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Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa - PubMed

  • ️Thu Jan 01 2015

doi: 10.7554/eLife.09560.

John Hawks  1 , Darryl J de Ruiter  1 , Steven E Churchill  1 , Peter Schmid  1 , Lucas K Delezene  1 , Tracy L Kivell  1 , Heather M Garvin  1 , Scott A Williams  1 , Jeremy M DeSilva  1 , Matthew M Skinner  1 , Charles M Musiba  1 , Noel Cameron  1 , Trenton W Holliday  1 , William Harcourt-Smith  1 , Rebecca R Ackermann  2 , Markus Bastir  1 , Barry Bogin  1 , Debra Bolter  1 , Juliet Brophy  1 , Zachary D Cofran  1 , Kimberly A Congdon  1 , Andrew S Deane  1 , Mana Dembo  1 , Michelle Drapeau  3 , Marina C Elliott  1 , Elen M Feuerriegel  1 , Daniel Garcia-Martinez  1 , David J Green  1 , Alia Gurtov  1 , Joel D Irish  1 , Ashley Kruger  1 , Myra F Laird  1 , Damiano Marchi  1 , Marc R Meyer  1 , Shahed Nalla  1 , Enquye W Negash  1 , Caley M Orr  1 , Davorka Radovcic  1 , Lauren Schroeder  1 , Jill E Scott  1 , Zachary Throckmorton  1 , Matthew W Tocheri  4 , Caroline VanSickle  1 , Christopher S Walker  1 , Pianpian Wei  1 , Bernhard Zipfel  1

Affiliations

Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa

Lee R Berger et al. Elife. 2015.

Abstract

Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.

Keywords: Dinaledi Chamber; Homo naledi; evolutionary biology; genomics; hominin; none; paleoanthropology.

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Conflict of interest statement

The authors declare that no competing interests exist.

Figures

Figure 1.
Figure 1.. Dinaledi skeletal specimens.

The figure includes approximately all of the material incorporated in this diagnosis, including the holotype specimen, paratypes and referred material. These make up 737 partial or complete anatomical elements, many of which consist of several refitted specimens. Specimens not identified to element, such as non-diagnostic long bone or cranial fragments, and a subset of fragile specimens are not shown here. The ‘skeleton’ layout in the center of the photo is a composite of elements that represent multiple individuals. This view is foreshortened; the table upon which the bones are arranged is 120-cm wide for scale. DOI:

http://dx.doi.org/10.7554/eLife.09560.003
Figure 2.
Figure 2.. Holotype specimen of Homo naledi, Dinaledi Hominin 1 (DH1).

U.W. 101-1473 cranium in (A) posterior and (B) frontal views (frontal view minus the frontal fragment to show calvaria interior). U.W. 101-1277 maxilla in (C) medial, (D) frontal, (E) superior, and (F) occlusal views. (G) U.W. 101-1473 cranium in anatomical alignment with occluded U.W. 101-1277 maxilla and U.W. 101-1261 mandible in left lateral view. U.W. 101-1277 mandible in (H) occlusal, (I) basal, (J) right lateral, and (K) anterior views. Scale bar = 10 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.019
Figure 3.
Figure 3.. Cranial paratypes.

(A) DH2, right lateral view. (B) DH5, left lateral view. (C) DH4, right lateral view. (D) DH4, posterior view. Scale bar = 10 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.005
Figure 4.
Figure 4.. Paratype DH3.

(A) Frontal view. (B) Left lateral view, with calvaria in articulation with the mandible (U.W. 101-361). (C) Basal view. Mandible in (D) medial view; (E) occlusal view; (F) basal view. DH3 was a relatively old individual at time of death, with extreme tooth wear. Scale bar = 10 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.006
Figure 5.
Figure 5.. U.W. 101-377 mandible.

(A) Lateral view; (B) medial view; (C) basal view; (D) occlusal view. (D) The distinctive mandibular premolar morphology with elongated talonids in unworn state. Scale bar = 2 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.007
Figure 6.
Figure 6.. Hand 1.

Palmar view on left; dorsal view on right. This hand was discovered in articulation and all bones are represented except for the pisiform. The proportions of digits are humanlike and visually apparent, as are the expanded distal apical tufts on all digits, the robust pollical ray, and the unique first metacarpal morphology. DOI:

http://dx.doi.org/10.7554/eLife.09560.008
Figure 7.
Figure 7.. U.W. 101-1391 paratype femur.

(A) Medial view; (B) posterior view; (C) lateral view; (D) anterior view. The femur neck is relatively long and anteroposteriorly compressed. The anteversion of the neck is evident in medial view. Scale bar = 2 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.009
Figure 8.
Figure 8.. U.W. 101-484 paratype tibia.

(A) Anterior view; (B) medial view; (C) posterior view; (D) lateral view. The tibiae are notably slender for their length. Scale bar = 10 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.010
Figure 9.
Figure 9.. Foot 1 in (A) dorsal view; and (B) medial view.

(C) Proximal articular surfaces of the metatarsals of Foot 1, shown in articulation to illustrate transverse arch structure. Scale bar = 10 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.011
Figure 10.
Figure 10.. Maximum tibia length in H. naledi and other hominins.

Maximum tibia length for U.W. 101-484, compared to other nearly complete hominin tibia specimens. Australopithecus afarensis represented by A.L. 288-1 and KSD-VP-1/1 (Haile-Selassie et al., 2010); Homo erectus represented by D3901 from Dmanisi and KNM-WT 15000; Homo habilis by OH 35; Homo floresiensis by LB1 and LB8 (Brown et al., 2004; Morwood et al., 2005). Chimpanzee and contemporary European ancestry humans from Cleveland Museum of Natural History (Lee, 2001); Andaman Islanders from Stock (2013). Vertical lines represent sample ranges; bars represent 1 standard deviation. DOI:

http://dx.doi.org/10.7554/eLife.09560.015
Figure 11.
Figure 11.. Virtual reconstruction of the endocranium of the larger composite cranium from DH1 and DH2 overlaid with the ectocranial surfaces.

