Search

Although many studies relating stature to foot length have been carried out, the relationship between foot size and body mass remains poorly understood. Here we investigate this relationship in 193 adult and 50 juvenile habitually unshod/minimally shod individuals from five different populations—Machiguenga, Daasanach, Pume, Hadzabe, and Samoans—varying greatly in body size and shape. Body mass is highly correlated with foot size, and can be predicted from foot area (maximum length × breadth) in the combined sample with an average error of about 10%. However, comparisons among populations indicate that body shape, as represented by the body mass index (BMI), has a significant effect on foot size proportions, with higher BMI samples exhibiting relatively smaller feet. Thus, we also derive equations for estimating body mass from both foot size and BMI, with BMI in footprint samples taken as an average value for a taxon or population, estimated independently from skeletal remains. Techniques are also developed for estimating body mass in juveniles, who have relatively larger feet than adults, and for converting between foot and footprint size. Sample applications are given for five Pliocene through Holocene hominin footprint samples from Laetoli (Australopithecus afarensis), Ileret (probable Homo erectus), Happisburgh (possible Homo antecessor), Le Rozel (archaic Homo sapiens), and Barcin Hoyuk (H. sapiens). Body mass estimates for Homo footprint samples appear reasonable when compared to skeletal estimates for related samples. However, estimates for the Laetoli footprint sample using the new formulae appear to be too high when compared to skeletal estimates for A. afarensis. Based on the proportions of A.L. 288-1, this is apparently a result of relatively large feet in this taxon. A different method using a ratio between body mass and foot area in A.L. 288-1 provides estimates more concordant with skeletal estimates and should be used for A. afarensis.

Bipedal trackways discovered in 1978 at Laetoli site G, Tanzania and dated to 3.66 million years ago are widely accepted as the oldest unequivocal evidence of obligate bipedalism in the human lineage1–3. Another trackway discovered two years earlier at nearby site A was partially excavated and attributed to a hominin, but curious affinities with bears (ursids) marginalized its importance to the paleoanthropological community, and the location of these footprints fell into obscurity3–5. In 2019, we located, excavated and cleaned the site A trackway, producing a digital archive using 3D photogrammetry and laser scanning. Here we compare the footprints at this site with those of American black bears, chimpanzees and humans, and we show that they resemble those of hominins more than ursids. In fact, the narrow step width corroborates the original interpretation of a small, cross-stepping bipedal hominin. However, the inferred foot proportions, gait parameters and 3D morphologies of footprints at site A are readily distinguished from those at site G, indicating that a minimum of two hominin taxa with different feet and gaits coexisted at Laetoli., Reanalysis of bipedal trackways from Laetoli site A in Tanzania suggest that the footprints were made by a hominin that coexisted with at least one other hominin species.

The Tianyuan 1 skeleton is among the oldest fossil specimens of our species from East Asia. Important in the original assessment of the skeleton as belonging to a modern human was its femoral structure, particularly its well-developed pilasters. Also noteworthy were its large femoral diaphyseal second moments of area, which were interpreted as evidence of a physically demanding lifestyle, and pathological bone growths on the posterior femoral shafts. Here, we build on previous studies of femoral morphology in Tianyuan 1 by analyzing its diaphyseal structure using micro-computed tomography coupled with a novel method of visualizing cortical bone thickness distributions along the shaft with color maps. Additionally, we calculate diaphyseal second moments of area in Tianyuan 1 with and without its pathological bone additions in order to evaluate whether these growths are the cause of its apparently high diaphyseal robusticity. Diaphyseal color maps and second moments of area of Tianyuan 1 femora are compared to those of three recent (Holocene) human comparative samples, as well as samples of Pleistocene humans. The results show that in terms of cortical thickness distribution, Tianyuan 1 femoral diaphyses are strikingly similar to those of recent humans, as well as Pleistocene early modern humans, yet distinct from the diaphyses of Neandertals. This provides additional support for the modern human status of Tianyuan 1. Analyses of second moments of area reveal that Tianyuan 1 is best classified as a Pleistocene early modern human rather than as a member of any of the recent samples when pathological bone additions are included in area moment calculations. Interestingly, however, once pathological growths are virtually removed, Tianyuan 1 cannot be assigned with confidence to any comparative group. This suggests that the relatively high anteroposterior femoral diaphyseal strength mid-distally in Tianyuan 1 is to some degree a product of its pathological condition.

