Homo naledi and Pleistocene hominin evolution in subequatorial Africa
Figures

African fossil sites from the Middle and earliest Late Pleistocene.
Sites discussed in the text are highlighted in pink here. Geological age estimations for each fossil hominin assemblage are given in Table 1, along with references.

Phylogenetic scenarios for H. naledi.
A simplified cladogram of Homo, with the possible placements of H. naledi indicated. The cladogram places A. africanus as an outgroup to the Homo + Au. sediba clade, as consistent with nearly all phylogenetic analyses of these species (Berger et al., 2010; Dembo et al., 2015, 2016). To simplify the tree, we have omitted H. antecessor, H. heidelbergensis and Neanderthals, which all phylogenetic analyses place as sisters to H. sapiens relative to H. erectus. There is no present consensus about the branching order among H. habilis, H. rudolfensis, H. floresiensis and Au. sediba (Dembo et al., 2015, 2016), and so these are depicted as a polytomy.
Tables
Significant hominin fossil remains from the Middle and Early Late Pleistocene of Africa. Included are those sites that have geological age estimates between 780,000 and 120,000 years ago, and some sites for which claims of Middle Pleistocene age have been made but without chronometric support. Sites denoted here with ‘no date’ are those for which no chronometric determinations based on samples of hominin material or securely associated faunal remains have been reported in the literature. Some chronometric determinations that were based only on morphology or associated fauna have given rise to broad age estimations; we omit the details of such determinations here. Some additional sites with fragmentary remains, especially isolated dental remains, are not listed. The first four entries (KNM-OL 45500, OH 12, Daka and Buia) are older than the beginning of the Middle Pleistocene but are included because they are discussed in text.
Site | Specimens | Location | Geological age (ka) | Source(s) |
---|---|---|---|---|
Olorgesailie (KNM-OL 45500) | Frontal | Kenya | 900–970 | Potts et al. (2004) |
Olduvai Gorge (OH 12)* | Partial calvaria | Tanzania | 780–1,200 | Tamrat et al. (1995); Mcbrearty and Brooks (2000) |
Daka | Calvaria, femur | Ethiopia | ~1,000 | Asfaw et al. (2002) |
Buia | Calvaria, postcranial fragments | Eritrea | ~1,000 | Abbate et al. (1998) |
Tighénif (Ternifine) | Three mandibles, skull fragment | Morocco | ~700 | Geraads et al. (1986) |
Elandsfontein (Saldanha) | Partial calvaria and mandible frag | South Africa | 600–1,000 | Klein et al. (2007a) |
Bodo | Partial calvaria, left parietal (found roughly 400 m from Bodo 1), distal humerus | Ethiopia | 550–640 | Conroy et al. (1978); Clark et al. (1994) |
Baringo (Kaphturin Formation) | Mandible, ulna | Kenya | 510–512 | Leakey et al. (1970); Deino and McBrearty (2002) |
Salé | Partial calvaria and upper dentition | Morocco | ~300 | Jaeger, 1975); Geraads (2012) |
Ndutu* | Partial calvaria | Tanzania | 370–990 | Tamrat et al. (1995); Mcbrearty and Brooks (2000) |
Berg Aukas | Partial femur | Namibia | No date | Grine et al. (1995) |
Kabwe | Calvaria, material from at least three individuals | Zambia | No date | Klein (1973) |
Florisbad | Partial cranium | South Africa | 224–294 | Grün et al. (1996) |
Cave of Hearths | Partial mandible | South Africa | No date | Cooke, 1962) |
Hoedjiespunt | Teeth, tibia | South Africa | No date | Berger and Parkington (1995); Stynder et al. (2001) |
Eliye Springs | Calvaria | Kenya | No date | Bräuer and Leakey (1986) |
Dinaledi Chamber (Rising Star) | Remains of at least 15 individuals | South Africa | 236–335 | Berger et al., 2015; Dirks et al. (2017) |
Lesedi Chamber (Rising Star) | Partial skeleton, remains of at least three individuals | South Africa | No date | Hawks et al., 2017) |
Omo Kibish | Two partial crania, partial skeleton | Ethiopia | 155–200 | McDougall et al. (2005); Aubert et al. (2012) |
Herto | Three partial crania | Ethiopia | 154–160 | White et al. (2003); Clark et al. (2003) |
Ileret (KNM-ER 3884) | Partial calvaria | Kenya | 162–∞ | Bräuer et al. (1997) |
Jebel Irhoud | Three calvaria, mandible, fragments of seven individuals | Morocco | 144–176 | Hublin (2001); Smith et al. (2007) |
Laetoli (Ngaloba Beds) | Cranium | Tanzania | 130 | Day et al. (1980); Hay (1987) |
Singa | Calvaria | Sudan | 131–135 | McDermott et al. (1996) |
Lake Eyasi | Calvaria | Tanzania | 88–132 | Mehlman (1984, 1987); Domínguez-Rodrigo et al., 2008 |
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*Many authors have studied the stratigraphy of Olduvai Gorge and nearby sites, resulting in varied dates being reported for these fossils. We report here the widest range as reviewed by Mcbrearty and Brooks (2000), based on the paleomagnetic sequence.