Modern humans, or Homo sapiens, are now the only living species in their genus. But as recently as 100,000 years ago, there were several other species that belonged to the genus Homo. Together with modern humans, these extinct human species, our immediate ancestors and their close relatives, are collectively referred to as ‘hominins’.

Now Berger et al. report the recent discovery of an extinct species from the genus Homo that was unearthed from deep underground in what has been named the Dinaledi Chamber, in the Rising Star cave system in South Africa. The species was named Homo naledi; ‘naledi’ means ‘star’ in Sotho (also called Sesotho), which is one of the languages spoken in South Africa.

The unearthed fossils were from at least 15 individuals and include multiple examples of most of the bones in the skeleton. Based on this wide range of specimens from a single site, Berger et al. describe Homo naledi as being similar in size and weight to a small modern human, with human-like hands and feet. Furthermore, while the skull had several unique features, it had a small braincase that was most similar in size to other early hominin species that lived between four million and two million years ago. Homo naledi's ribcage, shoulders and pelvis also more closely resembled those of earlier hominin species than those of modern humans.

The Homo naledi fossils are the largest collection of a single species of hominin that has been discovered in Africa so far and, in a related study, Dirks et al. describe the setting and context for these fossils. However, since the age of the fossils remains unclear, one of the next challenges will be to date the remains to provide more information about the early evolution of humans and their close relatives.

Fossil hominins were first recognized in the Dinaledi Chamber in the Rising Star cave system in October 2013. During a relatively short excavation, our team recovered an extensive collection of 1550 hominin specimens, representing nearly every element of the skeleton multiple times (Figure 1), including many complete elements and morphologically informative fragments, some in articulation, as well as smaller fragments many of which could be refit into more complete elements. The collection is a morphologically homogeneous sample that can be attributed to no previously-known hominin species. Here we describe this new species, Homo naledi. We have not defined H. naledi narrowly based on a single jaw or skull because the entire body of material has informed our understanding of its biology.

Figure 1

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Order Primates LINNAEUS 1758

Suborder Anthropoidea MIVART 1864

Superfamily Hominoidea GRAY 1825

Family Hominidae GRAY 1825

Tribe Hominini GRAY 1825

Genus Homo LINNAEUS 1758

Homo naledi sp. nov. urn:lsid:zoobank.org:pub:00D1E81A-6E08-4A01-BD98-79A2CEAE2411

Holotype, paratypes, and referred materials

Holotype

Dinaledi Hominin 1 (DH1) comprises the partial calvaria, partial maxilla, and nearly complete mandible of a presumed male individual, based on size and morphology within the sample (Figure 2; Supplementary file 1). The holotype was recovered in situ during excavations within the Dinaledi Chamber in March of 2014, embedded in unconsolidated fine clay matrix (Dirks et al., 2015). The holotype is housed in the Evolutionary Studies Institute at the University of the Witwatersrand, Johannesburg, South Africa.

Figure 2

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Paratypes

Dinaledi Hominin 2 (DH2) is a partial calvaria that preserves parts of the frontal, left and right parietals, right temporal, and occipital (Figure 3; Supplementary file 1). Dinaledi Hominin 3 (DH3) is a partial calvaria of a presumed female individual that preserves parts of the frontal, left parietal, left temporal, and sphenoid (Figure 4, Supplementary file 1). Dinaledi Hominin 4 (DH4) is a partial calvaria that preserves parts of the right temporal, right parietal, and occipital (Figure 3; Supplementary file 1). Dinaledi Hominin 5 (DH5) is a partial calvaria that preserves part of the left temporal and occipital (Figure 3; Supplementary file 1). U.W. 101-377 is a mandibular fragment that preserves dental anatomy in an unworn state; at present it cannot be definitively associated with any of these Dinaledi Hominin (DH) individuals, and indeed might represent another individual (Figure 5; Supplementary file 1). These cranial specimens agree closely in nearly all morphological details where they overlap in areas preserved except those we interpret as related to sex.

Figure 3

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Figure 4

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Figure 5

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Dinaledi hand 1 (H1) is a nearly complete (missing only the pisiform) right hand, found articulated in association, comprising specimens U.W. 101-1308 to −1311, −1318 to −1321, −1325 to −1329, −1351, −1464, and −1721 to −1732 (Figure 6; Supplementary file 1). U.W. 101-1391 is a proximal right femur preserving part of the head, the neck, some of the lesser and greater trochanter, and the proximal shaft (Figure 7; Supplementary file 1). U.W. 101-484 is a right tibial diaphysis missing only the proximal end (Figure 8; Supplementary file 1). Dinaledi foot 1 (F1) is a partial foot skeleton missing only the medial cuneiform and the phalanges of rays II–V. Foot 1 is composed of specimens U.W. 101-1322, −1417 to −1419, −1439, −1443, −1456 to −1458, −1551, −1553, −1562, and −1698 (Figure 9; Supplementary file 1).

