Seasonal shifts in community composition and proteome expression in a sulphur-cycling cyanobacterial mat.

Seasonal changes in light and physicochemical conditions have strong impacts on cyanobacteria, but how they affect community structure, metabolism, and biogeochemistry of cyanobacterial mats remains unclear. Light may be particularly influential for cyanobacterial mats exposed to sulphide by altering the balance of oxygenic photosynthesis and sulphide-driven anoxygenic photosynthesis. We studied temporal shifts in irradiance, water chemistry, and community structure and function of microbial mats in the Middle Island Sinkhole (MIS), where anoxic and sulphate-rich groundwater provides habitat for cyanobacteria that conduct both oxygenic and anoxygenic photosynthesis. Seasonal changes in light and groundwater chemistry were accompanied by shifts in bacterial community composition, with a succession of dominant cyanobacteria from Phormidium to Planktothrix, and an increase in diatoms, sulphur-oxidizing bacteria, and sulphate-reducing bacteria from summer to autumn. Differential abundance of cyanobacterial light-harvesting proteins likely reflects a physiological response of cyanobacteria to light level. Beggiatoa sulphur oxidation proteins were more abundant in autumn. Correlated abundances of taxa through time suggest interactions between sulphur oxidizers and sulphate reducers, sulphate reducers and heterotrophs, and cyanobacteria and heterotrophs. These results support the conclusion that seasonal change, including light availability, has a strong influence on community composition and biogeochemical cycling of sulphur and O2 in cyanobacterial mats.

Ecosystem engineering in the Quaternary of the West Coast of South Africa.

Despite advances in our understanding of the geographic and temporal scope of the Paleolithic record, we know remarkably little about the evolutionary and ecological consequences of changes in human behavior. Recent inquiries suggest that human evolution reflects a long history of interconnections between the behavior of humans and their surrounding ecosystems (e.g., niche construction). Developing expectations to identify such phenomena is remarkably difficult because it requires understanding the multi-generational impacts of changes in behavior. These long-term dynamics require insights into the emergent phenomena that alter selective pressures over longer time periods which are not possible to observe, and are also not intuitive based on observations derived from ethnographic time scales. Generative models show promise for probing these potentially unexpected consequences of human-environment interaction. Changes in the uses of landscapes may have long term implications for the environments that hominins occupied. We explore other potential proxies of behavior and examine how modeling may provide expectations for a variety of phenomena.

Increased ecological resource variability during a critical transition in hominin evolution.

Although climate change is considered to have been a large-scale driver of African human evolution, landscape-scale shifts in ecological resources that may have shaped novel hominin adaptations are rarely investigated. We use well-dated, high-resolution, drill-core datasets to understand ecological dynamics associated with a major adaptive transition in the archeological record ~24 km from the coring site. Outcrops preserve evidence of the replacement of Acheulean by Middle Stone Age (MSA) technological, cognitive, and social innovations between 500 and 300 thousand years (ka) ago, contemporaneous with large-scale taxonomic and adaptive turnover in mammal herbivores. Beginning ~400 ka ago, tectonic, hydrological, and ecological changes combined to disrupt a relatively stable resource base, prompting fluctuations of increasing magnitude in freshwater availability, grassland communities, and woody plant cover. Interaction of these factors offers a resource-oriented hypothesis for the evolutionary success of MSA adaptations, which likely contributed to the ecological flexibility typical of Homo sapiens foragers.

Fossil Cercopithecidae from the Early Pliocene Sagantole Formation at Gona, Ethiopia.

