The geological, isotopic, botanical, invertebrate, and lower vertebrate surroundings of Ardipithecus ramidus.

Sediments containing Ardipithecus ramidus were deposited 4.4 million years ago on an alluvial floodplain in Ethiopia's western Afar rift. The Lower Aramis Member hominid-bearing unit, now exposed across a > 9-kilometer structural arc, is sandwiched between two volcanic tuffs that have nearly identical 40Ar/39Ar ages. Geological data presented here, along with floral, invertebrate, and vertebrate paleontological and taphonomic evidence associated with the hominids, suggest that they occupied a wooded biotope over the western three-fourths of the paleotransect. Phytoliths and oxygen and carbon stable isotopes of pedogenic carbonates provide evidence of humid cool woodlands with a grassy substrate.

Macrovertebrate paleontology and the Pliocene habitat of Ardipithecus ramidus.

A diverse assemblage of large mammals is spatially and stratigraphically associated with Ardipithecus ramidus at Aramis. The most common species are tragelaphine antelope and colobine monkeys. Analyses of their postcranial remains situate them in a closed habitat. Assessment of dental mesowear, microwear, and stable isotopes from these and a wider range of abundant associated larger mammals indicates that the local habitat at Aramis was predominantly woodland. The Ar. ramidus enamel isotope values indicate a minimal C4 vegetation component in its diet (plants using the C4 photosynthetic pathway), which is consistent with predominantly forest/woodland feeding. Although the Early Pliocene Afar included a range of environments, and the local environment at Aramis and its vicinity ranged from forests to wooded grasslands, the integration of available physical and biological evidence establishes Ar. ramidus as a denizen of the closed habitats along this continuum.

A method for estimating the relative importance of characters in cladistic analyses.

The method of character importance ranking (CIR) is proposed here as a means for estimating the relative "importance" of characters in cladistic analyses, especially those based on morphological features. CIR uses the weighting variable to incrementally remove one character at a time from the analysis, and then evaluates the impact of the removal on the shape of the cladogram. The greater the impact, the more important the character. The CIR method for determining which characters drive the shape of a particular cladogram has several applications. It identifies the characters with the strongest (though not necessarily most accurate) signal in a cladistic analysis; it permits the informed prioritization of characters for further investigation via genetic, developmental, and functional approaches; and it highlights characters whose definition, scoring, independence, and variation should be reviewed with particular care. The application of CIR reveals that at least some cladograms depend entirely on a single character.

Natural selection on molar size in a wild population of howler monkeys (Alouatta palliata).

Dental traits have long been assumed to be under selection in mammals, based on the macroevolutionary correlation between dental morphology and feeding behaviour. However, natural selection acting on dental morphology has rarely, if ever, been documented in wild populations. We investigated the possibility of microevolutionary selection on dental traits by measuring molar breadth in a sample of Alouatta palliata (mantled howler monkey) crania from Barro Colorado Island (BCI), Panama. The age at death of the monkeys is an indicator of their fitness, since they were all found dead of natural causes. Howlers with small molars have significantly decreased fitness as they die, on average, at an earlier age (well before sexual maturity) than those with larger molars. This documents the existence of phenotypic viability selection on molar tooth size in the BCI howlers, regardless of causality or heritability. The selection is further shown to occur during the weaning phase of A. palliata life history, establishing a link between this period of increased mortality and selection on a specific morphological feature. These results provide initial empirical support for the long-held assumption that primate molar size is under natural selection.

Pleistocene Homo sapiens from Middle Awash, Ethiopia.

The origin of anatomically modern Homo sapiens and the fate of Neanderthals have been fundamental questions in human evolutionary studies for over a century. A key barrier to the resolution of these questions has been the lack of substantial and accurately dated African hominid fossils from between 100,000 and 300,000 years ago. Here we describe fossilized hominid crania from Herto, Middle Awash, Ethiopia, that fill this gap and provide crucial evidence on the location, timing and contextual circumstances of the emergence of Homo sapiens. Radioisotopically dated to between 160,000 and 154,000 years ago, these new fossils predate classic Neanderthals and lack their derived features. The Herto hominids are morphologically and chronologically intermediate between archaic African fossils and later anatomically modern Late Pleistocene humans. They therefore represent the probable immediate ancestors of anatomically modern humans. Their anatomy and antiquity constitute strong evidence of modern-human emergence in Africa.

Missing Omo L338y-6 occipital-marginal sinus drainage pattern: ground sectioning, computer tomography scanning, and the original fossil fail to show it.

The Omo L338y-6 occipital region has been recently studied by White and Falk (1999), who claim that it shows a readily identifiable enlarged left occipital-marginal sinus (O/M). These observations are contrary to the direct observations of previous investigators (Rak and Howell, 1978; Kimbel, 1984; Holloway, 1981; Holloway, 1988). White and Falk (1999) further argue that the presence of this enlarged O/M strongly suggests that the Omo L338y-6 hominid was indeed a "robust" Australopithecus. We used direct sectioning and CT scanning to analyze magnified sections of a high-quality first-generation cast of the newly cleaned original fossil. These methods fail to show any evidence of a morphological landmark that can be interpreted as an enlarged O/M, either as an eminence or a sulcus. In contrast, the same techniques used with both SK 1585 and OH5 ("robust" Australopithecus with an enlarged O/M) show extremely visible and palpable enlarged O/M's. Examination of the original Omo fossil confirms that it lacks an O/M. This evidence clearly shows that an enlarged O/M cannot be identified on either the original fossil or a first-generation cast, although this does not rule out the possibility that the Omo L338y-6 hominid was a "robust" Australopithecus. We believe that the differences between observers regarding this feature are most probably due to displacement caused by a crack and the different source materials employed, i.e., the difference between a first-generation cast of the original fossil and a third- or fourth-generation cast of the endocast made two decades ago.