E.S. Vrba and D. Panagos from bovid bones (the hippopotamus metacarpal was not located for sediments adhering to the northeastern cave wall of Gondolin does reexamination) at FLK N 6 “can be best explained as abrasion marks.” not contain hominin remains or artifacts (Watson, 1993b). More For present purposes, we analyze bovid mortality data from FLK N recent excavation of a test trench into an ex situ breccia dump, created 1–2, FLK NN 2, and FLK N 6, as samples of probable non- by early 20th century lime miners at the site, has yielded the isolated anthropogenic, natural background occurring on the Olduvai paleo- M2 of a robust australopithecine and an isolated M1 or M2 of a non- landscape during the Bed I time interval under study. This enables an robust hominin (Menter et al., 1999). No stone tools were recovered initial investigation for any resemblances or differences in bovid from the dump excavation, but one of us (TRP) did identify, in the mortality between these likely natural samples and the predomi- field, an ungulate long limb bone shaft fragment that preserves nantly anthropogenic sample from FLK Zinj. We do not regard these possible stone-tool cut marks. However, an exhaustively detailed data as “controlled,” simply because many of the specific events and study of the Gondolin faunas (Adams, 2006) supports Watson's processes involved are unknown or unknowable. We view all of these (1993b) earlier conclusion that the bovids from the Vrba–Panagos sites as natural history sites, or palimpsests (e.g., Isaac, 1984; sample were probably collected and modified exclusively by leopards. Domínguez-Rodrigo et al., 2007a), in which multiple taphonomic agents, including hominins, had varying contributions. For more Paleontological analyses controlled comparative analysis, we then rely on data from modern wildlife research in which the predation patterns of various large Because the Olduvai and South African bovid dental samples are carnivores are known (see below). differentially preserved we employed multiple analytical methods 398 H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404 that best matched the condition of each assemblage. For the Olduvai modern control samples, in which they are relatively common. samples, all excavated dental specimens that could be identified to Second, hypotheses of hominin hunting discern between “gathering” taxon (species or otherwise unrepresented bovid tribe or size group) of weak, slow and naïve baby animals and dispatching older animals were used to determine MNI and age, including partial and complete that possess adult or near-adult strength, speed and savvy. In hemimandibles (most MNIs were based on these), maxillary denti- contrast to young juveniles, subadult juveniles meet this latter tions, and isolated teeth. Tooth eruption sequence, homologous tooth description in physical and behavioral capabilities and we thus size, and the progressive loss of molar infundibula with advancing age restrict analysis of juveniles to this substage, defined dentally by (generally from mesial to distal), were all used in these determina- moderately to heavily worn or shed deciduous premolars and tions. In contrast, the bovid samples analyzed from Kromdraai A and erupting permanent premolars and third molars. Gondolin were restricted to non-antimeric hemimandibles. For each We use three categories, early prime, late prime and old, to classify hemimandible, individual teeth were first assigned an eruption/ adult animals. Early prime, is from ~20%–50% of PEL (in some studies, attritional score, and then the specimen as a whole was assigned to an this extends to ~60% of PEL); an early prime animal has full permanent age class (see Bunn and Pickering, 2010). The Swartkrans samples dentition showing light to moderate and substantial occlusal wear but were not available for study so we used Watson's (1993a) published data on bovid mortality for the site. Even with the inclusion of fragmentary specimens from Olduvai, sample sizes are small, ranging from MNIs of 1–9 per bovid species, whereas the South African samples are larger, ranging from MNIs of 10–27 per bovid species. To achieve usable sample sizes, some analytical pooling of species MNIs into size groups based on estimated live animal weights was employed.