The Early Pliocene Sagantole Fm. in the Gona Project area, Afar State, Ethiopia, is noted for discoveries of the early hominin Ardipithecus ramidus. A large series of fossil cercopithecid primates dated to between 4.8 and 4.3 Ma has also been collected from these sediments. In this paper, we use qualitative analysis and standard dental and postcranial measures to systematically describe the craniodental remains and tentatively allocate postcrania to taxa where we are able to. We then use these data to compare these specimens to fossil assemblages from contemporary sites, interpret their paleobiology, and discuss implications for the paleoecology of the Gona Sagantole Fm. We recognize three cercopithecid species in the Gona Sagantole Fm. Pliopapio alemui makes up approximately two-thirds of the identifiable specimens; nearly all of the rest are allocated to Kuseracolobus aramisi, and a single molar indicates the presence of a second, somewhat larger but morphologically distinct papionin. Among the Early Pliocene cercopithecids from Gona are also a number of postcranial elements. None of the postcranial remains are directly associated with any of the cranial material. Nonetheless, some of the distal humeri and proximal femora can be tentatively allocated to either Pl. alemui or K. aramisi based on a combination of size, as the latter is approximately 50% larger than the former, and morphology. If these assignments are correct, they suggest K. aramisi was primarily arboreal and similar to most extant colobines, whereas Pl. alemui was more mixed in its substrate use, being more terrestrially adapted than K. aramisi, but less so than extant Papio or Theropithecus. Thus, we interpret the predominance of Pl. alemui over K. aramisi is consistent with a somewhat more open environment at Gona than at Aramis.

Co-occurrence of Acheulian and Oldowan artifacts with Homo erectus cranial fossils from Gona, Afar, Ethiopia.

Although stone tools generally co-occur with early members of the genus Homo, they are rarely found in direct association with hominins. We report that both Acheulian and Oldowan artifacts and Homo erectus crania were found in close association at 1.26 million years (Ma) ago at Busidima North (BSN12), and ca. 1.6 to 1.5 Ma ago at Dana Aoule North (DAN5) archaeological sites at Gona, Afar, Ethiopia. The BSN12 partial cranium is robust and large, while the DAN5 cranium is smaller and more gracile, suggesting that H. erectus was probably a sexually dimorphic species. The evidence from Gona shows behavioral diversity and flexibility with a lengthy and concurrent use of both stone technologies by H. erectus, confounding a simple "single species/single technology" view of early Homo.

Age and context of mid-Pliocene hominin cranium from Woranso-Mille, Ethiopia.

A fossil hominin cranium was discovered in mid-Pliocene deltaic strata in the Godaya Valley of the northwestern Woranso-Mille study area in Ethiopia. Here we show that analyses of chemically correlated volcanic layers and the palaeomagnetic stratigraphy, combined with Bayesian modelling of dated tuffs, yield an age range of 3.804 ± 0.013 to 3.777 ± 0.014 million years old (mean ± 1σ) for the deltaic strata and the fossils that they contain. We also document deposits of a perennial lake beneath the deltaic sequence. Mammalian fossils associated with the cranium represent taxa that were widespread at the time and data from botanical remains indicate that the vegetation in the lake and delta catchment was predominantly dry shrubland with varying proportions of grassland, wetland and riparian forest. In addition, we report high rates of sediment accumulation and depositional features that are typical of a steep topographic relief and differ from younger Woranso-Mille fossil localities, reflecting the influence of active rift processes on the palaeolandscape.

Ardipithecus ramidus postcrania from the Gona Project area, Afar Regional State, Ethiopia.

Functional analyses of the 4.4 Ma hominin Ardipithecus ramidus postcrania revealed a previously unknown and unpredicted locomotor pattern combining arboreal clambering and a form of terrestrial bipedality. To date, all of the fossil evidence of Ar. ramidus locomotion has been collected from the Aramis area of the Middle Awash Research Project in Ethiopia. Here, we present the results of an analysis of additional early Pliocene Ar. ramidus fossils from the Gona Project study area, Ethiopia, that includes a fragmentary but informative partial skeleton (GWM67/P2) and additional isolated manual remains. While we reinforce the original functional interpretations of Ar. ramidus of having a mixed locomotor adaptation of terrestrial bipedality and arboreal clambering, we broaden our understanding of the nature of its locomotor pattern by documenting better the function of the hip, ankle, and foot. The newly recovered fossils document a greater adaptation to bipedality in the Ar. ramidus ankle and hallux than previously recognized. In addition, a newly discovered scaphoid bone with a fusing os centrale provides further evidence about the nature of hominin hand evolution.

Environmental dynamics during the onset of the Middle Stone Age in eastern Africa.

