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INTRODUCTION Inherent in traditional views of ape origins is the idea that, like living apes, early large-bodied apes lived in tropical forests. In response to constraints related to locomoting in forest canopies, it has been proposed that early apes evolved their quintessential upright torsos and acrobatic climbing and suspensory abilities, enhancing their locomotor versatility, to distribute their weight among small supports and thus reach ripe fruit in the terminal branches. This feeding and locomotor transition from a quadruped with a horizontal torso is thought to have occurred in the Middle Miocene due to an increasingly seasonal climate and feeding competition from evolving monkeys. Although ecological and behavioral comparisons among living apes and monkeys provide evidence for versions of terminal branch forest frugivory hypotheses, corroboration from the early ape fossil record has been lacking, as have detailed reconstructions of the habitats where the first apes evolved. RATIONALE The Early Miocene fossil site of Moroto II in Uganda provides a unique opportunity to test the predictions of terminal branch forest frugivory hypotheses. Moroto II documents the oldest [21 million years ago (Ma)] well-established paleontological record of ape teeth and postcranial bones from a single locality and preserves paleoecological proxies to reconstruct the environment. The following lines of evidence from Moroto II were analyzed: (i) the functional anatomy of femora and a vertebra attributed to the ape Morotopithecus ; (ii) dental traits, including molar shape and isotopic profiles of Morotopithecus enamel; (iii) isotopic dietary paleoecology of associated fossil mammals; (iv) biogeochemical signals from paleosols (ancient soils) that reflect local relative proportions of C 3 (trees and shrubs) and C 4 (tropical grasses and sedges that can endure water stress) vegetation as well as rainfall; and (v) assemblages of phytoliths, microscopic plant-derived silica bodies that reflect past plant communities. RESULTS A short, strong femur biomechanically favorable to vertical climbing and a vertebra indicating a dorsostable lower back confirm that ape fossils from Moroto II shared locomotor traits with living apes. Both Morotopithecus and a smaller ape from the site have elongated molars with well-developed crests for shearing leaves. Carbon isotopic signatures of the enamel of these apes and of other fossil mammals indicate that some mammals consistently fed on water-stressed C 3  plants, and possibly also C 4  vegetation, in a woodland setting. Carbon isotope values of pedogenic carbonates, paleosol organic matter, and plant waxes all point to substantial C 4 grass biomass on the landscape. Analysis of paleosols also indicates subhumid, strongly seasonal rainfall, and phytolith assemblages include forms from both arid-adapted C 4 grasses and forest-indicator plants. CONCLUSION The ancient co-occurrence of dental specializations for leaf eating, rather than ripe fruit consumption, along with ape-like locomotor abilities counters the predictions of the terminal branch forest frugivory hypotheses. The combined paleoecological evidence situates Morotopithecus in a woodland with a broken canopy and substantial grass understory including C 4 species. These findings call for a new paradigm for the evolutionary origins of early apes. We propose that seasonal, wooded environments may have exerted previously unrecognized selective pressures in the evolution of arboreal apes. For example, some apes may have needed to access leaves in the higher canopy in times of low fruit availability and to be adept at ascending and descending from trees that lacked a continuous canopy. Hominoid habitat comparisons. Shown are reconstructions of a traditionally conceived hominoid habitat ( A ) and the 21 Ma Moroto II, Uganda, habitat ( B ). 

Significance We have developed an Africa-wide synthesis of paleoenvironmental variability over the Plio-Pleistocene. We show that there is strong evidence for orbital forcing of variability during this time that is superimposed on a longer trend of increasing environmental variability, supporting a combination of both low- and high-latitude drivers of variability. We combine these results with robust estimates of mammalian speciation and extinction rates and find that variability is not significantly correlated with these rates. These findings do not currently support a link between environmental variability and turnover and thus fail to corroborate predictions derived from the variability selection hypothesis. 

Abstract Terrestrial‐marine dust fluxes, pedogenic carbonate δ¹³C values, and various paleovegetation proxies suggest that Africa experienced gradual cooling and drying across the Pliocene‐Pleistocene (Plio‐Pleistocene) boundary (2.58 million years ago [Ma]). However, the timing, magnitude, resolution, and relative influences of orbitally‐driven changes in high latitude glaciations and low latitude insolation differ by region and proxy. To disentangle these forcings and investigate equatorial eastern African climate across the Plio‐Pleistocene boundary, we generated a high‐resolution (∼3,000‐year) data set of compound‐specificn‐alkane leaf wax δ²H values—a robust proxy for atmospheric circulation and precipitation amount—from the HSPDP‐BTB13‐1A core, which spans a ∼3.3–2.6 Ma sequence in the Baringo‐Tugen Hills‐Barsemoi Basin of central Kenya. In combination with the physical sedimentology, our data indicate that precipitation varied strongly with orbital obliquity, not precession, during the late Pliocene, perhaps imparted by variations in the cross‐equatorial insolation gradient. We also observe a marked shift toward wetter conditions beginning ∼3 Ma that corresponds with global cooling, drying in western Australia, and a steepening of the west‐east zonal Indian Ocean (IO) sea surface temperature (SST) gradient. We propose that northward migration of the Subtropical Front reduced Agulhas current leakage, warming the western IO and causing changes in the IO zonal SST gradient at 3 Ma, a process that has been observed in the latest Pleistocene‐Holocene but not over longer timescales. Thus, the late Cenozoic moisture history of eastern Africa is driven by a complex mixture of low‐latitude insolation, the IO SST gradient, and teleconnections to distal high‐latitude cooling. 

The assembly of Africa’s iconic C₄grassland ecosystems is central to evolutionary interpretations of many mammal lineages, including hominins. C₄grasses are thought to have become ecologically dominant in Africa only after 10 million years ago (Ma). However, paleobotanical records older than 10 Ma are sparse, limiting assessment of the timing and nature of C₄biomass expansion. This study uses a multiproxy design to document vegetation structure from nine Early Miocene mammal site complexes across eastern Africa. Results demonstrate that between ~21 and 16 Ma, C₄grasses were locally abundant, contributing to heterogeneous habitats ranging from forests to wooded grasslands. These data push back the oldest evidence of C₄grass–dominated habitats in Africa—and globally—by more than 10 million years, calling for revised paleoecological interpretations of mammalian evolution. 

Paleoanthropologists have long speculated about the role of environmental change in shaping human evolution in Africa. In recent years, drill cores of late Neogene lacustrine sedimentary rocks have yielded valuable high-resolution records of climatic and ecosystem change. Eastern African Rift sediments (primarily lake beds) provide an extraordinary range of data in close proximity to important fossil hominin and archaeological sites, allowing critical study of hypotheses that connect environmental history and hominin evolution. We review recent drill-core studies spanning the Plio–Pleistocene boundary (an interval of hominin diversification, including the earliest members of our genus Homo and the oldest stone tools), and the Mid–Upper Pleistocene (spanning the origin of Homo sapiens in Africa and our early technological and dispersal history). Proposed drilling of Africa's oldest lakes promises to extend such records back to the late Miocene. ▪ High-resolution paleoenvironmental records are critical for understanding external drivers of human evolution. ▪ African lake basin drill cores play a critical role in enhancing hominin paleoenvironmental records given their continuity and proximity to key paleoanthropological sites. ▪ The oldest African lakes have the potential to reveal a comprehensive paleoenvironmental context for the entire late Neogene history of hominin evolution.