Modern African bovid samples

A long history of research and an extensive literature on modern African wildlife provide essential empirical data used in the comparative analyses herein. Long-term, field-based research in some notable ecosystems (e.g., the Serengeti) documents predator– prey dynamics and provides essential, large datasets on the age structure of various bovid populations and on the mortality structure of bovid prey samples from the major carnivore predators involved. Laboratory research on the degree of correlation between ontogenetic age and dental attrition provides data for a representative range of taxa. Unsurprisingly, these many independent projects have devel- oped a range of descriptive systems for defining age categories suited to their particular objectives. Some of this research employs cementum increment analysis in combination with loss of molar infundibula, yielding high-resolution age classes of 1–3 yr. Other studies use dental attrition in incisors, horn development, and changes in body form, which are convenient methods for aging of animals in the field but less applicable in paleontological analyses, based, of necessity, on attrition in the commonest molar teeth. As in Stiner's (1990) earlier research on this subject, we made a concerted effort to match the various degrees of temporal resolution and definitions of mortality class boundaries, to ensure that our age classes group fossil and modern animals of the same age range and that those classes are relevant to the hypotheses of hominin meat foraging that are being tested. Our accompanying methodological paper details the theoretical background and development of our categorization system (Bunn and Pickering, 2010), which we summarize here.

Analytical age and mortality classes for ungulates

Ungulates enter adulthood when they lose all deciduous teeth and begin to show occlusal wear of all permanent teeth. Prior to adulthood, an individual is considered a juvenile, a major stage of life history that is ~20% of its total lifespan or PEL. We divide the juvenile stage into young juvenile (newborn through yearling) and fi subadult juvenile substages. Bovids classi ed as young juveniles Figure 1. Fossil bovid dentitions from FLK Zinj illustrating occlusal wear stages for three possess light to moderately worn deciduous premolars and erupting adult age categories. Specimens D 41 and D 122 are the associated hemimandibles of a

M1 and sometimes M2. Young juveniles are excluded from our prime adult Kobus sigmoidalis; note the considerable wear on the occlusal surfaces of the analyses for two reasons. First, young juvenile teeth are usually teeth, but that all molar infundibula are still intact (a). Specimens D 42 and B 347 are the associated hemimandibles of an old adult (just beyond the late prime-old age destroyed when subjected to rigorous biostratinomic and diagenetic threshold) Parmularius altidens, which show the recent loss of the mesial infundibula forces (e.g., Munson, 2000; Munson and Garniewicz, 2003)andare from the M1s (b). Specimens G 294 and D 35 are the associated hemimandibles of an old thus relatively uncommon in paleontological samples compared to adult male Antidorcas recki, which show loss of all molar infundibula (c). H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404 399 no loss of molar infundibula (this categorization applies to size group Results 3 bovids, not to smaller taxa, which start losing lower molar infundibula earlier in life). Late prime is from ~50%–75% of PEL; a Table 1 shows the mortality data and the broad taxonomic late prime animal has moderate and substantial occlusal wear but no diversity among the fossil bovids from FLK Zinj. To enable compar- loss of mesial infundibulum from M1; (3) old is from ~75%–100% of ative analysis, data for the other Pleistocene assemblages from PEL; an old animal has heavy occlusal wear and progressive loss of Olduvai and from South Africa, and those for some of the best- both mesial and distal infundibula from the lower molars M1 and M2 documented case studies of modern predator–prey dynamics in the (this categorization applies more widely across bovid size groups). Serengeti ecosystem are also included in Table 1. The obvious We argue that although some late prime and old animals have passed disparity in sample sizes between the generally smaller fossil samples through a series of discrete life-history events that define them as and the larger wildlife samples is controlled statistically with the physiologically past prime, they still retain formidable capabilities to modified triangular graph program of Steele and Weaver (2002), evade and defend against predation. This position, developed fully in using the mortality data from the different contexts (Table 2). our accompanying paper (Bunn and Pickering, 2010), impacts our The introduction to this paper presented four predictions for bovid interpretation of the fossil bovid mortality patterns discussed below. mortality patterns should the ER-PH hypothesis or passive scavenging Figure 1 illustrates fossil bovid specimens from FLK Zinj representing hypothesis be true. Our results provide the tests of those predictions. the different adult age classes. First we consider test implications for large, size group 3 bovids, under each hypothesis. To restate:

