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1000002291104.Pdf Quaternary International 211 (2010) 4–13 Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint Morphological and morphometric analysis of variation in the Zhoukoudian Homo erectus brain endocasts Xiujie Wu a,b,*, Lynne A. Schepartz c, Christopher J. Norton d a Laboratory of Evolutionary Systematics of Vertebrates, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Xizhimenwaidajie 142, Box 643, Beijing 100044, China b State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Nanjing 210008, China c Department of Anthropology, Florida State University, Tallahassee, FL 32306-7772, USA d Department of Anthropology, University of Hawaii at Manoa, Honolulu, HI 96822-2223, USA article info abstract Article history: The six Zhoukoudian (ZKD) Locality 1 Homo erectus specimens derive from stratigraphic levels 11–3 with Available online 19 July 2009 a geochronological span of approximately 0.3 Ma. This paper introduces the history of the ZKD endocasts and presents data on their morphological features and linear dimensions in order to evaluate variability in the sample over time and in the broader context of human brain evolution using a comparative sample of African and other Asian H. erectus fossils and modern Chinese males. The ZKD brains are very similar in their morphological characteristics, but there are also significant but subtle changes involving expansion of the frontal and occipital lobe breadths that correlate with the geochronology. The same is not true for general endocranial volume. The ZKD brains, together with other Asian and African H. erectus specimens, have low height dimensions and short parietal chords that distinguish them from the modern Chinese. Furthermore, the lack of geographical patterning in the fossil sample, as determined by Principal Components Analysis, provides no support for arguments advocating the splitting of H. erectus into separate taxa. Ó 2009 Elsevier Ltd and INQUA. All rights reserved. 1. Introduction Homo erectus endocasts, a group of six specimens from a single locality, provide fundamental data for studying hominin brain The endocast is the impression taken from the inside of evolution. a cranium that retains the surface features of the brain. It does not For more than 70 years, a number of scholars conducted display the original brain anatomy, but instead reflects the external research on the ZKD endocasts (e.g., Black, 1932, 1933; Dubois, features of brain anatomy in detail. Studying endocasts of fossil 1933; Ariens-Kappers, 1934; Shellshear and Smith, 1934; Wei- hominins allows researchers to make inferences on functional denreich, 1936; Ariens-Kappers and Bouman, 1939; Broadfield anatomy, physiology, and phylogeny (Bruner, 2003). Thus, endo- et al., 2001; Holloway et al., 2004; Wu et al., 2009). These investi- casts from fossil crania are unquestionably the most important gations began in 1931 when Davidson Black manually recon- materials for analyzing and understanding human brain evolution structed the endocast of ZKD skull III (Locus E) using plaster. (Holloway, 1980; Falk, 1985; Begun and Walker, 1993; Broadfield He suggested the brain was essentially human in its form, that the et al., 2001; Bruner, 2004). Because hominin fossils are very rare, individual was right handed, and that there was evidence for the and their crania are often fragmentary, distorted, or not sufficiently neurological underpinnings of articulate speech (Black, 1932, 1933). prepared for endocasting, the number of hominin endocasts His work initiated the study of the actual brains of ancient humans available for analysis is quite limited and few are complete enough and apes. Shellshear and Smith (1934) argued that ZKD III displayed for morphometric comparisons. The Chinese Zhoukoudian (ZKD) primitive characters that did not appear in Pithecanthropus erectus such as a flat frontal lobe and a well-marked frontal keel. In the summer of 1930, a number of skull fragments from Locus D were refitted as the adult ZKD II. Weidenreich (1935) reconstructed the * Corresponding author. Laboratory of Evolutionary Systematics of Vertebrates, ZKD II endocast. Subsequently in 1936, ZKD X (Skull I of Locus L), Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of ZKD XI (Skull II of Locus L) and ZKD XII (Skull III of Locus L) were Sciences, Xizhimenwaidajie 142, Box 643, Beijing 100044, China. Tel.: þ86 10 88369124; fax: þ86 10 68337001. found. These three endocasts were reconstructed by Weidenreich E-mail address: [email protected] (X. Wu). (1938, 1941). The endocast of ZKD V was reconstructed in 1973 1040-6182/$ – see front matter Ó 2009 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2009.07.002 X. Wu et al. / Quaternary