Isimila is a Middle Pleistocene archaeological site located in southern Tanzania. The site is known for large surface assemblages of later Acheulean lithics such as hand axes, cleavers, scrapers and cores. While hominin remains have yet to be discovered at the site, Isimila offers a unique window into Middle Pleistocene Homo behaviour. Although Isimila has been studied extensively, the last published map of the site and surrounding area was made available in the 1970s. Here, we present an updated high-resolution map of Isimila. Data for the map were collected during aerial survey with an uncrewed(unmanned) aerial vehicle. With this map, we identified new archaeological localities, erosional patterns, newly exposed geological features and changes in site topography. The map demonstrates patterns of stone tool and raw material distribution that may support previous hypotheses of short-distance raw material transport into the area by hominins. This open-access map establishes a baseline for tracking changes to site topography in the future and serves as a unique tool to enable collaboration between researchers, museum personnel and local populations to better conserve Isimila. Significance: New potential archaeological localities and significant changes to erosional patterns at Isimila were identified. The open access map and associated raw data provided enable researchers to track seasonal and erosional changes and anthropogenic effects, and to develop protocols for conservation of this unique site. Open data set: https://doi.org/10.5281/zenodo.1470770

Isimila is a Middle Pleistocene archaeological site located in southern Tanzania. The site is known for large surface assemblages of later Acheulean lithics such as hand axes, cleavers, scrapers, and cores. While hominin remains have yet to be discovered at the site, Isimila offers a unique window into Middle Pleistocene Homo behavior. Although Isimila has been studied extensively, the last published map of the site and surrounding area was made available in the 1970s. Here, we present an updated high-resolution map of Isimila. Data for the map were collected during aerial survey with an uncrewed/unmanned aerial vehicle (UAV). With this map, we identify new archaeological localities, erosional patterns, newly exposed geological features, and changes in site topography. The map reveals patterns of stone tool and raw material distribution that may support previous hypotheses of raw material transport into the area by hominins. This open-access map establishes a baseline for tracking changes to site topography in the future and serves as a unique tool to enable collaboration between researchers, museum personnel, and local populations to better conserve Isimila.

The species Homo naledi was recently named from specimens recovered from the Dinaledi Chamber of the Rising Star cave system in South Africa. This large skeletal sample lacks associated faunal material and currently does not have a known chronological context. In this paper, we present comprehensive descriptions and metric comparisons of the recovered cranial and mandibular material. We describe 41 elements attributed to Dinaledi Hominin (DH1–DH5) individuals and paratype U.W. 101-377, and 32 additional cranial fragments. The H. naledi material was compared to Plio- Pleistocene fossil hominins using qualitative and quantitative analyses including over 100 linear measurements and ratios. We find that the Dinaledi cranial sample represents an anatomically homogeneous population that expands the range of morphological variation attributable to the genus Homo. Despite a relatively small cranial capacity that is within the range of australopiths and a few specimens of early Homo, H. naledi shares cranial characters with species across the genus Homo, including Homo habilis, Homo rudolfensis, Homo erectus, and Middle Pleistocene Homo. These include aspects of cranial form, facial morphology, and mandibular anatomy. However, the skull of H. naledi is readily distinguishable from existing species of Homo in both qualitative and quantitative assessments. Since H. naledi is currently undated, we discuss the evolutionary implications of its cranial morphology in a range of chronological frameworks. Finally, we designate a sixth Dinaledi Hominin (DH6) individual based on a juvenile mandible.

Actualistic observations on modern lions (Panthera leo) from Tarangire National Park (Tanzania) have expanded the reported range of carcass consumption behaviors by these felids. The present study confirms that lions efficiently deflesh small and medium-sized carcasses and they can even thoroughly deflesh carcasses heavier than 500 kg, such as those of buffaloes. Ecology plays a major role in the intensity with which lions deflesh their prey. The most intensive carcass consumption episodes in Tarangire were documented in alluvial environments near water. Bone damage is proportional to the intensity of carcass consumption and upper limb bones, usually the most defleshed elements, are also the most heavily damaged. Butchery experiments with stone tools modelling secondary access to lion kills yielded a low cut mark frequency with an anatomical distribution of cut marks occurring more frequently on intermediate than on upper limb bones and on ends and metadiaphyses than on mid-shafts. The combination of the damage inflicted by lions on bones and the occurrence of cut marks as the result of secondary-access butchery by humans provides a heuristic framework with which to understand similar patterns in the archaeological record.

