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Homo floresiensis-like fossils from the early Middle Pleistocene of Flores

Abstract

The evolutionary origin of Homo floresiensis, a diminutive hominin species previously known only by skeletal remains from Liang Bua in western Flores, Indonesia, has been intensively debated. It is a matter of controversy whether this primitive form, dated to the Late Pleistocene, evolved from early Asian Homo erectus and represents a unique and striking case of evolutionary reversal in hominin body and brain size within an insular environment1,2,3,4. The alternative hypothesis is that H. floresiensis derived from an older, smaller-brained member of our genus, such as Homo habilis, or perhaps even late Australopithecus, signalling a hitherto undocumented dispersal of hominins from Africa into eastern Asia by two million years ago (2 Ma)5,6. Here we describe hominin fossils excavated in 2014 from an early Middle Pleistocene site (Mata Menge) in the So’a Basin of central Flores. These specimens comprise a mandible fragment and six isolated teeth belonging to at least three small-jawed and small-toothed individuals. Dating to ~0.7 Ma, these fossils now constitute the oldest hominin remains from Flores7. The Mata Menge mandible and teeth are similar in dimensions and morphological characteristics to those of H. floresiensis from Liang Bua. The exception is the mandibular first molar, which retains a more primitive condition. Notably, the Mata Menge mandible and molar are even smaller in size than those of the two existing H. floresiensis individuals from Liang Bua. The Mata Menge fossils are derived compared with Australopithecus and H. habilis, and so tend to support the view that H. floresiensis is a dwarfed descendent of early Asian H. erectus. Our findings suggest that hominins on Flores had acquired extremely small body size and other morphological traits specific to H. floresiensis at an unexpectedly early time.

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Figure 1: SOA-MM4 mandible compared with a Liang Bua H. floresiensis specimen.
Figure 2: Isolated teeth from Mata Menge.
Figure 3: CT-based reconstruction of SOA-MM1 and the results of Elliptic Fourier Analysis of the molar crown contour.

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Acknowledgements

Funding for the So’a Basin project was provided by an Australian Research Council (ARC) Discovery grant (DP1093342) awarded to M.J.M. and A.B., and the project was directed by M.J.M. (2010-2013) and G.D.v.d.B. (2013-2015). The Japan Society for the Promotion of Science provided a grant (No. 24247044) to Y.K. Financial and technical support was provided by the Geological Survey Centre of Indonesia. The Indonesian State Ministry of Research and Technology granted permission to undertake this research, and we thank the successive directors of the Geological Survey Centre, Y. Kusumahbrata, A. Wibowo and A. Pribadi, the Heads of the Geological Agency (R. Sukyiar and Surono), and the successive directors of the Geology Museum in Bandung (S. Baskoro and O. Abdurahman) for facilitating and supporting this research. In addition, we acknowledge support and advice provided by I. Setiadi, D. Pribadi and Suyono. We also thank M. R. Puspaningrum, H. Insani, I. Sutisna, S. Sonjaya, U. P. Wibowo, A. Gunawan, A. M. Saiful, S. Hayes, B. Burhan, E. Sukandar, A. Rahman, A. Rahmadi and E. E. Laksmana for their assistance in the field (2014–2015), and G. Suwa, T. Djubiantono, F. Aziz, T. Jacob, E. Mbua, F. Schrenk, I. Tattersall, K. Mowbray, J. de Vos, P. Mennecier, F. Demeter, Nguyen Kim Thuy, and Nguyen Lan Cuong for access to the specimens in their care.

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Authors and Affiliations

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Contributions

G.D.v.d.B. is co-director of the So’a Basin project with A.B. and I.K. M.J.M. (deceased) was also a co-director of the So’a Basin project. The project was initiated by M.J.M. and F.A. G.D.v.d.B. and I.K. undertook initial identification and analyses of the hominin fossils. Y.K. conducted the morphological analysis together with R.T.K. Fieldwork was planned and directed by G.D.v.d.B., I.K., A.B. and E.S.. Y.K., G.D.v.d.B. and A.B. wrote the manuscript.

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Correspondence to Yousuke Kaifu.

