Disentangling Early Stone Age palimpsests: determining the functional independence of hominid- and carnivore-derived portions of archaeofaunas

Determining the extent to which hominid- and carnivore-derived components of fossil bone palimpsests formed independently of each other can provide valuable information to paleoanthropologists interested in reconstructing the foraging adaptations of hominids. Because stone tool cutmarks, hammerstone percussion marks, and carnivore tooth marks are usually only imparted on bone during nutrient extraction from a carcass, these bone surface modifications are particularly amenable to the types of analyses that might meet this goal. This study compares the percentage of limb bone specimens that preserve evidence of both hominid- and carnivore-imparted bone damage from actualistic control samples and several Plio-Pleistocene archaeofaunas, including new data from Swartkrans Member 3 (South Africa). We argue that this procedure, which elucidates the degree of hominid-carnivore independence in assemblage formation, will allow researchers to extract for focused analyses high integrity components (hominid and carnivore) from presumably low integrity sites. Comparisons suggest that the hominid- and carnivore-derived components from sites in Olduvai Gorge Bed II (Tanzania), the ST Site Complex at Peninj (Tanzania), and Swartkrans Member 3 formed largely independent of each other, while data from the FLK 22 Zinjanthropus (FLK Zinj) site (Olduvai Gorge Bed I) indicate significant interdependence in assemblage formation. This contrast suggests that some Early Stone Age assemblages (e.g., the Olduvai Gorge Bed II sites, the Peninj ST Site Complex, and Swartkrans Member 3) are probably more useful than others (e.g., FLK Zinj) for assessing the maximal carcass-acquiring abilities of early hominids; in such assemblages as those in the former set, sole hominid-contribution is more confidently discerned and isolated for analysis than in assemblages such as FLK Zinj.     

Meat-eating by early hominids at the FLK 22Zinjanthropussite, Olduvai Gorge (Tanzania): an experimental approach using cut-mark data

The meat-eating behavior of Plio-Pleistocene hominids, responsible for the bone accumulations at the earliest archaeological sites, is still a hotly-debated issue in paleoanthropology. In particular, meat-eating and bone marrow consumption are often presented as either complementary or opposing strategies of carcass exploitation. The presence of cut marks on fossil archeofauna is a potential source of information that has not been consistently used as evidence of carcass consumption by hominids. Some authors interpret cut marks as the result of hominids manipulating meat-bearing bones, while others argue that they can also be the result of hominids extracting marginal scraps of carcass flesh that have survived carnivores’ initial consumption. In this study, a referential framework concerning the interpretation of cut marks is presented, based on a set of experiments conducted by the author. It is suggested, according to these experiments and data drawn from the FLK “Zinj” site, that hominids processed meat-bearing bones (on which flesh was abundant) rather than defleshed carcasses from felid kills.     

Validation of bone surface modification models for inferring fossil hominin and carnivore feeding interactions, with reapplication to FLK 22, Olduvai Gorge, Tanzania

Resolving the issue of how Early Stone Age hominins acquired large mammal carcasses requires information on their feeding interactions with large carnivores. This ecological information and its behavioral and evolutionary implications are revealed most directly from the tooth, cut, and percussion marks on bone surfaces generated by hominin and carnivore feeding activities. This paper employs a bootstrap method, a form of random resampling with replacement, to refine published neotaphonomic models that use the assemblage-wide proportions of long bones bearing feeding traces to infer the sequences in which Plio-Pleistocene hominins and carnivores accessed flesh, marrow, and/or grease from carcasses. Results validate the sensitivity of the models for inferring hominin feeding ecology, which have been questioned on grounds shown here to be unfounded. The bootstrapped feeding trace models are applied to the late Pliocene larger mammal fossil assemblage from FLK 22 (Zinjanthropus site), Olduvai Gorge, Tanzania. High frequencies of tooth and percussion marking on long bone midshaft fragments from FLK 22 are most consistent with those feeding trace models that simulate hominin scavenging from carcasses defleshed by carnivores, while cut mark data indicate that hominins more often had access to upper forelimb flesh than upper hind limb flesh. Together, the bone surface modification data indicate that hominins typically gained secondary access to partially defleshed carnivore kills, but they also allow for the possibility of some carcasses being processed only by carnivores and only by hominins.     