The taphonomic signature of crocodiles as agents of bone modification has been previously identified by specific tooth mark types (e.g., bisected pits) and by a conspicuous presence of these marks: more than 75% of bones modified by crocodiles bear at least one of these distinctive marks. Therefore, crocodile tooth-marking would be notably prevalent in bone assemblages resulting from crocodile predation and active scavenging. The present study contributes to refine this diagnosis by showing greater variability of these types of marks, a different degree of tooth-marking intensity, and a somewhat different distribution of tooth-marked elements according to skeletal parts from previous experiments with crocodiles. Some of these differences are due to different experimental variables and conditions and this highlights the need to understand behavioral variability in crocodile ecological settings. This variability in crocodile tooth-marking probably results from several as-yet-inadequately measured behavioral and ecological factors, such as intensity of feeding competition and differences between male and female crocodile feeding behaviors, among others. Furthermore, this study also contributes to a better definition of the microscopic criteria that can be used to distinguish crocodile-inflicted marks from other types of bone surface modifications. In light of these and previous experimental frameworks, we reevaluate the application of these analogs to modifications documented in hominin fossils from Olduvai Gorge (OH8 and OH35) and the resulting inferences about the hazard posed by crocodiles on the paleolandscape where FLK North North and FLK Zinj (Bed I) were formed. The taphonomic analysis also shows that Olduvai OH8 and OH35 were probably not preyed upon by crocodiles. It is concluded that no tangible evidence can be used to support the interpretation that OH35 was modified by crocodiles and that the overall presence of crocodiles in FLK North North and FLK Zinj was rather marginal, based on the virtual absence of crocodile-modified bones in both archaeofaunal assemblages.

At 3.6 Ma, the Laetoli Pliocene hominin trackways are the earliest direct evidence of hominin bipedalism. Three decades since their discovery, not only is the question of their attribution still discussed, but marked differences in interpretation concerning the footprints’ qualitative features and the inferred nature of the early hominin foot morphology remain. Here, we establish a novel ichnotaxon, Praehominipes laetoliensis , for these tracks and clarify the distinctions of these footprints from those of later hominins, especially modern humans. We also contrast hominin, human, and ape footprints to establish morphological features of these footprints correlated with a midtarsal break versus a stiff longitudinal arch. Original photos, including stereo photographs, and casts of footprints from the 1978 Laetoli excavation, confirm midtarsal flexibility, and repeatedly indicate an associated midfoot pressure ridge. In contrast, the modern human footprint reflects the derived arched-foot architecture, combined with a stiff-legged striding gait. Fossilized footprints of unshod modern human pedestrians in Hawaii and Nicaragua unambiguously illustrate these contrasts. Some points of comparisons with ape footprints are complicated by a variable hallucal position and the distinct manner of ape facultative bipedalism. In contrast to the comparatively rigid platform of the modern human foot, midtarsal flexibility is present in the chimpanzee foot. In ape locomotion, flexion at the transverse tarsal joint, referred to as the “midtarsal break,” uncouples the respective functions of the prehensile forefoot and the propulsive hindfoot during grasp-climbing. At some point after the transition to habitual bipedalism, these grasp-climb adaptations, presumed to be present in the last common ancestor of apes and humans, were initially compromised by the loss of divergence of the hallux. An analogous trajectory is evident along an array of increasingly terrestrial extant ape species. However, a flexible midfoot was retained, presumably to spare lateral toes from bending stresses by concentrating push-off from the forefoot, beneath the metatarsals. Only later did the evolution of the longitudinal arch permit increased mechanical advantage of the plantarflexors for speed and improved economy of endurance distance walking and running.

Paleoecology allows construction of paleoenvironmental models, faunal changes and evolutionary trends of paleontological taxa using modern analogs. However, when linking modern analogs to paleontological taxa in paleoecological reconstruction, differential taxonomic preservation in the fossil record has to be taken into account. Paleontologists have known the biased nature of the fossil record since Efremov’s publication on taphonomy in 1940, yet many ecological models of habitats associated with hominins in paleontological and archaeological sites in Africa and elsewhere barely address the complexity of the fossil record. We use randomly sampled ungulates from modern biomes in a comparative taxonomic abundance to demonstrate how the combination of modern thanatocoenoses and taphocoenoses, when used in reference to habitat-specific biocoenosis, produce better inferences of past habitats in paleontological and archaeological sites than approaches currently used.

