A new African fossil caprin and a combined molecular and morphological Bayesian phylogenetic analysis of caprini (Mammalia: Bovidae).

Given that most species that have ever existed on Earth are extinct, no evolutionary history can ever be complete without the inclusion of fossil taxa. Bovids (antelopes and relatives) are one of the most diverse clades of large mammals alive today, with over a hundred living species and hundreds of documented fossil species. With the advent of molecular phylogenetics, major advances have been made in the phylogeny of this clade; however, there has been little attempt to integrate the fossil record into the developing phylogenetic picture. We here describe a new large fossil caprin species from ca. 1.9-Ma deposits from the Middle Awash, Ethiopia. To place the new species phylogenetically, we perform a Bayesian analysis of a combined molecular (cytochrome b) and morphological (osteological) character supermatrix. We include all living species of Caprini, the new fossil species, a fossil takin from the Pliocene of Ethiopia (Budorcas churcheri), and the insular subfossil Myotragus balearicus. The combined analysis demonstrates successful incorporation of both living and fossil species within a single phylogeny based on both molecular and morphological evidence. Analysis of the combined supermatrix produces superior resolution than with either the molecular or morphological data sets considered alone. Parsimony and Bayesian analyses of the data set are also compared and shown to produce similar results. The combined phylogenetic analysis indicates that the new fossil species is nested within Capra, making it one of the earliest representatives of this clade, with implications for molecular clock calibration. Geographical optimization indicates no less than four independent dispersals into Africa by caprins since the Pliocene.

Do species populations really start small? New perspectives from the Late Neogene fossil record of African mammals.

This analysis of all known African larger mammals of the past 10 Myr offers new perspectives on the geographical circumstances of speciation. Our central question is: does the fossil evidence support the idea that most new species start as small populations and, if true, how long is the average growth interval until species are established at their mean later size? This simple question is important to unravelling the competing claims of rival models of speciation. We approached it by direct use of fossil data, which, to our knowledge, has not been done previously. We compared the numbers of fossil site records, as a proxy for magnitude of geographical spread, between survivorship intervals across all species. The results show that the average mammal species has indeed started its life in a relatively small population, and thereafter increased rapidly in geographical spread to reach its long-term equilibrium abundance by about 1 million years after origin. Some theoretical implications of these results are discussed.

Environment and behavior of 2.5-million-year-old Bouri hominids.

The Hata Member of the Bouri Formation is defined for Pliocene sedimentary outcrops in the Middle Awash Valley, Ethiopia. The Hata Member is dated to 2.5 million years ago and has produced a new species of Australopithecus and hominid postcranial remains not currently assigned to species. Spatially associated zooarchaeological remains show that hominids acquired meat and marrow by 2.5 million years ago and that they are the near contemporary of Oldowan artifacts at nearby Gona. The combined evidence suggests that behavioral changes associated with lithic technology and enhanced carnivory may have been coincident with the emergence of the Homo clade from Australopithecus afarensis in eastern Africa.

Multiphasic growth models and the evolution of prolonged growth exemplified by human brain evolution.

New models for multiphasic growth are presented. They are illustrated by analysis of brain growth in humans and chimpanzees, and the results are used to test the hypothesis of evolution by proportional growth prolongation: that all descendant growth phases are extended by the same factor while each remains at the ancestral growth rate. The results are consistent with the hypothesis and imply that gross brain weight increase towards humans required change in only one growth parameter: prolongation of the nonlinear ancestral growth phases. The restricted and orderly nature of the developmental changes hints at a basis in few genetic changes. Proportional growth prolongation is of general evolutionary importance because it can reorganize body proportions.

A cladistic analysis of mitochondrial ribosomal DNA from the Bovidae.

