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Submitted by Leda Lopes (ledacplopes@hotmail.com) on 2022-06-02T18:38:37Z No. of bitstreams: 2 license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Tamires_Carolina_Campos_Dissertação.pdf: 16548320 bytes, checksum: 6457da3e9d937e5f9a5b8b32808e25ea (MD5) Approved for entry into archive by Aline Batista (alinehb.ufpel@gmail.com) on 2022-06-06T17:53:24Z (GMT) No. of bitstreams: 2 Tamires_Carolina_Campos_Dissertação.pdf: 16548320 bytes, checksum: 6457da3e9d937e5f9a5b8b32808e25ea (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Made available in DSpace on 2022-06-06T17:53:24Z (GMT). No. of bitstreams: 2 Tamires_Carolina_Campos_Dissertação.pdf: 16548320 bytes, checksum: 6457da3e9d937e5f9a5b8b32808e25ea (MD5) license_rdf: 0 bytes, checksum: d41d8cd98f00b204e9800998ecf8427e (MD5) Previous issue date: 2021-10-11 Desde a sua divulgação, em 1964, a espécie H. habilis, vem sofrendo com diversas críticas quanto ao seu lugar dentro da linhagem humana. Sua classificação, desencadeou dentro da Paleoantropologia, uma série de debates quanto a origem, e definição, do gênero Homo. Através da compilação de dados métricos da morfologia hominínia, retirados de diferentes publicações e planilhas disponíveis na web. Buscou-se realizar diferentes análises, a partir de uma abordagem fenético-comparativa, onde os dados métricos da morfologia de crânios, dentes e membros, foram transformados em matrizes de distância morfológica, e correlacionados, através de teste de Mantel, com matrizes geográficas e filogenéticas, a fim de testar sua validade. Como resultado, as matrizes de membros demonstraram as maiores correlações, sendo seguidas pelas matrizes de dentes mandibulares. Ambas relacionadas as matrizes filogenéticas. Esse resultado demonstrou, que os dados de membros e dentes mandibulares, são bons para a realização de inferências filogenéticas. Em contrapartida, as matrizes cranianas mostraram maior correlação com as matrizes geográficas. Em conclusão, todos os dendrogramas gerados, que apontaram maiores e significativas correlações (r ≥ 0,5 e p ≤ 0,05), evidenciaram relações diretas, entre H. habilis e H. rudolfensis, com membros de australopitecíneos. Através dos resultados obtidos, sugere-se que ambos os hominínios sejam reclassificados como pertencentes ao gênero Australopithecus. Since its disclosure in 1964, the species H. habilis has been suffering from several criticisms as to its place within the human lineage. Its classification, within Paleoanthropology, triggered a series of debates regarding the origin, and definition, of the genus Homo. Through the compilation of metric data of hominid morphology, taken from different publications and spreadsheets available on the web. We sought to carry out different analyses, from a pheneticcomparative approach, where the metric data of the morphology of skulls, teeth and limbs were transformed into morphological distance matrices, and correlated, through the Mantel test, with geographic and phylogenetic matrices in order to test their validity. As a result, limb matrices showed the highest correlations, followed by mandibular tooth matrices. Both related to phylogenetic matrices. This result showed that data from limbs and mandibular teeth are good for making phylogenetic inferences. On the other hand, cranial matrices showed greater correlation with geographic matrices, which shows their greater response to phenotypic plasticity. In conclusion, all the dendrograms generated, which showed higher and significant correlations (r ≥ 0.5 and p ≤ 0.05), showed direct relationships between H. habilis and H. rudolfensis, with members of australopithecines. Through the results obtained, it is suggested that both hominins be reclassified as belonging to the genus Australopithecus.

Temeljno pitanje kojim se ovaj rad bavio je mogu li se strategije preživljavanja i pribavljanje hrane kod vrsta Homo habilis i Homo rudolfensis gledati kao iskazi organiziranog i planiranog ponašanja ili spontanosti. Također su se istraživale razlike u strategijama preživljavanja između dvije spomenute vrste, kao i razlike između ranih pripadnika roda Homo i australopitecina. To je učinjeno na temelju analize sljedeća dva kriterija: funkcionalnosti i primjenjivosti oruđa olduvajske kulture te kriterija zasnovanog na međuvezi anatomije hominina i prehrane. Na temelju provedene analize došlo se do zaključka da je postojao neki oblik organiziranog ponašanja kod ranih pripadnika roda Homo koji se manifestirao u pogledu znanja, planiranja i provođenja spomenutih strategija. Primjera radi, olduvajska litika je izrađivana s predumišljajem, znanjem i sposobnošću za proizvodnju primjenjivog oruđa, dok su obrađivane vrste posjedovale niz anatomskih i bioloških preduvjeta koji su omogućavali organizirano uspješno preživljavanje i prikupljanje hrane.

