Understanding the extinction of Australopithecus and origins of Paranthropus and Homo in South Africa has been hampered by the perceived complex geological context of hominin fossils, poor chronological resolution, and a lack of well-preserved early Homo specimens. We describe, date, and contextualize the discovery of two hominin crania from Drimolen Main Quarry in South Africa. At ~2.04 million to 1.95 million years old, DNH 152 represents the earliest definitive occurrence of Paranthropus robustus, and DNH 134 represents the earliest occurrence of a cranium with clear affinities to Homo erectus. These crania also show that Homo, Paranthropus, and Australopithecus were contemporaneous at ~2 million years ago. This high taxonomic diversity is also reflected in non-hominin species and provides evidence of endemic evolution and dispersal during a period of climatic variability.

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Haasgat is a primate-rich fossil locality in the northeastern part of the Fossil Hominid Sites of South Africa UNESCO World Heritage Site. Here we report the first hominin identified from Haasgat, a partial maxillary molar (HGT 500), that was recovered from an ex situ calcified sediment block sampled from the locality. The in situ fossil bearing deposits of the Haasgat paleokarstic deposits are estimated to date to slightly older than 1.95 Ma based on magnetobiostratigraphy. This places the hominin specimen at a critical time period in South Africa that marks the last occurrence of Australopithecus around 1.98 Ma and the first evidence of Paranthropus and Homo in the region between ∼2.0 and 1.8 Ma. A comprehensive morphological evaluation of the Haasgat hominin molar was conducted against the current South African catalogue of hominin dental remains and imaging analyses using micro-CT, electron and confocal microscopy. The preserved occlusal morphology is most similar to Australopithecus africanus or early Homo specimens but different from Paranthropus. Occlusal linear enamel thickness measured from micro-CT scans provides an average of ∼2.0 mm consistent with Australopithecus and early Homo. Analysis of the enamel microstructure suggests an estimated periodicity of 7–9 days. Hunter–Schreger bands appear long and straight as in some Paranthropus, but contrast with this genus in the short shape of the striae of Retzius. Taken together, these data suggests that the maxillary fragment recovered from Haasgat best fits within the Australopithecus—early Homo hypodigms to the exclusion of the genus Paranthropus. At ∼1.95 Ma this specimen would either represent another example of late occurring Australopithecus or one of the earliest examples of Homo in the region. While the identification of this first hominin specimen from Haasgat is not unexpected given the composition of other South African penecontemporaneous site deposits, it represents one of the few hominin localities in the topographically-distinct northern World Heritage Site. When coupled with the substantial differences in the mammalian faunal communities between the northern localities (e.g., Haasgat, Gondolin) and well-sampled Bloubank Valley sites (e.g., Sterkfontein, Swartkrans, Kromdraai), the recovery of the HGT 500 specimen highlights the potential for further research at the Haasgat locality for understanding the distribution and interactions of hominin populations across the landscape, ecosystems and fossil mammalian communities of early Pleistocene South Africa. Such contextual data from sites like Haasgat is critical for understanding the transition in hominin representation at ∼2 Ma sites in the region from Australopithecus to Paranthropus and early Homo.

Objectives: Twenty-four dental specimens from the Drimolen Main Quarry (DMQ) are described. This increases the number of DMQ Paranthropus robustus specimens from 48 to 63 and DMQ Homo specimens from 8 to 12. This allows reassessment of the proposed differences between the DMQ P. robustus assemblage and that of Swartkrans. Analysis conducted assesses intraspecific and inter-locality variation.Materials and Methods: We examined the P. robustus and early Homo assemblages from South Africa. Morphology was observed using a hand lens and a binocular microscope. Mesiodistal and buccolingual measurements were taken using plastictipped calipers. Summary statistics were generated and patterns of variability in P. robustus were assessed through box plots and Mann-Whitney U tests.Results: Comparison between the expanded DMQ and Swartkrans P. robustus assemblages demonstrates overlap in size. Ten dental variables show statistically significant differences.Discussion: The expanded P. robustus sample allowed us to re-examine previous analyses of differences in tooth size between the samples. While analyses presented here show a high degree of overlap in the MD and BL dimensions of the two assemblages, significant differences were found in the mean values of these variables in the postcanine maxillary teeth-consistent with previous analyses. Two current hypotheses may explain this pattern: 1) dental size increase through the P. robustus lineage or 2) different sample composition between the two sites. Small sample sizes for all permanent dental classes in the DMQ assemblage represents a limitation on this analysis and interpretations thereof. Any addition to the DMQ or the Swartkrans samples may alter these results.

Classic depictions of human evolutionary ecology cast Homo as predator and other hominins, including Paranthropus robustus, as prey. Such hypotheses rest on a small number of fossils that exhibit evidence of carnivore predation, including the iconic SK 54 cranium from Swartkrans in South Africa. Here we demonstrate that the SK 54 cranium shares its closest affinities with H. erectus rather than P. robustus. Demonstrating that Homo was prey for leopards at Swartkrans weakens the historically significant hypothesis that Homo was better able to avoid predation because of being behaviourally and technologically advanced compared to Paranthropus. Subsequent ideas about hominin paleobiology derived from this hypothesis warrant reconsideration.

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