• Figure 1.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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.
    Download figureOpen in new tabFigure 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

  • Table 1.

    Cranial and mandibular measurements for H. naledi, early hominins, and modern humans

    DOI: http://dx.doi.org/10.7554/eLife.09560.012

    Measurement definitions as in Wood (1991)P. aethiopicusP. boiseiP. robustusAu. afarensisAu. africanusAu. sedibaH. nalediH. habilisH. rudolfensisH. erectusMP HomoH. sapiens
    Cranium
     Cranial capacity41048549345746742051361077686512661330
     Porion height6727486706781779094101112
     Posterior cranial length35847546044656070799981
     Bi-parietal breadth994989099100103107118129142132
     Bi-temporal breadth10110109108115104101107112126131146127
     Closest approach of temporal linescrest*crest*crest*crest*21565235517210196
     Supraorbital height index465350516056566459566271
     Minimum post-orbital breadth626670776770687578899697
     Superior facial breadth4910010710995868697113110124107
     Post-orbital constriction index6261646981797274818091
     EAM area (as an ellipse)778010370963876958561
     Root of zygomatic process originP4P4P3 to M1P4 to M1P4 to M1P4P3 to P4P4 to M1P4 to M1P4 to M1M1M1
     Petromedian angle13750455031335548525546
    Maxilloalveolar process
     Maxilloalveolar length87947869677163576568666955
     Maxilloalveolar breadth88837669686663716872707262
     Palate breadth91324035303629443840385640
     Palate depth at incisive fossa31110109105101311109
     Palate depth at M110371811111310101216151813
    Mandible
     Symphysis height141374942393732333137353434
     Symphysis width142262825202118182024181714
     Symphysis area at M1 (as an ellipse)1467571114835623606452467393723519474365
     Corpus height at M1150384236343230262936313128
     Corpus breadth at M1151252926202118162022191913
     Corpus area at M1 (as an ellipse)152742955736540539405326425631458469296
     Mental foramen height index§515054585350404649484850
    • At least in presumed males.

    • Post-orbital breadth/superior facial breadth × 100.

    • Following the formula (π × (corpus height/2) × (corpus breadth/2)).

    • Height of mental foramen from alveolar border relative to corpus height at the mental foramen.

    • MP, Middle Pleistocene.

    • Unless otherwise indicated measurements are defined as in Wood (1991). Chord distances are in mm. Data for H. naledi collected from original fossils or laser scans by DJdeR and HMG; comparative data collected by DJdeR on original fossils and casts and supplemented by data from Wood (1991).

  • Table 2.