(A) Lateral view. (B) Superior view. The resulting estimate of endocranial volume is 560cc. Scale bar = 10 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.016
Figure 12.
Figure 12.. Brain size and tooth size in hominins.

The buccolingual breadth of the first maxillary molar is shown here in comparison to endocranial volume for many hominin species. H. naledi occupies a position with relatively small molar size (comparable to later Homo) and relatively small endocranial volume (comparable to australopiths). The range of variation within the Dinaledi sample is also fairly small, in particular in comparison to the extensive range of variation within the H. erectus sensu lato. Vertical lines represent the range of endocranial volume estimates known for each taxon; each vertical line meets the horizontal line representing M1 BL diameter at the mean for each taxon. Ranges are illustrated here instead of data points because the ranges of endocranial volume in several species are established by specimens that do not preserve first maxillary molars. DOI:

http://dx.doi.org/10.7554/eLife.09560.017
Figure 13.
Figure 13.. Selected pelvic specimens of H. naledi.

U.W. 101-1100 ilium in (A) lateral view showing a weak iliac pillar relatively near the anterior edge of the ilium, with no cristal tubercle development; (B) anterior view, angled to demonstrate the degree of flare, which is clear in comparison to the subarcuate surface. U.W. 101-723 immature sacrum in (C) anterior view; and (D) superior view. U.W. 101-1112 ischium in (E) lateral view; and (F) anterior view, demonstrating relatively short tuberacetabular diameter. Scale bar = 2 cm. DOI:

http://dx.doi.org/10.7554/eLife.09560.018
Figure 14.
Figure 14.. First metacarpals of H. naledi.

Seven first metacarpals have been recovered from the Dinaledi Chamber. U.W. 101-1321 is the right first metacarpal of the associated Hand 1 found in articulation. U.W. 101-1282 and U.W. 101-1641 are anatomically similar left and right first metacarpals, which we hypothesize as antimeres, both were recovered from excavation. U.W. 101-007 was collected from the surface of the chamber, and exhibits the same distinctive morphological characteristics as all the first metacarpals in the assemblage. All of these show a marked robusticity of the distal half of the bone, a very narrow, ‘waisted’ appearance to the proximal shaft and proximal articular surface, prominent crests for attachment of M. opponens pollicis and M. first dorsal interosseous, and a prominent ridge running down the palmar aspect of the bone. The heads of these metacarpals are dorsopalmarly flat and strongly asymmetric, with an enlarged palmar-radial protuberance. These distinctive features are present among all the first metacarpals in the Dinaledi collection, and are absent from any other hominin sample. Their derived nature is evident in comparison to apes and other early hominins, here illustrated with a chimpanzee first metacarpal and the MH2 first metacarpal of Australopithecus sediba. DOI:

http://dx.doi.org/10.7554/eLife.09560.004
Figure 15.
Figure 15.. Posterior view of the virtual reconstruction of DH3.

The resultant mirror image is displayed in blue. The antimeres were aligned by the frontal crest and sagittal suture using the Manual Registration function in GeoMagic Studio 14.0. DOI:

http://dx.doi.org/10.7554/eLife.09560.020
Figure 16.
Figure 16.. Virtual reconstruction of (A) DH2 and (B) occipital portion of DH1.

The actual specimen displays its original coloration and the mirror imaged portion is illustrated in blue. DOI:

http://dx.doi.org/10.7554/eLife.09560.021
Figure 17.
Figure 17.. Postero-lateral view of the virtual reconstruction of a composite cranium from DH3 and DH4.

(A) The surface scan of DH3 was mirror imaged and merged as described in Supplementary Note 8. (B) The scan of DH4 was aligned to the DH3 model. (C) DH4 was then mirror imaged to complete the occipital contour (D). DOI:

http://dx.doi.org/10.7554/eLife.09560.022
Figure 18.
Figure 18.. Virtual reconstruction of a composite cranium from DH1 and DH2.

The surface model of DH2 (blue), consisting of the original scan merged with the mirror image, was then uploaded and aligned with the mirror-imaged DH1 model (pink). Note the similarity in size and shape between DH1 and DH2 observed in the posterior (A) anterior (B) lateral (C) and superior (D) views. DOI:

http://dx.doi.org/10.7554/eLife.09560.023
Figure 19.
Figure 19.. Virtual reconstruction of the endocranium of the composite cranium from DH3 and DH4.

(A) Lateral view. (B) Superior view. (C) Inferior view. In all views, anterior is to towards the left. DOI:

http://dx.doi.org/10.7554/eLife.09560.024
Figure 20.
Figure 20.. Virtual reconstruction of the endocranium of the composite cranium from DH3 and DH4 overlaid with the ectocranial surfaces.

(A) Lateral view. (B) Superior view. DOI:

http://dx.doi.org/10.7554/eLife.09560.025
Figure 21.
Figure 21.. Virtual reconstruction the DH3/DH4 cranial base using a model of Sts 19.

(A) Right lateral view. (B) Left lateral view. (C) Posterior view. (D) Inferior view. DOI:

http://dx.doi.org/10.7554/eLife.09560.026
Figure 22.
Figure 22.. Virtual reconstruction the DH3/DH4 endocranial volume using a cranial base model of Sts 19.

Right lateral view. DOI:

http://dx.doi.org/10.7554/eLife.09560.027

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