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Swanscombe, and from the Sima de los Huesos). The first DNA recovered from a fossil hominin was from the type specimen of H. neanderthalensis (Krings et al., 1997), and recently Green et al. (2008) sequenced the complete mtDNA of a specimen from Vindija. Briggs et al. (2009) reported the mtDNA sequences of five individuals and concluded that genetic diversity within H. neanderthalensis was substantially lower than that in modern humans.

The next fossil hominin taxon in this grade to be discovered was H. erectus (Dubois 1893) Weidenreich 1940. Its temporal range is ca. 1.8 Ma to ca. 30 ka. The initial discovery at Kedung Brubus was made in 1890, but the type specimen was recovered in 1891 from Trinil. H. erectus is known from sites in Indonesia (e.g., Trinil, Sangiran, and Sambungmachan), China (e.g., Zhoukoudian and Lantian), and Africa (e.g., Olduvai Gorge and Melka Kunturé). The hypodigm of H. erectus is dominated by cranial remains; there is some postcranial evidence but very few hand and foot fossils. Crania belonging to H. erectus have a low vault, a substantial more-or-less continuous torus above the orbits, and a sharply angulated occipital region, and the inner and outer tables of the cranial vault are thick. The body of the mandible is more robust than that of H. sapiens, it lacks a chin, and the mandibular tooth crowns are generally larger and the premolar roots more complicated than those of modern humans. The limb proportions of H. erectus are modern human-like, but the shafts of the long bones are robust and those of the lower limb are flattened (the femur from front to back and the tibia from side to side) relative to those of modern humans. Overall, the cortical bone of H. erectus is thicker than is the case in modern humans. All of the dental and cranial evidence points to a modern human-like diet for H. erectus, and the postcranial elements are consistent with an upright posture and obligate bipedalism.

The next taxon recognized within the genus Homo was H. heidelbergensis Schoetensack 1908. The initial discovery and the type specimen, the Mauer 1 adult mandible, was found in 1907 in a sand quarry near Heidelberg, Germany. Other evidence included in the taxon comes from sites in Europe (e.g., Petralona), the Near East (e.g., Zuttiyeh), Africa (e.g., Kabwe and Bodo), China (e.g., Dali, Jinniushan, Xujiayao, and Yunxian), possibly India (Hathnora), and depending on how inclusively H. neanderthalensis is interpreted, from the Sima de los Huesos at Atapuerca, Spain. The temporal range of H. heidelbergensis is ca. 600–100 ka. What sets this material apart from H. sapiens and H. neanderthalensis is the morphology of the cranium and the robusticity of the postcranial skeleton. Some H. heidelbergensis have endocranial volumes as large as those of some modern humans, but they are always more robustly built, with a thickened occipital region and a projecting face and with large separate ridges

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