Homo luzonensis Détroit &amp; Mijares &amp; Corny &amp; Daver &amp; Zanolli &amp; Dizon &amp; Robles &amp; Grün &amp; Piper 2019, sp. nov.
Creators: name: Détroit, Florent
name: Mijares, Armand Salvador
name: Corny, Julien
name: Daver, Guillaume
name: Zanolli, Clément
name: Dizon, Eusebio
name: Robles, Emil
name: Grün, Rainer
name: Piper, Philip J.
Publisher: name: Zenodo
Year: 2019
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Primates
Hominidae
Homo
Homo luzonensis

description: <i>Homo luzonensis</i> sp. nov. <b>Etymology.</b> The species name is derived from the island of Luzon, where the specimens were discovered. <b>Holotype.</b> CCH6 (CCH6-a to CCH6-e), maxillary right postcanine dentition of a single individual discovered on 24 August 2011. The repository is the National Museum of the Philippines, Manila. <i>Homo luzonensis</i> has been deposited in the ZooBank database (http://zoobank.org/) with Life Science Identifier urn:lsid:zoobank. org:act: 4F743862-662F-4E6B-9812-8 A05533 C1347. <b>Paratypes.</b> Recovered in 2007, 2011 and 2015 from the same excavation area and layer as the holotype: CCH1, a right third metatarsal 1; CCH2 and CCH5, two manual phalanges; CCH3 and CCH4, two pedal phalanges; CCH8, a left upper third or fourth premolar (P 3/4); and CCH9, a right M 3 (all specimens are housed at the National Museum of the Philippines, Manila). <b>Referred material.</b> CCH7, a femoral shaft that belonged to a juvenile individual (housed at the National Museum of the Philippines, Manila). <b>Locality.</b> The type locality is Callao Cave, in the Callao Limestone formation in the Peñablanca region of northern Luzon, the Philippines, at coordinates 17° 42′ 11.7″ N, 121° 49′ 25.5″ E. <b>Diagnosis.</b> Postcanine maxillary teeth of small size that are mesiodistally compressed, with a premolar:molar crown size ratio that is high compared to other species in the genus <i>Homo</i>. Upper premolars with two or three roots, a mesio-distally expanded lingual crown, strong buccal grooves, partial or continuous transverse crest, and an enameldentine junction (EDJ) shape that is distinct from that of <i>H. sapiens</i>, <i>Homo neanderthalensis</i> and Asian <i>Homo erectus</i>. Very small upper molars, with a M 1 &gt; M 2 &gt; M 3 crown size pattern, a simplified occlusal morphology with reduced metacone and hypocone, no crenulation on the EDJ, and EDJ shape affinities with that of <i>H. sapiens</i> and Asian <i>H. erectus</i>. Intermediate manual phalanx (rays 2–4) that is long and narrow (unlike all hominins except <i>H. sapiens</i>), with a longitudinally curved and dorso-palmarly compressed shaft, well-developed flexor sheath attachments and a strongly developed dorsal beak; it shares shape affinities with <i>Australopithecus</i>, <i>H. floresiensis</i> and—to a lesser extent— <i>H. sapiens</i>. Distal hand phalanx with proportions unlike 1 Département Homme &amp; Environnement, Muséum National d’Histoire Naturelle, UMR 7194, CNRS,Musée de l’Homme, Paris, France. 2 Archaeological Studies Program, University of the Philippines, Quezon City, The Philippines. 3 National Museum of the Philippines, Manila, The Philippines. 4 Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie (PALEVOPRIM), UMR 7262, CNRS, Université de Poitiers, Poitiers, France. 5 Laboratoire PACEA, UMR 5199 CNRS, Université de Bordeaux, Bordeaux, France. 6 Laboratoire AMIS, UMR 5288 CNRS, Université Toulouse III Paul Sabatier, Toulouse, France. 7 Australian Research Centre for Human Evolution, Environmental Futures Research Institute, Griffith University, Brisbane, Queensland, Australia. 8 Research School of Earth Sciences, Australian National University, Canberra, Australian Capital Territory, Australia. 