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Global and Local Perspectives on Cranial Shape Variation in Indonesian Homo Erectus

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Global and Local Perspectives on Cranial Shape Variation in Indonesian Homo Erectus ANTHROPOLOGICAL SCIENCE Vol. 125(2), 67–83, 2017 Global and local perspectives on cranial shape variation in Indonesian Homo erectus Karen L. BAAB1*, Yahdi ZAIM2 1Department of Anatomy, Midwestern University, Arizona College of Osteopathic Medicine, 19555 N. 59th Avenue, Glendale, AZ 85012, USA 2Department of Geology, Institut Technologi Bandung (ITB), Jl. Ganesa No. 10, Bandung–40132, Indonesia Received 5 December 2016; accepted 13 April 2017 Abstract Homo erectus is among the best-represented fossil hominin species, with a particularly rich record in Indonesia. Understanding variation within this sample and relative to other groups of H. erectus in China, Georgia, and Africa is crucial for answering questions about H. erectus migration, local adap- tation, and evolutionary history. Neurocranial shape is analyzed within the Indonesian sample, including representatives from Sangiran, Ngandong, Sambungmacan, and Ngawi, as well as a comparative sample of H. erectus from outside of Java, using three-dimensional geometric morphometric techniques. This study includes several more recently described Indonesian fossils, including Sambungmacan 4 and Skull IX, producing a more complete view of Indonesian variation than seen in previous shape analyses. While Asian fossils can be distinguished from the African/Georgian ones, there is not a single cranial Bauplan that distinguishes all Indonesian fossils from those in other geographic areas. Nevertheless, late Indone- sian H. erectus, from sites such as Ngandong, are quite distinct relative to all other H. erectus groups, including earlier fossils from the same region. It is possible that this pattern represents a loss of genetic diversity through time on the island of Java, coupled with genetic drift, although other interpretations are plausible. A temporal pattern of diachronic change was identified within Indonesia for the posterior neu- rocranium such that younger Sangiran fossils more closely approached the Ngandong/Sambungmacan/ Ngawi pattern, but there was not a linear trend of shape change from Sangiran to Sambungmacan to Ngandong, as has been suggested previously. The Sambungmacan 3 fossil, which often appears as a morphological outlier, fits the general pattern of late Indonesian vault shape, but has a more extreme expression of the shape trends for this group than other individuals. Key words: Homo erectus, Sangiran, Ngandong, cranial shape, geometric morphometrics dong, Sambungmacan, and Ngawi is generally considered to Introduction be Middle Pleistocene or even Late Pleistocene in age. The The Homo erectus fossil record from Java is extensive and Ngandong fossils, consisting of numerous partial and com- has wide-ranging implications for evolutionary questions plete calvaria and well as a few postcranial elements, were about the timing of early Homo migration, the effects of recovered from excavations on the high terraces of the Solo stochastic evolution on islands, local adaptation, and a po- River. However, the three Sambungmacan and single Ngawi tential speciation event that gave rise to Homo floresiensis, calvaria were recovered by local inhabitants from three dif- known from the nearby island of Flores (Brown et al., 2004; ferent sites along this river, and the exact stratigraphic prov- Baab, 2016a). Moreover, the time span, which may encom- enance of these fossils is uncertain. Most of these sites are pass more than a million years of evolution, allows us to situated along the Solo River in east Java, with the exception probe issues of diachronic change in this species. The oldest of the more eastern site of Modjokerto, which has yielded fossils (early Pleistocene) from Java are those from Trinil one young individual, and the more western site of Cisanca, and the Sangiran dome. The young individual recovered from which a single incisor was recovered (Kramer et al., from Modjokerto may also be Early Pleistocene in age, but 2005). this is less certain. A second group of fossils from Ngan- The Asian hominin record has been subject to a continu- ous stream of analyses beginning with the initial description of the Trinil fossils by Dubois (1894). Additional earlier * Correspondence to: Karen Baab, Department of Anatomy, Mid- western University, 19555 N. 59th Avenue, Glendale, AZ 85012, discoveries were also from the earlier time frame. The first USA. fossils recovered from the Middle/Late Pleistocene were E-mail: [email protected] from Ngandong in the 1930s. The Ngandong fossils were Published online 30 June 2017 initially placed in a distinct species, Homo (Javanthropus) in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.170413 soloensis (Oppenoorth, 