The newly described partial skeleton of Paranthropus boisei KNM-ER 47000 as well as the FwJj14E Ileret footprints provide new evidence on the paleobiology and diversity of hominins from the Okote Member of the Koobi Fora Formation at East Turkana about 1.5 Ma. To better understand the ecological context of the Okote hominins, it is necessary to broaden the geographical focus of the analysis to include the entire Omo-Turkana ecosystem, and the temporal focus to encompass the early Pleistocene. Previous work has shown that important changes in the regional vegetation occurred after 2 Ma, and that there was a peak in mammalian turnover and diversity close to 1.8 Ma. This peak in diversity included the Hominini, with the species P. boisei, Homo habilis, Homo rudolfensis, and Homo erectus co-occurring at around 1.8 Ma. There is considerable debate about whether H. habilis and H. rudolfensis indeed constitute separate species, but even if we consider them both as H. habilis sensu lato, the co-occurrence of three hominin species at any one time and place is rather unusually high diversity for hominin standards (even if not so for other mammalian groups such as suids, bovids, or cercopithecids). Here we use mammalian faunal abundance data to place confidence intervals on first and last appearances of hominin species in the early Pleistocene of the Omo-Turkana Basin, and use these estimates to discuss hominin diversity in the Okote Member. We suggest that in the early Pleistocene a wide range of depositional environments and vegetation types, along with a high frequency of volcanism, likely maintained high levels of environmental variability both in time and space across the Omo-Turkana region, and provided ecological opportunities for the coexistence of at least three hominin species alongside a diverse mammalian fauna. (C) 2018 Elsevier Ltd. All rights reserved. National Science FoundationNational Science Foundation (NSF) [BCS-0137235]

The craniomandibular morphology of Homo naledi shows variable resemblances with species across Homo, which confounds an easy assessment of its phylogenetic position. In terms of skull shape, H. naledi has its closest affinities with Homo erectus, while mandibular shape places it closer to early Homo. From a tooth crown perspective, the smaller molars of H. naledi make it distinct from early Homo and H. erectus. Here, we compare the mandibular molar root morphology of six H. naledi individuals from the Dinaledi Chamber to those of African and Eurasian Plio-Pleistocene fossil hominins (totalling 183 mandibular first, second and third molars). The analysis of five root metric variables (cervical plane area, root length, root cervix volume, root branch volume, and root surface area) derived from microCT reconstructions reveals that the molar roots of H. naledi are smaller than those of Homo habilis, Homo rudolfensis, and H. erectus, but that they resemble those of three Homo sp. specimens from Swartkrans and Koobi Fora in size and overall appearance. Moreover, though H. naledi molar roots are similar in absolute size to Pleistocene Homo sapiens, they differ from H. sapiens in having a larger root volume for a given cervical plane area and less taurodont roots; the root cervix-to-branch proportions of H. naledi are comparable to those of Australopithecus africanus and species of Paranthropus. H. naledi also shares a metameric root volume pattern (M2 > M3 > M1) with Australopithecus and Paranthropus but not with any of the other Homo species (M2 > M1 > M3). Our findings therefore concur with previous studies that found that H. naledi shares plesiomorphic features with early Homo, Australopithecus, and Paranthropus. While absolute molar root size aligns H. naledi with Homo from North and South Africa, it is distinguishable from these in terms of root volumetric proportions.

Herpes simplex virus 2 (HSV2) is a human herpesvirus found worldwide that causes genital lesions and more rarely causes encephalitis. This pathogen is most common in Africa, and particularly in central and east Africa, an area of particular significance for the evolution of modern humans. Unlike HSV1, HSV2 has not simply co-speciated with humans from their last common ancestor with primates. HSV2 jumped the species barrier between 1.4 and 3 MYA, most likely through intermediate but unknown hominin species. In this article, we use probability-based network analysis to determine the most probable transmission path between intermediate hosts of HSV2, from the ancestors of chimpanzees to the ancestors of modern humans, using paleo-environmental data on the distribution of African tropical rainforest over the last 3 million years and data on the age and distribution of fossil species of hominin present in Africa between 1.4 and 3 MYA. Our model identifies Paranthropus boisei as the most likely intermediate host of HSV2, while Homo habilis may also have played a role in the initial transmission of HSV2 from the ancestors of chimpanzees to P.boisei., SJU was funded by Oxford Brookes University. KK and CH were funded by the University of Cambridge. KK is a college research associate at King’s College, Cambridge. CH is a post-doctoral affiliate at Christ’s College, Cambridge.

