Homo naledi Berger, Hawks, Ruiter, Churchill, Schmid, Delezene, Kivell, Garvin, Williams, DeSilva, Skinner, Musiba, Cameron, Holliday, Harcourt-Smith, Ackermann, Bastir, Bogin, Bolter, Brophy, Cofran, Congdon, Deane, Dembo, Drapeau, Elliott, , Garcia-Martinez, Green, Gurtov, Irish, Kruger, Laird, Marchi, Meyer, Nalla, Negash, Orr, Radovcic, Schroeder, Scott, Throckmorton, Tocheri, VanSickle, Walker, Wei &amp; Zipfel, 2015, sp. nov.
Creators: name: Berger, Lee R
name: Hawks, John
name: De Ruiter, Darryl J
name: Churchill, Steven E
name: Schmid, Peter
name: Delezene, Lucas K
name: Kivell, Tracy L
name: Garvin, Heather M
name: Williams, Scott A
name: DeSilva, Jeremy M
name: Skinner, Matthew M
name: Musiba, Charles M
name: Cameron, Noel
name: Holliday, Trenton W
name: Harcourt-Smith, William
name: Ackermann, Rebecca R
name: Bastir, Markus
name: Bogin, Barry
name: Bolter, Debra
name: Brophy, Juliet
name: Cofran, Zachary D
name: Congdon, Kimberly A
name: Deane, Andrew S
name: Dembo, Mana
name: Drapeau, Michelle
name: Elliott, Marina C
name: , Elen M Feuerriegel
name: Garcia-Martinez, Daniel
name: Green, David J
name: Gurtov, Alia
name: Irish, Joel D
name: Kruger, Ashley
name: Laird, Myra F
name: Marchi, Damiano
name: Meyer, Marc R
name: Shahed Nalla
name: Enquye W Negash
name: Orr, Caley M
name: Radovcic, Davorka
name: Schroeder, Lauren
name: Scott, Jill E
name: Throckmorton, Zachary
name: Tocheri, Matthew W
name: VanSickle, Caroline
name: Walker, Christopher S
name: Pianpian Wei
name: Zipfel, Bernhard
Publisher: name: Zenodo
Year: 2015
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Primates
Hominidae
Homo
Homo naledi

description: <i>Homo naledi</i> sp. nov. urn:lsid:zoobank.org:pub:00D1E81A-6E08-4A01-BD98-79A2CEAE2411 Etymology The word <i>naledi</i> means ‘star’ in the Sotho language and refers to the Dinaledi Chamber’s location within the Rising Star cave system. Locality The Dinaledi chamber is located approximately 30 meters underground, within the Rising Star cave system at about 26&amp;ring;1 <i>′</i> 13 <i>′′</i> S; 27&amp;ring;42 <i>′</i> 43 <i>′′</i> E. The system lies within the Malmani dolomites, approximately 800 meters southwest of the well-known site of Swartkrans in the Cradle of Humankind World Heritage Site, Gauteng Province, South Africa. Horizon and associations The present sample of skeletal material from the Dinaledi Chamber was recovered during two field expeditions, in November 2013 and March 2014. Six specimens from an ex situ context can be identified as bird bones, and few fragmentary rodent remains have been recovered within the excavation area. Neither of these faunal constituents can presently be associated with the hominin fossil collection (<i>Dirks et al., 2015</i>). Aside from these limited faunal materials, the Dinaledi collection is entirely composed of hominin skeletal and dental remains. The collection so far comprises 1550 fossil hominin specimens, this number includes 1413 bone specimens and 137 isolated dental specimens; an additional 53 teeth are present in mandibular or maxillary bone specimens. Aside from the fragmentary rodent teeth, all dental crowns (n <b>=</b> 179) are hominin, recovered both from surface collection and excavation. Likewise, aside from the few bird elements, all morphologically informative bone specimens are clearly hominin. In all cases where elements are repeated in the sample, they are morphologically homogeneous, with variation consistent with body size and sex differences within a single population. These remains represent a minimum of 15 hominin individuals, as indicated by the repetition and presence of deciduous and adult dental elements. The geological age of the fossils is not yet known. Excavations have thus far recovered hominin material from Unit 2 and Unit 3 in the chamber (<i>Dirks et al., 2015</i>). Surface-collected hominin material from the present top of Unit 3, which includes material derived from both Unit 2 and Unit 3, represents a minority of the assemblage, and is morphologically indistinguishable from material excavated from in situ within Unit 3. In addition to general morphological homogeneity including cranial shape, distinctive morphological configurations of all the recovered first metacarpals, femora, molars, lower premolars and lower canines, are identical in both surface-collected and excavated specimens (see Figure 14 later in the text). These include traits not found in any other hominin species yet described. These considerations strongly indicate that this material represents a single species, and not a commingled assemblage. 