Laetoli, a paleoanthropological site in Northern Tanzania, is perhaps best known for its famous fossil hominid footprints that were discovered by Mary Leakey and her co-workers in 1978. The site not only preserves the hominid footprints but also trackways, which provide a snapshot of Pliocene faunal communities from East Africa and their inferred environments. Unlike the hominid footprints at site G, which have received tremendous attention, the animal trackways, especially at Localities 7, 8 and 10 have been neglected and are fast disappearing. In this paper, we discuss animal tracks at a newly discovered exposure and provide preliminary data on the tracks at this exposure and other sites. We also discuss the importance of the animal trackways as ecological indicators, which we have investigated as part of ongoing research and conservation efforts initiated by the Tanzania Field School in Paleoanthropology and the Associated Colleges of the Midwest (ACM) Tanzania Semester Abroad programs.

Homo rudolfensis Samples from Olduvai Gorge, East Lake Turkana, and Lake Malawi were included in this study. The East Lake Turkana fossils available prior to 2010 were examined first-hand, while for the Olduvai and Lake Malawi fossils and KNM-ER 60000, 62000, and 62003 we relied on original observations on fossils and casts as well as published reports (Schrenk et al., 1993; Blumenschine et al., 2003; Leakey et al., 2012). We include the following fossils in the hypodigm of H. rudolfensis: KNM-ER 819, KNM-ER 1470, KNM-ER 1482, KNM-ER 1483, KNM-ER 1590, KNM-ER 1801, KNM-ER 1802, KNM-ER 3732, KNM-ER 3891, KNM-ER 60000, KNM-ER 62000, KNM-ER 62003, OH 65, and UR 501. We do recognize that KNM-ER 60000 and KNM-ER 1802 present some conflicting anatomy that some authors have argued precludes them as conspecific specimens (Leakey et al., 2012); by considering both, we aim to be conservative as they encompass more variation within H. rudolfensis., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Australopithecus africanus The samples attributed to Au. africanus from Taung, Sterkfontein and Makapansgat were employed. Original specimens were examined first-hand by the authors., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Paranthropus boisei Samples from the Omo Shungura sequence, East Lake Turkana, Olduvai Gorge and Konso were included in this study. Original specimens from Olduvai Gorge and East Lake Turkana were examined first-hand, while casts and published reports (Tobias, 1967; Suwa et al., 1996, 1997; Do ḿınguez- Rodrigo et al., 2013) were used to study the Omo and Konso materials. Our postcranial considerations of P. boisei are very limited and we did not rely upon the association of KNM-ER 1500 (Grausz et al., 1988) to derive information about the postcranial skeleton of P. boisei. Paranthropus robustus The samples from Kromdraai, Swartkrans, Sterkfontein, Drimolen, Gondolin, and Coopers were included in this study. First-hand observations of original specimens from all localities were used with the exception of Drimolen fossils, which were compared using published reports (Keyser, 2000; Keyser et al., 2000)., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Paranthropus aethiopicus The cranium KNM-WT 17000 was examined first-hand for this study., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Homo erectus Samples from Buia, Chemeron, Daka, Dmanisi, East and West Lake Turkana, Gona, Hexian, Konso, Mojokerto, Olduvai Gorge, Sangiran, Swartkrans, Trinil, and Zhoukoudian were included in this study. South African material is of special interest in this comparison because of the geographic proximity, and because of the difficulty of clearly identifying Homo specimens within the large fossil sample from Swartkrans. In particular, the following specimens from Swartkrans are considered to represent H. erectus: SK 15, SK 18a, SK 27, SK 43, SK 45, SK 68, SK 847, SK 878, SK 2635, SKW 3114, SKX 257/258, SKX 267/ 2671, SKX 268, SKX 269, SKX 334, SKX 339, SKX 610, SKX 1756, SKX 2354, SKX 2355, SKX 2356, and SKX 21204. It has been suggested (Grine et al., 1993, 1996) that SK 847 and Stw 53 might represent the same taxon, and that this taxon is a currently undiagnosed species of Homo in South Africa. However, we agree with Clarke (1977; 2008) that SK 847 can be attributed to H. erectus, and that Stw 53 cannot. Because there is no clear indication that more than one species of Homo is represented in the Swartkrans sample, we consider all this material to belong to H. erectus. We considered ‘ Homo ergaster ’ (and also ‘ Homo aff. erectus ’ from Wood, 1991) to be synonyms of H. erectus for this study; Turkana Basin specimens that are attributed to H. erectus thus include KNM-ER 730, KNM-ER 820, KNM-ER 992, KNM- ER 1808, KNM-ER 3733, KNM-ER 3883, KNM-ER 42700, KNM-WT 15000. Olduvai specimens include OH 9, OH 12 and OH 28. Original fossil materials from Chemeron, Lake Turkana, Swartkrans, Trinil, and Dmanisi were examined first-hand by the authors, while the remainder were based on casts and published reports (Weidenreich, 1943; Wood, 1991; Anton´, 2003; Rightmire et al., 2006; Suwa et al., 2007). A large number of postcranial specimens have been collected from the Turkana Basin and appear consistent with the anatomical range otherwise found in Homo, and inconsistent with known samples of Australopithecus and Paranthropus from elsewhere. These include KNM-ER 1472, KNM-ER 1481, KNM- ER 3228, KNM-ER 737, and others. We may add other fossils from other sites lacking association with craniodental material, such as the partial BOU-VP 12/1 skeleton and even the Gona pelvis. These specimens attributable to Homo but not necessarily to a particular species did inform our understanding of variability within the genus, but for the most part these specimens do not inform our differential diagnosis of H. naledi relative to particular species. For example, the key element of femoral morphology of H. naledi in contrast to other species is the presence of two well-defined mediolaterally running pillars in the femoral neck; the isolated specimens of early Homo do not contradict this apparent autapomorphy. Likewise, no isolated specimens inform us about the humanlike aspects of foot morphology in H. naledi. In these cases, the lack of associations for this evidence actually is less important than the lack of specimens that replicate the distinctive features of the H. naledi morphology., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 27, DOI: 10.7554/eLife.09560