Recent fossil finds have highlighted extensive morphological diversity within our genus, Homo, and the co-existence of a number of species. However, little is known about the evolutionary processes responsible for producing this diversity. Understanding the action of these processes can provide insight into how and why our lineage evolved and diversified. Here, we examine cranial and mandibular variation and diversification from the earliest emergence of our genus at 2.8 Ma until the Late Pleistocene (0.126-0.0117 Ma), using statistical tests developed from quantitative genetics theory to evaluate whether stochastic (genetic drift) versus non-stochastic (selection) processes were responsible for the observed variation. Results show that random processes can account for species diversification for most traits, including neurocranial diversification, and across all time periods. Where selection was found to shape diversification, we show that: 1) adaptation was important in the earliest migration of Homo out of Africa; 2) selection played a role in shaping mandibular and maxillary diversity among Homo groups, possibly due to dietary differences; and 3) Homo rudolfensis is adaptively different from other early Homo taxa, including the earliest known Homo specimen. These results show that genetic drift, and likely small population sizes, were important factors shaping the evolution of Homo and many of its novel traits, but that selection played an essential role in driving adaptation to new contexts.

Mankind is thought to have separated from a common ancestor shared with chimpanzees approximately 7 million years ago.1 At the time, among our ancestors was a group that decided to come down from the trees to live in the savanna as desertification of Africa's forests progressed. Although the term “ape-man” was used for early mankind who walked on two feet, improvements in the precision of dating methods and excavations from the latter half of the 20th century made it clear that many types of human species coexisted. Thus, the term “ape-man” is now only rarely used. Bipedalism was likely something attempted by many groups, and although many theories exist, the underlying reason for this transition is unclear.2 However, the advantages are clear – it enabled viewing vast distances and the use of hands to carry food, and ultimately led to the expansion of brain size. Interestingly, it took our ancestors an additional 4 million years to expand their brain size. This evolutionary delay might have been caused by climate changes in Africa. The progressively drying climate led to reductions in forestland, and our ancestors who lived on the plains were subjected to a new type of natural selection pressure, causing changes in their physique and behavior. One possible reason that orangutans remained in the trees when transported to Asia by continental drift was that although Africa and the Northern hemisphere began to dry and cool from approximately 30 million to 2.5 million years ago, Asia's forests in the tropical regions remained intact. Evidence of brain size expansion and the use of tools can be found from approximately 2.5 million years ago. Mankind at this stage is referred to as Homo habilis or Homo rudolfensis. The concept of pre-adaptation refers to gradually altering physique in advance so that it works to one's advantage once subjected to a particular environment. For instance, the advantages of walking erect are thought to have played an important role in the evolution of mankind much later; that is, approximately 2.5 million years later. The expansion of brain size also brought about a revolution in food energy, which accompanied the use of stone tools – the ability to ingest high caloric and nutritious meat products. Although they were “meat-eating” and hunted herbivorous animals on occasion, their meat supply likely came from food left over by carnivorous animals. In addition to increasing meat-eating efficiency, the expansion of brain size was accompanied by acquisition of modern man's physique, an expanded range of activity and loss of body hair. Although many human species remained in Africa, one group, the Peking man or Java man (also known as Homo erectus) moved to Eurasia approximately 1.8 million years ago in an event referred to as the first African exodus. At least one group of Homo erectus began using fire, a monumental event in mankind's history. Homo erectus existed until 250 000 years ago. We enter the age of modern man from approximately 200 000 years ago. The Neanderthals were another species of man that coexisted with modern humans during this period. This period saw the use of symbols, language and original art-forms. One group of modern humans left Africa approximately 100 000 years ago, and eventually adapted to all global environments in an event famously known as the second African exodus. Many terrains were conquered, including grasslands, deserts and mountains, and humans began living under various climates (tropical, temperate and polar). The conquering of various terrains is evidenced by the existence of Eskimos in polar regions, Bushmen in the deserts, Pygmies in the tropical rain forests, and Tibetan highlanders. The spreading of humans was particularly fast, with humans reaching Europe, Asia, North and South America, and Australia within the span of tens of thousands of years. Mankind at the time formed small nomadic groups and lived as hunter-gatherers. From a modest population of approximately 100 000 about 1 million years ago, the global population dramatically increased to 500 000 about 200 000 years ago, and further to approximately 10 million with the advent of agriculture about 10 000 years ago.3

A Heady Find In the past two decades, excavations at the archaeological site at Dmanisi, Georgia, have revealed hominin fossils from the earliest Pleistocene, soon after the genus Homo first dispersed beyond Africa. Lordkipanidze et al. (p. 326 ; see the cover) now describe a fossil cranium from the site. Combined with mandibular remains that had been found earlier, this find completes the first entire hominin skull from this period.