Presentation of data Test prediction 1. If early Homo used ER-PH to obtain the size 3 bovids at FLK Zinj, then the mortality data should match what For visual comparison of different mortality samples we followed cursorial predators of large bovids are known to kill (which, methodology developed by Stiner (1990), who divided data into preferentially, is the relatively vulnerable young and old, more than juvenile, prime,orold groups, then converted those data to prime adults). percentages for each sample, and plotted each sample as a discrete point on a triangular graph. To control for differences in sample sizes Test prediction 2. If early Homo passively scavenged from large felids we used computer software (modified triangular graph program) to obtain the size 3 bovids at FLK Zinj, then the FLK Zinj mortality data created by Steele and Weaver (2002) that employs bootstrapping should match what lions (as the best available modern proxy for large statistics to produce a density contour around each graphed sample felid predators of size 3 bovids) are known to kill (i.e., closer to an point, approximating a 95% confidence interval. unselective, living-structure prey sample, with a relatively higher

Table 1 Age frequency distributions in African fossil and modern samples of bovid dental remains.a,b

Site/predatorc Size groupd Taxon MNI Age

Young juvenile Subadult juvenile Early prime Late prime Old

Olduvai FLK Zinj 1 Antidorcas recki 6114 2 Antilopini 1 1 1 and 2 Total 7 1 1 5 3a Parmularius altidens 6321 3b Connochaetes sp. 3 3 3b Kobus sigmoidalis 92 3 4 3b Oryx sp. 1 1 3 Total 19 2 3 8 4 2 4 Syncerus acoelotus 11 Olduvai FLK N 1–2 1 and 2 Total 14 3 10 1 3 Total 13 2 3 7 1 Olduvai FLK N 6 1 and 2 Total 6 1 4 1 3 Total 12 1 3 5 3 Olduvai FLK NN 2 3 Total 9 2 2 4 1 Swartkrans 1 1 Antidorcas sp. 11 5 1 3 1 1 Swartkrans 2 1 Antidorcas sp. 10 3 1 4 1 1 Swartkrans 3 1 Antidorcas sp. 18 4 2 7 2 3 Kromdraai A 1 Antidorcas sp. 10 4 5 1 2 Alcelaphini 23 6 11 6 1 and 2 Total 33 10 16 6 1 Gondolin 2 Redunca sp. 27 3 3 21 Leopard 1 Gazella thomsoni 30 9 4 15 2 Lion 1 Gazella thomsoni 204 67 12 82 11 32 3b Connochaetes taurinus 262 72 50 78 62 3b Kobus ellipsiprymnus (♂)9 2 3 1 3 1 Gazella thomsoni 192 124 2 36 8 22 Spotted hyena 1 Gazella thomsoni 98 42 10 25 21 3b Connochaetes taurinus 86 31 15 15 25 Wild dog 1 Gazella thomsoni 65 34 2 19 2 8

a Abbreviations: FLK Zinj=FLK 22 Zinjanthropus; Swartkrans 1=Swartkrans Member 1; Swartkrans 2=Swartkrans Member 2; Swartkrans 3=Swartkrans Member 3; MNI=minimum number of individuals. b All fossil data except those from Swartkrans (Watson, 1993a) were generated by the authors. See also Bunn and Pickering (2010-this issue) for details of the Gondolin mortality data. Modern wildlife data are from Schaller (1972), Kruuk (1972), and Spinage (1982). c Site/predator =for fossil assemblage of inferred, but ultimately unknown taphonomic derivation, the site name is provided, in contrast to the predator species that is provided for samples that formed under direct, modern observations. d Bovid size groups follow Bunn (1986); see text for discussion and additional references. 400 H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404