Taphonomic perspectives on hominid site use and foraging strategies during Bed II times at Olduvai Gorge, Tanzania

The faunal assemblages excavated by Mary Leakey in Bed II of Olduvai Gorge, Tanzania, have, like the more well-known Bed I assemblages, traditionally been interpreted as the result of hominid butchering activities in the lake margin and riverine settings of the paleo-Olduvai Basin. A reexamination of all of Leakey's Bed I sites has shown that hominids played little or no role in the formation of all but one of those faunal assemblages, a finding that prompted the reanalysis of the Bed II sites presented here. We expand upon a previous taphonomic study that provided systematic data for HWK East Levels 1–2, MNK Main, and BK. In addition to these assemblages, we provide data on HWK East Levels 3–5, FC West, TK, and SHK. Our data contradict previous interpretations of MNK Main as a hominid accumulation but uphold the contention that BK represents a primarily hominid accumulation reflecting early access to carcasses. The small and poorly preserved assemblages from FC West and TK are difficult to link unambiguously to either hominids or carnivores. Site MNK Main and HWK East Levels 3–5 appear to be death arenas where carcasses accumulated via natural deaths and/or serial predation. Site SHK is severely biased by selective retention and therefore little can be said of its formational history. Nevertheless, no hominid modifications were documented in this assemblage. Comparisons with other Olduvai sites indicate a more conspicuous hyena taphonomic signal during Bed II times than Bed I times, which appears to mirror the changing configuration of the large carnivore guild. These findings also beg the question of what activities were being carried out by hominids with the stone tools discarded at these sites. Although it seems clear that hominids were utilizing stone tools to carry out subsistence activities unrelated to carcass butchery, more excavation and techniques such as phytolith analysis should be employed to explore alternative explanations.     

Testing meat-eating in early hominids: an analysis of butchery marks on defleshed carcases

The diet of early hominids responsible for the bone accumulations at Plio-Pleistocene sites is still a controversial issue. Meateating and bone marrow consumption are often presented either as complementary or as opposing strategies of carcass exploitation. The occurrence of cut marks on fossil bones at early sites is a potential source of information that has not been consistently used as evidence of what products hominids obtained from carcasses. Some authors interpret them as the result of manipulating meat-bearing bones, whereas others believe that they can also be the result of extracting marginal scraps of flesh that have survived carnivores’ initial consumption of carcasses. In this study, a referential framework concerning the latter process is presented and it is concluded, according to the data drawn from the FLK “Zinj” site and the results obtained in the experiment, that hominids processed meat-bearing bones (in which flesh was abundant) and not defleshed carcasses from felid kills. This work constitutes a reference that can also be used for later Pleistocene sites and adds a further dimension to the hunting-versus-scavenging debate.     

Hominid carnivory and foraging strategies, and the socio-economic function of early archaeological sites.

New evidence for the tissue types exploited by early hominids from carcasses possibly acquired through scavenging is derived from the larger mammal bone assemblages from FLK I, level 22 (Zinjanthropus floor), and FLKN levels 1 and 2 from Bed I, Olduvai Gorge, Tanzania. Published skeletal part profiles from the two archaeological sites are evaluated using (i) modern observations on the sequence by which carnivores consume carcass parts in order to assess the timing of hominid access to carcasses, and (ii) measurements of flesh and marrow yields to assess the tissue types sought and acquired. These results suggest that the maximization of marrow (fat) yields, not flesh (protein) yields, was the criterion shaping decisions about carcass processing. Because of evidence for density-dependent destruction of some flesh-bearing parts by scavengers of the hominid-butchered assemblages, however, it is uncertain whether carcass parts were transported and acquired by hominids in a largely defleshed condition. The results on tissue types acquired are combined with a discussion of predation risk, feeding competition and the equipment needs of carcass processing in an attempt to identify archaeological test implications of competing hypotheses for the socio-economic function of the earliest archaeological sites.     

Crocodylian and mammalian carnivore feeding traces on hominid fossils from FLK 22 and FLK NN 3, Plio-Pleistocene, Olduvai Gorge, Tanzania