    Dental measures for H. naledi and comparative hominin species

    DOI: http://dx.doi.org/10.7554/eLife.09560.013

    Maxillary
    I1I2CP3P4M1M2M3
    MDLLMDLLMDLLMDBLMDBLMDBLMDBLMDBL
    Au. anamensisn352677653121010898
    mean10.88.77.311.010.69.912.68.913.611.512.913.014.412.514.2
    range9.1–12.48.2–9.37.0–7.59.9–12.39.1–11.88.2–11.810.1–14.37.2–12.112.6–14.27.8–14.39.0–16.710.9–16.312.9–16.111.1–15.713.0–15.7
    Au. afarensisn7899151512101812161310111111
    mean10.78.47.57.29.910.88.812.49.112.412.013.412.914.612.714.5
    range9.9–11.87.1–9.76.6–8.26.2–8.18.8–11.69.3–12.57.7–9.711.3–13.47.6–10.811.1–14.510.5–13.812.0–15.012.1–13.613.4–15.210.9–14.813.1–16.3
    Au. africanusn15151110161326252020212023242728
    mean10.78.36.96.89.910.39.212.79.513.412.913.914.115.714.216.0
    range9.4–12.57.4–9.15.8–8.05.6–7.98.8–11.08.7–12.08.5–10.210.7–14.57.2–11.012.4–15.311.7–14.412.9–15.312.1–16.312.8–17.911.2–16.913.1–18.6
    Au. sediban1111111111111122
    mean10.16.97.26.69.08.89.011.29.312.112.912.012.913.713.013.5
    range12.6–13.312.9–14.1
    H. naledin548109101077121311977
    mean9.46.56.66.28.18.68.010.58.111.011.611.712.212.811.612.4
    range8.8–9.86.3–7.06.3–7.05.8–6.67.3–8.98.0–9.67.7–8.49.8–11.07.7–8.710.5–11.210.5–12.411.2–12.411.0–13.011.9–13.611.0–12.711.4–13.4
    H. habilisn224423778813137777
    mean10.68.07.46.69.09.89.011.99.212.112.713.012.714.312.314.7
    range10.1–11.17.3–8.76.7–8.16.0–7.98.5–9.48.5–11.68.1–9.611.0–12.78.5–9.911.0–13.111.6–13.912.1–14.111.8–13.513.5–16.211.3–13.913.2–16.6
    H. rudolfensisn11111122222211
    mean12.37.711.512.510.513.610.212.514.014.014.315.813.313.5
    range9.7–10.711.1–13.813.9–14.213.3–14.814.1–14.614.1–17.6
    H. erectusn111266121227273029343222221616
    mean10.38.17.78.09.510.08.511.88.111.612.213.212.013.310.512.8
    range8.1–12.67.0–11.76.0–8.36.9–8.58.5–11.19.0–11.87.1–10.19.5–13.87.0–9.49.9–13.410.1–14.611.0–15.910.3–13.610.9–15.58.7–14.710.4–15.8
    H. neanderthalensisn28373541282916172119232427282221
    mean9.78.58.08.48.810.18.010.67.810.611.612.310.912.59.912.3
    range8.2–11.87.3–9.95.8–9.35.8–9.97.2–10.07.6–11.46.6–9.38.4–11.85.9–11.58.3–11.79.5–13.511.0–14.28.9–15.910.8–14.68.2–11.49.8–14.6
    H. heidelbergensisn21231921272925252223252424232627
    mean9.67.87.77.88.89.87.910.67.610.311.211.910.212.38.911.6
    range8.7–10.77.1–9.97.2–8.47.3–8.68.1–11.08.8–11.87.1–9.09.2–12.27.0–8.89.1–11.59.9–12.310.3–13.28.1–12.111.1–13.87.6–11.010.0–13.2
    MP/LP African Homon6678446610101414202099
    mean9.07.87.47.28.99.78.410.88.110.812.313.211.012.99.211.7
    range6.3–10.96.6–8.76.0–9.36.1–8.58.2–9.58.8–10.08.1–8.79.9–11.87.5–9.39.4–12.810.4–14.012.0–15.07.8–13.011.0–15.07.6–10.210.0–13.2
    Mandibular
    I1I2CP3P4M1M2M3
    MDLLMDLLMDLLMDBLMDBLMDBLMDBLMDBL
    Au. anamensisn21437788889107788
    mean6.97.47.88.310.010.412.49.29.111.312.912.314.013.415.313.4
    range6.8–6.96.6–8.77.9–8.66.6–13.99.2–11.411.3–13.48.6–10.07.4–9.89.6–13.211.6–13.810.2–14.813.0–15.912.3–14.913.7–17.012.1–15.2
    Au. afarensisn7876131627262421322631272623