Australopithecus garhi The cranium BOU-VP-12/130 from Bouri was included, with data taken from a published report (Asfaw et al., 1999)., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Australopithecus sediba The partial skeletons MH1 and MH2 from Malapa, South Africa were included in this study, based on examination of the original specimens by the authors., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Middle Pleistocene Homo Specimens from the latest Lower Pleistocene and MP of Europe and Africa that cannot be attributed to H. erectus were included in our comparisons. These include fossils that have been attributed to H. heidelbergensis, H. rhodesiensis, ‘archaic H. sapiens ’, or ‘evolved H. erectus ’ by a variety of other authors. Specimens attributed to MP Homo include materials from Eliye Springs, Arago, Atapuerca Sima de los Huesos, Bodo, Broken Hill, Cave of Hearths, Ceprano, Dali, Elandsfontein, Jinniushan, Kapthurin, Mauer, Narmada, Ndutu, Petralona, Reilingen-Schwetzingen, Solo, Steinheim, Swanscombe. This grouping includes the following specimens: KNM-ES 11693, Arago 2, Arago 13, Arago 21, Atapuerca 1, Atapuerca 2, Atapuerca 4, Atapuerca 5, Atapuerca 6, Cave of Hearths, SAM-PQ- EH1, Kabwe, Mauer, Ndutu, Sale´, Petralona, Reilingen-Schwetzingen, Steinheim.

© Berger et al. Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.

Acknowledgements We wish to express our gratitude to the National Geographic Society and the National Research Foundation of South Africa for funding the discovery, recovery, and analysis of the H. naledi material. The study reported here was also made possible by grants from the Social Sciences and Humanities Research Council of Canada, the Canada Foundation for Innovation, the British Columbia Knowledge Development Fund, the Canada Research Chairs Program, Simon Fraser University, the DST/NRF Centre of Excellence in Palaeosciences (COE-Pal), as well as by a Discovery Grant from the Natural Sciences and Engineering Research Council of Canada, a Young Scientist Development Grant from the Paleontological Scientific Trust (PAST), a Baldwin Fellowship from the L.S.B. Leakey Foundation, and a Seed Grant and a Cornerstone Faculty Fellowship from the Texas AM the Jacobs family for granting access; Wilma Lawrence, Bonita De Klerk, Merrill Van der Walt, and Justin Mukanku for their assistance during all phases of the project; Lucas Delezene for valuable discussion on the dental characters of H. naledi. We would also like to thank Peter Schmid for the preparation of the Dinaledi fossil material; Yoel Rak for explaining in detail some of the characters used in previous studies; William Kimbel for drawing our attention to the possibility that there might be a problem with Dembo et al.’s (2015) codes for the two characters related to the articular eminence; Will Stein for helpful discussion about the Bayesian analyses; Mike Lee for his comments on this manuscript; John Hawks for his support in organizing the Rising Star workshop; and the associate editor and three anonymous reviewers for their valuable comments. We are grateful to S. Potze and the Ditsong Museum, B. Billings and the School of Anatomical Sciences at the University of the Witwatersrand, and B. Zipfel and the Evolutionary Studies Institute at the University of the Witwatersrand for providing access to the specimens in their care; the University of the Witwatersrand, the Evolutionary Studies Institute, and the South African National Centre of Excellence in PalaeoSciences for hosting a number of the authors while studying the material; and the Western Canada Research Grid for providing access to the high-performance computing facilities for the Bayesian analyses. Last but definitely not least, we thank the head of the Rising Star project, Lee Berger, for his leadership and support, and for encouraging us to pursue the study reported here.