Taphonomic analysis of the Olduvai Hominid (OH) 8 left foot from FLK NN Level 3 and the OH 35 left leg from FLK Level 22 (Zinjanthropus level) in Middle Bed I, Olduvai Gorge, indicates that both were fed upon by crocodiles. Both bear extensive tooth marking, including bisected tooth marks diagnostic of crocodylian feeding. The location of the bisected tooth marks on the distal tibia and the talus indicates disarticulation of the foot by crocodiles. The broken proximal ends of the tibia and fibula are more typical of feeding by a leopard-like carnivore, as is damage to the OH 7 mandible and parietals that are associated with and may derive from the same individual as OH 8. Previous work showing a close articulation of the foot and the leg has been used to suggest that the two specimens belong to the same individual despite deriving from sites separated by 200 m and slightly different stratigraphic levels according to previous work. The location and agent of tooth marking and the nature of gross damage do not refute this hypothesis, but the punctures on the talus and distal tibia differ in size and sharpness. Recent work shows that the stratigraphic discrepancy between OH 8 and OH 35 is greater than previously thought, refuting the single-individual hypothesis. Although seemingly unlikely, this denotes that two hominids represented by rarely found leg and foot elements both lost their left foot to crocodiles at nearby sites within a 6,000 year interval. We cannot determine if the hominids were preyed upon by crocodiles or mammalian carnivores. However, the carnivore damage to them and associated faunal remains suggests that high predation risk constrained hominid activities involving discard of the stone artifacts found at these sites. This finding is inconsistent with the interpretation of the sites as home bases or living floors.     

Archaeological predictions for hominid land use in the paleo-Olduvai Basin, Tanzania, during lowermost Bed II times

We present a preliminary predictive model of Oldowan stone artefact and scavenged larger mammal bone assemblages for 11 landscape facets modeled earlier to occur across a large portion (>300 km²) of the paleo-Olduvai Basin during lowermost Bed II times. This second phase of model-building is based on our earlier characterizations of the basin's landscape ecostructure and the inter-facet distribution of key resources and hazards probably encountered by Late Pliocene hominids (Peters & Blumenschine, 1995, 1996). Our current extension of the model of hominid–landscape interactions specifies additional theoretical components, including: (1) the assumed capabilities of Oldowan hominids (presumablyHomo habilis, primarily); (2) the landscape-facet-specific tasks they carried out; (3) the immediate stone and bone task residues they produced; and (4) the predicted composition, condition, density, and clustering of stone artefact and butchered and unbutchered bone assemblages for each facet. We develop ecological linkages between these new and formerly reported modeling components, the most fundamental of which is the facet-specific degree of tree/shrub cover abundance, and the correlated degree of competition among larger carnivores and hominids for scavengeable larger mammal carcasses. These factors condition variability among landscape facets in scavenging opportunities encountered by hominids, which in our model is the major predictor of bone and stone artefact assemblage composition. The predictive value of scavenging reflects the bias of paleoanthropological traces toward technology and butchery in their landscape context, but the model is surprisingly insensitive to what are usually thought to be critical social components of hominid land use. The predictions for the traces of hominid–landscape interactions modeled herein can be tested in the future against the landscape archaeological sample being excavated from lowermost Bed II by the Olduvai Landscape Paleoanthropology Project.     

河南灵井许昌人遗址动物骨骼表面人工改造痕迹

河南灵井许昌人遗址发掘出土了距今约10—8万年前的古人类头骨化石, 与头骨化石同层发现的还有大量哺乳动物化石及石制品等文化遗物。本文是对该遗址2005—2006年出土动物化石骨骼表面人工改造痕迹的观察分析结果。灵井遗址中13%的动物骨骼表面有人工切割痕的产生, 其中切割痕位于长骨骨干部位的约占此类标本总数的98.45%; 同时,在具切割痕的长骨类化石材料中,属于食草动物上部和中部肢骨的分别为34%和41%, 属于下部肢骨的则仅为25%。此外, 灵井动物群中具敲击痕、火烧痕迹、人工使用痕迹的骨骼标本分别占全部观察例数的4.2%、1%、1.32%。总之, 通过对动物骨骼表面保留的上述人工改造痕迹的观察与统计分析, 并与埋藏学实验及其他考古遗址相关属性的对比, 表明古人类是这一遗址中大量脊椎动物肉食资源的初级获取者和利用者, 他们是导致这一动物群聚集形成的主要埋藏学因素。同时, 许昌人遗址中大量破碎动物骨骼的出现可能也与古人类敲骨取髓的取食行为有着非常紧密的联系。     

Cutmarked bones from Pliocene archaeological sites at Gona, Afar, Ethiopia: implications for the function of the world's oldest stone tools