Table 2 from FLK Zinj also contradict aggressive, power scavenging from lions, Age frequency distributions of bovid dental remains from modern carnivore kills and an alternative hypothesis developed, in the absence of evidence of Olduvai fossil sites grouped for modified triangular graph analyses (subadult juvenile, hunting by hominins, to account for the abundance of the very prime and old).a skeletal elements defleshed quickly by lions and the abundance of b Sample Subadult Prime Old MNI defleshing cut marks on them. The large bovid mortality data from FLK Juvenile n (%) n (%) Zinj correspond to Stiner's definition of a “prime-dominated” pattern, n(%) which she attributes exclusively to selective hunting by humans Small (size 1 and 2) (Stiner includes young juveniles, which are excluded here; including Leopard-killed Gazella thomsonic 4 (19) 15 (71) 2 (10) 21 Leopard-killed G. thomsonic 4 (5) 66 (84) 9 (11) 79 young juveniles increases the statistical separation between lion-killed Lion-killed G. thomsoni 12 (9) 93 (68) 32 (23) 137 wildebeest [Connochaetes spp.] and size-group 3 bovids at FLK Zinj). Cheetah-killed G. thomsoni 2 (3) 44 (65) 22 (32) 68 To add a comparative perspective, however, the mortality data for Spotted hyena-killed G. thomsoni 0 35 (63) 21 (38) 56 large bovids at FLK Zinj also show strong similarity to other Bed I Wild dog-killed G. thomsoni 2 (6) 21 (68) 8 (26) 31 Olduvai assemblages in which butchery evidence is uncommon or FLK Zinj bovid sizes 1 and 2 1 (14) 1 (14) 5 (71) 7 FLK N 1–2 bovid sizes 1 and 2 3 (21) 10 (71) 1 (7) 14 absent (Fig. 3), which warrants further discussion. FLK N 6 bovid sizes 1 and 2 1 (17) 4 (67) 1 (17) 6 The restated test implications for smaller, size-group 1–2 bovids Large (size 3) are: Lion-killed Connochaetes taurinus 50 (26) 78 (41) 62 (33) 190 Lion-killed Kobus ellipsiprymnus (♂) 2 (22) 4 (44) 3 (33) 9 Test prediction 3. If early Homo used ER-PH to acquire smaller, size Spotted hyena-killed C. taurinus 15 (27) 15 (27) 25 (45) 55 group 1–2 bovids at FLK Zinj, then the FLK Zinj mortality data should FLK Zinj K. sigmoidalis 0 7 (100) 7 FLK Zinj bovid size 3 3 (18) 12 (71) 2 (12) 17 match what cursorial predators are known to kill (again, the relatively FLK N 1–2 bovid size 3 3 (27) 7 (64) 1 (9) 11 vulnerable young and old). FLK N 6 bovid size 3 3 (27) 5 (45) 3 (27) 11 FLK NN 2 K. sigmoidalis 2 (33) 3 (50) 1 (17) 6 Test prediction 4. If early Homo passively scavenged from tree- Pooled Olduvai “background” 8 (28) 15 (54) 5 (18) 28 bovid size 3d stored leopard kills to obtain small bovids at FLK Zinj, then the FLK Zinj mortality data should match what leopards are known to kill (i.e., a Modern wildlife data are from Schaller (1972), Kruuk (1972) and Spinage (1982). a non-age-selective, living-structure sample). All fossil data were generated by the authors. Bovid size groups follow Bunn (1986); see text for discussion and additional references. b MNI=minimum number of individuals. As shown in Figure 4, the small-bovid data from FLK Zinj are c The first leopard-killed G. thomsoni sample reflects Schaller's (1972) observations dominated by old adult Antidorcas recki, which, based on their size and of gazelle age classes, the second sample includes Schaller's unaged adults (n=58), which we have assigned to age class based on proportions occurring in Schaller's rugosity, are male animals. The mortality distribution of small bovids smaller, aged sample. from FLK Zinj is statistically distinct from and, thus, contradicts the d The pooled Olduvai “background” sample is the combined data from FLK N 1–2, FLK test implications of passively scavenging from leopard or cheetah N 6 and FLK NN 2. (Acinonyx jubatus) kills. It is probable, at a 95% confidence level, that the small bovids whose butchered bones were found at FLK Zinj were percentage of prime adults than cursorial predators, such as spotted not acquired by passive scavenging from leopard kills. Small-bovid hyenas, take). mortality data from Swartkrans, Kromdraai A, and Gondolin, on the As revealed in Figure 2, the large-bovid data from FLK Zinj are other hand, do support the attribution of those remains to leopard (or dominated by prime adults and contradict directly the test implica- another felid ambush predator) kills, which has been argued tions of hypotheses that posit ER-PH or passive scavenging from lion previously from evidence of prey body size, skeletal part represen- kills. Both the ER-PH hypothesis and the passive scavenging tation and bone damage (e.g., Brain, 1981, 1993; Pickering et al., hypothesis are, thus, falsified by the FLK Zinj mortality data for large 2004b, 2007, 2008)(Fig. 5). Similarly, small-bovid mortality data from bovids, which were probably (at a 95% confidence level) derived from FLK N 1–2 and from FLK N 6 also exhibit a living-structure pattern that a different source than that predicted by ER-PH hunting or by passive is statistically indistinguishable from leopard prey, which contrasts scavenging from lions. Additionally, the large bovid mortality data