1932), and similarities to other Mid- © 2017 The Anthropological Society of Nippon 67 68 K.L. BAAB AND Y. ZAIM ANTHROPOLOGICAL SCIENCE dle Pleistocene hominins or late Pleistocene Homo sapiens descent” and Antón (2002: 319), who asserted that “the rela- from Australasia were noted. However, monographs on the tively small differences between the earlier and later Indone- Zhoukoudian (China) and Ngandong fossils by Weidenreich sian fossils must certainly be accepted as temporal changes were particularly influential (Weidenreich, 1943, 1951) in within a lineage, rather than indicative of a different line- establishing two common themes that have persisted until age.” Both workers identified non-metric features (e.g. fron- the present: the fossils from Trinil/Sangiran, Zhoukoudian, tal keel and anterior projection of the medial part of the su- and Ngandong share a common Bauplan but with regional praorbital torus) unique to this group, and Antón (2002) and temporal variations overprinted on this shared architec- further found similarity in size/shape of the cranial vault ture. Santa Luca’s (1980) in-depth analysis of the Ngandong based on quantitative measures. fossils further solidified this sample as part of the broader Nevertheless, there are differences of opinion regarding H. erectus type. Additional fossil discoveries have occurred the details of that evolutionary transition. In particular, re- in both the earlier time frame, including additional Sangiran cent analyses of 2D linear dimensions and non-metric fea- dome localities (Sartono and Tyler, 1993; Widianto et al., tures supported a linear transition from Sangiran/Trinil 1994; Widianto and Grimaud-Herve, 2000; Grimaud-Hervé through Sambungmacan/Ngawi and onto Ngandong (Baba et al., 2005; Zaim et al., 2011), and the later time frame, such et al., 2003; Kaifu et al., 2006, 2008, 2015), a view also em- as the three calvaria from Sambungmacan (Jacob et al., braced by Wolpoff (1999). This conclusion was based on 1978; Márquez et al., 2001; Baba et al., 2003) and the one observations that geochronologically younger Sangiran fos- from Ngawi (Sartono, 1991). Sambungmacan and Ngawi sils were more similar to later Indonesian fossils than were are considered to have particular affinities with Ngandong. older ones, and that both Ngawi and the three Sambung- More recent work that has focused on the cranial mor- macan specimens displayed intermediate morphologies be- phology of the Asian H. erectus fossil record includes evalu- tween Sangiran/Trinil and Ngandong. Durband (2006) sug- ations of the metric and non-metric diversity of Asian or gested near stasis in the vault shape in Indonesia for specifically the Indonesian fossil record (Antón, 2002; Kaifu 0.7–1.0 Myr based on canonical variate analysis of size- et al., 2008), cranial shape variation in the Asian fossils corrected neurocranial length and breadth dimensions, but (Baab, 2010; Zeitoun et al., 2010), and analyses focused on identified a handful of non-metric features in the later Indo- the affinities of particular fossils (Delson et al., 2001; nesian sample not present in Sangiran/Trinil. He attributed Widianto et al., 2001; Antón et al., 2002; Widianto and this combined pattern to geographic and genetic isolation of Zeitoun, 2003; Durband, 2006; Kaifu et al., 2006, 2015; these populations on Java. In contrast, analysis of vault Indriati and Antón, 2010). It was demonstrated that Asian, shape via 3D landmarks demonstrated distinct neurocranial along with numerous African and Georgian fossils assigned shape in the Sambungmacan/Ngawi, Ngandong, and Trinil/ to H. erectus, were distinct from or displayed minimal over- Sangiran groups, and the differences among them were lap with other archaic Homo species on the basis of three- non-linear (Baab, 2010; Zeitoun et al., 2010). At least some dimensional (3D) neurocranial shape (Zeitoun et al., 2010; of the differences between the early Indonesian and the Baab, 2016b). There is substantial variation within this sam- Ngandong fossils appear to relate to the increased size of the ple that spans three continents and more than a million years latter (Antón, 2002; Baab, 2016b). A small number of work- of evolutionary change. Some of this variation is attributable ers argue that the morphological distance between the Trinil/ to diachronic change (Wolpoff et al., 1994; Antón, 2003; Sangiran and later Indonesian fossils should perhaps be rec- Kaifu et al., 2005, 2008), and to geography (perhaps related ognized at the species level (Schwartz and Tattersall, 2005; to genetic drift or local adaptation) (Antón, 2002; Kidder Zeitoun et al., 2010), with a further possible split between and Durband, 2004; Durband, 2006; Baab, 2010), but is also Sambungmacan/Ngawi and Ngandong (Tattersall and influenced by cranial size (Santa Luca, 1980; Antón et al., Schwartz, 2009), but with the recognition
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