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Craniofacial Morphology of Homo Floresiensis: Description, Taxonomic

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Craniofacial Morphology of Homo Floresiensis: Description, Taxonomic Journal of Human Evolution 61 (2011) 644e682 Contents lists available at SciVerse ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol Craniofacial morphology of Homo floresiensis: Description, taxonomic affinities, and evolutionary implication Yousuke Kaifu a,b,*, Hisao Baba a, Thomas Sutikna c, Michael J. Morwood d, Daisuke Kubo b, E. Wahyu Saptomo c, Jatmiko c, Rokhus Due Awe c, Tony Djubiantono c a Department of Anthropology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki Prefecture Japan b Department of Biological Sciences, The University of Tokyo, 3-1-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan c National Research and Development Centre for Archaeology, Jl. Raya Condet Pejaten No 4, Jakarta 12001, Indonesia d Centre for Archaeological Science, School of Earth and Environmental Sciences, University of Wollongong, Wollongong, NSW 2522, Australia article info abstract Article history: This paper describes in detail the external morphology of LB1/1, the nearly complete and only known Received 5 October 2010 cranium of Homo floresiensis. Comparisons were made with a large sample of early groups of the genus Accepted 21 August 2011 Homo to assess primitive, derived, and unique craniofacial traits of LB1 and discuss its evolution. Prin- cipal cranial shape differences between H. floresiensis and Homo sapiens are also explored metrically. Keywords: The LB1 specimen exhibits a marked reductive trend in its facial skeleton, which is comparable to the LB1/1 H. sapiens condition and is probably associated with reduced masticatory stresses. However, LB1 is Homo erectus craniometrically different from H. sapiens showing an extremely small overall cranial size, and the Homo habilis Cranium combination of a primitive low and anteriorly narrow vault shape, a relatively prognathic face, a rounded Face oval foramen that is greatly separated anteriorly from the carotid canal/jugular foramen, and a unique, tall orbital shape. Whereas the neurocranium of LB1 is as small as that of some Homo habilis specimens, it exhibits laterally expanded parietals, a weak suprameatal crest, a moderately flexed occipital, a marked facial reduction, and many other derived features that characterize post-habilis Homo. Other craniofacial characteristics of LB1 include, for example, a relatively narrow frontal squama with flattened right and left sides, a marked frontal keel, posteriorly divergent temporal lines, a posteriorly flexed anteromedial corner of the mandibular fossa, a bulbous lateral end of the supraorbital torus, and a forward protruding maxillary body with a distinct infraorbital sulcus. LB1 is most similar to early Javanese Homo erectus from Sangiran and Trinil in these and other aspects. We conclude that the craniofacial morphology of LB1 is consistent with the hypothesis that H. floresiensis evolved from early Javanese H. erectus with dramatic island dwarfism. However, further field discoveries of early hominin skeletal remains from Flores and detailed analyses of the finds are needed to understand the evolutionary history of this endemic hominin species. Ó 2011 Elsevier Ltd. All rights reserved. Introduction Westaway et al., 2009). The unusual combination of extremely small brain size, short stature, and other unique physical traits of Homo floresiensis is a small-bodied, hominin species that lived H. floresiensis have led some to argue that the skeletal remains on the Indonesian island of Flores in the late Pleistocene. Skeletal represent a population of pathological modern humans. However, remains of this species are currently only known from Liang Bua, such proponents have been unable to indicate a specific syndrome a limestone cave, where they are dated to between 74 and 17 kyr. At that fully explains these traits, and there is now growing support least 14 individuals are represented by these remains, which for the hypothesis that H. floresiensis was a late-surviving species of include LB1, an almost complete skeleton and the species holotype, pre-modern Homo (reviewed in Aiello, 2010). popularly known as ‘Hobbit’ (Brown et al., 2004; Morwood and The origins of this novel species still remain highly controversial Jungers, 2009; Morwood et al., 2009; Roberts et al., 2009; despite lively debate and further studies following the initial reports (Brown et al., 2004; Morwood et al., 2004, 2005). In fact, fl * Corresponding author. nominated candidates for ancestral species of H. oresiensis include E-mail address: [email protected] (Y. Kaifu). Javanese Homo erectus and pre-erectus grade hominins such as 0047-2484/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2011.08.008 Y. Kaifu et al. / Journal of Human Evolution 61 (2011) 644e682 645 Homo habilis or even Australopithecus (e.g., Brown et al., 2004; description of the external cranial morphology of LB1, and assess its Argue et al., 2009; Brown and Maeda, 