Figure 3. Modified triangular graph comparing mortality data for large (size group 3) Figure 2. Modified triangular graph comparing modern Serengeti lion-killed wildebeest bovids (excluding young juveniles; see text for definition) from FLK Zinj and from the and FLK Zinj large (size group 3) bovid mortality data. Data for lion-killed wildebeest “background” sample at Olduvai, pooled from the FLK N 1–2, FLK NN 2, and FLK N 6 from Schaller, 1972. assemblages. H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404 401

Figure 4. Modified triangular graph comparing modern Serengeti leopard-killed Figure 6. Modified triangular graph comparing mortality data for modern Serengeti Thomson's gazelles and FLK Zinj smaller (size groups 1–2, Antidorcas recki and leopard-killed Thomson's gazelles, for smaller (size groups 1–2) bovids from FLK Zinj Antilopini size group 2) bovid mortality data. Data for leopard-killed gazelle from and for the “background” sample at Olduvai, pooled from the FLK N 1–2 and FLK N 6 Schaller, 1972. assemblages. Data for leopard-killed gazelles from Schaller, 1972.

with earlier reconstructions of FLK N 1–2 as an occupation floor Discussion produced by hominins but is consistent with recent attribution of small bovids there as leopard prey based on patterns of skeletal Given the abundance of large-bovid long-limb bone specimens representation and bone damage (Domínguez-Rodrigo et al., 2007a,b; with defleshing cut marks and the quantity of the very skeletal Domínguez-Rodrigo and Pickering, 2010; contra Blumenschine et al., elements eaten first by carnivores at their kills, which implies early 2007)(Fig. 6). Bovid remains at FLK N 6 have always been considered access by hominins to intact carcasses (see detailed discussions in a natural background, rather than food refuse of hominins; the Domínguez-Rodrigo, 2002; Domínguez-Rodrigo and Pickering, 2003; mortality data on small bovids presented here are consistent with Pickering and Domínguez-Rodrigo, 2006), the simple conclusion from evidence of skeletal representation and bone damage, as reported by the statistically distinct, prime-dominated mortality profile for FLK Domínguez-Rodrigo et al. (2007a), in specifying leopard predation as Zinj is, or could be, that early Homo, as ambush predators, killed the a likely dominant factor. large bovids. The mortality distribution of large bovids from FLK Zinj, In contrast, the old-male-dominated pattern of A. recki mortality in however, also shows strong similarity and no statistical separation the FLK Zinj assemblage is not inconsistent with the ER-PH from the large bovids from FLK N 1–2, FLK NN 2, and FLK N 6. These hypothesis. This is the single prediction tested in this study that is latter bovid assemblages exhibit little (FLK N 1–2, FLK NN 2) or met. In that context, we still seriously question the plausibility of ER- possibly no (FLK N 6) butchery evidence, and based on bone damage PH as a regular carcass acquisition strategy for early hominins, a point patterns, they have recently been attributed mainly to predation and we expand upon below. feeding by large felids (Domínguez-Rodrigo et al., 2007a). It is possible that ambush predation by large felids was dominant in all three of these “background” Olduvai assemblages (and, unsurpris- ingly, in the Olduvai paleo-community in general), with the wealth of butchery evidence at FLK Zinj reflecting both hunting success by early Homo and access to complete and fairly complete carcasses by power scavenging from felid predators. In any case, the prime-dominated mortality profile for large bovids from FLK Zinj falsifies ER-PH as a credible explanation for the zooarchaeological and taphonomic patterns observed in the assemblage. Those patterns also argue strongly against passive scavenging by hominins as a significant factor in creating the FLK Zinj large bovid subassemblage. Using mortality data to evaluate this assertion is complicated by the fact that the comparative fossil samples from Olduvai “background” sites (FLK N 1–2, FLK NN 2, FLK N 6) are small and thus the 95% confidence contours for those probable felid-derived assemblages are large. If instead the “background” samples are pooled it emphasizes their more even, living-structure age distribution, in contrast to the prime-dominated pattern at FLK Zinj. The prime- dominated pattern at FLK Zinj also trends away from the