Homo naledi and Pleistocene hominin evolution in subequatorial Africa

  1. Lee R Berger  Is a corresponding author
  2. John Hawks
  3. Paul HGM Dirks
  4. Marina Elliott
  5. Eric M Roberts
  1. University of the Witwatersrand, South Africa
  2. University of Wisconsin, United States
  3. James Cook University, Australia
2 figures and 1 table


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.



Table 1

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.

SiteSpecimensLocationGeological age (ka)Source(s)
(KNM-OL 45500)
FrontalKenya900–970Potts et al. (2004)
Olduvai Gorge
(OH 12)*
Partial calvariaTanzania780–1,200Tamrat et al. (1995); Mcbrearty and Brooks (2000)
DakaCalvaria, femurEthiopia~1,000Asfaw et al. (2002)
BuiaCalvaria, postcranial fragmentsEritrea~1,000Abbate et al. (1998)
Tighénif (Ternifine)Three mandibles, skull fragmentMorocco~700Geraads et al. (1986)
Elandsfontein (Saldanha)Partial calvaria and mandible fragSouth Africa600–1,000Klein et al. (2007a)
BodoPartial calvaria, left parietal (found roughly 400 m from Bodo 1), distal humerusEthiopia550–640Conroy et al. (1978); Clark et al. (1994)
Baringo (Kaphturin Formation)Mandible, ulnaKenya510–512Leakey et al. (1970); Deino and McBrearty (2002)
SaléPartial calvaria and upper dentitionMorocco~300Jaeger, 1975); Geraads (2012)
Ndutu*Partial calvariaTanzania370–990Tamrat et al. (1995); Mcbrearty and Brooks (2000)
Berg AukasPartial femurNamibiaNo dateGrine et al. (1995)
KabweCalvaria, material from at least three individualsZambiaNo dateKlein (1973)
FlorisbadPartial craniumSouth Africa224–294Grün et al. (1996)
Cave of HearthsPartial mandibleSouth AfricaNo dateCooke, 1962)
HoedjiespuntTeeth, tibiaSouth AfricaNo dateBerger and Parkington (1995); Stynder et al. (2001)
Eliye SpringsCalvariaKenyaNo dateBräuer and Leakey (1986)
Dinaledi Chamber (Rising Star)Remains of at least 15 individualsSouth Africa236–335Berger et al., 2015; Dirks et al. (2017)
Lesedi Chamber (Rising Star)Partial skeleton, remains of at least three individualsSouth AfricaNo dateHawks et al., 2017)
Omo KibishTwo partial crania, partial skeletonEthiopia155–200McDougall et al. (2005); Aubert et al. (2012)
HertoThree partial craniaEthiopia154–160White et al. (2003); Clark et al. (2003)
Ileret (KNM-ER 3884)Partial calvariaKenya162–∞Bräuer et al. (1997)
Jebel IrhoudThree calvaria, mandible, fragments of seven individualsMorocco144–176Hublin (2001); Smith et al. (2007)
(Ngaloba Beds)
CraniumTanzania130Day et al. (1980); Hay (1987)
SingaCalvariaSudan131–135McDermott et al. (1996)
Lake EyasiCalvariaTanzania88–132Mehlman (1984, 1987); Domínguez-Rodrigo et al., 2008
  1. *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.

Download links

A two-part list of links to download the article, or parts of the article, in various formats.

Downloads (link to download the article as PDF)

Open citations (links to open the citations from this article in various online reference manager services)

Cite this article (links to download the citations from this article in formats compatible with various reference manager tools)

  1. Lee R Berger
  2. John Hawks
  3. Paul HGM Dirks
  4. Marina Elliott
  5. Eric M Roberts
Homo naledi and Pleistocene hominin evolution in subequatorial Africa
eLife 6:e24234.