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      Characterizing the Evolutionary Path(s) to Early Homo

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          Abstract

          Numerous studies suggest that the transition from Australopithecus to Homo was characterized by evolutionary innovation, resulting in the emergence and coexistence of a diversity of forms. However, the evolutionary processes necessary to drive such a transition have not been examined. Here, we apply statistical tests developed from quantitative evolutionary theory to assess whether morphological differences among late australopith and early Homo species in Africa have been shaped by natural selection. Where selection is demonstrated, we identify aspects of morphology that were most likely under selective pressure, and determine the nature (type, rate) of that selection. Results demonstrate that selection must be invoked to explain an Au. africanusAu. sedibaHomo transition, while transitions from late australopiths to various early Homo species that exclude Au. sediba can be achieved through drift alone. Rate tests indicate that selection is largely directional, acting to rapidly differentiate these taxa. Reconstructions of patterns of directional selection needed to drive the Au. africanusAu. sedibaHomo transition suggest that selection would have affected all regions of the skull. These results may indicate that an evolutionary path to Homo without Au. sediba is the simpler path and/or provide evidence that this pathway involved more reliance on cultural adaptations to cope with environmental change.

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          Most cited references29

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          Generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution.

          Fossils and molecular data are two independent sources of information that should in principle provide consistent inferences of when evolutionary lineages diverged. Here we use an alternative approach to genetic inference of species split times in recent human and ape evolution that is independent of the fossil record. We first use genetic parentage information on a large number of wild chimpanzees and mountain gorillas to directly infer their average generation times. We then compare these generation time estimates with those of humans and apply recent estimates of the human mutation rate per generation to derive estimates of split times of great apes and humans that are independent of fossil calibration. We date the human-chimpanzee split to at least 7-8 million years and the population split between Neanderthals and modern humans to 400,000-800,000 y ago. This suggests that molecular divergence dates may not be in conflict with the attribution of 6- to 7-million-y-old fossils to the human lineage and 400,000-y-old fossils to the Neanderthal lineage.
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            Australopithecus sediba: a new species of Homo-like australopith from South Africa.

            Despite a rich African Plio-Pleistocene hominin fossil record, the ancestry of Homo and its relation to earlier australopithecines remain unresolved. Here we report on two partial skeletons with an age of 1.95 to 1.78 million years. The fossils were encased in cave deposits at the Malapa site in South Africa. The skeletons were found close together and are directly associated with craniodental remains. Together they represent a new species of Australopithecus that is probably descended from Australopithecus africanus. Combined craniodental and postcranial evidence demonstrates that this new species shares more derived features with early Homo than any other australopith species and thus might help reveal the ancestor of that genus.
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              The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme.

              An adaptationist programme has dominated evolutionary thought in England and the United States during the past 40 years. It is based on faith in the power of natural selection as an optimizing agent. It proceeds by breaking an oragnism into unitary 'traits' and proposing an adaptive story for each considered separately. Trade-offs among competing selective demands exert the only brake upon perfection; non-optimality is thereby rendered as a result of adaptation as well. We criticize this approach and attempt to reassert a competing notion (long popular in continental Europe) that organisms must be analysed as integrated wholes, with Baupläne so constrained by phyletic heritage, pathways of development and general architecture that the constraints themselves become more interesting and more important in delimiting pathways of change than the selective force that may mediate change when it occurs. We fault the adaptationist programme for its failure to distinguish current utility from reasons for origin (male tyrannosaurs may have used their diminutive front legs to titillate female partners, but this will not explain why they got so small); for its unwillingness to consider alternatives to adaptive stories; for its reliance upon plausibility alone as a criterion for accepting speculative tales; and for its failure to consider adequately such competing themes as random fixation of alleles, production of non-adaptive structures by developmental correlation with selected features (allometry, pleiotropy, material compensation, mechanically forced correlation), the separability of adaptation and selection, multiple adaptive peaks, and current utility as an epiphenomenon of non-adaptive structures. We support Darwin's own pluralistic approach to identifying the agents of evolutionary change.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2014
                3 December 2014
                : 9
                : 12
                : e114307
                Affiliations
                [1 ]Department of Archaeology, University of Cape Town, Rondebosch, 7701, South Africa
                [2 ]Department of Anthropology, University of Illinois, Urbana-Champaign, Illinois, United States of America
                [3 ]Department of Biology, Loyola University, Chicago, Illinois, United States of America
                Duke University School of Medicine, United States of America
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: LS RRA CCR JMC. Performed the experiments: LS. Analyzed the data: LS CCR. Wrote the paper: RRA LS.

                Article
                PONE-D-14-38513
                10.1371/journal.pone.0114307
                4255019
                25470780
                467bdd54-374a-4ae6-b0dd-16f306bdbce8
                Copyright @ 2014

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 29 August 2014
                : 7 November 2014
                Page count
                Pages: 20
                Funding
                Funding provided by the Palaeontological Scientific Trust (LS), the L.S.B. Leakey Foundation (Baldwin Fellowship; LS), the South African National Research Foundation (LS, RRA), and the University Research Council at the University of Cape Town (RRA). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Evolutionary Biology
                Evolutionary Processes
                Evolutionary Theory
                Organismal Evolution
                Paleontology
                Paleoanthropology
                Physical Anthropology
                Theoretical Biology
                Social Sciences
                Anthropology
                Custom metadata
                The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are within the paper and its Supporting Information files.

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