Homo erectus and later humans have enlarged body sizes, reduced sexual dimorphism, elongated lower limbs, and increased encephalization compared to Australopithecus , together suggesting a distinct ecological pattern. The mosaic expression of such features in early Homo , including Homo habilis , Homo rudolfensis , and some early H. erectus , suggests that these traits do not constitute an integrated package. We examined the evidence for body mass, stature, limb proportions, body size and dental size dimorphism, and absolute and relative brain size in Homo naledi as represented in the Dinaledi Chamber sample. H. naledi stature and body mass are low compared to reported values for H. erectus , with the exception of some of the smaller bodied Dmanisi H. erectus specimens, and overlap with larger Australopithecus and early Homo estimates . H. naledi endocranial volumes (465–560 cc) and estimates of encephalization quotient are also similar to Australopithecus and low compared to all Homo specimens, with the exception of Homo floresiensis (LB1) and the smallest Dmanisi H. erectus specimen (D4500). Unlike Australopithecus , but similar to derived members of genus Homo , the Dinaledi assemblage of H. naledi exhibits both low levels of body mass and dental size variation, with an estimated body mass index of sexual dimorphism less than 20%, and appears to have an elongated lower limb. Thus, the H. naledi bauplan combines features not typically seen in Homo species (e.g., small brains and bodies) with those characteristic of H. erectus and more recent Homo species (e.g., reduced mass dimorphism, elongated lower limb).

Since its discovery in 1972 (ref. 1), the cranium KNM-ER 1470 has been at the centre of the debate over the number of species of early Homo present in the early Pleistocene epoch of eastern Africa. KNM-ER 1470 stands out among other specimens attributed to early Homo because of its larger size, and its flat and subnasally orthognathic face with anteriorly placed maxillary zygomatic roots. This singular morphology and the incomplete preservation of the fossil have led to different views as to whether KNM-ER 1470 can be accommodated within a single species of early Homo that is highly variable because of sexual, geographical and temporal factors, or whether it provides evidence of species diversity marked by differences in cranial size and facial or masticatory adaptation. Here we report on three newly discovered fossils, aged between 1.78 and 1.95 million years (Myr) old, that clarify the anatomy and taxonomic status of KNM-ER 1470. KNM-ER 62000, a well-preserved face of a late juvenile hominin, closely resembles KNM-ER 1470 but is notably smaller. It preserves previously unknown morphology, including moderately sized, mesiodistally long postcanine teeth. The nearly complete mandible KNM-ER 60000 and mandibular fragment KNM-ER 62003 have a dental arcade that is short anteroposteriorly and flat across the front, with small incisors; these features are consistent with the arcade morphology of KNM-ER 1470 and KNM-ER 62000. The new fossils confirm the presence of two contemporary species of early Homo, in addition to Homo erectus, in the early Pleistocene of eastern Africa.

We review the fossil and genetic evidence that relate to evolution in the genus Homo. We focus on the origin of Homo and on the evidence for taxonomic diversity at the beginning of the evolutionary history of Homo and in the last 200,000 years. We set out the arguments for recognizing a second early Homo taxon, Homo rudolfensis, and the arguments for and against including Homo habilis sensu stricto and Homo rudolfensis within Homo. We end by reviewing recent genomic evolution within Homo. The challenge of the upcoming decades is to meld innovations in molecular genetic methods and technology with evidence from the fossil record to generate hypotheses about the developmental bases of the phenotypic and behavioral developments we see within the genus Homo.

The origin of the genus Homo in Africa signals the beginning of the shift from increasingly bipedal apes to primitive, large-brained, stone tool-making, meat-eaters that traveled far and wide. This early part of the human genus is represented by three species: Homo habilis, Homo rudolfensis, and Homo erectus. H. habilis is known for retaining primitive features that link it to australopiths and for being the first stone tool makers. Little is known about H. rudolfensis except that it had a relatively large brain and large teeth compared to H. habilis and that it overlapped in time and space with other early Homo. Our understanding of the paleobiology and evolution of the larger-brained H. erectus is enhanced due to its rich fossil record. H. erectus was the first obligate, fully committed biped, and with a body adapted for modern striding locomotion, it was also the first in the human lineage to disperse outside of Africa. The early members of the genus Homo are the first to tip the scale from the more apish side of our evolutionary history toward the more human one.