Figure 6

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Figure 7

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Figure 8

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Figure 9

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Referred material

Referred material is also listed in Supplementary file 1. We refer to H. naledi all hominin material from the Dinaledi collection that can be identified to element; in total, the holotypes, paratypes and referred material comprise 737 partial or complete anatomical elements.

Specimen numbers in the collection are assigned at the point of excavation. Later laboratory analyses allowed us to refit specimens into more complete elements, which we have used as units of anatomical study. Here we refer to refitted elements by only a single specimen number; either the number of the most constitutive specimen, or the first diagnostic part to be discovered. DH designations are reserved for clearly associated individuals; at this time these are limited to the five partial crania designated above. Future excavation and analyses will certainly uncover more refits among specimens. As refits are found, all numbers assigned to refitted elements will remain stable, and all numbers in Supplementary file 1 will be retained.

The collection is morphologically homogeneous in all duplicated elements, except for those anatomical features that normally reflect body size or sex differences in other primate taxa. Therefore, although we refer to the holotype and the paratypes for differential diagnoses; the section describing the overall anatomy encompasses all morphologically informative specimens.

This comprehensive differential diagnosis is based upon cranial, dental and postcranial characters. The hypodigms used for other species are detailed in the ‘Materials and methods’. We examined original specimens for most species, except where indicated in the ‘Materials and methods’; when we relied on other sources for anatomical observations we indicate this. A summary of traits of H. naledi in comparison to other species is presented in Supplementary file 2. Comparative cranial and mandibular measures are presented in Table 1, and comparative dental measures are provided in Table 2.

H. naledi exhibits anatomical features shared with Australopithecus, other features shared with Homo, with several features not otherwise known in any hominin species. This anatomical mosaic is reflected in different regions of the skeleton. The morphology of the cranium, mandible, and dentition is mostly consistent with the genus Homo, but the brain size of H. naledi is within the range of Australopithecus. The lower limb is largely Homo-like, and the foot and ankle are particularly human in their configuration, but the pelvis appears to be flared markedly like that of Au. afarensis. The wrists, fingertips, and proportions of the fingers are shared mainly with Homo, but the proximal and intermediate manual phalanges are markedly curved, even to a greater degree than in any Australopithecus. The shoulders are configured largely like those of australopiths. The vertebrae are most similar to Pleistocene members of the genus Homo, whereas the ribcage is wide distally like Au. afarensis.

H. naledi has a range of body mass similar to small-bodied modern human populations, and is similar in estimated stature to both small-bodied humans and the largest known australopiths. We estimated body mass from eight femoral specimens for which subtrochanteric diameters can be measured (‘Materials and methods’), with results ranging between 39.7 kg and 55.8 kg (Table 3). No femur specimen is sufficiently complete to measure femur length accurately, but the U.W. 101-484 tibia preserves nearly its complete length, allowing a tibia length estimate of 325 mm (Figure 10). Estimates for the stature of this individual based on African human population samples range between 144.5 and 147.8 mm. Again, this stature estimate is similar to small-bodied modern human populations. It is within the range estimated for Dmanisi postcranial elements (Lordkipanidze et al., 2007), and slightly smaller than estimated for early Homo femoral specimens KNM-ER 1472 and KNM-ER 1481. Some large australopiths also had long tibiae and presumably comparably tall statures, as evidenced by the KSD-VP 1/1 skeleton from Woranso-Mille (Haile-Selassie et al., 2010).

Figure 10

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The endocranial volume of all H. naledi specimens is clearly small compared to most known examples of Homo. We combined information from the most complete cranial vault specimens to arrive at an estimate of endocranial volume for both larger (presumably male) and smaller (presumably female) individuals (larger composite depicted in Figure 11). The resulting estimates of approximately 560cc and 465cc, respectively, overlap entirely with the range of endocranial volumes known for australopiths. Within the genus Homo, only the smallest specimens of H. habilis, one single H. erectus specimen, and H. floresiensis overlap with these values.