Development of the African Middle Stone Age (MSA) before 300,000 years ago raises the question of how environmental change influenced the evolution of behaviors characteristic of early Homo sapiens We used temporally well-constrained sedimentological and paleoenvironmental data to investigate environmental dynamics before and after the appearance of the early MSA in the Olorgesailie basin, Kenya. In contrast to the Acheulean archeological record in the same basin, MSA sites are associated with a markedly different faunal community, more pronounced erosion-deposition cycles, tectonic activity, and enhanced wet-dry variability. Aspects of Acheulean technology in this region imply that, as early as 615,000 years ago, greater stone material selectivity and wider resource procurement coincided with an increased pace of land-lake fluctuation, potentially anticipating the adaptability of MSA hominins.

Aridity and hominin environments.

Aridification is often considered a major driver of long-term ecological change and hominin evolution in eastern Africa during the Plio-Pleistocene; however, this hypothesis remains inadequately tested owing to difficulties in reconstructing terrestrial paleoclimate. We present a revised aridity index for quantifying water deficit (WD) in terrestrial environments using tooth enamel δ18O values, and use this approach to address paleoaridity over the past 4.4 million years in eastern Africa. We find no long-term trend in WD, consistent with other terrestrial climate indicators in the Omo-Turkana Basin, and no relationship between paleoaridity and herbivore paleodiet structure among fossil collections meeting the criteria for WD estimation. Thus, we suggest that changes in the abundance of C4 grass and grazing herbivores in eastern Africa during the Pliocene and Pleistocene may have been decoupled from aridity. As in modern African ecosystems, other factors, such as rainfall seasonality or ecological interactions among plants and mammals, may be important for understanding the evolution of C4 grass- and grazer-dominated biomes.

Dentognathic remains of Australopithecus afarensis from Nefuraytu (Woranso-Mille, Ethiopia): Comparative description, geology, and paleoecological context.

Australopithecus afarensis is the best-known and most dimorphic species in the early hominin fossil record. Here, we present a comparative description of new fossil specimens of Au. afarensis from Nefuraytu, a 3.330-3.207 million-years-old fossil collection area in the Woranso-Mille study area, central Afar, Ethiopia. These specimens include NFR-VP-1/29, one of the most complete mandibles assigned to the species thus far and among the largest mandibles attributed to Au. afarensis, likely representing a male individual. NFR-VP-1/29 retains almost all of the distinctive archaic features documented for Au. afarensis. These features include a posteriorly sloping symphysis, a low and rounded basally set inferior transverse torus, anterosuperiorly opening mental foramen, a lateral corpus hollow bound anteriorly by the C/P3 jugae and posteriorly by the lateral prominence, and the ascending ramus arising high on the corpus. Dental morphology and metrics of the Nefuraytu specimens also falls within the range of Au. afarensis. The presence of this species at Woranso-Mille between 3.330 and 3.207 million years ago confirms the existence of this species in the area in close spatial and temporal proximity to other middle Pliocene hominin taxa such as the one represented by the Burtele foot (BRT-VP-2/73) and the recently named species Australopithecus deyiremeda. This has important implications for our understanding of middle Pliocene hominin diversity.

Dietary change among hominins and cercopithecids in Ethiopia during the early Pliocene.

The incorporation of C4 resources into hominin diet signifies increased dietary breadth within hominins and divergence from the dietary patterns of other great apes. Morphological evidence indicates that hominin diet became increasingly diverse by 4.2 million years ago but may not have included large proportions of C4 foods until 800 thousand years later, given the available isotopic evidence. Here we use carbon isotope data from early to mid Pliocene hominin and cercopithecid fossils from Woranso-Mille (central Afar, Ethiopia) to constrain the timing of this dietary change and its ecological context. We show that both hominins and some papionins expanded their diets to include C4 resources as early as 3.76 Ma. Among hominins, this dietary expansion postdates the major dentognathic morphological changes that distinguish Australopithecus from Ardipithecus, but it occurs amid a continuum of adaptations to diets of tougher, harder foods and to committed terrestrial bipedality. In contrast, carbon isotope data from cercopithecids indicate that C4-dominated diets of the earliest members of the Theropithecus oswaldi lineage preceded the dental specialization for grazing but occurred after they were fully terrestrial. The combined data indicate that the inclusion of C4 foods in hominin diet occurred as part of broader ecological changes in African primate communities.

