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Abstract and Figures

A hominin third metatarsal discovered in 2007 in Callao Cave (Northern Luzon, the Philippines) and dated to 67 thousand years ago provided the earliest direct evidence of a human presence in the Philippines. Analysis of this foot bone suggested that it belonged to the genus Homo, but to which species was unclear. Here we report the discovery of twelve additional hominin elements that represent at least three individuals that were found in the same stratigraphic layer of Callao Cave as the previously discovered metatarsal. These specimens display a combination of primitive and derived morphological features that is different from the combination of features found in other species in the genus Homo (including Homo floresiensis and Homo sapiens) and warrants their attribution to a new species, which we name Homo luzonensis. The presence of another and previously unknown hominin species east of the Wallace Line during the Late Pleistocene epoch underscores the importance of island Southeast Asia in the evolution of the genus Homo.
Inventory of the fossil elements attributed to H. luzonensis and detailed views of the dental remains a, The hominin fossils recovered from Callao Cave. R, right; L, left; P, premolar; M, molar. b, Three-dimensional rendering of the postcanine maxillary teeth CCH6-b to CCH6-e (M²–P³): occlusal (top row) and buccal (bottom row) aspects. Enamel is shown in dark blue, dentine and cement in light brown and pulp cavity in dark grey. In all views, mesial is to the right, distal to the left. c, CCH6-a, right M³: occlusal, buccal, lingual, mesial and distal aspects (from top to bottom and left to right). Occlusal view: mesial is to the right, distal to the left. d, CCH6-a to CCH6-e, right M³–P³: photograph of occlusal aspect. Mesial is to the right, distal to the left. The numbers indicate the locations of the detailed views of the inter-proximal contact facets (IPCFs): P³ (CCH6-e), mesial IPCF 1: note the small size of this IPCF, indicating that the canine was probably not large; 2–5: note the perfect match between corresponding pairs of mesial (top row) and distal (bottom row, mirrored images) IPCFs, from the P³ (CCH6-e) to the M³ (CCH6-a). e, CCH6-c, right M¹: distal aspect, showing the partially fused lingual and disto-buccal roots. Lingual is to the left, buccal to the right. f, CCH8, left P³ or P⁴, photograph of the original fossil (occlusal view) and three-dimensional rendering: occlusal (top row), buccal (middle row) and mesio-buccal (bottom row) aspects. Enamel is shown in dark blue, dentine and cement in light brown and pulp cavity in dark grey. g, CCH9, right M³: occlusal, buccal, lingual, mesial and distal aspects (from top to bottom and left to right). Occlusal view: mesial is to the right, distal to the left; captures of the three-dimensional surface model. Scale bars, 10 mm (IPCF views in d (1–5) are not to scale); the mirrored image is indicated by an asterisk.
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Dental metrics a, b, d–f, Bivariate scatter plots for mesio-distal (MD) versus bucco-lingual (BL) diameters of P³ (a), P⁴ (b), M¹ (d), M² (e) and M³ (f). c, Key for a, b, d–g, j, k. Sample sizes for a, b, d, e, f, respectively: Australopithecus, n = 23, 23, 21, 26, 26; Paranthropus, n = 22, 20, 26, 23, 20; African and European early Homo, n = 23, 19, 37, 21, 17; Asian early Homo, n = 16, 17, 17, 13, 11; H. neanderthalensis, n = 23, 26, 27, 30, 20; H. sapiens, n = 57, 61, 86, 70, 33; H. floresiensis, n = 1, 1, 1, 1, 0; H. luzonensis, n = 2∗, 2∗, 1, 1, 2 (∗CCH8 is a P³ or a P⁴). A detailed list of specimens can be found in Supplementary Table 4. g–i, bgPCA of the log-shape ratios of bucco-lingual and mesio-distal diameters of four postcanine maxillary teeth (P³, P⁴, M¹ and M²), CCH6 was treated as a supplementary individual and was plotted a posteriori. g, Scatter plot of specimens for bgPC1 versus bgPC2, with convex hulls for all groups, except H. floresiensis and H. luzonensis. Sample sizes: Australopithecus, n = 6; Paranthropus, n = 5; African and European early Homo, n = 13; Asian early Homo, n = 5; H. neanderthalensis, n = 12; H. sapiens, n = 47; H. floresiensis, n = 1; H. luzonensis, n = 1. A detailed list of specimens can be found in Supplementary Table 4. h, Variable scores for bgPC1 versus bgPC2 (correlation circle; log-shape ratios of variables). i, Bar plot of eigenvalues (%) of bgPC1–bgPC5. j, k, Bivariate scatter plot of the summed square root of computed occlusal surface areas of premolars versus molars (j) and bgPCA of the log-shape ratios of bucco-lingual and mesio-distal diameters of four postcanine maxillary teeth (k) similar to the analyses presented in Fig. 3b and in g, respectively, but with ‘Negritos’ treated as a separate group (same sample sizes as in g, except for: H. sapiens, n = 38 and H. sapiens ‘Negritos’, n = 9).