9 School of Archaeology and Anthropology, Australian National University, Canberra, Australian Capital Territory, Australia. <i>*</i> e-mail: florent.detroit@mnhn.fr; mandy24_us@yahoo.com those of early <i>Homo</i> / <i>Paranthropus</i>, <i>Homo naledi</i>, <i>H. floresiensis</i> and <i>H. neanderthalensis</i>. The third metatarsal has a base that is very small relative to bone length, dorso-plantarly short and with a marked dorsoplantar convexity of the proximal articular facet. The proximal pedal phalanx (rays 2–4) shares shape affinities with <i>Australopithecus</i>: marked longitudinal curvature, parallel medial and lateral sides in dorsal view, a small bicondylar head and weak dorsal canting of the proximal articular surface (unlike African and European early <i>Homo</i>, <i>H. naledi</i>, <i>H. neanderthalensis</i>, <i>H. floresiensis</i> and <i>H. sapiens</i>). Description and comparison of the fossil elements Of the seven posterior maxillary teeth, six are from the right side (CCH6-a to CCH6-e and CCH9) and one (CCH8) is from the left (Fig. 2a, f, g, Extended Data Fig. 1 and Supplementary Information). The M 2 (CCH6-b) and one of the M 3 s (CCH9) are complete; the other teeth have well-preserved crowns, and at least one—but sometimes all—roots are partly broken. Micro-fissures are present in the enamel, dentine and cementum, and the pulp cavity is generally filled by calcium carbonate; however, these post-depositional modifications do not affect the size, shape or morphological attributes of the teeth. In comparison to <i>Australopithecus</i>, <i>Paranthropus</i> and other species of the genus <i>Homo</i>, the maxillary postcanine teeth of <i>H. luzonensis</i> are small and mesio-distally compressed, and there is a marked contrast between the relative size and shape of the premolars and molars (Figs. 2–4 and Extended Data Figs. 1–4). The upper molars of <i>H. luzonensis</i> have small crown sizes with a M 1 &gt; M 2 &gt; M 3 pattern and simplified occlusal surface morphology with a reduced number of cusps and an absence of marked crenulations at the EDJ (Fig. 3 and Extended Data Fig. 1). These derived features are found in other Late Pleistocene species of the genus <i>Homo</i> (that is, <i>H. neanderthalensis</i>, <i>H. floresiensis</i> and <i>H. sapiens</i>), but in this respect <i>H. luzonensis</i> molars more closely resemble those of <i>H. sapiens</i> 4, 5. However, the M 1 and M 2 of <i>H. luzonensis</i> have smaller crown sizes than those of <i>H. sapiens</i> (Fig. 3 and Extended Data Fig. 2d, e). Compared to the molars of <i>H. floresiensis</i>, <i>H. luzonensis</i> molars are smaller and the M 1 is not more mesio-distally compressed than the other postcanine teeth 6, 7 (Extended Data Figs. 2–4). For their diminutive size and simplified crown morphology, <i>H. luzonensis</i> molars also differ considerably from Asian <i>H. erectus</i> 4, 5, 8, 9 and Denisovans 10, although the EDJ of <i>H. luzonensis</i> M 1 shows some shape affinities with several Indonesian <i>H. erectus</i> specimens (Extended Data Fig. 4f, h). <i>H. luzonensis</i> premolars are noticeably large relative to the molarseven more so than seen in <i>H. floresiensis</i> and unlike the pattern seen in other hominins, except <i>Paranthropus</i> (Fig. 3)—and also display several primitive features. For instance, the P 3 and P 4 crowns are asymmetric with a large and mesially displaced lingual cusp, as in early <i>Homo</i> 11, and complete or partial mesial transverse ridges are present on the upper premolars, similar to those of <i>H. floresiensis</i> 6, 7. Developed mesial and distal vertical grooves are also evident on the buccal aspects of the P 3 and P 4; features that are more frequent in early <i>Homo</i> than in <i>H. sapiens</i> 12 and that are absent in <i>H. floresiensis</i> 6, 7. Moreover, <i>H. luzonensis</i> premolars have multiple roots (P 3 has three, P 4 has two) that are robust and highly divergent, an archaic condition typically found in <i>Australopithecus</i>, <i>Paranthropus</i> and early <i>Homo</i> 13. Multiple robust and divergent premolar roots also occur in Asian <i>H</i>. <i>erectus</i> (Early Pleistocene Indonesian and Middle Pleistocene Chinese specimens), but are rare in <i>H. sapiens</i> and absent in <i>H. floresiensis</i> 4, 6, 9, 12 – 17. CCH8 (Fig. 2f and Extended Data Fig. 1f), an isolated left upper premolar, exhibits three roots (typically a P 3 feature) with a relatively symmetric crown (typically a P 4 feature). If CCH8 is a P 4, a similar three-rooted condition has not been reported for any Pleistocene <i>Homo</i> from Asia and is rarely found in modern humans (1 to 3%) 13, 14. In terms of absolute tooth size and premolar–molar proportions, <i>H. luzonensis</i> shows a pattern that is not seen elsewhere in the genus <i>Homo</i> (Fig. 3 and Extended Data Fig. 2). Although <i>Paranthropus</i> shows similarly large premolars relative to molars (Fig. 3), the maxillary postcanine teeth of <i>H. luzonensis</i> differ markedly from those of <i>Paranthropus</i> in absolute size and shape (Extended Data Fig. 2). Multivariate statistical analysis of P 3 –M 2 dental diameters results in <i>H. luzonensis</i> clustering with Asian <i>H. erectus</i>. Both species are characterized by almost similarly sized first and second molars, large premolars relative to molars and postcanine teeth that are mesio-distally compressed (Extended Data Fig. 2g, h). The mesio-distally compressed shape of the M 1 of <i>H. luzonensis</i> is also visible in the results of the elliptic Fourier analysis of the crown contour, in which <i>H. luzonensis</i> plots at the margins of the range of variation for <i>H. sapiens</i> (which is characterized by more squared or rhomboidal M 1 s) (Extended Data Fig. 3). <i>H. luzonensis</i> differs from <i>H. floresiensis</i>, which displays an M 1 crown contour shape that is even more mesio-distally compressed and has a more developed protocone (Extended Data Fig. 3c, f). Threedimensional geometric morphometric analyses of the P 3 and P 4 EDJs discriminate CCH6 and CCH8 from fossil and extant <i>H. sapiens</i>, <i>H. neanderthalensis</i> and Asian <i>H. erectus</i>, with <i>H. luzonensis</i> occupying its own area of the morphospace or sharing it with <i>H. floresiensis</i> when the latter is included (Fig. 4 and Extended Data Fig. 4d, e). The 3D geometric morphometric analyses of M 1 and M 2 EDJs are less clear, with <i>H. luzonensis</i> clustering with extant <i>H. sapiens</i> and Asian <i>H. erectus</i> (Extended Data Fig. 4f), with extant <i>H. sapiens</i> only (Extended Data Fig. 4g), with Asian <i>H. erectus</i> only (Extended Data Fig. 4h), or by itself (Extended Data Fig. 4i). However, the overall pattern of morphology seen in the dental remains of <i>H. luzonensis</i> is easily distinguishable from all previously described hominin species. CCH2 is an intermediate manual phalanx with a total length of 32.5 mm (interarticular length, 31 mm; Fig. 2b, Supplementary Information and Supplementary Table 2). The bone is complete and well-preserved, but it was recovered broken just distal of the midshaft in two refitting parts (Extended Data Fig. 5a). On the basis of the asymmetry of the proximal and distal ends, it belongs to rays 2, 3 or 4 of the left hand. CCH2 displays primitive features such as an elongated and dorso-palmarly compressed shaft and well-developed flexor sheath attachments (Extended Data Fig. 5b). All these features are observed in <i>Australopithecus</i> 18 – 20 and to a lesser extent in the <i>Homo habilis</i> holotype OH7 21 (but see a previous publication 22), but not in other <i>Homo</i> species that show more derived morphologies 23 – 26 (Extended Data Fig. 5c), although a primitive morphology is reported for <i>H. naledi</i> 27. Metric (Extended Data Fig. 5 d–f) and 3D geometric morphometric analyses (Extended Data Fig. 6) show that the intermediate manual phalanx of <i>H. luzonensis</i> is long, narrow medio-laterally (base, shaft and head) and markedly curved. These shape characteristics are shared with <i>Australopithecus</i> and <i>H. floresiensis</i>, but not with <i>H. naledi</i> (which shares shape affinities with <i>Paranthropus</i> /early <i>Homo</i> specimens from Swartkrans), and are seen occasionally in <i>H. sapiens</i> (Extended Data Fig. 6a, d, e). <i>H. floresiensis</i> differs from <i>H. luzonensis</i> and most other hominins by the shape of the phalangeal head, which—in lateral or medial