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Unique Dental Morphology of Homo Floresiensis and Its Evolutionary Implications Yousuke Kaifu National Museum of Nature and Science, Japan, [email protected]

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Unique Dental Morphology of Homo Floresiensis and Its Evolutionary Implications Yousuke Kaifu National Museum of Nature and Science, Japan, Kaifu@Kahaku.Go.Jp University of Wollongong Research Online Faculty of Science, Medicine and Health - Papers Faculty of Science, Medicine and Health 2015 Unique dental morphology of Homo floresiensis and its evolutionary implications Yousuke Kaifu National Museum of Nature and Science, Japan, [email protected] Reiko T. Kono National Museum of Nature and Science, Japan Thomas Sutikna University of Wollongong, [email protected] E Wahyu Saptomo University of Wollongong - Jatmiko University of Wollongong See next page for additional authors Publication Details Kaifu, Y., Kono, R. T., Sutikna, T., Saptomo, E. Wahyu., Jatmiko, & Due Awe, R. (2015). Unique dental morphology of Homo floresiensis and its evolutionary implications. PLoS One, 10 (11), e0141614-1-e0141614-27. Research Online is the open access institutional repository for the University of Wollongong. For further information contact the UOW Library: [email protected] Unique dental morphology of Homo floresiensis and its evolutionary implications Abstract Homo floresiensis is an extinct, diminutive hominin species discovered in the Late Pleistocene deposits of Liang Bua cave, Flores, eastern Indonesia. The an ture and evolutionary origins of H. floresiensis’ unique physical characters have been intensively debated. Based on extensive comparisons using linear metric analyses, crown contour analyses, and other trait-by-trait morphological comparisons, we report here that the dental remains from multiple individuals indicate that H. floresiensis had primitive canine-premolar and advanced molar morphologies, a combination of dental traits unknown in any other hominin species. The primitive aspects are comparable to H. erectus from the Early Pleistocene, whereas some of the molar morphologies are more progressive even compared to those of modern humans. This evidence contradicts the earlier claim of an entirely modern human-like dental morphology of H. floresiensis, while at the same time does not support the hypothesis that H. floresiensis originated from a much older H. habilis or Australopithecus-like small-brained hominin species currently unknown in the Asian fossil record. These results are however consistent with the alternative hypothesis that H. floresiensis derived from an earlier Asian Homo erectus population and experienced substantial body and brain size dwarfism in an isolated insular setting. The dentition of H. floresiensis is not a simple, scaled-down version of earlier hominins. Publication Details Kaifu, Y., Kono, R. T., Sutikna, T., Saptomo, E. Wahyu., Jatmiko, & Due Awe, R. (2015). Unique dental morphology of Homo floresiensis and its evolutionary implications. PLoS One, 10 (11), e0141614-1-e0141614-27. Authors Yousuke Kaifu, Reiko T. Kono, Thomas Sutikna, E Wahyu Saptomo, - Jatmiko, and Rokus Awe Due This journal article is available at Research Online: http://ro.uow.edu.au/smhpapers/3733 RESEARCH ARTICLE Unique Dental Morphology of Homo floresiensis and Its Evolutionary Implications Yousuke Kaifu1,2*, Reiko T. Kono1, Thomas Sutikna3,4, Emanuel Wahyu Saptomo3,4, Jatmiko3,4, Rokus Due Awe3,4† 1 Department of Anthropology, National Museum of Nature and Science, Tokyo, Japan, 2 Department of Biological Sciences, The University of Tokyo, Tokyo, Japan, 3 Centre for Archaeological Science, University of Wollongong, Wollongong, Australia, 4 The National Research and Development Centre for Archaeology, Jakarta, Indonesia † Deceased. * [email protected] Abstract Homo floresiensis is an extinct, diminutive hominin species discovered in the Late Pleisto- cene deposits of Liang Bua cave, Flores, eastern Indonesia. The nature and evolutionary OPEN ACCESS origins of H. floresiensis’ unique physical characters have been intensively debated. Based Citation: Kaifu Y, Kono RT, Sutikna T, Saptomo EW, on extensive comparisons using linear metric analyses, crown contour analyses, and other Jatmiko, Due Awe R (2015) Unique Dental trait-by-trait morphological comparisons, we report here that the dental remains from multi- Morphology of Homo floresiensis and Its Evolutionary ple individuals indicate that H. floresiensis had primitive canine-premolar and advanced Implications. PLoS ONE 10(11): e0141614. molar morphologies, a combination of dental traits unknown in any other hominin species. doi:10.1371/journal.pone.0141614 The primitive aspects are comparable to H. erectus from the Early Pleistocene, whereas Editor: Christopher Bae, University of Hawaii at some of the molar morphologies are more progressive even compared to those of modern Manoa, UNITED STATES humans. This evidence contradicts the