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Elliptic Fourier analysis of M¹ crown contour CCH6-c compared to the holotype of H. floresiensis (LB1) and large samples of archaeological and recent H. sapiens individuals. a, PCA of shape data for all specimens, scatter plot of individual scores for PC1 versus PC2 (see Methods; elliptic Fourier descriptors applied to Procrustes-aligned outlines, ten harmonics included). LGM, Last Glacial Maximum. Sample sizes: H. luzonensis, n = 1; H. floresiensis, n = 2; pre-LGM, n = 2; pre-Neolithic post-LGM, n = 12; Neolithic/post-Neolithic, n = 232; recent ‘Negritos’, n = 19. A detailed list of specimens can be found in Supplementary Table 5. b, Bar plot of eigenvalues (%) of PC1–PC6. c, Extreme shape variations along PC1 and PC2. The scores of H. luzonensis M¹ along PC1 and PC2 reflects a crown outline shape that is mesio-distally compressed, but not as much as that of H. floresiensis (two versions of the LB1 right M¹). d, Right M¹ of the holotype of H. floresiensis LB1 showing the two different versions of the crown outline (see Methods): the original contour (V1; in blue) published in a previous study⁷, and the contour (V2; in red) drawn by J.C. differ in the compensation of the mesial IPCF. These two versions differ minimally in the results of the elliptic Fourier analysis (see d and h). e, PCA of Fourier descriptors for the means of 16 groups of H. sapiens (sample sizes in brackets, see details in Supplementary Table 5), H. luzonensis CCH6-c and H. floresiensis LB1 (V1 and V2 treated as 2 groups): scatter plot of mean scores for PC1 versus PC2 with a superimposed minimum spanning tree indicating distances between groups. f, Extreme shape variations along PC1 and PC2: H. floresiensis differs from H. luzonensis in having a M¹ crown contour shape that is more compressed mesio-distally, with a more developed protocone.
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ARTICLE https://doi.org/10.1038/s41586-019-1067-9
A new species of Homo from the Late
Pleistocene of the Philippines
Florent Détroit1*, Armand Salvador Mijares2,3*, Julien Corny1, Guillaume Daver4, Clément Zanolli5,6, Eusebio Dizon3,
Emil Robles2, Rainer Grün7,8 & Philip J. Piper3,9
A hominin third metatarsal discovered in 2007 in Callao Cave (Northern Luzon, the Philippines) and dated to 67 thousand
years ago provided the earliest direct evidence of a human presence in the Philippines. Analysis of this foot bone suggested
that it belonged to the genus Homo, but to which species was unclear. Here we report the discovery of twelve additional
hominin elements that represent at least three individuals that were found in the same stratigraphic layer of Callao Cave
as the previously discovered metatarsal. These specimens display a combination of primitive and derived morphological
features that is different from the combination of features found in other species in the genus Homo (including Homo
floresiensis and Homo sapiens) and warrants their attribution to a new species, which we name Homo luzonensis. The
presence of another and previously unknown hominin species east of the Wallace Line during the Late Pleistocene epoch
underscores the importance of island Southeast Asia in the evolution of the genus Homo.