view—has a very small diameter and is only slightly palmarly deviated relative to the proximo-distal axis of the shaft (Extended Data Fig. 6b, c). An idiosyncratic feature that differentiates <i>H. luzonensis</i> from all other species of <i>Homo</i> is the strong development and proximal projection of the dorsal beak and this may have limited extension at the interphalangeal joint. CCH5 is a complete and well-preserved distal manual phalanx 15.9 mm in total length (interarticular length, 15.6 mm; Fig. 2c, Extended Data Fig. 7, Supplementary Information and Supplementary Table 2). Its laterality and ray cannot be determined with certainty, but its proximal articular surface is too small to articulate with CCH2. The proportions of CCH5 (apical tuft expansion and robusticity indices) are within the ranges of variation for <i>H. sapiens</i> and <i>Australopithecus</i> and outside those of <i>H. neanderthalensis</i> and <i>H. floresiensis</i> 23, 28 (Extended Data Fig. 7d, e). CCH4 is a complete and well-preserved right proximal pedal phalanx of rays 2, 3 or 4 (Fig. 2d, Extended Data Fig. 8, Supplementary Information and Supplementary Table 2) that is 27.4 mm long (interarticular length, 26.5 mm). It shows primitive— <i>Australopithecus</i> -likefeatures, including pronounced longitudinal curvature of the shaft in lateral view, parallel medial and lateral diaphyseal margins in dorsal view and well-developed flexor sheath attachments in the distal two-thirds of the shaft, bordered by marked plantar crests (Extended Data Fig. 8b, c). The base is small, with a circular proximal articular surface that displays dorsal canting and articular angles of 94.5° and 89.6°, respectively. These angular values are outside of the ranges of variation for <i>H. sapiens</i>, <i>H. neanderthalensis</i>, <i>H. naledi</i> and <i>H. floresiensis</i>, but within those of <i>Australopithecus afarensis</i> and <i>Australopithecus africanus</i> 20, 29 – 31 (Extended Data Fig. 8f, g). The head is relatively small, bicondylar and proximo-dorsally to disto-plantarly compressed, with a plantar breadth that is larger than its dorsal one. All of these features are typically found in combination in <i>Australopithecus</i> 19, 20, 29, 31. The morphology of the proximal pedal phalanges of <i>H. floresiensis</i> has also been described as <i>Australopithecus</i> -like in some respects (for example, absence of an hourglass shape) 32, 33; however, the combination of features found in CCH4 is essentially indistinguishable from the features of <i>A. afarensis</i> and <i>A. africanus</i> as shown by a 3D geometric morphometric shape analysis (Fig. 5). Although proximal pedal phalanges are missing from the Asian <i>H. erectus</i> fossil record, those known for African and European species (for example, <i>H. naledi</i>, <i>Homo antecessor</i> and <i>H. neanderthalensis</i>) suggest that the morphology of <i>H. luzonensis</i> is unique among the genus <i>Homo</i> 26, 31, 34. <b>Fig. 4</b> | <b>Premolar EDJ of <i>H. luzonensis</i>. a</b> , EDJ of the P 3 of <i>H. luzonensis</i> (CCH6-e) compared to the EDJs of <i>H. floresiensis</i> (Liang Bua 1 (LB1)), <i>H. sapiens</i>, <i>H. erectus</i> (Sangiran 4) and <i>H. neanderthalensis</i> (KRD 53). Horns of dentine were reconstructed for CCH6-e, LB1 and Sangiran 4; see Methods. Scale bar, 5 mm. <b>b</b>, <b>c</b>, Between-group principal component analyses (bgPCAs) of the three-dimensional landmark Procrustes- registered shape coordinates of the P 3 s (<b>b</b>) and P 4 s (<b>c</b>). Sample sizes for <b>b</b>, <b>c</b>, respectively: <i>H. erectus</i>, <i>n</i> = 2, 3; <i>H. neanderthalensis</i>, <i>n</i> = 5, 6; fossil <i>H. sapiens</i>, <i>n</i> = 3, 3; extant <i>H. sapiens</i>, <i>n</i> = 8, 9; <i>H. floresiensis</i>, <i>n</i> = 1, 0; <i>H. luzonensis</i>, <i>n</i> = 2, 2. A detailed list of specimens can be found in Supplementary Table 6. CCH3 is a complete and well-preserved intermediate pedal phalanx with a total length of 16.3 mm (interarticular length, 14.9 mm; Fig. 2e, Extended Data Fig. 9, Supplementary