Figure 11

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Despite its small vault size, the cranium of H. naledi is structurally similar to those of early Homo. Frontal bossing is evident, as is a marked degree of parietal bossing. There is no indication of metopic keeling, though there is slight parasagittal keeling between bregma and lambda, and some prelambdoidal flattening. The cranial vault bones are generally thin, becoming somewhat thicker in the occipital region. The supraorbital torus is well developed, though weakly arched, and is bounded posteriorly by a well-developed supratoral sulcus. The lateral corners of the supraorbital torus are rounded and relatively thin. The temporal lines are widely spaced, and there is no indication of a sagittal crest or temporal/nuchal cresting. The temporal crest is positioned on the posterior aspect of the lateral supraorbital torus, rather than on the superior aspect as in australopiths. At the posteroinferior extent of the temporal lines, they curve anteroinferiorly presenting a well-developed angular torus. The crania have a pentagonal outline in posterior view, with the greatest vault breadth located in the supramastoid region. The nuchal region exhibits sexually dimorphic development of nuchal muscle markings and the external occipital protuberance, and there is a clear indication of a tuberculum linearum in addition to the external occipital protuberance. In superior view the vault tapers from posterior to anterior, though post-orbital constriction is slight. The squamosal suture is low and gently curved, and parietal striae are well defined. The lateral margins of the orbits face laterally. A small zygomaticofacial foramen is typically present near the center of the zygomatic bone. The root of the zygomatic process of the maxilla is anteriorly positioned, at the level of the P³ or the P⁴. There is no indication of a zygomatic prominence, and the zygomatic arches do not flare laterally to any extent. The root of the zygomatic process of the temporal is angled downwards approximately 30° relative to FH. The root of the zygomatic process of the temporal begins to laterally expand above the level of the mandibular fossa, rather than above the level of the EAM as in australopiths. The mandibular fossa is somewhat large, and moderately deep. The articular eminence of the mandibular fossa is saddle-shaped, and oriented posteroinferiorly. Almost the entire mandibular fossa is positioned medial to the temporal squama. The entoglenoid process is elongated and faces primarily laterally. The postglenoid process is small and closely appressed to the tympanic, forming part of the posterior wall of the fossa. The petrotympanic is distinctly coronally oriented. The vaginal process is small but distinct. The crista petrosa is weakly developed and not notably sharpened. There is a strong Eustachian process. The external auditory meatus is small, oval-shaped, and obliquely oriented, and a distinct suprameatal spine is present. The mastoid region is slightly laterally inflated. The mastoid process is triangular in cross-section, with a rounded apex and a mastoid crest. The digastric groove is deep and narrow, alongside a marked juxtamastoid eminence. The canine juga are weakly developed and there is no indication that anterior pillars would have been present. A shallow, ill-defined canine fossa is indicated. The nasoalveolar clivus is flat and square-shaped. The parabolic-shaped palate is broad and anteriorly shallow, becoming deeper posteriorly.

The mandibular dentition of H. naledi is arranged in a parabolic arch. The alveolar and basal margins of the corpus diverge slightly. A single, posteriorly opening mental foramen is positioned slightly above the mid-corpus level, between the position of the P₃ and the P₄. The mandibular corpus is relatively gracile, with a well-developed lateral prominence whose maximum extent is typically at the M₂. A slight supreme lateral torus (of Dart) weakly delineates the extramolar sulcus from the lateral corpus. The superior lateral torus is moderately developed, running anteriorly to the mental foramen where it turns up to reach the P₃ jugum. The marginal torus is moderately developed, and defines a moderate intertoral sulcus. The posterior and anterior marginal tubercles are indicated only as slight roughenings of bone. The gracile mandibular symphysis is vertically oriented. A well-developed mental protuberance and weak lateral tubercles are delineated by a slight mandibular incisure, thereby presenting a weak mentum osseum. The post-incisive planum is steeply inclined and not-shelf-like. There is no superior transverse torus, while a weak, basally oriented inferior transverse torus is present. The anterior and posterior subalveolar fossae are continuous and deep, overhung by a well-developed alveolar prominence. The extramolar sulcus is moderately wide. The root of the ramus of the mandible originates high on the corpus at the level of the M₂. Strong ectoangular tuberosities are indicated. A large mandibular foramen is present, with a diffusely defined mylohyoid groove.

Like the skull, the dentition of H. naledi compares most favorably to early Homo samples. Yet compared to samples of H. habilis, H. rudolfensis, and H. erectus, the teeth of H. naledi are comparatively quite small, similar in dimensions to much later samples of Homo. With both small post-canine teeth and a small endocranial volume, H. naledi joins Au. sediba and H. floresiensis in an area distinct from the general hominin relation of smaller post-canine teeth in species with larger brains (Figure 12).

