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A new species of Homo from the Late Pleistocene of the Philippines

  • April 2019
  • Nature 568(7751):181-186
DOI:10.1038/s41586-019-1067-9
Authors:
Florent Détroit at Muséum National d'Histoire Naturelle
Florent Détroit
Armand Salvador Belano Mijares at University of the Philippines
Armand Salvador Belano Mijares
Julien Corny
Julien Corny
Guillaume Daver at Université de Poitiers
Guillaume Daver
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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.
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.
… 
CCH7, femoral shaft of a juvenile individual of H. luzonensis a, Photograph of the original specimen CCH7 (posterior aspect). b, Three-dimensional rendering of CCH7. From left to right: anterior, medial, posterior and lateral aspects. Scale bar, 20 mm. c, Transverse micro-CT slices of CCH7 at proximal diaphysis (top), midshaft (middle) and distal diaphysis (bottom), and posterior aspect of the three-dimensional rendering of the femoral shaft, during the segmentation process (orientation of slices: anterior is up, posterior is down, lateral is left, medial is right).
CCH7, femoral shaft of a juvenile individual of H. luzonensis a, Photograph of the original specimen CCH7 (posterior aspect). b, Three-dimensional rendering of CCH7. From left to right: anterior, medial, posterior and lateral aspects. Scale bar, 20 mm. c, Transverse micro-CT slices of CCH7 at proximal diaphysis (top), midshaft (middle) and distal diaphysis (bottom), and posterior aspect of the three-dimensional rendering of the femoral shaft, during the segmentation process (orientation of slices: anterior is up, posterior is down, lateral is left, medial is right).
… 
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).
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).
… 
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.
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.
… 
Procrustes analyses of the premolar and molar EDJ a, Landmarks placed on the main dentine horns (in red) and semilandmarks (in white) positioned along the marginal ridges of the premolar EDJ. b, Landmarks placed on the main dentine horns (in red) and the semilandmarks (in white) positioned along the marginal ridges and on the oblique crest of the molar EDJ. c, Key for d–i. d–g, PCAs of the three-dimensional landmarks Procrustes-registered shape coordinates of the P³s (d), P⁴s (e), M¹s (f) and M²s (g). h, i, bgPCAs of the three-dimensional landmarks Procrustes-registered shape coordinates of the M¹s (h) and M²s (i). Sample sizes for d, e, f and h, and g and i, respectively: H. erectus, n = 2, 3, 5, and 3; H. neanderthalensis, n = 5, 6, 5, and 6; fossil H. sapiens, n = 3, 3, 4, and 3; extant H. sapiens, n = 8, 9, 7, and 9; H. floresiensis, n = 1, 0, 0, and 0; H. luzonensis, n = 2, 2, 1, and 1. A detailed list of specimens can be found in Supplementary Table 6.
+10
Procrustes analyses of the premolar and molar EDJ a, Landmarks placed on the main dentine horns (in red) and semilandmarks (in white) positioned along the marginal ridges of the premolar EDJ. b, Landmarks placed on the main dentine horns (in red) and the semilandmarks (in white) positioned along the marginal ridges and on the oblique crest of the molar EDJ. c, Key for d–i. d–g, PCAs of the three-dimensional landmarks Procrustes-registered shape coordinates of the P³s (d), P⁴s (e), M¹s (f) and M²s (g). h, i, bgPCAs of the three-dimensional landmarks Procrustes-registered shape coordinates of the M¹s (h) and M²s (i). Sample sizes for d, e, f and h, and g and i, respectively: H. erectus, n = 2, 3, 5, and 3; H. neanderthalensis, n = 5, 6, 5, and 6; fossil H. sapiens, n = 3, 3, 4, and 3; extant H. sapiens, n = 8, 9, 7, and 9; H. floresiensis, n = 1, 0, 0, and 0; H. luzonensis, n = 2, 2, 1, and 1. A detailed list of specimens can be found in Supplementary Table 6.
… 
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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

Supplementary resources (2)

Détroit_etal_2019_cover.jpg
Data
May 2019
Florent Détroit · Armand Salvador Belano Mijares · Julien Corny · Guillaume Daver · Philip Piper
Détroit_etal_2019_SuppInfo.pdf
Data
April 2019
Florent Détroit · Armand Salvador Belano Mijares · Julien Corny · Guillaume Daver · Philip Piper
... 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). ...