There is a huge data base of genetic information for the domestic artiodactyl species Bos taurus (cow), Ovis aries (sheep), and Capra hircus (goat). However, the phylogenetic relationships of these economically critical taxa and their close relatives, family Bovidae, remain for the most part unresolved. In this report, we aligned new mitochondrial (mt) 12S and 16S ribosomal (r) DNA sequences from 26 bovid taxa with published sequences. Phylogenetic analyses of the more than 64 kilobases of mt rDNA from 57 taxa support a basal division in the Bovidae that separates Bos and its close relatives from Capra, Ovis, and their kin. As suggested by previous molecular and morphological studies, "antelopes" are a paraphyletic assemblage. Caprinae (sheep, goats, goat antelopes, and musk oxen) groups consistently with hippotragine and alcelaphine antelopes, while Bovini (cattle and buffaloes) clusters with tragelaphine and boselaphine antelopes. The traditional tribal subdivisions of Bovidae are supported in most cases, but there are exceptions within Caprinae and Antilopinae (gazelles and close relatives). The rDNA data consistently place the enigmatic genera Pelea, Pantholops, and Saiga, but the origin of Aepyceros, the impala, remains obscure. Combined phylogenetic analyses of the rDNA data with the skeletal characters of Gentry (1992) were used to assess the stability of the molecular results.

African Homo erectus: old radiometric ages and young Oldowan assemblages in the Middle Awash Valley, Ethiopia.

Fossils and artifacts recovered from the middle Awash Valley of Ethiopia's Afar depression sample the Middle Pleistocene transition from Homo erectus to Homo sapiens. Ar/Ar ages, biostratigraphy, and tephrachronology from this area indicate that the Pleistocene Bodo hominid cranium and newer specimens are approximately 0.6 million years old. Only Oldowan chopper and flake assemblages are present in the lower stratigraphic units, but Acheulean bifacial artifacts are consistently prevalent and widespread in directly overlying deposits. This technological transition is related to a shift in sedimentary regime, supporting the hypothesis that Middle Pleistocene Oldowan assemblages represent a behavioral facies of the Acheulean industrial complex.

New discoveries of Australopithecus at Maka in Ethiopia.

The taxonomy of Australopithecus afarensis, the oldest known hominid species, has been a matter of debate since its description in 1978 (ref. 1). Some authorities regard all specimens assigned to A. afarensis as belonging to a single taxon whereas others regard the Tanzanian and Ethiopian specimens as each representing a different species. Further controversy surrounds the issues of sexual dimorphism and locomotion among these hominids. Resolution of these problems would shed light on hominid phylogeny in general and on the ancestry of later Australopithecus and Homo. Fossils discovered in the Afar of Ethiopia in 1990 constitute the first major addition to the 3-4 million year (Myr) hominid record since the 1970s. We report here the discovery of new fossils from Maka, dated to 3.4 Myr ago, which provide powerful support for the interpretation of A. afarensis as a single, ecologically diverse, sexually dimorphic, bipedal Pliocene primate species whose known range encompassed Ethiopia and Tanzania.

Phylogeny of the Bovidae (Artiodactyla, Mammalia), based on mitochondrial ribosomal DNA sequences.

Portions of the 12S and 16S mitochondrial ribosomal genes for 16 species representing nine tribes in the mammal family Bovidae were compared with six previously published orthologous sequences. Phylogenetic analysis of variable nucleotide positions under different constraints and weighting schemes revealed no robust groupings among tribes. Consensus trees support previous hypotheses of monophyly for four clades, including the traditional subfamily Bovinae. However, the basal diversification of bovid tribes, which was largely unresolved by morphological, immunodiffusion, allozyme, and protein sequence data, remains unresolved with the addition of DNA sequence data. The intractability of this systematic problem is consistent with a rapid radiation of the major bovid groups. Several analyses of our data show that monophyly of the Bovidae, which was weakly supported by previous morphological and molecular work, is questionable.

Macroevolutionary trends: new perspectives on the roles of adaptation and incidental effect.

Trends, long-term directional tendencies in evolution, are traditionally interpreted as selected for and adaptive. Alternatively, trends may be unselected effects of characters and processes within species: the effect hypothesis. Thus adaptations of organisms, varying among species, were selected for immediate fitness, but they may also incidentally determine different speciation and extinction rates and trends.

Australopithecine enamel prism patterns.

Following a recent suggestion that tooth enamel prism shape differs within Hominoidea, the teeth of a number of extinct and extant hominoid species were analyzed by scanning electron microscopy. The enamel prism patterns of some gracile and robust australopithecine specimens from Sterkfontein, Swartkrans, and Kromdraai are recorded. The characteristic arrangements of enamel prisms in all modern and extinct hominoid species were found to be essentially similar. The implications of enamel prisms for phylogenetic deduction in Hominoidea are discussed.