Figure 12

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In comparison to H. habilis, H. rudolfensis, and H. erectus, the teeth of H. naledi are not only small, but also markedly simple in crown morphology. Maxillary and mandibular molars lack extensive crenulation, secondary fissures and supernumerary cusps. The M¹ has an equal-sized metacone and paracone, and has a slight expression of Carabelli's trait represented by a small cusp or shallow pit. I¹ exhibits slight occlusal curvature with trace marginal ridges and variably small tuberculum dentale. I² exhibits greater occlusal curvature and tuberculum dentale expression but neither upper incisor has double shovelling or interruption groove. The mandibular canines of H. naledi have a small occlusal area, and have a distal marginal cuspule as a topographically distinct expression of the cingular margin. The P₃ is double-rooted, fully bicuspid with metaconid and protoconid of approximately equal height and occlusal area separated by a distinct longitudinal groove, has a distally extensive talonid, and an occlusal outline approximately symmetrical with respect to the mesiodistal axis. P₄ likewise has a distally extensive talonid and approximately symmetrical occlusal outline (Figure 5). M₁ and M₂ lack cusp 6 and cusp 7, except for very slight expression in a small fraction of specimens, and have a very faint subvertical depression rather than a distinct or extensive protostylid. Like australopiths and some early Homo specimens, H. naledi has an increasing molar size gradient in the mandibular dentition (M1 < M2 < M3).

The lower limb of H. naledi is defined not only by a unique combination of primitive and derived traits, but also by the presence of unique features in the femur and tibia. Like all other bipedal hominins, H. naledi possesses a valgus knee and varus ankle. The femoral neck is long, anteverted, and anteroposteriorly compressed. Muscle insertions for the M. gluteus maximus are strong and the femur has a well-marked linea aspera with pilaster variably present. The patella is relatively anteroposteriorly thick. The tibia is mediolaterally compressed with a rounded anterior border, a large proximal attachment for the M. tibialis posterior, and a thin medial malleolus. The fibula is gracile with laterally oriented lateral malleolus, a relatively circular neck and a convex surface for the proximal attachment of the M. peroneus longus. Unique features in the lower limb of H. naledi include a depression in the superior aspect of the femoral neck that results in two mediolaterally oriented pillars inferoposteriorly and superoanteriorly, and a strong distal attachment of the pes anserinus on the tibia.

The foot and ankle of H. naledi are largely humanlike (Figure 9). The tibia stands orthogonally upon the talus, which is moderately wedged, with a mediolaterally flat trochlea having medial and lateral margins at even height, a form distinct from the strong keeling seen in OH 8 (H. habilis) and several tali from Koobi Fora. The talar head and neck exhibit strong, humanlike torsion; the horizontal angle is higher than in most humans, similar to that found in australopiths. The calcaneus is only moderately robust, but possesses the plantar declination of the retrotrochlear eminence and plantarly positioned lateral plantar process found in both modern humans and Au. afarensis. The peroneal trochlea is small, unlike that found in australopiths and similar only to that in H. sapiens and Neanderthals. The talonavicular, subtalar joints and calcaneocuboid joints are humanlike in possessing minimal ranges of motion and evidence for a locking, rigid midfoot. The intermediate and lateral cuneiforms are proximodistally elongated. The hallucal tarsometatarsal joint is flat and proximodistally aligned indicating that H. naledi possessed an adducted, non-grasping hallux. The head of the first metatarsal is mediolaterally expanded dorsally, indicative of a humanlike windlass mechanism. The foot possesses humanlike metatarsal lengths, head proportions, and torsion.

The phalanges are moderately curved, slightly more so than in H. sapiens. The only primitive anatomies found in the foot of H. naledi are the talar head and neck declination and sustentaculum tali angles, suggestive of a lower arched foot with a more plantarly positioned and horizontally inclined medial column than typically found in modern humans (Harcourt-Smith et al., 2015).

The axial skeleton presents a combination of derived (mainly aspects of the vertebrae) and seemingly primitive (mainly the ribs) traits. The preserved adult T10 and T11 vertebrae are proportioned similarly to Pleistocene Homo, with transverse process morphology most similar to Neandertals. The neural canals of these vertebrae are large in comparison to the diminutive overall size of the vertebrae, proportionally recalling Dmanisi H. erectus, Neandertals, and modern humans. The 11th rib is straight (uncurved), similar to Au. afarensis, and the shape of the upper rib cage appears narrow, as assessed from first and second rib fragments, suggesting that the thorax was pyramidal in shape. The 12th rib presents a robust shaft cross-section most similar to Neandertals. This combination is not found in other hominins and might reflect allometric scaling at a small trunk size.

The Dinaledi iliac blade is flared and shortened anteroposteriorly, resembling Au. afarensis or Au. africanus. The ischium is short with a narrow tuberoacetabular sulcus, and the ischiopubic and iliopubic rami are thick, resembling Au. sediba and H. erectus. This combination of iliac and ischiopubic features has not been found in other fossil hominins (Figure 13).