Dietary changes of large herbivores in the Turkana Basin, Kenya from 4 to 1 Ma.

A large stable isotope dataset from East and Central Africa from ca. 30 regional collection sites that range from forest to grassland shows that most extant East and Central African large herbivore taxa have diets dominated by C4 grazing or C3 browsing. Comparison with the fossil record shows that faunal assemblages from ca. 4.1-2.35 Ma in the Turkana Basin had a greater diversity of C3-C4 mixed feeding taxa than is presently found in modern East and Central African environments. In contrast, the period from 2.35 to 1.0 Ma had more C4-grazing taxa, especially nonruminant C4-grazing taxa, than are found in modern environments in East and Central Africa. Many nonbovid C4 grazers became extinct in Africa, notably the suid Notochoerus, the hipparion equid Eurygnathohippus, the giraffid Sivatherium, and the elephantid Elephas. Other important nonruminant C4-grazing taxa switched to browsing, including suids in the lineage Kolpochoerus-Hylochoerus and the elephant Loxodonta. Many modern herbivore taxa in Africa have diets that differ significantly from their fossil relatives. Elephants and tragelaphin bovids are two groups often used for paleoecological insight, yet their fossil diets were very different from their modern closest relatives; therefore, their taxonomic presence in a fossil assemblage does not indicate they had a similar ecological function in the past as they do at present. Overall, we find ecological assemblages of C3-browsing, C3-C4-mixed feeding, and C4-grazing taxa in the Turkana Basin fossil record that are different from any modern ecosystem in East or Central Africa.

New species from Ethiopia further expands Middle Pliocene hominin diversity.

Middle Pliocene hominin species diversity has been a subject of debate over the past two decades, particularly after the naming of Australopithecus bahrelghazali and Kenyanthropus platyops in addition to the well-known species Australopithecus afarensis. Further analyses continue to support the proposal that several hominin species co-existed during this time period. Here we recognize a new hominin species (Australopithecus deyiremeda sp. nov.) from 3.3-3.5-million-year-old deposits in the Woranso-Mille study area, central Afar, Ethiopia. The new species from Woranso-Mille shows that there were at least two contemporaneous hominin species living in the Afar region of Ethiopia between 3.3 and 3.5 million years ago, and further confirms early hominin taxonomic diversity in eastern Africa during the Middle Pliocene epoch. The morphology of Au. deyiremeda also reinforces concerns related to dentognathic (that is, jaws and teeth) homoplasy in Plio-Pleistocene hominins, and shows that some dentognathic features traditionally associated with Paranthropus and Homo appeared in the fossil record earlier than previously thought.

Late Miocene hominin teeth from the Gona Paleoanthropological Research Project area, Afar, Ethiopia.

Since 2000, significant collections of Latest Miocene hominin fossils have been recovered from Chad, Kenya, and Ethiopia. These fossils have provided a better understanding of earliest hominin biology and context. Here, we describe five hominin teeth from two periods (ca. 5.4 Million-years-ago and ca. 6.3 Ma) that were recovered from the Adu-Asa Formation in the Gona Paleoanthropological Research Project area in the Afar, Ethiopia that we assign to either Hominina, gen. et sp. indet. or Ardipithecus kadabba. These specimens are compared with extant African ape and other Latest Miocene and Early Pliocene hominin teeth. The derived morphology of the large, non-sectorial maxillary canine and mandibular third premolar links them with later hominins and they are phenetically distinguishable and thus phyletically distinct from extant apes.

Measurement of δ18O, δ17O, and 17O-excess in water by off-axis integrated cavity output spectroscopy and isotope ratio mass spectrometry.