Hominin systematics, encompassing both taxonomy and phylogeny (Strait, 2013), has significant implications for how the evolution of species and traits are understood and communicated. Following a recent influx of fossils (e.g., Brown et al., 2004; Lordkipanidze et al., 2013; Villmoare et al., 2015a; Berger et al., 2015) the amount of diversity in fossil morphology has increased correspondingly. As researchers do not yet approach diversity in a uniform manner, the literature has been flooded with conflicting theories and methodologies (Strait, 2013). Particularly volatile has been the study of the origin of the genus Homo and the extent of variation therein: much controversy arises from conflicting views of the number of valid species subsumed within ‘early Homo’ given unspecified definitions of species and genera. Additionally, there is still a lack of understanding of the extent of and mechanism behind variation, especially within Hominina. The first section of the following paper addresses ‘how can species be identified?’ and ‘how should species be classified into higher taxa?’ The second section reviews the prevalent arguments used to systematise fossils frequently classified as ‘early Homo.’ It considers: the validity of Homo rudolfensis; the morphological, spatial & temporal overlap of earlier Homo with Homo ergaster; the systematic significance of the recently discovered LD 350-1; and finally, the appropriateness of ‘early Homo’ as an adaptive grade.

The Plio-Pleistocene site of Dmanisi, Georgia, has yielded a rich fossil and archaeological record documenting an early presence of the genus Homo outside Africa. Although the craniomandibular morphology of early Homo is well known as a result of finds from Dmanisi and African localities, data about its postcranial morphology are still relatively scarce. Here we describe newly excavated postcranial material from Dmanisi comprising a partial skeleton of an adolescent individual, associated with skull D2700/D2735, and the remains from three adult individuals. This material shows that the postcranial anatomy of the Dmanisi hominins has a surprising mosaic of primitive and derived features. The primitive features include a small body size, a low encephalization quotient and absence of humeral torsion; the derived features include modern-human-like body proportions and lower limb morphology indicative of the capability for long-distance travel. Thus, the earliest known hominins to have lived outside of Africa in the temperate zones of Eurasia did not yet display the full set of derived skeletal features.

Sites in eastern Africa have shed light on the emergence and early evolution of the genus Homo. The best known early hominin species, H. habilis and H. erectus, have often been interpreted as time-successive segments of a single anagenetic evolutionary lineage. The case for this was strengthened by the discovery of small early Pleistocene hominin crania from Dmanisi in Georgia that apparently provide evidence of morphological continuity between the two taxa. Here we describe two new cranial fossils from the Koobi Fora Formation, east of Lake Turkana in Kenya, that have bearing on the relationship between species of early Homo. A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. The discovery of a particularly small calvaria of H. erectus indicates that this taxon overlapped in size with H. habilis, and may have shown marked sexual dimorphism. The new fossils confirm the distinctiveness of H. habilis and H. erectus, independently of overall cranial size, and suggest that these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years.

The Quaternary, which is sometimes also termed the Anthropogene Period, often is linked to the beginning of humankind. But what exactly is the basis in the fossil hominin record for this link, in particular what is the early Homo evidence? Since 1999, debate concerning the definition of the genus Homo and the identification of its first members has intensified. At the centre of recent debates, there are four hypotheses: (1) all early Homo specimens belong to the same species: Homo habilis sensu lato; (2) the early Homo hypodigm is heterogeneous, two species could be defined: Homo habilis sensu stricto and Homo rudolfensis; (3) these species do not belong to the genus Homo but to the genus Australopithecus; or (4) it would be more appropriate to put the specimens of Homo rudolfensis into the genus Kenyanthropus. The earliest appearance of the genus Homo will differ depending upon the favoured hypothesis: 2.45 Myr (hypotheses 1 and 2); 1.9 Myr (hypothesis 3); 1.9 or 2.4 Myr (hypothesis 4, depending on the genus attribution for habilis (2.4 Myr if Homo; 1.9 Myr if Australopithecus)). Our study, based on palaeoanthropological data, is focused on the 2.6-1.6 Myr period. In this paper, we propose to review and assess knowledge of early Homo taxonomy in the context of the definition of the Quaternary boundary.

The nature of the human fossil record is less than ideal for the generation of precise correlations between environmental variables and patterns of evolution in specific lineages. Nonetheless, a critical look at what can and cannot be said from individual fossil morphology and the correlation of specific environmental proxies with specific hominin fossils may lead to a greater understanding of the degree of certainty with which we should embrace environmental hypotheses for the evolution of Homo. Climate shifts have been implicated in both the origin of the genus and its dispersal from Africa. Here, I consider three areas in which a climatic influence has been posited to explain evolutionary shifts in the genus Homo : the origin and dispersal of the genus from Africa; geography, climate and body size in early Homo, and the influence of climate-induced sea level rise on morphological isolation in H. erectus. Each of the data sets is far from ideal, and interpretations of each of the data sets are fraught with issues of equifinality. Of the three hypotheses discussed, the clearest link is seen between latitudinal variation (and presumably temperature) and body size in H. erectus. Similarly, climate-induced sea level change seems a reasonable isolating mechanism to explain the pattern of cranial variation in later Asian H. erectus, but the distribution could also reflect incompletely sampled clinal variation. Alternatively, only equivocal support is found for the influence of climate on the differentiation of H. erectus from H. habilis (as proxied by body/brain size scaling), and therefore the dispersal of the genus Homo cannot be as clearly linked to changes in body size and shape as it has been in the past. These preliminary data suggest that an emphasis on understanding local adaptation before looking at global (and specific) level change is critical to elucidating the importance of climatic factors on the evolution of the genus Homo .