Figure 13

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The shoulder of H. naledi is configured with the scapula situated high and lateral on the thorax, short clavicles, and little or no torsion of the humerus. The humerus is notably slender for its length, with prominent greater and lesser tubercles bounding a deep bicipital groove, with a small, non-projecting humeral deltoid tuberosity and brachioradialis crest. Distally, the humerus has a wide lateral distodorsal pillar and narrow medial distodorsal pillar, and a medially-displaced olecranon fossa with septal aperture. The Dinaledi radius and ulna diaphyses exhibit little curvature. The radius has a globular radial tuberosity, prominent pronator quadratus crest, and reduced styloid process.

The hand shares many derived features of modern humans and Neandertals in the thumb, wrist, and palm, but has relatively long and markedly curved fingers (Kivell et al., 2015). The thumb is long relative to the length of the other digits, and includes a robust metacarpal with well-developed intrinsic (M. opponens pollicis and M. first dorsal interosseous) muscle attachments (Figure 6). The pollical distal phalanx is large and robust with a well-developed ridge along the distal border of a deep proximal palmar fossa for the attachment of flexor pollicis longus tendon. Ungual spines also project proximopalmarly from a radioulnarly expanded apical tuft with a distinct area for the ungual fossa. The wrist includes a boot-shaped trapezoid with an expanded non-articular palmar surface, an enlarged and palmarly-expanded trapezoid-capitate joint, and a trapezium-scaphoid joint that extends further onto the scaphoid tubercle. Overall, carpal shapes and articular configurations are very similar to those of modern humans and Neandertals, and unlike those of great apes and other extinct hominins. However, the H. naledi wrist lacks a third metacarpal styloid process, has a more radioulnarly oriented capitate-Mc2 joint, and has a relatively small trapezium-Mc1 joint compared to humans and Neandertals. Moreover, the phalanges are long (relative to the palm) and more curved than most australopiths.

The overall morphology of H. naledi places it within the genus Homo rather than Australopithecus or other early hominin genera. The shared derived features that connect H. naledi with other members of Homo occupy most regions of the H. naledi skeleton and represent distinct functional systems, including locomotion, manipulation, and mastication. Locomotor traits shared with Homo include the absolutely long lower limb, with well-marked linea aspera, strong M. gluteus maximus insertions, gracile fibula and generally humanlike ankle and foot. These aspects of the lower limb suggest enhanced locomotor performance for a striding gait. The H. naledi hand shares aspects of Homo morphology in the wrist, thumb and palm, pointing to enhanced object manipulation ability relative to australopiths, including Au. sediba (Kivell et al., 2011; Kivell et al., 2015). H. naledi lacks the powerful mastication that typifies Australopithecus and Paranthropus, with generally small teeth across the dentition, gracile mandibular corpus and symphysis, laterally-positioned temporal lines, slight postorbital constriction and non-flaring zygomatic arches. The upper limb, shoulder and ribcage have a more primitive morphological pattern, but do not preclude affiliating H. naledi with Homo, particularly considering that postcranial remains of H. habilis appear to reflect an australopith-like body plan (Johanson et al., 1986). Locomotor, manipulatory, and masticatory systems have both historical and current importance in defining Homo (Wood and Collard, 1999; Holliday, 2012; Antón et al., 2014), and H. naledi fits within our genus in these respects.

The structural configuration of the H. naledi cranium, beyond the functional aspects of mastication, is likewise shared with Homo. As in many specimens of H. erectus and H. habilis, the H. naledi vault includes a well-developed and moderately arched supraorbital torus, marked from the frontal squama by a continuous supratoral sulcus, frontal bossing. Further, as in many H. erectus crania, H. naledi exhibits a marked angular torus and occipital torus. The H. naledi face includes a flat and squared nasoalveolar clivus, comparable to H. rudolfensis (Leakey et al., 2012), and weak canine fossae. While its anatomy places it unambiguously within Homo, the H. naledi cranium and dentition lack many derived features shared by MP and LP Homo and H. sapiens. The australopith-like features of the postcranium, including the ribcage, shoulder, proximal femur, and relatively long, curved fingers, also depart sharply from the morphology present in MP humans and H. sapiens. The similarities of H. naledi to earlier members of Homo, including H. habilis, H. rudolfensis, and H. erectus, suggest that this species may be rooted within the initial origin and diversification of our genus.