Continued excavations in Callao Cave (Fig.1) that originally yielded
the hominin third metatarsal
1,2
(which we here call CCH1 for ‘Callao
Cave Hominin 1’) have produced another twelve hominin elements
(Extended Data Fig.1a) from the same stratigraphic layer (layer 14):
seven postcanine maxillary teeth (CCH6-a to CCH6-e, CCH8, CCH9;
Fig.2a, f, g), two manual phalanges (CCH2 and CCH5; Fig.2b, c), two
pedal phalanges (CCH3 and CCH4; Fig.2d, e) and a femoral shaft
(CCH7; Fig.2h). CCH1 and CCH6-a are directlydated by U-series
analysis to minimum ages of 67thousand years (kyr)1 and 50kyr3,
respectively. Crown morphology, grade of occlusal wear and exact
correspondences of interproximal contact facets demonstrate that five
of the upper right teeth belonged to a single individual (CCH6-a to
CCH6-e; Extended Data Fig.1b–g and Supplementary Information).
The presence of two right upper third molars (M
3
; CCH6-a and CCH9)
and a juvenile femoral shaft (CCH7) indicates that at least three indi-
viduals are represented. On the basis of the unique mosaic of primi-
tive (that is, Australopithecus-like) and derived (that is, H. sapiens-like)
morphological features observed on these specimens, we assign them
to a new species, H. luzonensis.
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 luzonensis sp. nov.
Etymology. The species name is derived from the island of Luzon,
where the specimens were discovered.
Holotype. CCH6 (CCH6-a to CCH6-e), maxillary right postca-
nine dentition of a single individual discovered on 24 August 2011.
The repository is the National Museum of the Philippines, Manila.
Homo luzonensis has been deposited in the ZooBank database
(http://zoobank.org/) with Life Science Identifier urn:lsid:zoobank.
org:act:4F743862-662F-4E6B-9812-8A05533C1347.
Paratypes. 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 pha-
langes; CCH8, a left upper third or fourth premolar (P3/4); and CCH9,
a right M
3
(all specimens are housed at the National Museum of the
Philippines, Manila).
Referred material. CCH7, a femoral shaft that belonged to a juvenile
individual (housed at the National Museum of the Philippines, Manila).
Locality. The type locality is Callao Cave, in the Callao Limestone for-
mation in the Peñablanca region of northern Luzon, the Philippines,
at coordinates 17°4211.7N, 121°4925.5E.
Diagnosis. Postcanine maxillary teeth of small size that are mesio-
distally compressed, with a premolar:molar crown size ratio that is high
compared to other species in the genus Homo. Upper premolars with
two or three roots, a mesio-distally expanded lingual crown, strong
buccal grooves, partial or continuous transverse crest, and an enamel–
dentine junction (EDJ) shape that is distinct from that of H. sapiens,
Homo neanderthalensis and Asian Homo erectus. Very small upper
molars, with a M
1
>M
2
>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 H. sapiens and Asian
H. erectus. Intermediate manual phalanx (rays 2–4) that is long and
narrow (unlike all hominins except H. sapiens), 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 Australopithecus, H. floresiensis and—to a lesser
extent—H. sapiens. Distal hand phalanx with proportions unlike
1Département Homme & Environnement, Muséum National d’Histoire Naturelle, UMR 7194, CNRS, Musée de l’Homme, Paris, France. 2Archaeological Studies Program, University of the
Philippines, Quezon City, The Philippines. 3National Museum of the Philippines, Manila, The Philippines. 4Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie (PALEVOPRIM),
UMR 7262, CNRS, Université de Poitiers, Poitiers, France. 5Laboratoire PACEA, UMR 5199 CNRS, Université de Bordeaux, Bordeaux, France. 6Laboratoire AMIS, UMR 5288 CNRS, Université
Toulouse III Paul Sabatier, Toulouse, France. 7Australian Research Centre for Human Evolution, Environmental Futures Research Institute, Griffith University, Brisbane, Queensland, Australia.