Newly recorded archaeological sites at Gona (Afar, Ethiopia) preserve both stone tools and faunal remains. These sites have also yielded the largest sample of cutmarked bones known from the time interval 2.58-2.1 million years ago (Ma). Most of the cutmarks on the Gona fauna possess obvious macroscopic (e.g., deep V-shaped cross-sections) and microscopic (e.g., internal microstriations, Herzian cones, shoulder effects) features that allow us to identify them confidently as instances of stone tool-imparted damage caused by hominid butchery. In addition, preliminary observations of the anatomical placement of cutmarks on several of the recovered bone specimens suggest that Gona hominids may have eviscerated carcasses and defleshed the fully muscled upper and intermediate limb bones of ungulates--activities that further suggest that Late Pliocene hominids may have gained early access to large mammal carcasses. These observations support the hypothesis that the earliest stone artifacts functioned primarily as butchery tools and also imply that hunting and/or aggressive scavenging of large ungulate carcasses may have been part of the behavioral repertoire of hominids by c. 2.5 Ma, although a larger sample of cutmarked bone specimens is necessary to support the latter inference.     

Taphonomic analysis of the early Pleistocene (2.4 Ma) faunal assemblage from A.L. 894 (Hadar, Ethiopia)

The A.L. 894 site (Hadar, Ethiopia) is, together with OGS 7 (Gona, Ethiopia), one of the oldest archaeological sites documenting a spatial association of stone tools and bones retrieved from an in situ excavation. In contrast with OGS 7, the better preservation of the bone assemblage at A.L. 894 allows the identification of taphonomic processes of bone breakage, thanks to abundant green bone fractures. The presence of tooth marks and the lack of hominin-produced bone modifications together argue against hominins as the responsible agents for bone accumulation and modification. This taphonomic study of A.L. 894 shows lack of evidence for functional associations between stone tools and bones, a pattern documented in several other early Pleistocene sites. Such a pattern underscores the complex phenomena involved in site formation processes, especially in the earliest archaeological assemblages     

A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania

Neotaphonomic studies have determined the patterns of bone damage created by larger mammalian carnivores when consuming mammalian carcasses. Typically, mammalian carnivores gnaw and break bones to various degrees in order to access marrow, grease, and brain tissue. In contrast, crocodiles attempt to swallow whole parts of mammal carcasses, inflicting in the process tooth marks and other feeding traces on some of the bones they are unable to ingest. Although crocodiles are major predators of larger mammals along the margins of protected tropical rivers and lakes, their feeding traces on bone have received little systematic attention in neotaphonomic research. We present diagnostic characteristics of Crocodylus niloticus damage to uningested mammal bones resulting from a series of controlled observations of captive crocodile feeding. The resulting bone assemblages are composed of primarily complete elements from articulating units, some of which bear an extremely high density of shallow to deep, transversely to obliquely oriented tooth scores over often large areas of the bone, along with shallow to deep pits and punctures. Some of the tooth marks (bisected pits and punctures, hook scores) have a distinctive morphology we have not observed to be produced by mammalian carnivores. The assemblages are also characterized by the retention of both low- and high-density bone portions, an absence of gross gnawing, and minimal fragmentation. Together, the damage characteristics associated with feeding by crocodiles are highly distinctive from those produced by mammalian carnivores. Modern surface bone assemblages along the Grumeti River in Tanzania's Serengeti National Park contain a mixture of specimens bearing damage characteristic of crocodiles and mammalian carnivores. Comparison of Plio-Pleistocene fossil bones from Olduvai Gorge, Tanzania, to bones damaged by captive and free-ranging Nile crocodiles reveals direct evidence of fossil crocodilian feeding from larger mammal bones associated with Oldowan stone artifacts.     

Systematic butchering of fallow deer (Dama) at the early middle Pleistocene Acheulian site of Gesher Benot Ya'aqov (Israel)

Three assemblages of fallow deer (Dama sp.) bones excavated from the early middle Pleistocene (oxygen isotope stage 18) layers of the Acheulian site of Gesher Benot Ya'aqov, Israel, furnish evidence of systematic and repeated exploitation of complete carcasses by hominins. The excellent state of preservation of the bones and the presence of only minimal signs of carnivore involvement permit an investigation of the role of hominins as the primary agents responsible for the damage to these bones. Hominin expertise in dealing with fallow deer carcasses is manifested by cut marks, percussion marks, and hack marks on the bones. The archaeozoological analysis of the anatomical position and frequency of these marks suggests that carcass processing followed systematic practices that reflect an in-depth knowledge of fallow deer anatomy and a consistent behavioral strategy. These assemblages represent one of the earliest examples of methodological butchering practices in Eurasia. The evidence of carcass processing observed at Gesher Benot Ya'aqov resembles that seen in late Pleistocene sites in Israel, which were inhabited by modern humans. We interpret the Gesher Benot Ya'aqov data as indicating that the Acheulian hunters at the site (1) were proficient communicators and learners and (2) possessed anatomical knowledge, considerable manual skill, impressive technological abilities, and foresight.     