The fossil record of early Homo and Homo-like australopiths has rapidly increased during the last 15 years, and this accumulating evidence has changed our perspective on the rise of our genus. Many skeletal and behavioral features observed to separate later Homo from earlier hominins were formerly argued to have arisen as a single adaptive package, including increased brain size, tool manipulation, increased body size, smaller dentition, and greater commitment to terrestrial long-distance walking or running (Wood and Collard, 1999; Hawks et al., 2000). But we now recognize that such features appeared in different combinations in different fossil samples (Antón et al., 2014). The Dmanisi postcranial sample (Lordkipanidze et al., 2007) and additional cranial remains of H. erectus from Dmanisi (Gabunia et al., 2000; Vekua et al., 2002; Lordkipanidze et al., 2013) and East Africa (Spoor et al., 2007; Leakey et al., 2012), demonstrate that larger brain size and body size did not arise synchronously with improved locomotor efficiency and adaptations to long-distance walking or running in H. erectus (Holliday, 2012; Antón et al., 2014). Further, the discovery of Au. sediba showed that a mosaic of Homo-like hand, pelvis and aspects of craniodental morphology can occur within a species with primitive body size, limb proportions, lower limb and foot morphology, thorax shape, vertebral morphology, and brain size (Berger et al., 2010; Carlson et al., 2011; Kivell et al., 2011; Churchill et al., 2013; DeSilva et al., 2013; Schmid et al., 2013). H. naledi presents yet a different combination of traits. This species combines a humanlike body size and stature with an australopith-sized brain; features of the shoulder and hand apparently well-suited for climbing with humanlike hand and wrist adaptations for manipulation; australopith-like hip mechanics with humanlike terrestrial adaptations of the foot and lower limb; small dentition with primitive dental proportions. In light of this evidence from complete skeletal samples, we must abandon the expectation that any small fragment of the anatomy can provide singular insight about the evolutionary relationships of fossil hominins.

A recent phylogenetic analysis of fossil hominins based on craniodental morphology placed Au. sediba at the base of the genus Homo (Dembo et al., 2015), in agreement with earlier analyses of this species (Berger et al., 2010). The cranial and dental affinities identified between Au. sediba and Homo include many features shared by H. naledi. But H. naledi and Au. sediba share different postcranial features with other species of Homo. Resolving the phylogenetic placement of H. naledi will require both postcranial and craniodental evidence to be integrated together. Such integration poses a challenge because of the poor representation of several key species both within and outside of Homo, most notably H. habilis, for which postcranial evidence is slight, and H. rudolfensis for which no associated postcranial remains are known. We propose the testable hypothesis that the common ancestor of H. naledi, H. erectus, and H. sapiens shared humanlike manipulatory capabilities and terrestrial bipedality, with hands and feet like H. naledi, an australopith-like pelvis and the H. erectus-like aspects of cranial morphology that are found in H. naledi. Enlarged brain size was evidently not a necessary prerequisite for the generally human-like aspects of manipulatory, locomotor, and masticatory morphology of H. naledi.

Although it contains an unprecedented wealth of anatomical information, the Dinaledi deposit remains undated (Dirks et al., 2015). Considering that H. naledi is a morphologically primitive species within our genus, an age may help elucidate the ecological circumstances within which Homo arose and diversified. If the fossils prove to be substantially older than 2 million years, H. naledi would be the earliest example of our genus that is more than a single isolated fragment. The sample would illustrate a model for the relation of adaptive features of the cranium, dentition and postcranium during a critical time interval that is underrepresented by fossil evidence of comparable completeness. A date younger than 1 million years ago would demonstrate the coexistence of multiple Homo morphs in Africa, including this small-brained form, into the later periods of human evolution. The persistence of such a species with clear adaptations for manipulation and grip, alongside MP humans or perhaps even alongside modern humans, would challenge many assumptions about the development of the archaeological record in Africa.

The depth of evidence of H. naledi may provide a perspective on the variation to be expected within fossil hominin taxa (Lordkipanidze et al., 2013; Bermúdez de Castro et al., 2014). The entire Dinaledi collection is remarkably homogeneous. There is very little size variation among adult elements within the collection. Eight body mass estimates from the femur (Table 2) have a standard deviation of only 4.3 kilograms, for a body mass coefficient of variation (CV) of only 9%. The CV of body mass within most human populations is substantially higher than this, with an average near 15% (McKellar and Hendry, 2009). Likewise, the size variation of cranial and dental elements is minimal. With 11 mandibular first molars, the CV of buccolingual breadth is only 3.2% and for 13 maxillary first molars the CV of buccolingual breadth is only 2.0% (buccolingual breadth is used because it is not subject to variance from interproximal wear). Not only size, but also anatomical shape and form are homogeneous within the sample. Almost every aspect of the morphology of the dentition, including the distinctive form of the lower premolars, the distal accessory cuspule of the mandibular canines, and the expression of nonmetric features that normally vary in human populations, is uniform in every specimen from the collection. The distinctive aspects of cranial morphology are repeated in every specimen, and even the aspects that normally vary among individuals of different body size or between sexes exhibit only slight variation among the Dinaledi remains. One of the most unique aspects of H. naledi is the morphology of the first metacarpal; the derived aspects of this anatomy are present in every one of seven first metacarpal specimens in the collection (Figure 14). Unlike any other fossil hominin site in Africa, the Dinaledi Chamber seems to preserve a large number of individuals from a single population, one with variation equal to or less than that found within local populations of modern humans.