8Research School of Earth Sciences, Australian National University, Canberra, Australian Capital Territory, Australia. 9School of Archaeology and Anthropology, Australian National University,
Canberra, Australian Capital Territory, Australia. *e-mail: florent.detroit@mnhn.fr; mandy24_us@yahoo.com
11 APRIL 2019 | VOL 568 | NATURE | 181
... Given recent fossil hominin discoveries (Morwood et al., , 2005Van den Bergh et al., 2016;Ingicco et al., 2018), the diversity of the Asian, and particularly Southeast Asian, Homo taxa increasingly reflects the high biodiversity of the region's modern fauna. Attention has focussed on ISEA, due to the discovery of Homo floresiensis and, more recently, Homo luzonensis (Détroit et al., 2019). Discoveries of Denisovan fossils on the Eurasian mainland and the evidence for episodic genetic exchange among hominin groups, including modern humans (Section 2.3.3), ...
... Both the Philippines (excepting Palawan) and Wallacea remained isolated from Eurasia throughout the Pleistocene (Sections 2.2.1 and 2.2.3), with strong north-south currents through Wallacea acting as a further barrier to dispersal (Morwood, 2014). Apparently persisting from the middle Pleistocene into the late Pleistocene, Homo floresiensis and Homo luzonensis were species of archaic hominin endemic to Flores in Wallacea and Luzon in the Philippines, respectively (Morwood, 2014;Van den Bergh et al., 2016;Détroit et al., 2019). Their presence indicates multiple, successful sea crossings by archaic Homo, but the high levels of endemism within the region's Pleistocene hominins suggest that regional populations remained largely isolated from other hominin groups throughout most of the Pleistocene (Westaway, 2019;Teixeira and Cooper, 2019). ...
... Their presence indicates multiple, successful sea crossings by archaic Homo, but the high levels of endemism within the region's Pleistocene hominins suggest that regional populations remained largely isolated from other hominin groups throughout most of the Pleistocene (Westaway, 2019;Teixeira and Cooper, 2019). The archaic traits of Wallacean hominin species (Tocheri et al., 2007;Détroit et al., 2019) and teeth provisionally identified as Homo habilis from an early Pleistocene context in southern China (Li et al., 2017a) have led some to propose a hominin dispersal prior that of H. erectus, but this remains speculative based on the current dataset (Tocheri, 2019 (Sutikna et al., 2018;Veatch et al., 2019). ...
Thesis
Full-text available
The Pleistocene archaeological record of mainland Southeast Asia (MSEA) is difficult to interpret, due to a sparsity of dated sites and a lithic record that is ill-suited to typological analysis. These challenges are compounded by the poorly constrained effects of tropical environments upon the deposition, preservation and degradation of archaeological cave sediments. These uncertainties restrict the interpretative potential of archaeological investigations, but the development of a rigorous, geoarchaeological framework of interpretation that is tailored to tropical cave sites offers an opportunity to improve research outcomes in MSEA and in tropical zones worldwide. Con Moong Cave (henceforth CMC), a Pleistocene archaeological site in North Vietnam, provided a small-scale example with which to explore the effects of tropical conditions upon archaeological site formation processes, and the potential of micro-geoarchaeological methods to overcome the difficulties of site interpretation in tropical zones. Research at CMC produced a depositional history for the site that revealed a correlation between sedimentation, the intensity of human occupation and changes in regional precipitation, as recognised in speleothem records in the published literature. Mineral suites from CMC’s guano deposits did not conform to established models of guano-driven diagenetic change and suggested that fluctuating hydrological conditions had led to diachronous episodes of mineral authigenesis. A shortage of reference data relevant to these processes meant it was not possible to conclusively relate these results to sedimentary palaeoenvironments or to understand their effects on assemblage taphonomy. Geo-ethnoarchaeological experiments were conducted to generate such a dataset, to enable hypothesis testing of the models of environmental changes at CMC and provide an assessment of the effects of tropical climates on sediment diagenesis.