Beyond leopards: tooth marks and the contribution of multiple carnivore taxa to the accumulation of the Swartkrans Member 3 fossil assemblage

The ca. 1.0 myr old fauna from Swartkrans Member 3 (South Africa) preserves abundant indication of carnivore activity in the form of tooth marks (including pits) on many bone surfaces. This direct paleontological evidence is used to test a recent suggestion that leopards, regardless of prey body size, may have been almost solely responsible for the accumulation of the majority of bones in multiple deposits (including Swartkrans Member 3) from various Sterkfontein Valley cave sites. Our results falsify that hypothesis and corroborate an earlier hypothesis that, while the carcasses of smaller animals may have been deposited in Swartkrans by leopards, other kinds of carnivores (and hominids) were mostly responsible for the deposition of large animal remains. These results demonstrate the importance of choosing appropriate classes of actualistic data for constructing taphonomic inferences of assemblage formation. In addition, they stress that an all-encompassing model of assemblage formation for the hominid-bearing deposits of the Sterkfontein Valley is inadequate and that each must be evaluated individually using not just analogical reasoning but also incorporating empirical data generated in the preserved fossil samples.     

Zooarchaeological and taphonomic analysis of the Die Kelders Cave 1 layers 10 and 11 Middle Stone Age larger mammal fauna

Middle Stone Age (MSA) and Middle Paleolithic (MP) faunal assemblages have gained widespread attention due to their relevance to the debate over the modernity of hominid behavior during the MSA/MP. A recent critique of the scavenging argument for MSA/MP behavior drew on a summary presentation of the skeletal abundance and surface modification data from Die Kelders Cave 1 Layer 10 (Marean, 1998). This paper provides a more complete presentation of those data, adds the smaller Layer 11 sample, and provides a detailed analysis of the taphonomic history of both samples.Bone fragment density is higher in Layer 10 than in Layer 11. Bone densities vary horizontally as well, with Layer 10 showing greater deposition in the exposed areas of the cave. An analysis of long bone breakage patterns indicates that non-nutritive breakage on the Layers 10 and 11 samples was present but not intense. Size 1 mammals were predominantly accumulated by owls and/or other large raptors, not hominids, in Layer 10. Hominids were the predominant accumulator of Sizes 2-4 mammals in Layers 10 and 11 as indicated by the frequency of hammer-stone percussion marks and carnivore toothmarks. After discard by hominids, a significant portion of these remains were discovered and scavenged by carnivores. Overall, the larger mammal fauna of Layer 10 is dominated by Sizes 3 and 4 bovids, mostly young and adult eland, and thus hominids were focusing on the high-ranked prey items. Shaft portions of long bones, the portions with the most flesh, have the highest frequencies of cutmarks. A comparison of the Layers 10 and 11 cutmark frequencies to Selvaggio's (1998) scavenging model shows that the frequencies are significantly outside the range of variation documented in Selavaggio's scavenging sample.     

Role of carnivores in the accumulation of the Sterkfontein Member 4 hominid assemblage: a taphonomic reassessment of the complete hominid fossil sample (1936-1999)

New taphonomic data on the Sterkfontein Member 4 (South Africa) fossil hominid assemblage are presented. The previous estimate of hominid individuals represented in the deposit (45) is increased to 87. New minimum numbers of hominid skeletal elements are provided, and incidences of bone surface damage inflicted by prehistoric biological agents are summarized. The hominid sample from Member 4 is composed predominately of gnathic remains and has a paucity of postcrania. This dearth of postcrania limits, to some extent, inferences about the formation of the Sterkfontein assemblage. However, carnivore tooth marks on some fossil specimens and an overall broad similarity in patterns of skeletal part representation between Sterkfontein and primate bone assemblages created by extant carnivores suggest that carnivores did have some involvement in the accumulation of the fossil hominid assemblage. Thus, this study provides support for the “carnivore‐collecting hypothesis” of Brain (Brain [ 1981 ] The Hunters or the Hunted? Chicago: University of Chicago Press), which implicates large carnivores as prominent collecting agents of hominid body parts in Sterkfontein Member 4. Evidence of bone surface damage is, however, too scant to make confident inferences about specific carnivore taxon/taxa involved in hominid bone collection at the site. Am J Phys Anthropol, 2004. © 2004 Wiley‐Liss, Inc.     