International 211 (2010) 4–13 5 based on five fragments of cranium found in Locus H III. Two fragments, consisting of the temporal and adjoining parietal and occipital bones, were excavated in 1934 and described by Wei- denreich (1935). More fragments of the cranium, including the frontal, parietal, sphenoid and occipital bones, were found in 1966. The endocranial volume was determined (Qiu et al., 1973), but no detailed analysis of the endocast was undertaken until recently (Wu et al., 2009). In addition to the works on the ZKD endocasts mentioned above, some broader comparative studies have included the ZKD speci- mens. For example, Begun and Walker (1993) compared the endocast of KNM-WT 15000 with ZKD II, III, X and XII. They found that the overall morphology of those specimens was quite similar to KNM-WT 15000. However, they also found that the four ZKD endocasts can be distinguished from KNM-WT 15000 by the development of the superior sagittal sinus and the apparently well- developed connection between the sphenoparietal sinus and the middle meningeal veins. Broadfield et al. (2001) found that the endocast of Sambungmacan (Sm) 3 has a mosaic of features that are shared with both other Indonesian H. erectus and ZKD III. Wu et al. (2006) compared the endocast of Hexian with ZKD (II, III, X, XI and XII), Sm 3 and KNM-WT 15000, and determined that Hexian is most similar to the ZKD H. erectus specimens. The ZKD hominins included in this study all derive from the principal locality at Zhoukoudian designated Locality 1; a cave complex located 50 km southwest of Beijing, China. The strati- graphic sequence was extensively studied and its designations were revised by the different excavators, most notably Teilhard de Chardin, Davidson Black and Jia Lanpo. ZKD III was found in Black’s Layer 11 (Pei, 1929; Black et al., 1933; Lin, 2004) or Jia’s Layer 10 (Jia, Fig. 1. Composite section of West Profile, ZKD Locality 1 showing the stratigraphic 1959). ZKD X, XI and XII are from Layer 8/9 (Black et al., 1933; Jia, positions of the cranial discoveries (Adapted from Jia, 1959). 1959). ZKD II is from Black’s Layer 8/9 or Jia’s Layer 10. ZKD V was found in Layer 3 (Jia, 1959). Fig. 1 illustrates the approximate spatial relationships of the six crania we used in this paper. ZKD V was 2. Materials and methods found in Layer 3; ZKD II, X, XI and XII are from Layer 8/9, and ZKD III was found in Layer 11 (Lin, 2004). 2.1. Endocasts A series of different dating studies have been conducted to determine the absolute ages of the ZKD strata. Electron spin reso- Table 1 is the catalogue of endocasts used in this study. The nance (ESR) dating of mammal teeth collected from Layers 3, 4, 6–9, specimens include representatives of Asian H. erectus (ZKD, Hexian, 11 and 12 suggests an age range of 0.28–0.58 Ma for the fossils Trinil II, Sangiran 2, 10, 12, 17, and Sm 3) and an outgroup of African (Huang et al., 1993; Gru¨ n et al., 1997) with ZKD III at 0.58 Ma, ZKD II, presapiens populations (KNM-ER 3733, 3883, KNM-WT 15000 and X, XI and XII all at approximately 0.42 Ma. The stratigraphically Sale´). Endocasts of ZKD III, ZKD II, ZKD X, ZKD XI, ZKD XII, ZKD V, highest specimen is ZKD V at 0.28 Ma. However, thermal ionization Hexian and Trinil II are from the collections of the Institute of 230 234 mass spectrometric (TIMS) Th/ U dating on intercalated spe- Vertebrate Paleontology and Paleoanthropology, Chinese Academy leothem samples suggests that the age of the ZKD fossils is between of Sciences. The six ZKD endocasts are excellent first-generation 0.4 and 0.8 Ma (Shen et al., 2001). As a result, the age of ZKD V is casts prepared from the original fossils. They therefore preserve >0.4 Ma, possibly in the range of 0.4–0.5 Ma. The age of the hom- more detailed information than can be observed on the later inin fossils from the lower strata is at least 0.6 Ma, and possibly generation casts that are available to most researchers conducting 0.8 Ma or older (Shen et al., 2001). Shen et al.’s (2009) new dating comparative endocast studies. Endocasts of 31 modern Chinese 26 10 study, based upon cosmogenic Al/ Be burial dating on quartz males were made by the first author (Wu et al., 2006). They provide grains, suggests that the ages of Layers 7–10 of ZKD are between a dataset representing general aspects of modern human brain 0.68 and 0.78 Ma (Shen et al., 2009). The new age estimates imply variation. Data for the endocasts of Sm 3, Sangiran 2, 10, 12, San- that the early ZKD specimens are probably contemporaneous with giran 17, KNM-ER 3733, 3883, KNM-WT 15000 and Sale´ are from Sangiran 17 that may date to 0.7 Ma (Swisher et al.,1994). While the published studies (see Table 1).
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