Stable isotopes of water have long been used to improve understanding of the hydrological cycle, catchment hydrology, and polar climate. Recently, there has been increasing interest in measurement and use of the less-abundant (17)O isotope in addition to (2)H and (18)O. Off-axis integrated cavity output spectroscopy (OA-ICOS) is demonstrated for accurate and precise measurements δ(18)O, δ(17)O, and (17)O-excess in liquid water. OA-ICOS involves no sample conversion and has a small footprint, allowing measurements to be made by researchers collecting the samples. Repeated (514) high-throughput measurements of the international isotopic reference water standard Greenland Ice Sheet Precipitation (GISP) demonstrate the precision and accuracy of OA-ICOS: δ(18)OVSMOW-SLAP = -24.74 ± 0.07‰ (1σ) and δ(17)OVSMOW-SLAP = -13.12 ± 0.05‰ (1σ). For comparison, the International Atomic Energy Agency (IAEA) value for δ(18)OVSMOW-SLAP is -24.76 ± 0.09‰ (1σ) and an average of previously reported values for δ(17)OVSMOW-SLAP is -13.12 ± 0.06‰ (1σ). Multiple (26) high-precision measurements of GISP provide a (17)O-excessVSMOW-SLAP of 23 ± 10 per meg (1σ); an average of previously reported values for (17)O-excessVSMOW-SLAP is 22 ± 11 per meg (1σ). For all these OA-ICOS measurements, precision can be further enhanced by additional averaging. OA-ICOS measurements were compared with two independent isotope ratio mass spectrometry (IRMS) laboratories and shown to have comparable accuracy and precision as the current fluorination-IRMS techniques in δ(18)O, δ(17)O, and (17)O-excess. The ability to measure accurately δ(18)O, δ(17)O, and (17)O-excess in liquid water inexpensively and without sample conversion is expected to increase vastly the application of δ(17)O and (17)O-excess measurements for scientific understanding of the water cycle, atmospheric convection, and climate modeling among others.

A new hominin foot from Ethiopia shows multiple Pliocene bipedal adaptations.

A newly discovered partial hominin foot skeleton from eastern Africa indicates the presence of more than one hominin locomotor adaptation at the beginning of the Late Pliocene epoch. Here we show that new pedal elements, dated to about 3.4 million years ago, belong to a species that does not match the contemporaneous Australopithecus afarensis in its morphology and inferred locomotor adaptations, but instead are more similar to the earlier Ardipithecus ramidus in possessing an opposable great toe. This not only indicates the presence of more than one hominin species at the beginning of the Late Pliocene of eastern Africa, but also indicates the persistence of a species with Ar. ramidus-like locomotor adaptation into the Late Pliocene.

Stable isotope ecology in the Omo-Turkana Basin.

Stable isotopes provide an independent assessment of paleoenvironments in the Omo-Turkana Basin. Stable isotopes track the flow of oxygen and carbon through ecosystems and accordingly are not directly related to changes in mammalian faunal composition or sedimentology. Therefore, isotope studies give insight into the paleoenvironmental conditions in which human evolutionary trends have been recorded. The development of stable isotopes as indicators of continental environmental conditions has proceeded in parallel with questions about the conditions of human environment. What was the vegetation? How hot was it? How dry? What were the diets of animals living among early humans? And most persistently, how important were "savannas" to early hominids? In this review, we take the opportunity to provide extensive background on the use of isotopes in anthropological sites. The application of stable isotope ecology to anthropological sites in the Turkana Basin has a long history, but in many ways the Omo-Turkana Basin has been a proving ground for the development of new proxy methods for understanding tropical terrestrial environments in the Neogene and Quaternary. For that reason, we also describe some of the fundamental aspects of isotope ecology that developed outside the field of paleoanthropology.

Woody cover and hominin environments in the past 6 million years.

The role of African savannahs in the evolution of early hominins has been debated for nearly a century. Resolution of this issue has been hindered by difficulty in quantifying the fraction of woody cover in the fossil record. Here we show that the fraction of woody cover in tropical ecosystems can be quantified using stable carbon isotopes in soils. Furthermore, we use fossil soils from hominin sites in the Awash and Omo-Turkana basins in eastern Africa to reconstruct the fraction of woody cover since the Late Miocene epoch (about 7 million years ago). (13)C/(12)C ratio data from 1,300 palaeosols at or adjacent to hominin sites dating to at least 6 million years ago show that woody cover was predominantly less than ∼40% at most sites. These data point to the prevalence of open environments at the majority of hominin fossil sites in eastern Africa over the past 6 million years.