earlier claim of an entirely modern human-like dental Received: July 15, 2015 morphology of H. floresiensis, while at the same time does not support the hypothesis that Accepted: October 9, 2015 H. floresiensis originated from a much older H. habilis or Australopithecus-like small-brained Published: November 18, 2015 hominin species currently unknown in the Asian fossil record. These results are however Copyright: © 2015 Kaifu et al. This is an open consistent with the alternative hypothesis that H. floresiensis derived from an earlier Asian access article distributed under the terms of the Homo erectus population and experienced substantial body and brain size dwarfism in an Creative Commons Attribution License, which permits isolated insular setting. The dentition of H. floresiensis is not a simple, scaled-down version unrestricted use, distribution, and reproduction in any of earlier hominins. medium, provided the original author and source are credited. Data Availability Statement: All relevant data are within the paper and its Supporting Information file. Funding: This work was supported by funding from the Japan Society for the Promotion of Science Introduction (https://www.jsps.go.jp/english/), No. 24247044, YK; National Museum of Nature and Science, YK. The Previous studies showed that Homo floresiensis exhibits unusually small body and brain sizes funders had no role in study design, data collection for a Late Pleistocene Homo [1], a H. erectus-like cranial shape [2,3], an Australopithecus-like and analysis, decision to publish, or preparation of upper vs. lower limb proportion [4–7], and other primitive, advanced, and unique skeletal fea- the manuscript. tures [8–14]. Researchers agree that this unique mosaic has significant evolutionary meaning, Competing Interests: The authors have declared but disagree on what it is. Some anticipate that H. floresiensis evolved from an Australopithecus that no competing interests exist. or H. habilis-like, primitive, small-brained hominin currently unknown in the Asian fossil PLOS ONE | DOI:10.1371/journal.pone.0141614 November 18, 2015 1/27 Dental Morphology of Homo floresiensis record and thus signals unexpectedly early hominin dispersal into this region [14–19], whereas others suggest that the species is an example of substantial body and brain size insular dwarf- ism from Asian H. erectus [1,2,20,21]. In this paper, we present the first comprehensive analy- ses of their dental morphology to further contribute this debate. Teeth are one of the most informative elements in hominin evolutionary studies [22], and this is no exception in the case of H. floresiensis. Unlike postcranial elements, abundant com- parative fossil specimens are available for the teeth. Indeed, only one cranium of H. floresiensis currently exists (LB1), but dental remains represent up to three individuals for each tooth. Nev- ertheless, interpretations of the H. floresiensis dental morphology has been controversial, with opinion divided if their teeth are primitive [15] or modern like H. sapiens [23]. “How primi- tive” is another unresolved question. For example, there is a claim that the mesiodistally elon- gated P3 of H. floresiensis represents a very primitive hominin condition not seen in H. erectus [15], but more detailed analyses of this and other dental traits are needed to assess its taxo- nomic affinity [24]. The currently available H. floresiensis dental sample is comprised of one nearly complete maxillary dentition (LB1), two almost complete mandibular dentitions (LB1, LB6/1), and four 1 isolated teeth (I [LB15/2], I1 [LB6/14], P3 [LB2/2], P4 [LB15/1]) (Fig 1). All of these specimens are housed at the National Research and Development Centre for Archaeology, Jakarta. Full morphological descriptions of these specimens are available elsewhere along with a reassess- ment of the previous studies [24]. The purpose of the present study is to examine general dental evolutionary trends in Early Pleistocene Homo with particular emphasis on H. floresiensis’ two major ancestral candidates, the earliest Homo from Africa (H. habilis sensu lato: >2.3―1.6 Ma [million years ago]) and early Javanese H. erectus (>1.2―0.8 Ma) [2], and evaluate H. floresien- sis within this framework. Comparisons are also made with a sample of global modern humans including local prehistoric populations (N = 490) to examine dental morphological modernity in H. floresiensis. Three analytical approaches are employed: 1) traditional metric analyses based on crown length and breadth data, 2) comparisons of crown contour using normalized Elliptic Fourier Analysis (EFA), and 3) non-metric and linear metric comparisons of individual morphological traits not captured by the above two analyses. Materials and Methods Comparative samples Comparative fossil Homo samples are from the Early Pleistocene of Africa (H. habilis s. l., H. ergaster), Caucasia (Dmanisi Homo), and Indonesia (early Javanese H. erectus)
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