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

A fifth hominin skull (cranium D4500 and mandible D2600) from Dmanisi is massively constructed, with a large face and a very small brain. Traits documented for the first time in a basal member of the Homo clade include the uniquely low ratio of endocranial volume to basicranial width, reduced vertex height, angular vault profile, smooth nasal sill coupled with a long and sloping maxillary clivus, elongated palate, and tall mandibular corpus. The convex clivus and receding symphysis of skull 5 produce a muzzle-like form similar to that of Australopithecus afarensis. While the Dmanisi cranium is very robust, differing from OH 13, OH 24, and KNM-ER 1813, it resembles Homo habilis specimens in the “squared off” outline of its maxilla in facial view, maxillary sulcus, rounded and receding zygomatic arch, and flexed zygomaticoalveolar pillar. These characters distinguish early Homo from species of Australopithecus and Paranthropus. Skull 5 is unlike Homo rudolfensis cranium KNM-ER 1470. Although it appears generally primitive, skull 5 possesses a bar-like supraorbital torus, elongated temporal squama, occipital transverse torus, and petrotympanic traits considered to be derived for Homo erectus. As a group, the Dmanisi crania and mandibles display substantial anatomical and metric variation. A key question is whether the fossils document age-related growth and sex dimorphism within a single population, or whether two (or more) distinct taxa may be present at the site. We use the coefficient of variation to compare Dmanisi with Paranthropus boisei, H. erectus, and recent Homo sapiens, finding few signals that the Dmanisi sample is excessively variable in comparison to these reference taxa. Using cranial measurements and principal components analysis, we explore the proposal that the Dmanisi skulls can be grouped within a regionally diverse hypodigm for H. erectus. Our results provide only weak support for this hypothesis. Finally, we consider all available morphological and paleobiological evidence in an attempt to clarify the phyletic relationship of Dmanisi to Homo species evolving >2.0 to 1.0 Ma.

© 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.

Recent geologic study shows that all hominins and nearly all other published mammalian fossils from Paleontological Collection Area 123, Koobi Fora, Kenya, derive from levels between the KBS Tuff (1.87+/-0.02 Ma) and the Lower Ileret Tuff (1.53+/-0.01 Ma). More specifically, the fossils derive from 53 m of section below the Lower Ileret Tuff, an interval in which beds vary markedly laterally, especially those units containing molluscs and algal stromatolites. The upper Burgi Member (approximately 2.00-1.87 Ma) crops out only in the southwestern part of Area 123. Adjacent Area 110 contains larger exposures of the member, and there the KBS Tuff is preserved as an airfall ash in lacustrine deposits and also as a fluvially redeposited ash. We observed no mammalian fossils in situ in this member in Area 123, but surface specimens have been documented in some monographic treatments. Fossil hominins from Area 123 were attributed to strata above the KBS Tuff in the 1970s, but later they were assigned to strata below the KBS Tuff (now called the upper Burgi Member). This study definitively places the Area 123 hominins in the KBS Member. Most of these hominins are between 1.60 and 1.65 myr in age, but the youngest may date to only 1.53 Ma, and the oldest, to 1.75 Ma. All are 0.15-0.30 myr younger than previously estimated. The new age estimates, in conjunction with published taxonomic attributions of fossils, suggest that at least two species of Homo coexisted in the region along with A. boisei until at least 1.65 Ma. Comparison of crania KNM-ER 1813 and KNM-ER 1470, which were believed to be of comparable age, is at the focus of the debate over whether Homo habilis sensu lato is in fact composed of two species: Homo habilis and Homo rudolfensis. These two crania are separated in time by approximately 0.25 myr, and therefore, arguments for their conspecificity no longer need to confront the issue of unusually high contemporaneous variation within a single species.

