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Diet provides critical information about the ecology and environment of herbivores. Hence, understanding the dietary strategies of fossil herbivores and the associated temporal changes is one aspect of inferring paleoenvironmental conditions. Here, we present carbon isotope data from more than 1,050 fossil teeth that record the dietary patterns of nine herbivore families in the late Pliocene and early Pleistocene (3.6 to 1.05 Ma) from the Shungura Formation, a hominin-bearing site in southwestern Ethiopia. An increasing trend toward C(4) herbivory has been observed with attendant reductions in the proportions of browsers and mixed feeders through time. A high proportion of mixed feeders has been observed prior to 2.9 Ma followed by a decrease in the proportion of mixed feeders and an increase in grazers between 2.7 and 1.9 Ma, and a further increase in the proportion of grazers after 1.9 Ma. The collective herbivore fauna shows two major change points in carbon isotope values at ∼2.7 and ∼2.0 Ma. While hominin fossils from the sequence older than 2.7 Ma are attributed to Australopithecus, the shift at ∼2.7 Ma indicating the expansion of C(4) grasses on the landscape was concurrent with the first appearance of Paranthropus. The link between the increased C(4) herbivory and more open landscapes suggests that Australopithecus lived in more wooded landscapes compared to later hominins such as Paranthropus and Homo, and has implications for key morphological and behavioral adaptations in our lineage.

The East African Rift System (EARS) has played a central role in our understanding of human origins and vertebrate evolution in the late Cenozoic of Africa. However, the distribution of fossil sites along the rift is highly biased towards its northern extent, and the types of paleoenvironments are primarily restricted to fluvial and lacustrine settings. Here we report the discovery of the first fossil sites from the Urema Rift at Gorongosa National Park (central Mozambique) at the southern end of the EARS, and reconstruct environmental contexts of the fossils. In situ and surface fossils from the lower member of the Mazamba Formation, estimated to be of Miocene age, comprise mammals, reptiles, fishes, invertebrates, palms, and dicot trees. Fossil and geological evidence indicates a coastal-plain paleoenvironmental mosaic of riverine forest/woodland and estuarine habitats that represent the first coastal biomes identified in the Neogene EARS context. Receiving continental sediment from source terranes west of today's Urema Graben, estuarine sequences accumulated prior to rifting as compound incised-valley fills on a low-gradient coastal plain following transgression. Modern environmental analogues are extremely productive habitats for marine and terrestrial fauna, including primates. Thus, our discoveries raise the possibility that the Miocene coastal landscapes of Gorongosa were ecologically-favorable habitats for primates, providing relatively stable maritime climate and ecosystem conditions, year-round freshwater availability, and food both from terrestrial and marine sources. The emerging fossil record from Gorongosa is beginning to fill an important gap in the paleobiogeography of Africa as no fossil sites of Neogene age have previously been reported from the southernmost part of the EARS. Furthermore, this unique window into past continental-margin ecosystems of central Mozambique may allow us to test key paleobiogeographic hypotheses during critical periods of primate evolution. National Geographic SocietyNational Geographic Society [GEFNE169-16] John Fell Fund, University of Oxford [153/086] Philip Leverhulme Prize [PLP-2016-114] Foundation for Science and Technology (FCT-Portugal) DFGGerman Research Foundation (DFG) [LU 2199/1-1] St. Hugh's College, University of Oxford

New approaches to the study of early hominin diets have refreshed interest in how and when our diets diverged from those of other African apes. A trend toward significant consumption of C(4) foods in hominins after this divergence has emerged as a landmark event in human evolution, with direct evidence provided by stable carbon isotope studies. In this study, we report on detailed carbon isotopic evidence from the hominin fossil record of the Shungura and Usno Formations, Lower Omo Valley, Ethiopia, which elucidates the patterns of C(4) dietary utilization in the robust hominin Paranthropus. The results show that the most important shift toward C(4) foods occurred at ∼2.37 Ma, within the temporal range of the earliest known member of the genus, Paranthropus aethiopicus, and that this shift was not unique to Paranthropus but occurred in all hominins from this fossil sequence. This uptake of C(4) foods by hominins occurred during a period marked by an overall trend toward increased C(4) grazing by cooccurring mammalian taxa from the same sequence. However, the timing and geographic patterns of hominin diets in this region differ from those observed elsewhere in the same basin, where environmental controls on the underlying availability of various food sources were likely quite different. These results highlight the complexities of dietary responses by hominins to changes in the availability of food resources.