New insights into the Pleistocene archaeology of North Vietnam: understanding tropical cave sediments through microstratigraphic investigations and geo-ethnoarchaeological experiments
Thesis
Full-text available
  • Jan 2022
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). ...
Preceramic riverside hunter-gatherers and the arrival of Neolithic farmers in northern Luzon
Article
  • Jun 2022
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. ...
A Middle Pleistocene Denisovan molar from the Annamite Chain of northern Laos
Article
Full-text available
  • May 2022
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). ...
Insights on the Early Pleistocene Hominin Population of the Guadix-Baza Depression (SE Spain) and a Review on the Ecology of the First Peopling of Europe
Article
Full-text available
  • Apr 2022
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; ...
Enhancing the learning of evolutionary anthropology skills by combining student‐active teaching with actual and virtual immersion of Master's students in fieldwork, laboratory practice, and dissemination
Article
Full-text available
  • Apr 2022
Higher education in evolutionary anthropology involves providing students with in‐depth knowledge of biological and cultural heritage sites and collections that are frequently inaccessible. Indeed, most sites, fossils, and archaeological remains can be visited or manipulated only rarely and solely by specialists with extensive experience. Owing to the development of 3D and medical imaging techniques, this fragile heritage is now more widely accessible, and in a dynamic way. However, exclusive adoption of virtual teaching and learning has a negative impact on student engagement and, naturally, on exchanges with instructors, and thus cannot be used without some reservations. In the ITAP (Immersion dans les Terrains de l’Anthropologie biologique et de la Préhistoire) project of the higher education STEP (Soutien à la Transformation et à l’Expérimentation Pédagogiques) transformation program at the University of Bordeaux, we combine student‐active teaching with Master's students fully immersed in ongoing fieldwork, laboratory study, and dissemination of research results in order to develop more individually shaped learning curricula and to foster both professional and new interdisciplinary skills. Here, we present examples of experiments conducted in the ITAP project using both authentic and virtual collections of archaeological, experimental, and reference materials that help to break down the barriers between research activities and higher education, as well as providing a more general appraisal of the appropriate use of virtual tools in higher education by combining them with real‐life situations. The article describes examples of novel teaching methods that we have developed in our Master's program in evolutionary anthropology, where we combine student‐active teaching with Master's students fully immersed in ongoing fieldwork, laboratory study, and dissemination of research results.
... 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]. ...
Tropical forests in the deep human past
Article
Full-text available
  • Apr 2022
Since Darwin, studies of human evolution have tended to give primacy to open ‘savannah’ environments as the ecological cradle of our lineage, with dense tropical forests cast as hostile, unfavourable frontiers. These perceptions continue to shape both the geographical context of fieldwork as well as dominant narratives concerning hominin evolution. This paradigm persists despite new, ground-breaking research highlighting the role of tropical forests in the human story. For example, novel research in Africa's rainforests has uncovered archaeological sites dating back into the Pleistocene; genetic studies have revealed very deep human roots in Central and West Africa and in the tropics of Asia and the Pacific; an unprecedented number of coexistent hominin species have now been documented, including Homo erectus , the ‘Hobbit’ ( Homo floresiensis ), Homo luzonensis , Denisovans, and Homo sapiens . Some of the earliest members of our own species to reach South Asia, Southeast Asia, Oceania and the tropical Americas have shown an unexpected rapidity in their adaptation to even some of the more ‘extreme’ tropical settings. This includes the early human manipulation of species and even habitats. This volume builds on these currently disparate threads and, for the first time, draws together a group of interdisciplinary, agenda-setting papers that firmly places a broader spectrum of tropical environments at the heart of the deep human past. This article is part of the theme issue ‘Tropical forests in the deep human past’.
... 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]. ...
Fossils, fish and tropical forests: prehistoric human adaptations on the island frontiers of Oceania
Article
Oceania is a key region for studying human dispersals, adaptations and interactions with other hominin populations. Although archaeological evidence now reveals occupation of the region by approximately 65–45 000 years ago, its human fossil record, which has the best potential to provide direct insights into ecological adaptations and population relationships, has remained much more elusive. Here, we apply radiocarbon dating and stable isotope approaches to the earliest human remains so far excavated on the islands of Near and Remote Oceania to explore the chronology and diets of the first preserved human individuals to step across these Pacific frontiers. We demonstrate that the oldest human (or indeed hominin) fossil outside of the mainland New Guinea-Aru area dates to approximately 11 800 years ago. Furthermore, although these early sea-faring populations have been associated with a specialized coastal adaptation, we show that Late Pleistocene–Holocene humans living on islands in the Bismarck Archipelago and in Vanuatu display a persistent reliance on interior tropical forest resources. We argue that local tropical habitats, rather than purely coasts or, later, arriving domesticates, should be emphasized in discussions of human diets and cultural practices from the onset of our species' arrival in this part of the world. This article is part of the theme issue ‘Tropical forests in the deep human past’.