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 (<i>Figure 2</i>; <i>Supplementary file 1</i>). The holotype was recovered in situ during excavations within the Dinaledi Chamber in March of 2014, embedded in unconsolidated fine clay matrix (<i>Dirks et al., 2015</i>). The holotype is housed in the Evolutionary Studies Institute at the University of the Witwatersrand, Johannesburg, South Africa. Paratypes Dinaledi Hominin 2 (DH2) is a partial calvaria that preserves parts of the frontal, left and right parietals, right temporal, and occipital (<i>Figure 3</i>; <i>Supplementary file 1</i>). 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 (<i>Figure 4</i>, <i>Supplementary file 1</i>). Dinaledi Hominin 4 (DH4) is a partial calvaria that preserves parts of the right temporal, right parietal, and occipital (<i>Figure 3</i>; <i>Supplementary file 1</i>). Dinaledi Hominin 5 (DH5) is a partial calvaria that preserves part of the left temporal and occipital (<i>Figure 3</i>; <i>Supplementary file 1</i>). 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 (<i>Figure 5</i>; <i>Supplementary file 1</i>). These cranial specimens agree closely in nearly all morphological details where they overlap in areas preserved except those we interpret as related to sex. 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 (<i>Figure 6</i>; <i>Supplementary file 1</i>). 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 (<i>Figure 7</i>; <i>Supplementary file 1</i>). U.W. 101-484 is a right tibial diaphysis missing only the proximal end (<i>Figure 8</i>; <i>Supplementary file 1</i>). 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 (<i>Figure 9</i>; <i>Supplementary file 1</i>). Referred material Referred material is also listed in <i>Supplementary file 1</i>. We refer to <i>H. naledi</i> 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 <i>Supplementary file 1</i> 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. Differential diagnosis 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 <i>H. naledi</i> in comparison to other species is presented in <i>Supplementary file 2</i>. Comparative cranial and mandibular measures are presented in <i>Table 1</i>, and comparative dental measures are provided in <i>Table 2</i>. Cranium, mandible, and dentition (DH1, DH2, DH3, DH4, DH5, U.W. 101-377) The cranium of <i>H. naledi</i> does not have the well-developed crest patterns that characterize <i>Australopithecus garhi</i> (<i>Asfaw et al., 1999</i>) and species of the genus <i>Paranthropus</i>, nor the derived facial morphology seen in the latter genus. The mandible of <i>H. naledi</i> is notably more gracile than those of <i>Paranthropus</i>. Although maxillary and mandibular incisors and canines of <i>H. naledi</i> overlap in size with those of <i>Paranthropus</i>, the post-canine teeth are notably smaller than those of <i>Paranthropus</i> and <i>Au. garhi</i>, with mandibular molars that are buccolingually narrow. <i>H. naledi</i> differs from <i>Australopithecus afarensis</i> and <i>Australopithecus africanus</i> in having a pentagonal-shaped cranial vault in posterior view, sagittal keeling, widely spaced temporal lines, an angular torus, a deep and narrow digastric fossa, an external occipital protuberance, an anteriorly positioned root of the zygomatic process of the maxilla, a broad palate, and a small canine jugum lacking anterior pillars. The anterior and lateral vault of <i>H. naledi</i> differs from <i>Au. afarensis</i> and <i>Au. africanus</i> in exhibiting only slight post-orbital constriction, frontal bossing, a well-developed supraorbital torus with a well-defined supratoral sulcus, temporal lines that are positioned on the posterior rather than the superior aspect of the supraorbital torus, a root of the zygomatic process of the temporal that is angled downwards approximately 30&amp;ring; relative to the Frankfort Horizontal (FH) and which begins its lateral expansion above the mandibular fossa rather than the EAM, a mandibular fossa that is positioned medial to the wall of the temporal squame, a small postglenoid process that contacts the tympanic, a coronally oriented petrous, and a small and obliquely oriented EAM. The <i>H. naledi</i> mandible exhibits a more gracile symphysis and corpus, a more vertically inclined symphysis, a slight mandibular incurvation delineating a faint mental trigon, and a steeply inclined posterior face of the mandibular symphysis without a post incisive planum. The incisors of <i>H. naledi</i> overlap in size with some specimens of <i>Au. africanus</i>, though the canines and post-canine dentition are notably smaller, with relatively narrow buccolingual dimensions of the mandibular molars. The maxillary I1 lacks a median lingual ridge and exhibits a broad and uninflated lingual cervical prominence, the lingual