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Abstract
There are thousands of fossils of hominins, but no fossil chimpanzee has yet been
reported. The chimpanzee (Pan) is the closest living relative to humans. Chimpanzee
populations today are confined to wooded West and central Africa, whereas most hominin
fossil sites occur in the semi-arid East African Rift Valley. This situation has fuelled
speculation regarding causes for the divergence of the human and chimpanzee lineages
five to eight million years ago. Some investigators have invoked a shift from wooded
to savannah vegetation in East Africa, driven by climate change, to explain the apparent
separation between chimpanzee and human ancestral populations and the origin of the
unique hominin locomotor adaptation, bipedalism. The Rift Valley itself functions
as an obstacle to chimpanzee occupation in some scenarios. Here we report the first
fossil chimpanzee. These fossils, from the Kapthurin Formation, Kenya, show that representatives
of Pan were present in the East African Rift Valley during the Middle Pleistocene,
where they were contemporary with an extinct species of Homo. Habitats suitable for
both hominins and chimpanzees were clearly present there during this period, and the
Rift Valley did not present an impenetrable barrier to chimpanzee occupation.
Consensus on the evolutionary relationships of humans, chimpanzees, and gorillas has not been reached, despite the existence of a number of DNA sequence data sets relating to the phylogeny, partly because not all gene trees from these data sets agree. However, given the well-known phenomenon of gene tree-species tree mismatch, agreement among gene trees is not expected. A majority of gene trees from available DNA sequence data support one hypothesis, but is this evidence sufficient for statistical confidence in the majority hypothesis? All available DNA sequence data sets showing phylogenetic resolution among the hominoids are grouped according to genetic linkage of their corresponding genes to form independent data sets. Of the 14 independent data sets defined in this way, 11 support a human-chimpanzee clade, 2 support a chimpanzee-gorilla clade, and one supports a human-gorilla clade. The hypothesis of a trichotomous speciation event leading to Homo; Pan, and Gorilla can be firmly rejected on the basis of this data set distribution. The multiple-locus test (Wu 1991), which evaluates hypotheses using gene tree-species tree mismatch probabilities in a likelihood ratio test, favors the phylogeny with a Homo-Pan clade and rejects the other alternatives with a P value of 0.002. When the probabilities are modified to reflect effective population size differences among different types of genetic loci, the observed data set distribution is even more likely under the Homo-Pan clade hypothesis. Maximum-likelihood estimates for the time between successive hominoid divergences are in the range of 300,000-2,800,000 years, based on a reasonable range of estimates for long-term hominoid effective population size and for generation time. The implication of the multiple-locus test is that existing DNA sequence data sets provide overwhelming and sufficient support for a human-chimpanzee clade: no additional DNA data sets need to be generated for the purpose of estimating hominoid phylogeny. Because DNA hybridization evidence (Caccone and Powell 1989) also supports a Homo-Pan clade, the problem of hominoid phylogeny can be confidently considered solved.
Bipedality, the definitive characteristic of the earliest hominids, has been regarded as an adaptive response to a transition from forested to more-open habitats in East Africa sometime between 12 million and 5 million years ago. Analyses of the stable carbon isotopic composition (delta(13)C) of paleosol carbonate and organic matter from the Tugen Hills succession in Kenya indicate that a heterogeneous environment with a mix of C3 and C4 plants has persisted for the last 15.5 million years. Open grasslands at no time dominated this portion of the rift valley. The observed delta(13)C values offer no evidence for a shift from more-closed C3 environments to C4 grassland habitats. If hominids evolved in East Africa during the Late Miocene, they did so in an ecologically diverse setting.
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