    mean6.77.16.78.08.810.49.610.69.811.013.112.614.313.415.313.5
    range5.6–7.75.6–8.05.0–8.06.7–8.87.5–11.78.0–12.47.9–12.68.9–13.87.7–11.49.8–12.810.1–14.811.0–14.012.1–16.511.1–15.213.4–18.111.3–15.3
    Au. africanusn11121213232520212523293238383435
    mean6.26.77.27.99.410.19.711.510.411.614.013.015.714.516.314.6
    range4.8–6.95.7–7.95.6–8.16.6–9.28.5–10.78.2–12.18.8–11.09.9–13.98.7–12.39.3–13.212.4–15.811.2–15.114.2–17.712.8–16.813.5–18.512.2–16.8
    Au. sediban11221111222222
    mean5.96.67.78.08.19.28.89.713.111.414.512.814.913.2
    range7.3–8.07.4–8.613.1–13.111.3–11.514.4–14.512.3–13.214.9–14.912.5–13.6
    H. naledin7756779106611119965
    mean6.15.46.95.97.17.19.08.88.79.112.210.713.311.213.412.1
    range5.7–7.05.3–5.96.6–7.45.9–6.06.4–7.56.9–7.48.2–9.48.2–9.78.3–9.08.5–10.211.3–12.710.3–11.412.3–14.010.7–12.212.9–13.711.7–12.8
    H. habilisn2222324433554444
    mean6.46.87.47.68.79.09.69.69.910.513.711.915.013.515.413.3
    range6.4–6.56.7–7.07.2–7.77.6–7.67.6–9.67.9–10.09.0–10.68.6–11.19.0–10.59.9–11.013.0–14.810.9–12.814.2–15.712.0–15.114.8–15.912.4–14.4
    H. rudolfensisn1113366556533
    mean5.46.78.39.911.110.111.414.012.716.013.716.414.1
    range9.0–10.79.5–12.38.8–11.89.8–12.212.8–15.211.4–13.214.0–18.312.7–14.915.6–17.013.1–14.6
    H. erectusn11121416141630302526434341402627
    mean6.26.477.28.79910.18.710.112.711.913.312.512.711.7
    range4.8–7.45.8–7.15.3–8.16.4–8.57.0–10.38.0–10.47.0–12.08.2–12.07.2–10.38.0–12.59.9–14.810.1–13.311.3–15.310.8–14.310.0–15.210.0–14.2
    H. neanderthalensisn9162331364120212325384026271820
    mean5.67.26.87.87.88.87.99.17.89.411.811.112.111.312.011.0
    range4.2–6.45.2–8.85.9–7.56.8–9.06.7–8.86.8–10.36.6–9.18.0–10.36.5–9.48.5–10.510.1–13.610.2–12.99.3–14.08.8–12.411.2–13.99.9–12.2
    H. heidelbergensisn21221920232422222626292929293232
    mean5.66.76.57.37.68.77.98.97.28.711.310.611.210.511.510.0
    range4.8–6.56.0–7.56.0–7.26.6–8.06.9–9.07.3–10.07.2–9.07.6–11.66.6–8.87.2–11.710.4–13.89.6–13.09.7–14.68.5–13.99.7–13.28.6–12.5
    MP/LP African Homon55888888129161620201313
    mean6.06.86.87.28.89.68.69.88.610.313.111.812.511.712.411.5
    range5.7–6.46.1–7.25.6–8.36.4–8.07.8–10.08.8–10.37.7–9.08.6–11.26.9–9.69.3–11.410.7–14.210.0–13.010.8–15.09.2–13.610.6–13.59.9–12.7
    • MP, Middle Pleistocene and LP, Late Pleistocene.

  • Table 3.

    Dinaledi body mass estimates from femur specimens preserving subtrochanteric diameters

    DOI: http://dx.doi.org/10.7554/eLife.09560.014

    Specimen IDSideAP subtrochanteric breadthML subtrochanteric breadthMass (a)Mass (b)
    U.W. 101-002R18.523.640.044.7
    U.W. 101-003R21.631.454.555.8
    U.W. 101-018R18.123.839.744.4
    U.W. 101-226L19.124.041.345.7
    U.W. 101-1136R16.925.539.744.4
    U.W. 101-1391R18.823.940.845.3
    U.W. 101-1475L18.829.046.549.7
    U.W. 101-1482L20.728.949.752.1
    • Regression equations described in ‘Materials and methods’. Mass (a) based on forensic statures from European individuals. Mass (b) based on multiple population sample. The two estimates diverge somewhat for smaller femora.