Information and Supplementary Table 2). Because the shape, size and morphology of this element is highly variable in <i>H. sapiens</i> and other hominins, little can be said at this time about its comparative morphology; however, it visually more closely resembles the intermediate pedal phalanges of <i>H. sapiens</i> and <i>H. floresiensis</i> than it does those of <i>A. afarensis</i> (Extended Data Fig. 9c). The previously published third metatarsal (CCH1) 1, 2 has a proximal base that is particularly small compared to the total length of the bone. It is further characterized by a pronounced triangular shape with a short dorso-plantar height and a medio-laterally expanded dorsal aspect, and a marked dorso-plantar convexity of the proximal articular facet for the lateral cuneiform. The latter feature, which is very uncommon in hominins, has been reported (since the description of CCH1) in a fourth metatarsal of <i>Australopithecus sediba</i> 35. The referred specimen, CCH7, is a partial left femur from which both the proximal and distal ends are missing (Fig. 2h, Extended Data Fig. 10 and Supplementary Information). Transverse slices of micro-computed tomography (micro-CT) scans show a high density of Haversian canals in the cortical bone that is characteristic of a growing juvenile individual (Extended Data Fig. 10c). Implications of <i>H. luzonensis</i> for hominin evolution The Late Pleistocene hominin fossils from Callao Cave show a combination of dental and postcranial (hand and foot) features that is distinct from currently known species of the genus <i>Homo</i> (Supplementary Table 3). The premolars of <i>H. luzonensis</i> combine size and shape features seen in other Late Pleistocene species of the genus <i>Homo</i> (for example, <i>H. neanderthalensis</i>, <i>H. floresiensis</i> and <i>H. sapiens</i>) with primitive morphological features of the crown, EDJ and roots, which are typically found in earlier hominins, including <i>Australopithecus</i> and <i>Paranthropus</i> (Supplementary Table 3). The molars are extremely small and their external morphology resembles those of <i>H. sapiens</i> and—to a lesser extent—those of <i>H. floresiensis</i>, whereas some of their EDJ shapes show affinities with those of Asian <i>H. erectus</i>. Notable similarities are observed between <i>H. luzonensis</i> and <i>Australopithecus</i> in the anatomy of their manual and pedal elements. The morphologies of the hands and feet of <i>Australopithecus</i>, which are generally described as an intermediate between the morphologies of great apes and modern humans, are typically interpreted either as indicating adaptations to various degrees of bipedalism and climbing and/or suspension 36 or as reflecting the retention of plesiomorphic features in obligatory bipeds 35, 37. However, the partial and fragmentary nature of the <i>H. luzonensis</i> postcranial elements presently limits further interpretation of its locomotor and manipulative abilities. The origin of <i>H. luzonensis</i>, as well as its phylogenetic relationships with other hominins present in eastern Asia at around the same timeincluding <i>H. sapiens</i> 38, <i>H. floresiensis</i> 39 and Denisovans 10, and hominins recently discovered in China 40 —remains to be determined. As is the case for most hominins recovered from tropical Asia 41, attempted DNA extraction from <i>H. luzonensis</i> fossils has been unsuccessful. Direct dates on <i>H. luzonensis</i> tooth and bone samples indicate it was present on Luzon before 50 kyr ago 1, 3 and the recent discovery 42 of stone tools and a butchered rhinoceros in the nearby Cagayan Valley indicates t
description: Published as part of <i>Détroit, Florent, Mijares, Armand Salvador, Corny, Julien, Daver, Guillaume, Zanolli, Clément, Dizon, Eusebio, Robles, Emil, Grün, Rainer &amp; Piper, Philip J., 2019, A new species of Homo from the Late Pleistocene of the Philippines, pp. 181-186 in Nature 568</i> on pages 1-5, DOI: 10.1038/s41586-019-1067-9, http://zenodo.org/record/2640603
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