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Extended data figures and tables

Extended Data Figure 1 CT-based images of the SOA-MM4 mandible and photos of the SOA-MM6 incisor.

a–i, SOA-MM4 mandible. Surface-rendered images of superior (a), lateral (b), inferior (c), lingual (d), anterior (e), and posterior (f) views. Sagittal (h) and horizontal (i). CT sections at the plane indicated by the green (g) and red (h) lines. aMF, a branch of the mandibular foramen; ARR, anterior ramus root; LP, lateral prominence; M1, M1 alveolus; M2, M2 alveolus, M3, M3 alveolus; MC, mandibular canal; Mas, line for the masseter muscle attachment; MHL, mylohyoid line; pbMC, posterior branch of the mandibular canal; SLT, superior lateral torus. jk, SOA-MM6 mandibular incisor (I1/2) fragments. The crown (j, SOA-MM6a) and a root (k, SOA-MM6b) fragments were used for laser ablation uranium-series dating. The specimen was deposited before at least 0.55 Ma7. Note the bevelled occlusal wear surface (arrow). Scale bar, 5 mm.

Extended Data Figure 2 Linear metric comparisons of the mandibles and permanent teeth.

ae, Scatter plots of the mandibular corporal dimensions (a, b) and permanent tooth crown diameters (ce). We identify SOA-MM1 as M1 (e), but there remains a slight possibility that this tooth is M2 (f). Metric data of SOA-MM4: corpus height at M2, 18 mm; corpus height at M2/3, 18.5 mm; corpus width at M2, 12.5 mm; corpus width at M2/3, 13 mm.

Extended Data Figure 3 Mandibular comparisons.

am, H. habilis sensu lato: OH 13 (a, late adolescent), OH 37 (b, horizontally flipped image), KNM-ER 1802 (c, late adolescent), KNM-ER 3734 (d, horizontally flipped image), KNM-ER 60000 (e, horizontally flipped image) (photo by F. Spoor, copyright National Museums of Kenya); early Javanese H. erectus, Sangiran 1b (f), Sangiran 9 (g), Sangiran 22 (h), Sb 8103 (i), Sangiran 21 (j); Liang Bua H. floresiensis: LB1 (k), LB6/1 (l, horizontally flipped image; the corpus is distorted); (m) SOA-MM4. Scale bar, 30 mm. Note that the H. habilis mandibles tend to exhibit a thicker corpus, the position of the basal ramus (filled arrow) that is shifted laterally relative to the corpus midline axis, a prominent posterior part of the alveolar prominence (filled triangle), and a wider extramolar sulcus between the anterior ramus root (open arrow) and the molar row. The early Javanese H. erectus sample is variable but includes specimens with weaker expressions in these traits. The Liang Bua H. floresiensis and the SOA-MM4 mandibles share such derived features with early Javanese H. erectus.

Extended Data Figure 4 Comparisons of the hominin mandibles and teeth from So’a Basin (Mata Menge) and H. floresiensis from Liang Bua.

a, SOA-MM4 mandible. b, c, Right lateral and left lateral (horizontally flipped) views of LB1. d, Right lateral view of LB6/1. e, f, The SOA-MM3 and LB1 P3s, respectively. g, SOA-MM1 M1. hj, Occlusal views of SOA-MM4 (h), LB1 (i), and LB6/1 (j) mandibles. Scale bar, 10 mm.

Extended Data Figure 5 Principal component analyses of the four size-standardized mandibular measurements.

a, Scatter plot of the PC scores. b, Component loading of each PC. PC1 does not distinguish Homo from Au. afarensis, but Au. afarensis and post-habilis Homo are relatively well-separated in PC2. SOA-MM4 belongs to the cluster of Homo in this PC. SOA-MM4 occupies the space in between the two Liang Bua H. floresiensis mandibles, suggesting their shared lateral corporal shape.

Extended Data Figure 6 Metric analyses of mandibular deciduous canines.

ae, Comparisons of the crown length and breadth (a), and relative crown height (b). Results of the PCA based on size-adjusted five crown diameters (c, d) and the component loadings of each PC (e). ‘Crown size’ = geometric mean of the five crown diameters used. Au. afarensis and H. sapiens are indistinguishable in crown size (d) but they are discriminated from each other by PC1 (P < 0.00002, t-test). SOA-MM7 occupies an intermediate position between Au. afarensis and H. sapiens, suggesting its moderately primitive crown configuration. The other PCs did not discriminate Au. afarensis and H. sapiens.

Extended Data Table 1 2014 Hominin fossil collection from Mata Menge
Extended Data Table 2 Hominin teeth from Mata Menge as compared to those of Liang Bua H. floresiensisa
Extended Data Table 3 Comparative fossil samples
Extended Data Table 4 Comparative Homo sapiens dental sample

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van den Bergh, G., Kaifu, Y., Kurniawan, I. et al. Homo floresiensis-like fossils from the early Middle Pleistocene of Flores. Nature 534, 245–248 (2016). https://doi.org/10.1038/nature17999

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