living- structure patterns evident in modern lion kills. Yet another way to increase sample sizes analytically is to include Figure 5. Modified triangular graph comparing modern Serengeti leopard-killed young juveniles in the comparison of FLK Zinj and the pooled Thomson's gazelle and smaller fossil bovid (size groups 1–2, Antidorcas, Redunca, and “background” sites, and our subdivision of mortality data into Alcelaphini size group 2) mortality data pooled from Swartkrans, Kromdraai A and Gondolin. Data on leopard-killed gazelle from Schaller, 1972. Swartkrans data from multiple age classes is designed to enable and encourage such Watson, 1993. flexibility in analytical approach. The presence of relatively fragile, 402 H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404 young juvenile specimens in all of these assemblages argues against ambush predator, alert to all animal movements in these relatively severe, density-mediated biasing from biostratinomic (carnivore vegetated settings. scavenging) or diagenetic (profile compaction) processes. At FLK At the risk of being branded ethnographic tyrants again (and Zinj, for example, in addition to very young juvenile (or fetal) bovids, falsely), we recall our empirical observations that small bovids can be there are also teeth of a young juvenile suid and hemimandibles of a walked to exhaustion—not requiring ER—by modern, highly skilled young juvenile giraffe, and in each of the background assemblages, Kua hunters at a significant physical cost to them, in the sparsely multiple young juvenile suid individuals are also represented. Figure 7, vegetated, soft sandy substrate of the Kalahari that is the ideal context which includes young juveniles and subadult juveniles in the juvenile for using sophisticated tracking skills. In the more heavily vegetated class, shifts the distributions toward the juvenile side and shows more savanna–bush–woodlands that characterize the East African Rift clearly the living-structure pattern of the “background sites.” If the Valley today, and paleoenvironmental reconstructions of the habitats young juveniles bovids at FLK Zinj are actually fetal (these deciduous in which early Homo evolved in the Pleistocene, even greater tracking upper premolars are essentially unworn, and Spinage (1967) reports skills beyond those possessed by modern foragers would be required to this condition at birth but not at six months of age in modern succeed at ER hunting. Despite long-term ethnographic research, the waterbuck [Kobus ellipsiprymnus]) and excluded (as in Fig. 3) because Hadza have not been observed either attempting, much less they were not directly hunted and killed, then there is an even clearer succeeding at ER hunting in the more vegetated East African Rift separation between the prime-dominated mortality at FLK Zinj and Valley. This demonstrates a likely cause–effect relationship between the living-structure mortality at the “background” sites. This likely vegetation cover and viability of tracking, and it casts doubt on the indicates a derivation from a different source: ambush hunting by likelihood of highly sophisticated, successful tracking by early Homo. early Homo at FLK Zinj, rather than any form of scavenging from felids This, of course, does not mean that ER-PH never happened in the or ER-PH. Since the initial writing of this paper, two additional Pleistocene. It simply illustrates that a different ethnographic example maxillary and mandibular specimens of waterbuck have been (i.e., hunting with bow-and-arrow) emphasized by Lieberman et al. identified by HTB in the original fossil assemblage from FLK Zinj. (2007: 442), when viewed in broader and better-matched ethno- These specimens (catalog numbers B 4 and C [no number, so it was graphic and ecological perspective, does not provide quite the likely recovered from the screen]) represent two additional indivi- epiphany about the evolution of ER-PH hunting that is desired: “In duals that died younger than the two fetal or neonatal individuals fact, without projectiles, it is hard to imagine how early Homo in the previously known (as discussed above) and are clearly fetal (unworn [Early Stone Age] would have either scavenged or hunted safely or upper and lower deciduous premolars). This raises an intriguing effectively unless they employed ER to some extent.” Ironically, possibility in relation to the small