... These sites contain cultural sequences from the Preceramic through to the Metal Age, and document continuing interaction between hinterland foragers and riverine agriculturalists, described ethnographically for this region by Jean Peterson (1978). Callao Cave is the largest and oldest of these sites; excavations here have recovered evidence of Homo luzonensis, a small-bodied hominin species of the Late Pleistocene (Détroit et al. 2019). ...
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The most westerly Pacific island chain, running from Taiwan southwards through the Philippines, has long been central in debates about the origins and early migrations of Austronesian-speaking peoples from the Asian mainland into the islands of Southeast Asia and Oceania. Focusing on the Cagayan Valley of northern Luzon in the Philippines, the authors combine new and published radiocarbon dates to underpin a revised culture-historical synthesis. The results speak to the initial contacts and long-term relationships between Indigenous hunter-gatherers and immigrant Neolithic farmers, and the question of how the early speakers of Malayo-Polynesian languages spread into and through the Philippines.
... We first ran the sample as a normal pIR-IRSL 50,270 to observe the dose response of a saturated sample to ensure that the fitting procedure could accommodate these high dosing values. ...
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The Pleistocene presence of the genus Homo in continental Southeast Asia is primarily evidenced by a sparse stone tool record and rare human remains. Here we report a Middle Pleistocene hominin specimen from Laos, with the discovery of a molar from the Tam Ngu Hao 2 (Cobra Cave) limestone cave in the Annamite Mountains. The age of the fossil-bearing breccia ranges between 164-131 kyr, based on the Bayesian modelling of luminescence dating of the sedimentary matrix from which it was recovered, U-series dating of an overlying flowstone, and U-series-ESR dating of associated faunal teeth. Analyses of the internal structure of the molar in tandem with palaeoproteomic analyses of the enamel indicate that the tooth derives from a young, likely female, Homo individual. The close morphological affinities with the Xiahe specimen from China indicate that they belong to the same taxon and that Tam Ngu Hao 2 most likely represents a Denisovan.
... These features make sense in the context of a highly isolated inbred population with small genetic variation (as expected from the founder effect of the island colonizers), a population that was adaptively constrained. It should be noted, however, that a similar combination of primitive and modern characters is also present in H. luzonensis, the dwarfed hominin of the larger island of Luzon (Philippines), which probably descended also from an initial population of H. erectus (Détroit et al., 2019). ...
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The chronology and environmental context of the first hominin dispersal in Europe have been subject to debate and controversy. The oldest settlements in Eurasia (e.g., Dmanisi, ∼1.8 Ma) suggest a scenario in which the Caucasus and southern Asia were occupied ∼0.4 Ma before the first peopling of Europe. Barranco León (BL) and Fuente Nueva 3 (FN3), two Early Pleistocene archeological localities dated to ∼1.4 Ma in Orce (Guadix-Baza Depression, SE Spain), provide the oldest evidence of hominin presence in Western Europe. At these sites, huge assemblages of large mammals with evidence of butchery and marrow processing have been unearthed associated to abundant Oldowan tools and a deciduous tooth of Homo sp. in the case of BL. Here, we: (i) review the Early Pleistocene archeological sites of Europe; (ii) discuss on the subsistence strategies of these hominins, including new estimates of resource abundance for the populations of Atapuerca and Orce; (iii) use cartographic data of the sedimentary deposits for reconstructing the landscape habitable in Guadix-Baza; and (iv) calculate the size of the hominin population using an estimate of population density based on resource abundance. Our results indicate that Guadix-Baza could be home for a small hominin population of 350–280 individuals. This basin is surrounded by the highest mountainous reliefs of the Alpine-Betic orogen and shows a limited number of connecting corridors with the surrounding areas, which could have limited gene flow with other hominin populations. Isolation would eventually lead to bottlenecks, genetic drift and inbreeding depression, conditions documented in the wild dog population of the basin, which probably compromised the viability of the hominin population in the medium to long term. This explains the discontinuous nature of the archeological record in Guadix-Baza, a situation that can also be extrapolated to the scarcity of hominin settlements for these ancient chronologies in Europe.