2009; Lyras et al., 2009; morphological affinities. Morwood and Jungers, 2009). All these possibilities have major implications for our understanding of the evolution of genus Homo. Background and the scope of this study If H. floresiensis evolved from a habiline-like ancestor on Flores, then H. erectus sensu lato (H. erectus s. l.) was not the first hominin The LB1 skeleton is that of an adult individual whose sex is species to disperse into Eurasia, as assumed in the current Out of presumed to be female on the basis of pelvic morphology (Brown Africa 1 hypothesis (Morwood and Jungers, 2009). It would also et al., 2004; Jungers et al., 2009b). The cranium is almost imply that H. erectus and another more primitive form of Homo complete (Reference number LB1/1; Figs. 1 and 2) and is the only coexisted in Southeast Asia for a substantial period. Alternatively, if example of a H. floresiensis cranium yet recovered (Morwood and H. floresiensis originated from Asian H. erectus, then insular Jungers, 2009). In this section, we review the published studies dwarfing to an unparalleled degree has been a significant factor in on its morphological affinities. early hominin evolution on Flores (Brown et al., 2004). In the original reports of H. floresiensis, Brown et al. (2004) Skeletal evidence of the first hominins to colonize Flores would and Morwood et al. (2005) described “a mosaic of primitive, provide direct and conclusive evidence for the evolutionary history unique and derived features not recorded for any other hominin” of H. floresiensis, but further study of the Liang Bua hominin in the cranium and other skeletal parts of LB1. For instance, they remains is also essential. In this paper, we provide a detailed found that the endocranial capacity is small and comparable to Figure 1. Facial, posterior, right lateral, left lateral, superior, and basal views of LB1/1 oriented based on the Frankfurt Horizontal. Scale ¼ 5 cm. 646 Y. Kaifu et al. / Journal of Human Evolution 61 (2011) 644e682 Australopithecus; the face was said to be Homo-like, lacking a series most probably evolved from an ancestral H. erectus population on of characteristic morphologies of Australopithecus such as a great Flores as a result of long-term isolation and insular dwarfing facial height, marked prognathism, large tooth crown size, and an (Brown et al., 2004). With the recovery of additional H. floresiensis anteriorly oriented infraorbital region; the cranial vault is similar to postcranial remains, however, the Liang Bua research team were those of H. erectus s. l. in height-breadth relationships, bone less certain about the genealogy of H. floresiensis e noting the thicknesses, and some basicranial traits; and the frontal resembles species “is not just an allometrically scaled-down version of those of early African and Dmanisi Homo in exhibiting a strong H. erectus” (Morwood et al., 2005: 1016). midsagittal curvature. Furthermore, principal component analysis Subsequently, two studies employed multivariate analyses of (PCA) based on 5 cranial vault measurements (Howells’ GOL, WFB, linear cranial measurements to further investigate LB1’s morpho- XCB [SMCB of us], ASB, and VRR: see Table 1 and Howells, 1973) also logical affinities. Argue et al. (2006) conducted canonical variate showed that the vault shape of LB1/1 is, among extant and various analyses (CVA). Their Analysis 3 is based on 5 cranial vault fossil hominin crania, most similar to KNM-ER 3733, KNM-ER 3883, measurements (Howells’ GOL, XCB, BBH, AUB, ASB: data of LB1 Sangiran 2 and another unspecified Indonesian H. erectus (Brown cited from Brown et al., 2004) and includes a recent modern human et al., 2004: SOM Fig. 1). sample (Howells’ data), as well as a small sample of australopith- On the basis of the location and age of the find, as well as some ecine and early Homo specimens (Sts 5; OH 24; KNM-ER 406, 1813, morphological traits, it was initially suggested that H. floresiensis 3733, 3883; D2280; Sangiran 17; five Ngandong H. erectus). Another Figure 2. Surface rendered CT images of LB1/1. The orientations and scale same as in Fig. 1. Table 1 Craniofacial measurements of LB1. Abb. This studya Brown et al. Definition [M57, H73, K08]b (2004) Cranial vault length Maximum cranial length GOL (139) (143) Glabellaeopisthocranion [1, GOL, 1] Cranial vault breadth Postorbital breadth POBB 71 Min. transverse breadth across the frontal squama [9(1), e,4] Maximum frontal breadth XFB 84 Max. transverse breadth across the frontal squama [10, XFB, 5] Minimum frontal breadth WFRB 61 67 Measured between the superior lines when the temporal line is split into the superior and inferior branches [z9, e,6] Bi-stephanic breadth BSTB 64 Stephanionestephanion. As above [z10b, zSTB, 7] Bi-temporal line breadth on parietal BTLB 64 Min. breadths between the superior temporal lines on the parietals [e, e, e] Squamosal suture breadth SQSB 110 The posterior end of the squamosal suture is defined at the posterior tip of the supramastoid crest [8c, e,8] Maximum bi-parietal breadth XBPB 110 Max.
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