We describe the physical context of the Dinaledi Chamber within the Rising Star cave, South Africa, which contains the fossils of Homo naledi. Approximately 1550 specimens of hominin remains have been recovered from at least 15 individuals, representing a small portion of the total fossil content. Macro-vertebrate fossils are exclusively H. naledi, and occur within clay-rich sediments derived from in situ weathering, and exogenous clay and silt, which entered the chamber through fractures that prevented passage of coarser-grained material. The chamber was always in the dark zone, and not accessible to non-hominins. Bone taphonomy indicates that hominin individuals reached the chamber complete, with disarticulation occurring during/after deposition. Hominins accumulated over time as older laminated mudstone units and sediment along the cave floor were eroded. Preliminary evidence is consistent with deliberate body disposal in a single location, by a hominin species other than Homo sapiens, at an as-yet unknown date. DOI: http://dx.doi.org/10.7554/eLife.09561.001, eLife digest Modern humans, or Homo sapiens, are now the only living species in their genus. But as recently as 20,000 years ago there were other species that belonged to the genus Homo. Together with modern humans, these extinct human species, our immediate ancestors and their close relatives are collectively referred to as ‘hominins’. Now, Dirks et al. describe an unusual collection of hominin fossils that were found within the Dinaledi Chamber in the Rising Star cave system in South Africa. The fossils all belong to a newly discovered hominin species called Homo naledi, which is described in a related study by Berger et al. The unearthed fossils are the largest collection of hominin fossils from a single species ever to be discovered in Africa, and include the remains of at least 15 individuals and multiple examples of most of the bones in the skeleton. Dirks et al. explain that the assemblage from the Dinaledi Chamber is unusual because of the large number of fossils discovered so close together in a single chamber deep within the cave system. It is also unusual that no other large animal remains were found in the chamber, and that the bodies had not been damaged by scavengers or predators. The fossils were excavated from soft clay-rich sediments that had accumulated in the chamber over time; it also appears that the bodies were intact when they arrived in the chamber, and then started to decompose. Dirks et al. discuss a number of explanations as to how the remains came to rest in the Dinaledi Chamber, which range from whether Homo naledi lived in the caves to whether they were brought in by predators. Most of the evidence obtained so far is largely consistent with these bodies being deliberately disposed of in this single location by the same extinct hominin species. However, a number of other explanations cannot be completely ruled out and further investigation is now needed to uncover the series of events that resulted in this unique collection of hominin fossils. DOI: http://dx.doi.org/10.7554/eLife.09561.002

A geomorphological, sedimentological, stratigraphic, and geometric study of 30 trenches excavated around FLK Zinj (Bed I, Olduvai Gorge) has enabled the partial reconstruction of the paleolandscape surrounding this site for a radius of similar to 1000 m. This is the largest sample of geological and archaeological information yet available to reconstruct the topography, ecology, and geomorphology of the Zinj paleosurface and the hominin activities preserved within it. Contrary to previous interpretations, which place FLK Zinj on an isolated and narrow peninsula, it appears that the site was located on the edge of an elevated platform traceable for hundreds of meters. Hominins created FLK Zinj (and other sites, such as the recently discovered PTK and AMK) within the wooded habitats of this platform rather than the more open and grassy environments situated on lower portions of the lacustrine floodplain. Input areas, probably in the form of alluvial fans, existed to the south, following a North-South direction. These input areas are partially responsible for changes in the type sequence. Restricted erosion documented on the wooded platform was mostly caused by runoff processes. An archaeological study of the excavated trenches reveals a sharp contrast in fossil and stone tool density between FLK Zinj and the surrounding landscape, further supporting the contention that the site may have acted as a "central place" where repeated carcass transport, butchery, and consumption took place. Taphonomic studies indicate that at this stage of human evolution, hominins had primary access to carcasses and were not dependent on other carnivores for obtaining meat.