International audience; To better understand the environmental conditions that prevailed in the Plio-Pleistocene Shungura Formation, Lower Omo Valley, Ethiopia, we analyze the feeding preferences of Tragelaphini, the third most common tribe of Shungura bovids. Molar mesowear and dental microwear texture analyses were applied to three species (Tragelaphus rastafari, T. nakuae, and T. gaudryi) and body mass estimates were calculated for the T. rastafari-nakuae lineage to test whether dietary shifts were linked to body mass changes. We compare our results with previous work on stable carbon isotopes (δ13C) of enamel, which indicated that tragelaphins from the Shungura Formation possessed dietary flexibility. We found that the both the T. rastafari-nakuae lineage and T. gaudryi maintained a mixed feeding dietary niche, with varying proportions of C3 versus C4 inputs, from ~3.6 to >2 Ma. Our results show that T. rastafari consumed more browse than its descendant, T. nakuae, which was a mixed feeder consuming a greater proportion of C4 dicots ~ 2.8 Ma. The T. rastafari-T. nakuae dietary shift may reflect environmental changes in the Shungura Formation during this time, but appears to be offset from body mass increases in this lineage, which occurred gradually through the Plio-Pleistocene. This study highlights the importance of a multi-proxy approach to precisely determine the dietary ecologies of extinct bovids and points to how each proxy offers a slightly different perspective on the ecology of fossil organisms.

Preservation of the stable carbon isotopic composition of fossil tooth enamel enables us to estimate the relative proportion of C3 versus C4 vegetation in an animal's diet, which, combined with analysis of faunal abundance, may provide complementary methods of paleoenvironmental reconstruction. To this end, we analyzed stable carbon isotopic composition (δ(13)C values) of tooth enamel from four bovid tribes (Tragelaphini, Aepycerotini, Reduncini, and Alcelaphini) derived from six members of the Shungura Formation (Members B, C, D, F, G, and L; ages from ca. 2.90-1.05 Ma (millions of years ago) in the Lower Omo Valley of southwestern Ethiopia. The bovids show a wide range of δ(13)C values within taxa and stratigraphic members, as well as temporal changes in the feeding strategies of taxa analyzed throughout the middle to late Pliocene and early Pleistocene. Such variation suggests that the use of actualistic approaches for paleoenvironmental reconstruction may not always be warranted. Alcelaphini was the only taxon analyzed that retained a consistent dietary preference throughout the sequence, with entirely C4-dominated diets. Reduncini had a mixed C3/C4 to C4-dominated diet prior to 2.4 Ma, after which this taxon shifted to a largely C4-dominated diet. Aepycerotini generally showed a mixed C3/C4 diet, with a period of increased C4 diet from 2.5 to 2.3 Ma. Tragelaphini showed a range of mixed C3/C4 diets, with a median value that was briefly nearer the C4 end member from 2.9 to 2.4 Ma but was otherwise towards the C3 end member. These isotopic results, combined with relative abundance data for these bovids, imply that the environment of the Lower Omo Valley consisted of a mosaic of closed woodlands, with riverine forests and open grasslands. However, our data also signify that the overall environment gradually became more open, and that C4 grasses became more dominant. Finally, these results help document the range and extent of environments and potential diets that were available to the four hominin species encountered in the Shungura sequence.

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

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., 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 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

Australopithecus afarensis The samples attributed to Au. afarensis from Hadar, Laetoli, the Middle Awash, Woranso-Mille and Dikika were utilized. For this taxon we relied upon published reports (Johanson et al., 1982; Kimbel et al., 2004; Drapeau et al., 2005; Alemseged et al., 2006; Haile-Selassie et al., 2010; Ward et al., 2012), in addition to our own observations on original fossils and casts., 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

Homo floresiensis Specimens from Liang Bua, Flores as described by Brown et al., 2004; Morwood et al., 2005, Jungers et al., 2009a, Jungers et al., 2009b, and Falk et al., 2005 were included in 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 27, 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

Homo habilis Samples from Olduvai Gorge, East Lake Turkana, the Omo Shungura sequence, Hadar, and Sterkfontein were included in this study. Original Olduvai Gorge and East Lake Turkana fossils were examined first-hand, while for the Omo and Hadar materials we relied on our original observations on casts and originals and published reports (Boaz and Howell, 1977; Tobias, 1991; Kimbel et al., 1997). We include the following fossils in the hypodigm of H. habilis: A.L. 666-1, KNM-ER 1478, KNM- ER 1501, KNM-ER 1502, KNM-ER 1805, KNM-ER 1813, KNM-ER 3735, OH 4, OH 6, OH 7, OH 8, OH 13, OH 15, OH 16, OH 21, OH 24, OH 27, OH 31, OH 35, OH 37, OH 39, OH 42, OH 44, OH 45, OH 62, OMO-L894-1, and Stw 53. We recognize that some authors (including some of the authors of this paper) prefer to classify OH 62, Stw 53 and A.L. 666-1 outside of H. habilis, (e.g., as Homo gautengensis which we do not recognize as valid), or even outside the genus Homo; these specimens expand the morphological and temporal variability encompassed within H. habilis., 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.