prime adult waterbuck individuals Lieberman et al. base that claim on their assertion that because of (MNI=7 adults, of which three are smaller, early prime females and technology, hunting strategies of recent foragers are not useful four are larger, late prime males) from FLK Zinj: the targeted prey of analogs for early Homo, which they then justify using survey results hominin ambush hunting was likely pregnant or recently pregnant of ethnographic hunting technology (e.g., spear efficiency). Lieberman female and somewhat older, perhaps non-territorial male waterbucks. et al. confuse and conflate the concepts of hunting efficiency and Even though ER-PH was argued to be most productive for hunting capability. In all likelihood, hunting efficiency is significantly hominins when used against large, size group 3 prey, we reiterate increased in recent contexts with bows and arrows, relative to the that the old-male-dominated pattern in A. recki mortality at FLK Zinj Pleistocene, but it does not follow from relative efficiency, that does not contradict the prediction of the ER-PH hypothesis for small hunting capability in the absence of bows and arrows was lacking bovids. We question, however, the relative merits and adaptive value (except by ER). of targeting small, old gazelles in their preferred bush–woodland Logically extending such hyperbole to include actual scientific habitat by jogging with eyes fixed to the ground in a most challenging evidence, would mean that in the vast array of paleoecological contexts tracking exercise right through the prime hunting habitat of large in which hunting is reconstructed—from Gona to the most recent hunter felids, when the same bovids could have been exploited by Homo as missing his bow and arrows, from flat, sandy terrain to steep, wooded, rocky slopes, and from tropical and temperate regions—premodern or past human hunting (and scavenging) without bows and arrows reduces to ER (or to hopeless inefficiency?). To learn about past human foraging strategies, including all forms of hunting and scavenging, it is not justifiable scientifically to assume and attribute to early Homo,orto their immediate ancestors, the sophisticated tracking skills of modern humans or skills beyond those of modern humans—essential to the success of ER-PH—when the evolution of hunting capabilities is the very topic being investigated. Rudimentary tracking skills would not enable successful, much less, efficient ER, in the vegetated East African Rift Valley paleoenvironments in which early Homo evolved. And it is not reasonable to assume sophisticated tracking skills in early Homo, while at the same time discounting any intellectual ability to ambush prey. In the absence of direct evidence, we do not know how early Homo would have hunted large prey (sensu Bunn and Kroll (1986), who for that reason raised the alternative of aggressive, power scavenging; contra Lieberman et al.'s characterization), but there is compelling evidence that Pleistocene Homo had early access to intact, prime adult, large bovids. A plausible method of ambush hunting would require that hominins had the ability to get close enough to prey animals to inflict Figure 7. Modified triangular graph comparing mortality data for large (size group 3) a mortal wound and that they had the technology to do so. One likely bovids (including young juveniles; see text for definition) from FLK Zinj and from the “background” sample at Olduvai, pooled from the FLK N 1–2, FLK NN 2, and FLK N 6 method of achieving exceedingly close encounters with prey animals assemblages. that is known to be effective in modern contexts is climbing into a tree H.T. Bunn, T.R. Pickering / Quaternary Research 74 (2010) 395–404 403 along a game trail and simply waiting for animals to walk by with hominin carcass foraging by addressing the following substantive their gaze fixed horizontally to guard against terrestrial predators. As points: for lethal weaponry, wooden spears are a distinct possibility, although we emphasize that there is currently no direct evidence of such 1. The falsification of the ER hunting hypotheses by bovid–mortality technology. We pose three questions: 1. Was there a wood-chopping evidence; function for Oldowan choppers whose edges are battered through use 2. The probable implications for Pleistocene Homo of a likely cause– as tools (e.g., Hayden 2008), comparable to the woodworking effect relationship between vegetational habitats and success rate activities reconstructed by Domínguez-Rodrigo et al. (2001) from of human tracking; wood phytoliths on Acheulean tools at Peninj (Tanzania)? 