KLIMATICKÉ ZMENY V KENOZOIKU A ICH VPLYV NA EVOLÚCIU FAUNY SO ZAMERANÍM NA ŠUPINATÉ PLAZY
Book
Full-text available
  • Sep 2021
Taking Human Origins Research Into the Next Decade
Article
  • Mar 2022
Человек разумный и все-все-все... : пособие Homo sapiens and all relatives...: manual
Book
Full-text available
  • Mar 2022
До недавнего времени считалось, что одновременно на Земле сосуществовали два вида рода Homo: неандерталец и кроманьонец – человек анатомически современного типа. Молекулярно-генетические исследования древней ДНК конца XX – начала XXI века вскрыли наличие еще одной ветви древнего человека – денисовца. В 2003 году в Индонезии были открыты несколько скелетов флореcского человека, или хоббита – гипотетического вида карликовых людей. Исследования древней ДНК у вымерших видов и современного человека позволяют предположить существование ещё по крайней мере одного вида, внёсшего вклад в геном современного человека. Пособие разработано для практического использования в рамках ряда дисциплин у магистрантов-биологов и археологов. Оно может также быть использовано в учебном процессе для преподавателей, магистрантов, студентов высших учебных заведений биологического профиля. Until recently, it was believed that two species of the genus Homo coexisted simultaneously on Earth: the Neanderthal and the Cro-Magnon, an anatomically modern human. Molecular genetic studies of ancient DNA of the late 20th - early 21st century revealed the presence of another branch of ancient man - the Denisovan. In 2003, several skeletons of a Floresian man, or hobbit, a hypothetical species of dwarf people, were discovered in Indonesia. Studies of ancient DNA in extinct species and modern humans suggest the existence of at least one other species that contributed to the modern human genome. The manual has been developed for practical use in a number of disciplines by undergraduates in biology and archaeology. It can also be used in the educational process for teachers, undergraduates, students of higher educational institutions of a biological profile.
Momocs : Outline Analysis Using R
Article
Full-text available
  • Jan 2014
We introduce here Momocs, a package intended to ease and popularize modern mor- phometrics with R, and particularly outline analysis, which aims to extract quantitative variables from shapes. It mostly hinges on the functions published in the book entitled Modern Morphometrics Using R by Claude (2008). From outline extraction from raw data to multivariate analysis, Momocs provides an integrated and convenient toolkit to students and researchers who are, or may become, interested in describing the shape and its variation. The methods implemented so far in Momocs are introduced through a simplistic case study that aims to test if two sets of bottles have different shapes.
The Middle Pleistocene (MIS 12) human dental remains from Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), Italy. A comparative high resolution endostructural assessment
Article
Full-text available
The penecontemporaneous Middle Pleistocene sites of Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), set c. 450 km apart in central and northeastern Italy, respectively, have yielded some among the oldest human fossil remains testifying to a peopling phase of the Italian Peninsula broadly during the glacial MIS 12, a stage associated with one among the harshest climatic conditions in the Northern hemisphere during the entire Quaternary period. Together with the large samples from Atapuerca Sima de los Huesos, Spain, and Caune de l’Arago at Tautavel, France, the remains from Fontana Ranuccio and Visogliano are among the few mid-Middle Pleistocene dental assemblages from Western Europe available for investigating the presence of an early Neanderthal signature in their inner structure. We applied two- three-dimensional techniques of virtual imaging and geometric morphometrics to the high-resolution X-ray microtomography record of the dental remains from these two Italian sites and compared the results to the evidence from a selected number of Pleistocene and extant human specimens/samples from Europe and North Africa. Depending on their preservation quality and on the degree of occlusal wear, we comparatively assessed: (i) the crown enamel and radicular dentine thickness topographic variation of a uniquely represented lower incisor; (ii) the lateral crown tissue proportions of premolars and molars; (iii) the enamel-dentine junction, and (iv) the pulp cavity morphology of all available specimens. Our analyses reveal in both samples a Neanderthal-like inner structural signal, for some aspects also resembling the condition shown by the contemporary assemblage from Atapuerca SH, and clearly distinct from the recent human figures. This study provides additional evidence indicating that an overall Neanderthal morphological dental template was preconfigured in Western Europe at least 430 to 450 ka ago.