An Update on Human Evolution Brain Dance: New Discoveries about Human Origins and Brain Evolution Dean Falk Revised and Expanded Edition, University Press of Florida 2004 This updated version of a popular book seeks to trace human evolution from pre-hominoid ancestors through to Homo sapiens, with prime reference to that supremely human characteristic, the human brain. It is useful to students coming to grips with palaeontology for the first time, or the curious, intelligent layman. It also contains up-to-date observations on the Out-of-Africa debate. Dean Falk, a professor of anthropology at Florida State University, begins by pointing out that at the time of the first (1992) edition of her text, "most paleontologists agreed that the first African Homo to appear was Homo habilis, and that this species gave rise to Homo erectus who in turn gave rise to Homo sapiens. Furthermore, at that time workers believed that the transition between Homo habilis and Homo erectus had occurred rapidly (some thought between 1.8 and 1.6 million years ago) and exclusively in Africa." However, since then, she observes, "the picture has become murkier." Classifying Hominin Fossils This is because, as more fossil evidence becomes available, paleontologists have been virtually overwhelmed by the variety. Remains that were originally grouped together as Homo habilis have been split into categories such as Homo habilis sensu styncto and Homo rudolfensis, and yet others reclassified as Australopithecines. In short, while classification is a vital step toward what may one day be a fairly accurate picture of hominoid and hominid evolution, the evidence is as yet too scanty to allow us to construct with any degree of certainty what happened among evolving and increasingly advanced varieties of intelligent primates over the past two or three million years. Paleontologists can be divided into "splitters and lumpers," those who tend to be affected by what Russ Tuttle has described as "splitomania," regarding almost every new fossil as representative of a different species, and those who are overly keen to see differences as purely idiosyncratic, and who therefore tend to lump a wide range of fossils into the same group. The latter argue that fossils that appear to be more or less contemporaneous should be regarded as belonging to the same species, and play down the possibility that different species of hominins might have been residing on the earth at the same time, even on the same continent. Yet there is sufficient evidence that different species of hominins did in fact inhabit the earth at various times. It is generally accepted that different species of Australopithecines lived in Africa at the same time, and more recently we have the evidence of Homo florensi, seemingly akin to Homo erectus, surviving in an age when essentially modern men had already populated many parts of the Old World. Falk's book is not only interestingly written, but attempts to introduce the reader to the wide variety of terminological inventions that we find in the literature. Terminology is a necessary step toward scientific classification but it is not itself bedrock science. The guesswork involved in much terminology is illustrated by the fact that is has recently been suggested that Homo habilis should have been classified as an Australopithicine rather than as Homo. Falk is acutely aware of the danger of classificatory excess, and asks: "can two fossils that are identical (not to mention contemporaneous) reasonably be assigned to two separate species? I do not think so," while at the same time warning that in cladistic analysis, a family tree is only as good as the traits the researcher selects when seeking to construct it. The Evolution of the Human Brain But the main purpose of Falk's book is to summarize what we now know about the growth of the human brain. It is now established that chimpanzees are the closest surviving relatives of Homo, and much work has been done to compare the workings of the chimpanzee brain with that of modern humans, in the hope that this may provide clues to the evolutionary history of the human brain. …

Currently, it is widely accepted that only one hominin genus, Homo, was present in Pleistocene Asia, represented by two species, Homo erectus and Homo sapiens. Both species are characterized by greater brain size, increased body height and smaller teeth relative to Pliocene Australopithecus in Africa. Here we report the discovery, from the Late Pleistocene of Flores, Indonesia, of an adult hominin with stature and endocranial volume approximating 1 m and 380 cm3, respectively--equal to the smallest-known australopithecines. The combination of primitive and derived features assigns this hominin to a new species, Homo floresiensis. The most likely explanation for its existence on Flores is long-term isolation, with subsequent endemic dwarfing, of an ancestral H. erectus population. Importantly, H. floresiensis shows that the genus Homo is morphologically more varied and flexible in its adaptive responses than previously thought.

New hominid teeth from the Kaitio member (1.65–1.9 Myr) in West Turkana (Kenya). New hominid teeth have been recovered from the archaeological sites of Kokiselei 1 and Naiyena Engol 1. These two sites are located in the west side of the Turkana Basin and belong to the Kaitio member of the Nachukui Formation. They are dated between 1.65-1.79 and 1.7-1.8 Myr respectively. The four teeth (left maxillary canine and first molar, right maxillary third molar and left mandibular third molar) discovered in Kokiselei 1 are attributed to Australopithecus boisei. The right mandibular first premolar found in Naiyena Engol 1 is referred to Homo sp. aff. ergaster. To cite this article: S. Prat et al., C. R. Palevol 2 (2003). © 2003 Academie des sciences. Publie par Elsevier SAS. Tous droits reserves. Mots cles : Hominides ; dents ; Australopithecus boisei ; Homo ; Kenya ; Pleistocene ancien

Excavation in the previously little-explored western portion of Olduvai Gorge indicates that hominid land use of the eastern paleobasin extended at least episodically to the west. Finds included a dentally complete Homo maxilla (OH 65) with lower face, Oldowan stone artifacts, and butchery-marked bones dated to be between 1.84 and 1.79 million years old. The hominid shows strong affinities to the KNM ER 1470 cranium from Kenya ( Homo rudolfensis ), a morphotype previously unrecognized at Olduvai. ER 1470 and OH 65 can be accommodated in the H. habilis holotype, casting doubt on H. rudolfensis as a biologically valid taxon.