mesial and distal marginal ridges do not merge onto the cervical prominence in the maxillary I2, the mandibular canine exhibits only a weak lingual median ridge and a broad and uninflated lingual cervical prominence, and the buccal grooves on the maxillary premolars are only weakly developed. <i>H. naledi</i> exhibits a small and isolated Carabelli’s feature in the maxillary molars, unlike the more prominent and extensive Carabelli’s feature of <i>Australopithecus</i>. Moreover, the <i>H. naledi</i> mandibular molars possess small, mesiobuccally restricted protostylids that do not intersect the buccal groove, differing from the typically enlarged, centrally positioned protostylids that intersect the buccal groove in <i>Australopithecus</i>. The cranium of <i>H. naledi</i> differs from <i>Australopithecus sediba</i> (<i>Berger et al., 2010</i>) in exhibiting sagittal keeling, a more pronounced supraorbital torus and supratoral sulcus, a weakly arched supraorbital contour with rounded lateral corners, an angular torus, a well-defined supramastoid crest, a curved superior margin of the temporal squama, a root of the zygomatic process of the temporal that is angled downwards approximately 30&amp;ring; relative to FH, a flattened nasoalveolar clivus, weak canine juga, an anteriorly positioned root of the zygomatic process of the maxilla, and a relatively broad palate that is anteriorly shallow. The <i>H. naledi</i> mandible (DH1) has a mental foramen positioned superiorly on the corpus that opens posteriorly, unlike the mid-corpus height, more laterally opening mental foramen of <i>Au. sediba</i>. The maxillary and mandibular teeth of <i>H. naledi</i> are smaller than those of <i>Au. sediba</i>, with mandibular molars that are buccolingually narrow. The lingual mesial and distal marginal ridges do not merge onto the cervical prominence in the maxillary I2, the paracone of the maxillary P3 is equal in size to the protocone, the protoconid and metaconid of the mandibular molars are equally mesially positioned, and the lingual cusps of the molars are positioned at the occlusobuccal margin while the buccal cusps are positioned slightly lingual to the occlusobuccal margin. Also, <i>Au. sediba</i> shares with other australopiths a protostylid that is centrally located and which intersects the buccal groove of the lower molars, unlike the small and mesiobuccally restricted protostylid that does not intersect the buccal groove in <i>H. naledi</i>. The cranium of <i>H. naledi</i> differs from <i> <i>Homo</i> <i>habilis</i> in exhibiting sagittal keeling, a weakly arched supraorbital contour, temporal lines that are positioned on the posterior rather than the superior aspect of the supraorbital torus, an angular torus, an occipital torus, only slight postorbital constriction, a curved superior margin of the temporal squama, a suprameatal spine, a weak crista petrosa, a prominent Eustachian process, a small EAM, weak canine juga, and an anteriorly positioned root of the zygomatic process of the maxilla. Mandibles attributed to <i> <i>H.</i> <i>habilis</i> show a weakly inclined, shelf-like post incisive planum with a variably developed superior transverse torus, unlike the steeply inclined posterior face of the mandibular symphysis of <i> <i>H.</i> <i>naledi</i> , which lacks both a post incisive planum or superior transverse torus. The <i>H. naledi</i> mandible (DH1) has a mental foramen positioned superiorly on the corpus that opens posteriorly, while the mental foramen of <i>H. habilis</i> is at mid-corpus height and opens more laterally. The maxillary and mandibular dentitions of DH1 are smaller than typical for <i>H. habilis</i>. The mandibular P3 of <i>H. naledi</i> is more molarized and lacks the occlusal simplification seen in <i>H. habilis</i>; it has a symmetrical occlusal outline, and multiple roots (two roots: mesiobuccal and distal) not seen in <i> <i>H.</i> <i>habilis</i> . The molars of <i>H. naledi</i> lack crenulation, secondary fissures, and supernumerary cusps that are common to <i>H. habilis</i>. The protoconid and metaconid of the mandibular molars are equally mesially positioned.