Figure 14

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The Dinaledi collection is the richest assemblage of associated fossil hominins ever discovered in Africa, and aside from the Sima de los Huesos collection and later Neanderthal and modern human samples, it has the most comprehensive representation of skeletal elements across the lifespan, and from multiple individuals, in the hominin fossil record. The abundance of evidence from this assemblage supports our emerging understanding that the genus Homo encompassed a variety of evolutionary experiments (Antón et al., 2014), with diversity now evident for fossil Homo in each of the few intensively explored parts of Africa (Leakey et al., 2012). But as much as it advances our knowledge, H. naledi also highlights our ignorance about ancient Homo across the vast geographic span of the African continent. The tree of Homo-like hominins is far from complete: we have missed key transitional forms and lineages that persisted for hundreds of thousands of years. With an increasing pace of discovery from the field and the laboratory, more light will be thrown on the origin of humans.

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Author details

1. Lee R Berger

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. School of Geosciences, University of the Witwatersrand, Johannesburg, South Africa

   Contribution

   LRB, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   For correspondence

   Lee.Berger@wits.ac.za

   Competing interests

   The authors declare that no competing interests exist.

2. John Hawks

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. Department of Anthropology, University of Wisconsin-Madison, Madison, United States

   Contribution

   JH, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

3. Darryl J de Ruiter

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. Department of Anthropology, Texas A&M University, College Station, United States

   Contribution

   DJR, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

4. Steven E Churchill

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. Department of Evolutionary Anthropology, Duke University, Durham, United States

   Contribution

   SEC, Conception and design, Acquisition of data, Analysis and interpretation of data, Drafting or revising the article, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

5. Peter Schmid

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. Anthropological Institute and Museum, University of Zurich, Zurich, Switzerland

   Contribution

   PS, Conception and design, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

6. Lucas K Delezene

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. Department of Anthropology, University of Arkansas, Fayetteville, United States

   Contribution

   LKD, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

7. Tracy L Kivell

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. School, University of Kent, Canterbury, United Kingdom
   3. Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany

   Contribution

   TLK, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

8. Heather M Garvin

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. Department of Anthropology/Archaeology and Department of Applied Forensic Sciences, Mercyhurst University, Erie, United States

   Contribution

   HMG, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

9. Scott A Williams

   1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
   2. Center for the Study of Human Origins, Department of Anthropology, New York University, New York, United States
   3. New York Consortium in Evolutionary Primatology, New York, United States

   Contribution

   SAW, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

   Competing interests

   The authors declare that no competing interests exist.

10. Jeremy M DeSilva

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, Dartmouth College, Hanover, United States

    Contribution

    JMDS, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

11. Matthew M Skinner

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. School, University of Kent, Canterbury, United Kingdom
    3. Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany

    Contribution

    MMS, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

12. Charles M Musiba

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, University of Colorado Denver, Denver, United States

    Contribution

    CMM, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

13. Noel Cameron

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. School of Sport, Exercise and Health Sciences, Loughborough University, Loughborough, United Kingdom

    Contribution

    NC, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

14. Trenton W Holliday

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, Tulane University, New Orleans, United States

    Contribution

    TWH, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

15. William Harcourt-Smith

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, Lehman College, Bronx, United States
    3. Division of Paleontology, American Museum of Natural History, New York, United States

    Contribution

    WH-S, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

16. Rebecca R Ackermann

    Department of Archaeology, University of Cape Town, Rondebosch, South Africa

    Contribution

    RRA, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

17. Markus Bastir

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Paleoanthropology Group, Museo Nacional de Ciencias Naturales, Madrid, Spain

    Contribution

    MB, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

18. Barry Bogin

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. School of Sport, Exercise and Health Sciences, Loughborough University, Loughborough, United Kingdom

    Contribution

    BB, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

19. Debra Bolter

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, Modesto Junior College, Modesto, United States

    Contribution

    DB, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

20. Juliet Brophy

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Geography and Anthropology, Louisiana State University, Baton Rouge, United States

    Contribution

    JB, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

21. Zachary D Cofran

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. School of Humanities and Social Sciences, Nazarbayev University, Astana, Kazakhstan