For many years, the management of cultural heritage sites and the designation of some of them as World Heritage Sites in Africa were based on European ideas of conservation and this disconnected many African local communities from their cultural heritage sites. As a result, local African communities living near cultural heritage sites were not involved in their conservation and management. Discourses on the administration of cultural heritage sites in many African countries, such as Botswana South Africa, Kenya, Tanzania, and Zimbabwe, are now making it possible to engage local people in the management of these sites (see, for example, Pwiti 1997; Musiba and Mabulla 2003). Not only do these discussions bring great prestige to the African States Parties whose sites are designated as World Heritage properties but they are also now making it possible for the local communities to economically benefit from them. Part of the strategy of sustainable management of cultural World Heritage Sites now includes creating opportunities for the local communities to be involved in tourism activities so as to economically empower them and improve their lives. In this chapter, I discuss the administration of cultural World Heritage Sites in Africa and their contribution to the economic empowerment of local communities which subsist near these sites. I use the planned development of paleoanthropological sites of Laetoli and Olduvai Gorge in Tanzania as examples to show how local communities can benefit from cultural World Heritage Sites.

Data on footprints and gait of 54 Hadzabe, 6-70 years of age expand understanding of pedal morphology of unshod people and assist the development of ideas about the evolution of hominid bipedality and upright posture. Contrary to published data on gaits and pedal morphology of unshod populations, Hadzabe (also known as Hadza) from northern Tanzania exhibit values of stride length, relative stride length, and walking speeds that exceed those of rural and small-town populations. In all observable plantar features, including foot indices, an interdigital space between the hallux and second toe, fanning of the foot anteriorly, and foot angles (in-toeing and out-toeing), Hadzabe feet are comparable with those of never-shod Machiguengas in Peru ´ . On average, Hadzabe hallucal gaps and ball widths are narrower than those of Machiguengas and other unshod short people. Hadzabe feet are also charac- terized by valgus halluces versus the varus halluces of never-shod Machiguen- gas and certain West Africans. Although characterized by a valgus toe, Had- zabe hallucal angles, which do not exceed 208, are lower than those of Northern Hemispheric urbanites and shod rural populations of the Southern and North- ern Hemispheres. Hadzabe also exhibit less medial and lateral rotation of the hip joint than Machiguengas do. The heel and the longitudinal arch impressions of the Hadzabe footprints closely resemble those of the Laetoli bipeds in the manner of weight distribution during locomotion. The striking similarity of footprint impressions, especially the heel and the longitudinal arch, between Hadzabe and Laetoli hominid footprints clearly imply that the pedal features of the Laetoli printmakers are remarkably humanoid. Am. J. Hum. Biol. 9:303-321, 1997. q 1997 Wiley-Liss, Inc.

International audience; Recent excavations in Level 4 at BK (Bed II, Olduvai Gorge, Tanzania) have yielded nine hominin teeth, a distal humerus fragment, a proximal radius with much of its shaft, a femur shaft, and a tibia shaft fragment (cataloged collectively as OH 80). Those elements identified more specifically than to simply Hominidae gen. et sp. indet are attributed to Paranthropus boisei. Before this study, incontrovertible P. boisei partial skeletons, for which postcranial remains occurred in association with taxonomically diagnostic craniodental remains, were unknown. Thus, OH 80 stands as the first unambiguous, dentally associated Paranthropus partial skeleton from East Africa. The morphology and size of its constituent parts suggest that the fossils derived from an extremely robust individual who, at 1.338±0.024 Ma (1 sigma), represents one of the most recent occurrences of Paranthropus before its extinction in East Africa.

International audience; Recent excavations at FLK North (Olduvai Gorge, Tanzania) have produced new information on the orientation of archaeological materials at various levels of the site. This information includes the uniform distribution of material azimuths, which contrasts with previous inferences of highly patterned orientations of materials in the Bed I archaeological sites. Those previous inferences of patterned material orientations are based on Mary Leakey's 50-year-old drawings of artifact and fossil bone distribution, but are not verified by our precise measurements of archaeological objects made in situ. Nor do those previous results agree with the general lack of geological, geomorphological, and/or taphonomic data that would indicate significant post-depositional movement of archaeological materials in the sites. We argue here that Leakey's drawings are incomplete (only portions of each assemblage were drawn) and inaccurate in their representation of the original locations, shapes and orientations of most archaeological specimens. This argument is supported by several important mismatches in object representations between a photograph taken of a small portion of the FLK 22 Zinjanthropus site floor before the removal of the archaeological items, and the sketch of the same area drawn by Leakey. Thus, we conclude that primary orientation data of excavations (i.e., direct measurements taken from items) generated prior to object removal are the only valid indicators of the relative isotropy or anisotropy of these important paleoanthropological assemblages.