</i></i></i></i> The cranium of <i>H. naledi</i> differs from <i> <i>Homo</i> <i>rudolfensis</i> by its smaller cranial capacity, and by exhibiting frontal bossing, a post-bregmatic depression, sagittal keeling, a well-developed supraorbital torus delineated by a distinct supratoral sulcus, temporal lines that are positioned on the posterior rather than the superior aspect of the supraorbital torus, an occipital torus, an external occipital protuberance, only slight post-orbital constriction, a small postglenoid process, a weak crista petrosa, a laterally inflated mastoid process, a canine fossa, incisors that project anteriorly beyond the bi-canine line, and a shallow anterior palate. As in <i>H. habilis</i>, mandibles attributed to <i>H. rudolfensis</i> show a weakly inclined, shelf-like post incisive planum with a variably developed superior transverse torus, unlike the steeply inclined posterior face of the mandibular symphysis of DH1, the latter of which lacks either a post incisive planum or superior transverse torus. The mandibular symphysis and corpus of <i>H. naledi</i> are more gracile than those attributed to <i>H. rudolfensis</i>, and the <i>H. naledi</i> mandible (DH1) has a mental foramen positioned superiorly on the corpus that opens posteriorly, unlike the mid-corpus height, more laterally opening mental foramen of <i>H. rudolfensis</i>. The maxillary and mandibular dentition of <i>H. naledi</i> is smaller than that of most specimens of <i>H. rudolfensis</i>, with only KNM-ER 60000 and KNM-ER 62000 appearing similar in size for some teeth (<i>Leakey et al., 2012</i>). The molars of <i>H. naledi</i> lack crenulation, secondary fissures, or supernumerary cusps common in <i>H. rudolfensis</i>. The buccal grooves of the maxillary premolars are weak in <i>H. naledi</i>, and the protoconid and metaconid of the mandibular molars are equally mesially positioned.</i> <i>H. naledi</i> lacks the typically distinctive long and low cranial vault of <i>Homo erectus</i>, as well as the metopic keeling that is typically present in the latter species. <i>H. naledi</i> also differs from <i>H. erectus</i> in having a distinct external occipital protuberance in addition to the tuberculum linearum, a laterally inflated mastoid process, a flat and squared nasoalveolar clivus, and an anteriorly shallow palate. The parasagittal keeling that is present between bregma and lambda in <i>H. naledi</i> (DH1 and DH3) is less marked than often occurs in <i>H. erectus</i>, including in small specimens such as KNM-ER 42700 and the Dmanisi cranial sample. Also unlike most specimens of <i>H. erectus</i>, <i>H. naledi</i> has a small vaginal process, a weak crista petrosa, a marked Eustachian process, and a small EAM. The mandible of <i>H. erectus</i> shows a moderately inclined, shelf-like post incisive planum terminating in a variably developed superior transverse torus, differing from the steeply inclined posterior face of the <i>H. naledi</i> mandibular symphysis, which lacks both a post incisive planum or a superior transverse torus. The mental foramen is positioned superiorly and opens posteriorly in DH1, unlike the mid-corpus height, more laterally opening mental foramen of <i>H. erectus</i>. The maxillary and mandibular incisors and canines of <i>H. naledi</i> are smaller than typical of <i>H. erectus</i>. The mandibular P3 of <i>H. naledi</i> is more molarized and lacks the occlusal simplification seen in <i>H. erectus</i>, they reveal a symmetrical occlusal outline, and multiple roots (2R: MB <b>+</b> D) not typically seen in <i>H. erectus</i>. Furthermore, the molars of <i>H. naledi</i> lack crenulation, secondary fissures, or supernumerary cusps common in <i>H. erectus</i>. <i>H. naledi</i> lacks the reduced cranial height of <i>Homo floresiensis</i>, and displays a marked angular torus and parasagittal keeling between bregma and lambda that is absent in the latter species. <i>H. naledi</i> further has a flat and squared nasoalveolar clivus, unlike the pronounced maxillary canine juga and prominent pillars of <i>H. floresiensis</i>. The mandible of <i>H. floresiensis</i> shows a posteriorly inclined post incisive planum with superior and inferior transverse tori, differing from the steeply inclined posterior face of the <i>H. naledi</i> mandibular symphysis, which lacks both a post incisive planum or a superior transverse torus. Dentally, <i>H. naledi</i> is distinguishable from <i>H. floresiensis</i> by the mesiodistal elongation and extensive talonid of the mandibular P4, and the lack of Tomes’ root on the mandibular premolars. The molar size gradient of <i>H. naledi</i> follows the M1 <b>&lt;</b> M2 <b>&lt;</b> M3 pattern, unlike the M3 <b>&lt;</b> M2 <b>&lt;</b> M1 pattern in <i>H. floresiensis</i>, and the mandibular molars are relatively mesiodistally long and buccolingually narrow compared to those of <i>H. floresiensis</i>. *At least in presumed males. †Post-orbital breadth/superior facial breadth <b>×</b> 100. ‡Following the formula (π <b>×</b> (corpus height/2) <b>×</b> (corpus breadth/2)). <b>§&lt;</b>
description: Published as part of <i>Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei &amp; Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4</i> on pages 3-24, DOI: 10.7554/eLife.09560