Tooth Marks of Mammalian Incisors on Rocky Substrate in Brazil: Evidence of Geophagy in the Cerrado Biome

Tooth marks on sandstone in an area of the Cerrado Biome are reported, indicating geophagy. The tooth marks were found on reddish sandstones cropping out in a pasture environment with typical components of the Cerrado Biome, in the Municipality of Campina Verde, Triângulo Mineiro region (west Minas Gerais State, Brazil). Studies have shown that the soil of the Cerrado is acid, with a low concentration of nutrients and minerals (also present in the plants living on this environment), which usually produce an alimentary deficiency in herbivorous animals. Therefore, these tooth marks indicate geophagy, in order to extract extra minerals from these sandstone levels, which have a high concentration of calcium carbonate and iron. The tooth marks consist of two parallel concave grooves and a medial prominent crest which results from the action of the incisors of mammals. Although the identification of the gnawing species for these sets of tooth marks are estimates at best, after wide comparisons we tentatively suggest that the tooth marks are most likely the result of the action of the incisors of rodents, such as Dasyprocta or Coendou.     

Scavenging or Hunting in Early Hominids: Theoretical Framework and Tests

Evidence from Bed I, Olduvai, supports the hypothesis that scavenging, not hunting, was the major meat-procurement strategy of hominids between 2 and 1.7 million years ago. Data used to evaluate the hunting and scavenging hypotheses are derived from studying cut marks on Bed I bovids, comparing adaptations necessary for scavenging with those of early hominids, and a pa-leoecological reconstruction of Bed I carcass biotnass, carnivore guild, and hominidforaging area.     

Growth layers and incremental markings in hard tissues; a review of the literature and some preliminary observations about enamel structure in Paranthropus boisei

The general factors underlying the formation of growth layers and incremental markings in hard tissues are reviewed with particular reference to fossil hominid tooth enamel. The experimental and circumstantial evidence that point to a slowing of enamel matrix secretion in a daily (circadian) and near weekly (circaseptan) mode during tooth formation is also reviewed. Data from previous studies in which the number of daily increments between adjacent striae of Retzius have been recorded in primates are reviewed and new data are presented for this repeat interval in fossil hominids. The factors likely to influence the number of striae of Retzius beneath the cuspal regions of anterior teeth are outlined and the limitations of employing surface incremental features to obtain estimates for age at death of an individual are also discussed. It is concluded that there is good evidence to support the hypothesis that perikymata are near weekly incremental phenomena with a likely periodicity of 7,8 or 9 days in fossil hominids. It can also be concluded that at present, better estimates for the age at death of an individual during early phases of the growth period can be obtained from studies of perikymata than by any other non-destructive technique.     

Environments and hominin activities across the FLK Peninsula during Zinjanthropus times (1.84 Ma), Olduvai Gorge, Tanzania

We establish through 13 excavations the landscape context and nature of hominin activities across the Zinjanthropus land surface from which the Leakeys recovered the FLK 22 and FLK NN 1 paleoanthropological assemblages. The land surface was created by fluvial incision of the eastern margin of paleo-Lake Olduvai following a major lake withdrawal. Erosion was uneven, leaving a peninsula bounded by a river channel, the FLK Fault, and a freshwater wetland. This FLK Peninsula supported groves of trees that attracted hominins and carnivores, and that preserved the dense concentrations of carcass remains and stone tools they left behind, including those at FLK 22. Some carcasses appear to have been acquired at the ecotone of the Peninsula and Wetland, where another dense artifact and bone assemblage accumulated. A lesser topographic high at the edge of a Typha marsh in the Wetland was the site of FLK NN 1 and a scatter of large stone tools used possibly for rootstock processing.Our landscape reconstruction delimits the vegetation mosaic indicated by previous work and provides a topographical explanation for the existence of FLK 22 and FLK NN 1. Both are unexpected if the FLK area was the flat, featureless lake margin terrain typical of lake basins similar to paleo-Olduvai. The results show that the Leakeys’ sites were not isolated occupation floors but rather parts of a land surface utilized intensively by hominins. Although commonly considered to have been home bases, their likely high predation risk, evidenced by large carnivore feeding traces and the remains of four hominin individuals, suggests visits to them were brief and limited to feeding. Finally, stratigraphic observations confirm that FLK NN 3 accumulated on an older land surface, refuting the hypothesis that the OH 8 foot found there is the same individual as the OH 35 leg from FLK 22.