    Contribution

    ZDC, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

22. Kimberly A Congdon

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Pathology and Anatomical Sciences, University of Missouri, Columbia, United States

    Contribution

    KAC, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

23. Andrew S Deane

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anatomy and Neurobiology, University of Kentucky College of Medicine, Lexington, United States

    Contribution

    ASD, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

24. Mana Dembo

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Human Evolutionary Studies Program and Department of Archaeology, Simon Fraser University, Burnaby, Canada

    Contribution

    MD, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

25. Michelle Drapeau

    Department d'Anthropologie, Université de Montréal, Montréal, Canada

    Contribution

    MD, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

26. Marina C Elliott

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Human Evolutionary Studies Program and Department of Archaeology, Simon Fraser University, Burnaby, Canada

    Contribution

    MCE, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

27. Elen M Feuerriegel

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. School of Archaeology and Anthropology, Australian National University, Canberra, Australia

    Contribution

    EMF, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

28. Daniel Garcia-Martinez

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Paleoanthropology Group, Museo Nacional de Ciencias Naturales, Madrid, Spain
    3. Faculty of Sciences, Biology Department, Universidad Autònoma de Madrid, Madrid, Spain

    Contribution

    DG-M, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

29. David J Green

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anatomy, Midwestern University, Downers Grove, United States

    Contribution

    DJG, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

30. Alia Gurtov

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, University of Wisconsin-Madison, Madison, United States

    Contribution

    AG, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

31. Joel D Irish

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Research Centre in Evolutionary Anthropology and Palaeoecology, Liverpool John Moores University, Liverpool, United Kingdom

    Contribution

    JDI, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

32. Ashley Kruger

    Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa

    Contribution

    AK, Acquisition of data, Analysis and interpretation of data

    Competing interests

    The authors declare that no competing interests exist.

33. Myra F Laird

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Center for the Study of Human Origins, Department of Anthropology, New York University, New York, United States
    3. New York Consortium in Evolutionary Primatology, New York, United States

    Contribution

    MFL, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

34. Damiano Marchi

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Biology, University of Pisa, Pisa, Italy

    Contribution

    DM, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

35. Marc R Meyer

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, Chaffey College, Rancho Cucamonga, United States

    Contribution

    MRM, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

36. Shahed Nalla

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Human Anatomy and Physiology, University of Johannesburg, Johannesburg, South Africa

    Contribution

    SN, Acquisition of data, Analysis and interpretation of data

    Competing interests

    The authors declare that no competing interests exist.

37. Enquye W Negash

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Center for the Advanced Study of Human Paleobiology, George Washington University, Washington, United States

    Contribution

    EWN, Acquisition of data, Analysis and interpretation of data

    Competing interests

    The authors declare that no competing interests exist.

38. Caley M Orr

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Cell and Developmental Biology, University of Colorado School of Medicine, Aurora, United States

    Contribution

    CMO, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

39. Davorka Radovcic

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Geology and Paleontology, Croatian Natural History Museum, Zagreb, Croatia

    Contribution

    DR, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

40. Lauren Schroeder

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Archaeology, University of Cape Town, Rondebosch, South Africa

    Contribution

    LS, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

41. Jill E Scott

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, University of Iowa, Iowa City, United States

    Contribution

    JES, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

42. Zachary Throckmorton

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anatomy, DeBusk College of Osteopathic Medicine, Lincoln Memorial University, Harrogate, United States

    Contribution

    ZT, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

43. Matthew W Tocheri

    1. Human Origins Program, Department of Anthropology, National Museum of Natural History, Smithsonian Institution, Washington, United States
    2. Department of Anthropology, Lakehead University, Thunder Bay, Canada

    Contribution

    MWT, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

44. Caroline VanSickle

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Anthropology, University of Wisconsin-Madison, Madison, United States
    3. Department of Gender and Women's Studies, University of Wisconsin-Madison, Madison, United States

    Contribution

    CVS, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

45. Christopher S Walker

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Evolutionary Anthropology, Duke University, Durham, United States

    Contribution

    CSW, Acquisition of data, Analysis and interpretation of data, Contributed unpublished essential data or reagents

    Competing interests

    The authors declare that no competing interests exist.

46. Pianpian Wei

    1. Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa
    2. Department of Paleoanthropology, Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China

    Contribution

    PW, Acquisition of data, Analysis and interpretation of data

    Competing interests

    The authors declare that no competing interests exist.

47. Bernhard Zipfel

    Evolutionary Studies Institute and Centre of Excellence in PalaeoSciences, University of the Witwatersrand, Johannesburg, South Africa

    Contribution

    BZ, Acquisition of data, Analysis and interpretation of data

    Competing interests

    The authors declare that no competing interests exist.

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