The development of East African savannas is crucial for the origin and evolution of early hominins. These ecosystems, however, vary widely in their fraction of woody cover and today range from closed woodland to open grassland savanna. Here, we present the first Plio-Pleistocene long-term carbon isotope (δ13C) record from pedogenic carbonate and Suidae teeth in the southern East African Rift (EAR). These δ13C data from the Chiwondo and Chitimwe Beds (Karonga Basin, Northern Malawi) represent a southern hemisphere record in the EAR, a key region for reconstructing vegetation patterns in today's Zambezian Savanna, and permit correlation with data on the evolution and migration of early hominins in today's Somali-Masai Endemic Zone. The sediments along the northwestern shore of Lake Malawi contain fossils attributed to Homo rudolfensis and Paranthropus boisei. The associated hominin localities (Uraha, Malema) are situated between the well-known hominin bearing sites of the Somali-Masai Endemic Zone in the Eastern Rift and the Highveld Grassland in southern Africa, and fill an important geographical gap for hominin research. Persistent δ13C values around −9‰ from pedogenic carbonate and suid enamel covering the last ∼4.3 Ma indicate a C3-dominated closed environment with regional patches of C4-grasslands in the Karonga Basin. The overall fraction of woody cover of 60–70% reflects significantly higher canopy density in the Malawi Rift than the Eastern Rift through time. The discrepancy between the two savanna types originated in the Late Pliocene, when the Somali-Masai ecosystem started to show increasing evidence for open, C4-dominated landscapes. Based on the Malawi δ13C data, the evolution of savanna ecosystems in Eastern Africa followed different patterns along the north-south extent of the EAR. The appearance of C4-grasses is considered a driver of evolutionary faunal shifts, but despite the difference of ecosystem evolution in the north, similar hominins and suids occurred in both landscapes, pointing to distinct habitat flexibility and also nutritional versatility.

Four human remains: one mandible, two skulls and one metatarsus were discovered between 1991 and 1999 at the open-air site of Dmanisi, Georgia, in a precise stratigraphic, palaeontological and archaeological context, in volcanic ashes dated to 1.81 ± 0.05 Ma. The first studies of these fossils enable the authors to compare them with the morphology of archaic African Homo erectus , ascribed to Homo ergaster , and to ascertain hominid presence at the gates of Europe 300 000 years earlier than the classical scenario forecasted. In September 2000, the discovery of a second more complete and robust mandible D 2600 presents a threefold interest: palaeontological, functional and pathological. A comparison with Homo habilis and Homo erectus leads to the recognition of a new Homo species: H. georgicus sp. nov. The morphofunctional characteristics and the antiquity of H. georgicus characterise the root of a long Eurasian line.

The genesis, evolution and fate of Homo erectus have been explored palaeontologically since the taxon's recognition in the late nineteenth century. Current debate is focused on whether early representatives from Kenya and Georgia should be classified as a separate ancestral species ('H. ergaster'), and whether H. erectus was an exclusively Asian species lineage that went extinct. Lack of resolution of these issues has obscured the place of H. erectus in human evolution. A hominid calvaria and postcranial remains recently recovered from the Dakanihylo Member of the Bouri Formation, Middle Awash, Ethiopia, bear directly on these issues. These approximately 1.0-million-year (Myr)-old Pleistocene sediments contain abundant early Acheulean stone tools and a diverse vertebrate fauna that indicates a predominantly savannah environment. Here we report that the 'Daka' calvaria's metric and morphological attributes centre it firmly within H. erectus. Daka's resemblance to Asian counterparts indicates that the early African and Eurasian fossil hominids represent demes of a widespread palaeospecies. Daka's anatomical intermediacy between earlier and later African fossils provides evidence of evolutionary change. Its temporal and geographic position indicates that African H. erectus was the ancestor of Homo sapiens.

Remains of earlyHomo andParanthropus have been recovered from two contemporaneous sites (Uraha and Malema) in the “Hominid Corridor” in Northern Malawi (Chiwondo Beds). Faunal dating suggests an age of 2.5–2.3 Ma for both hominids. The two specimens, a mandible attributed toHomo rudolfensis (UR 501 from Uraha), and a maxillary fragment ofParanthropus boisci. (RC 911 from Malema) known only from eastern Africa, represent the southernmost known distribution of these taxa. The biogeographic significance of these hominids from the Malawi-Rift lay in their association with the eastern African endemic animal group. Biogeographic variation in south-eastern Africa may be linked to habitat change occurring due to climate change, with maximum change occurring around 2.5 Ma.