... In particular, photogrammetry and confocal microscopy have been widely used to quantify alterations preserved at micro-and nano-scopic scales on the surface of materials. The applications of these techniques permit, for instance, (1) study of cut marks and taphonomic alterations on bone surfaces, (2) identification of the functional history of stone and bone tools and personal ornaments through use-wear analysis, (3) reconstruction of the diet and environment of past human and non-human animal groups through dental macro-and microwear analysis, and (4) elucidation of biological processes, such as growth and its disturbances through the study of growth increments in teeth and bones(Calandra et al., 2019;Galland et al., 2018;Li et al., 2020;Magnani et al., 2020;Maté-González et al., 2017;McGrath et al., 2021;Rosso et al., 2017;Schmidt et al., 2020;Stemp, 2014;Yravedra Sainz de los Terreros et al., 2019).Furthermore, other 3D techniques such as X-ray, synchrotron radiation, and neutron microtomography have been widely used to non-invasively study the internal structure of bones and teeth, which holds a significant amount of valuable information for providing taxonomic identification, reconstructing phylogenetic relationships, and assessing adaptive strategies in past groups (among others, see studies byBayle et al., 2010;Beaudet et al., 2016;Benazzi et al., 2011;Crevecoeur et al., 2014;Détroit et al., 2019; ...
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... Far from being uninhabited by hominins, African tropical forest habitats seem to have been integral to our hominin ancestors [23], and Homo erectus notably reached Southeast Asia 1.2 million years ago (Ma), at a time when it has been argued that tropical forest was widespread ( [24,25]although see [26]). These environments likely formed at least part of the backdrop of local trajectories of evolution, as manifested in species such as Homo floresiensis and Homo luzonensis [27][28][29]. However, in the history of our genus, it was Homo sapiens that went on to most intensively inhabit and exploit tropical forests [6,15]. ...
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... This article is part of the theme issue 'Tropical forests in the deep human past'. regard [3][4][5][6][7]. The earliest human arrival in New Guinea, the Aru Islands, mainland Australia and Tasmania, which then formed the single continent of Sahul (ca 65 ka [5]), has been frequently prominently tied into wider discussions of the timing and number of our species' dispersals beyond Africa [8]. ...
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This study provides new original data, including the endostructure of most Zhoukoudian H. erectus teeth preserved to date, since the publication of Black in 1927 and Weidenreich in 1937. The new evidence ratifies the similarities of Zhoukoudian with other East Asian mid-Middle Pleistocene hominins such as Hexian and Yiyuan, and allows defining a dental pattern potentially characteristic of this population commonly referred to as classic H. erectus. Given the possible chronological overlaps of classic H. erectus with other archaic Homo, the characterization of this group becomes a key issue when deciphering the taxonomy and evolutionary scenario of the Middle Pleistocene hominins in East Asia. Internally, the most remarkable feature of Zhoukoudian teeth is the highly crenulated enamel-dentine junction (EDJ) and its imprint on the roof of the pulp cavity. So far, this "dendrite-like" EDJ has been found only in East Asia Middle Pleistocene hominins although a large group of samples were assessed, and it could be useful to dentally define classic H. erectus in China. The crenulated EDJ surface, together with the stout roots and the taurodontism could be a mechanism to withstand high biomechanical demand despite a general dentognathic reduction, particularly of the crowns, in these populations.
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All humans share certain components of tooth structure, but show variation in size and morphology around this shared pattern. This book presents a worldwide synthesis of the global variation in tooth morphology in recent populations. Research has advanced on many fronts since the publication of the first edition, which has become a seminal work on the subject. This revised and updated edition introduces new ideas in dental genetics and ontogeny and summarizes major historical problems addressed by dental morphology. The detailed descriptions of 29 dental variables are fully updated with current data and include details of a new web-based application for using crown and root morphology to evaluate ancestry in forensic cases. A new chapter describes what constitutes a modern human dentition in the context of the hominin fossil record.