Earliest known hominin activity in the Philippines by 709 thousand years ago
Article
Full-text available
Over 60 years ago, stone tools and remains of megafauna were discovered on the Southeast Asian islands of Flores, Sulawesi and Luzon, and a Middle Pleistocene colonization by Homo erectus was initially proposed to have occurred on these islands. However, until the discovery of Homo floresiensis in 2003, claims of the presence of archaic hominins on Wallacean islands were hypothetical owing to the absence of in situ fossils and/or stone artefacts that were excavated from well-documented stratigraphic contexts, or because secure numerical dating methods of these sites were lacking. As a consequence, these claims were generally treated with scepticism. Here we describe the results of recent excavations at Kalinga in the Cagayan Valley of northern Luzon in the Philippines that have yielded 57 stone tools associated with an almost-complete disarticulated skeleton of Rhinoceros philippinensis, which shows clear signs of butchery, together with other fossil fauna remains attributed to stegodon, Philippine brown deer, freshwater turtle and monitor lizard. All finds originate from a clay-rich bone bed that was dated to between 777 and 631 thousand years ago using electron-spin resonance methods that were applied to tooth enamel and fluvial quartz. This evidence pushes back the proven period of colonization of the Philippines by hundreds of thousands of years, and furthermore suggests that early overseas dispersal in Island South East Asia by premodern hominins took place several times during the Early and Middle Pleistocene stages. The Philippines therefore may have had a central role in southward movements into Wallacea, not only of Pleistocene megafauna, but also of archaic hominins.
The fossil teeth of the Peking Man
Article
Full-text available
  • Feb 2018
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.
The search for ancient DNA heads east
Article
  • Jul 2018
Ancient teeth, phenetic affinities, and African hominins: Another look at where Homo naledi fits in
Article
A new species of Homo, Homo naledi, was described in 2015 based on the hominin skeletal remains from the Dinaledi Chamber of the Rising Star cave system, South Africa. Subsequent craniodental comparative analyses, both phenetic and cladistic, served to support its taxonomic distinctiveness. Here we provide a new quantitative analysis, where up to 78 nonmetric crown and root traits of the permanent dentition were compared among samples of H. naledi (including remains from the recently discovered Lesedi Chamber) and eight other species from Africa: Australopithecus afarensis, Australopithecus africanus, Paranthropus boisei, Paranthropus robustus, Homo habilis, Homo erectus, Middle Pleistocene Homo sp., and Pleistocene and Holocene Homo sapiens. By using the mean measure of divergence distance statistic, phenetic affinities were calculated among samples to evaluate interspecific relatedness. The objective was to compare the results with those previously obtained, to assess further the taxonomic validity of the Rising Star hominin species. In accordance with earlier findings, H. naledi appears most similar dentally to the other African Homo samples. However, the former species is characterized by its retention and full expression of features relating to the main cusps, as well as the root numbers, with a near absence of accessory traits-including many that, based on various cladistic studies, are plesiomorphic in both extinct and extant African hominins. As such, the present findings provide additional support for the taxonomic validity of H. naledi as a distinct species of Homo.
Cladistics and insular evolution, an unfortunate marriage? Another tangle in the Deinogalerix analysis of Borrani et al. (2017)
Article
  • Mar 2018
Insular taxa, such as the late Miocene giant erinaceid Deinogalerix and the insular hominin Homo floresiensis, display a surprisingly primitive placement in cladistic analyses of their respective groups. This has led to speculations of early dispersal onto islands, not corroborated by biogeographical evidence. Insular evolution is notorious for vast phenotypic changes in at least part of the island fauna. These changes encompass typical apomorphies, but, as they may also include evolutionary reversals, a parsimony‐based tree will inevitably yield a more basic position in cladograms. Therefore, we need to be cautious in drawing hasty conclusions from phylogenetic trees including insular taxa. Scrutiny is required to monitor possible evolutionary reversals and scenarios should be tested against biogeography to test their viability.
The Anthropology of Modern Human Teeth: Dental Morphology and Its Variation in Recent and Fossil Homo sapien</I
Book
  • Mar 2018
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.
Effect of X-ray irradiation on ancient DNA in sub-fossil bones – Guidelines for safe X-ray imaging
Article
  • Dec 2016
vegan: Community Ecology Package. R package version 2.0-8
Article
  • Jan 2013