2. What 3. The logic of assuming sophisticated tracking skills in early Homo was the hominin task that produced microwear evidence of wood and their immediate ancestors, while discounting any intellectual sawing or scraping on Oldowan stone flakes at Koobi Fora (Kenya) ability to ambush prey. (Keeley and Toth 1981)? 3. If a chimpanzee level of cognitive ability can yield a sharpened branch stabbing weapon for killing small George Schaller (1972: 195) stated that “Predation is exceedingly mammal prey (Pruetz and Bertolani, 2007), then how capable would complex.” So is the reconstruction of ancient predation from fossil more encephalized early Homo have been at fashioning wooden teeth. Paleoanthropologists are well-advised to keep both declara- weapons and tools? A fourth question is directed to any who would tions of counsel in mind when constructing models of early hominin discount the very idea of large mammal hunting with wooden spears carcass foraging. by Pleistocene Homo: How can science ever determine when this behavior actually began, if we allow our investigations to be guided by Acknowledgments ideology rather than evidence and if we do not ask such provocative questions and then explore ways of answering them? HTB's research was supported by the National Science Foundation Moving beyond hunting, in creating a diurnal scavenging niche for (USA), the L.S.B. Leakey Foundation, and the University of Wisconsin- early Homo made possible by ER, Bramble and Lieberman (2004) Madison. HTB thanks the Tanzania Commission for Science and reason that ER would have aided early Homo in competitive Technology (COSTECH) for permission to study the Olduvai collec- scavenging against hyenas, wild dogs (Lycaon pictus), and other tions from Mary Leakey's research and to conduct research with hominins. Yet, Lieberman et al. (2007: 441) seek to enhance their ER Hadzabe foragers, and the National Museums of Kenya for access to scavenging niche by misrepresenting the timing of scavenging by Olduvai collections. TRP thanks his family for their support and various carnivores, including spotted hyenas, as an exclusively Francis Thackeray, Stephany Potze, and the Transvaal Museum for nighttime activity, or at least discounting their ability to run and permission to study the Gondolin and Kromdraai faunas. TRP's scavenge effectively during the day, stating that “With ER capabilities, research was supported by the National Science Foundation (USA), hominins may have had a previously unrecognized advantage the LSB Leakey Foundation (USA), the Palaeontology Scientific Trust scavenging in open habitats during the day when other scavengers (South Africa) and the University of Wisconsin-Madison (through a are prevented from running long distances because of thermoregu- Vilas Fellowship). We thank Manuel Domínguez-Rodrigo, José latory constraints (hyenas confine their running to dawn, dusk and Yravedra, Gail Ashley, and Kathryn Remer for suggestions that night). Scavenged meat is always an ephemeral resource, requiring improved this paper. We also thank Teresa Steele for providing the speed.” Ironically, what started as endurance running now requires Modified Triangular Graph software used in the analysis. Special speed running, and hunting/scavenging toward flying vultures by thanks to Gary Haynes and an anonymous reviewer for their insightful cursorial carnivores is now prevented during the day, according to comments. this modified view—a view that comprehends the natural world not as one of complexities, but as one of simple absolutes. In this case, References diurnal scavenging becomes a tortoise-and-hare race to the carcass, with hominins and the various carnivores in equal numbers and Adams, J.W., 2006. Taphonomy and paleoecology of the Gondolin Pleistocene cave site, South Africa. Ph.D. dissertation. Washington University. distribution on the landscape, but with only hominins able to persist Binford, L.R., 1981. Bones: Ancient Men and Modern Myths. Academic Press, New York. via diurnal running to the finish line. Binford, L.R., 1985. Human ancestors: changing views of their behavior. 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