International audience; Meat-eating was an important factor affecting early hominin brain expansion, social organization and geographic movement. Stone tool butchery marks on ungulate fossils in several African archaeological assemblages demonstrate a significant level of carnivory by Pleistocene hominins, but the discovery at Olduvai Gorge of a child's pathological cranial fragments indicates that some hominins probably experienced scarcity of animal foods during various stages of their life histories. The child's parietal fragments, excavated from 1.5-million-year-old sediments, show porotic hyperostosis, a pathology associated with anemia. Nutritional deficiencies, including anemia, are most common at weaning, when children lose passive immunity received through their mothers' milk. Our results suggest, alternatively, that (1) the developmentally disruptive potential of weaning reached far beyond sedentary Holocene food-producing societies and into the early Pleistocene, or that (2) a hominin mother's meat-deficient diet negatively altered the nutritional content of her breast milk to the extent that her nursing child ultimately died from malnourishment. Either way, this discovery highlights that by at least 1.5 million years ago early human physiology was already adapted to a diet that included the regular consumption of meat.

Data on footprints and gait of 54 Hadzabe, 6-70 years of age expand understanding of pedal morphology of unshod people and assist the development of ideas about the evolution of hominid bipedality and upright posture. Contrary to published data on gaits and pedal morphology of unshod populations, Hadzabe (also known as Hadza) from northern Tanzania exhibit values of stride length, relative stride length, and walking speeds that exceed those of rural and small-town populations. In all observable plantar features, including foot indices, an interdigital space between the hallux and second toe, fanning of the foot anteriorly, and foot angles (in-toeing and out-toeing), Hadzabe feet are comparable with those of never-shod Machiguengas in Perú. On average, Hadzabe hallucal gaps and ball widths are narrower than those of Machiguengas and other unshod short people. Hadzabe feet are also characterized by valgus halluces versus the varus halluces of never-shod Machiguengas and certain West Africans. Although characterized by a valgus toe, Hadzabe hallucal angles, which do not exceed 20°, are lower than those of Northern Hemispheric urbanites and shod rural populations of the Southern and Northern Hemispheres. Hadzabe also exhibit less medial and lateral rotation of the hip joint than Machiguengas do. The heel and the longitudinal arch impressions of the Hadzabe footprints closely resemble those of the Laetoli bipeds in the manner of weight distribution during locomotion. The striking similarity of footprint impressions, especially the heel and the longitudinal arch, between Hadzabe and Laetoli hominid footprints clearly imply that the pedal features of the Laetoli printmakers are remarkably humanoid. Am. J. Hum. Biol. 9:303-321, 1997. © 1997 Wiley-Liss, Inc.

1. The definition of habitat (on page 2) and ecosystem is incomplete. For habitat it currently reads; a place where a population of animals and plants live, as well as their surroundings, both living and non-living. This should read as follows: A portion of the biome, which shows distinctive faunal, vegetational, trophic and lithological properties. For ecosystem it currently reads; An ecosystem here is considered as part of ecology consisting of the environment, its living parts, and the non-living factors that affect it. It should read: An ecosystem here is considered as part of ecology within a defined pluviometric regime consisting of the environment (with its range of habitats), its living parts, and the non-living factors that affect it. 2. Page 4 of the current pdf, first column, first paragraph, line 12. It reads: in this case lacustrine and alluvial habitats). It should read: (most of the time, alluvial habitats). 3. Page 7 of current pdf. First column, 4th paragraph. Here, the reference to Figure 2 is incorrect as it refers to a figure that was removed during revisions. This figure 2 does not exist in this paper. 4. Table 1 caption should be modified to: Minimum number of individuals (MNI) according to taxa reported in the Rwenzori National park for the whole ecosystem, the alluvial portion of the ecosystem and the thaphocoenosis documented in the latter. 5. Table 2 caption should be modified to: Number of animals collected throughout the alluvial woodland habitats sampled areas 1–6 in the Tarangire Park by Lamprey (1964). The areas sampled are: Riverine grassland, Acacia tortilis woodland, Gall Acacia woodland, drainage line woodland, Acacia-Commiphora woodland, Acacia mellifera-Commiphora woodland. 6. Figure 2 caption. The first sentence should be expanded as follows: Hierarchical agglomerative cluster analyses of several African national parks representing wooded biomes and more open savanna landscapes compared to two Pliocene hominin localities in Ethiopia. Data on bovid tribe.....