Athena Review Vol. 5, no. 1
Records of Life: Fossils as Original Sources
46. Homo erectus 3: China
China, along with Java, was the first place outside of Africa where evidence of early hominins was found.
The "Peking Man" discoveries at Zhoukoudian in the 1920s and 30s established that early humans had travelled far out of Africa during the Early and Middle Pleistocene. Correlation of the fossil remains with those of Java led by the 1940s to the realization that Homo erectus represents both a long-lived, and a very wide-ranging stage of human development.
Fig.1: Locations of Homo erectus sites in China (map: Athena Review).
Later work in China has established a series of further revelations, both of the wide variety of physical traits of Asian Homo erectus, and its great age in China. The recent redating of the Lantian skull to about 1.62 mya places it on an Early Pleistocene temporal footing with the earliest Homo erectus migrants out of Africa, at Dmanisi in Georgia. In the Middle Pleistocene time range of about 800,000-400,000 years ago, both the Zhoudoudian and Yunxian skulls are comparable to varous examples from Java. The youngest hominin fossils attributed to Homo e
rectus
in China, such as Dali from about 200,000 years ago, have some features
comparable to both Solo in Java, and various "archaic" Homo sapiens in
Europe, but retain other traits that seem unique to Chinese Homo erectus.Interestingly, one of the most controversial aspects of Chinese Homo erectus has been the identification of the long-lived species with the Chinese physical character. Weidenreich, whose basic interpretation of the Zhoukoudian fossils is of first importance, also believed in the significance of geographic variation. Some of Weidenreich's (1946) findings on the association between Homo erectus and Homo sapiens may have been influential in both Chinese palaeoanthropology and the multiregional school of modern human origins (Wolpoff 1989; Brown xxxx).
In association with the excavation of new sites there has been an intensive effort to date the Chinese sequence. Debate over the Early Pleistocene age of cave localities such as Longgupo or Yuanmou often hinges on palaeomagnetic interpretations of Matuyama or Bruhnes epoch associations (Wei Dong 2016).
Fig.2: Time line of Homo erectus in China, correlated with Pleistocene stages.
Zhoukoudian
In
the 1920s the famous “Dragon Bone Hill” (Longgushan) quarry site at
Zhoukoudian (fig.3), located 50 km south of Peking (Beijing), yielded hominid fossils identified by Davidson Black as
Sinanthropus pekinensis, or “Peking Man”. The Longgushan limestone
deposits, originally mined for fossils called “dragon bones,” which
were ground up and sold by druggists as aphrodisiacs, eventually
provided the largest known sample of Homo erectus fossils. Fig.3: The quarry at Zhoukoudian called Dragon Bone Hill (Longghushan) (photo: R.Ciochon, in Athena Review 4(1), 2004).
Davidson Black and Franz Weidenreich: The Discovery of "Peking Man"
In 1923, a molar and premolar excavated in 1921-1923 at Dragon Bone Hill in Zhoukoudian by paleontologist Otto Zdansky were sent to Canadian anatomist Davidson Black (fig.4) for identification. Black and geologist J. Gunnar Andersson then organized field work led by Birger Bohlin, who found another molar in 1927, enabling Black to make a positive identification., He identified the fossils as belonging to a new species, Sinanthropus pekinensis, informally called "Peking Man."
Expanded excavations by the Chinese paleontologists Z. Yang and W.C. Pei in 1928-1929 found jaw fragments and a well-preserved skull at Zhoukoudian’s Locality 1. When freed of its limestone matrix, the skull was clearly that of a human ancestor, resembling the Trinil 2 skullcap, and vindicating Black’s original identification.
.
Black continued to work on the Zhoukoudian material in his lab at Peking Union Medical College until his premature death in 1934. His work was taken over by the anatomist Franz Weidenreich (1875-1948; fig.5), who made casts of the growing number of hominid fossils. With Japanese invasions in 1937 he stored the fossils in a bank vault, but was not permitted to remove them from China when World War II broke out.
Fig.4: Davidson Black (1884-1934).
By 1937, when work halted at Zhoukoudian due to Chinese civil war, excavations under W.C. Pei, Lanpo Jia, and colleagues such as Teilhard de Chardin had produced six skullcaps, eleven mandibles, a mixed assortment of facial bones, and about 150 teeth. This sample, while notably lacking in long bones and extremities, represented over 40 individuals of both sexes and different ages, with cranial capacities from 915-1225 cc (Boaz and Ciochon 2004).
Fig.5: Franz Weidenreich (1875-1948) (Photo: Frankfurt Am Main Archive).
All but a few teeth were subsequently lost. The accurate casts made by Weidenrich, however, included a relatively large sample of six skulls (figs.6,7), and bones from numerous individuals. These casts are at the American Museum of Natural History and the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing. Weidenreich, already familiar with some of the Pithecanthropus finds from Java,eventually completed a detailed analysis of the Zhoukoudian hominids in 1943. Recognizing many similarities with the Java fossil hominids after consultations with von Koenigswald, Weidenreich nevertheless retained in his publications the original, regionally distinct taxa of Sinanthropus pekinensis and Pithecanthropus erectus. He supported, however, combining both as Homo erectus as part of a single, evolving Homo lineage. Similarities between Java Man and Peking Man led Ernst Mayr to rename both as Homo erectus (Mayr 1950), which has remained in current usage.
Just as the date ranges of the Java sites are being refined, Zhoukoudian’s date range has also been under under revision, along with that of several other Middle Pleistocene Homo erectus sites in China (Etler 2004). In 1985, uranium isotope dating gave absolute dates on hominid-bearing layers at Zhoukoudian’s Localities 1 and 2 from 290,000 - 230,000 BP. Yet the upper strata have recently been redated to be substantially earlier, at 410,000 BP, while the lowest hominid layer is redated at 670,000 BP.
Homo erectus: cranial capacity
The Chinese skulls resembled those from Java, but tended to be later, and to have a higher cranial capacity averaging 1000-1100 cc, compared to about 800-900 cc for Pithecanthropus.
Fig.6: Homo erectus skullcap, from 1920s-30s excavations (American Museum of Natural History, cast; photo: Athena Review)
Zhoukoudian ZKD V is a fossil cranium from Zhoukoudian with a relatively well-preserved brain endocast. The cranial capacity is estimated by Wu et al. (2010) to be 1140 cubic centimenters (cc), based on the brain endocast (fig.7). This compares with a range of 915 to 1225 cc for a total of six crania measured for the Zhoukoudian Homo erectus specimens. The ZKD V cranium is dated between 500,000 and 400,000 years ago, possibly closer to 415,000 years ago according to Wu et al. (2010). The endocast, while missing some areas, clearly shows several of the main lobes of the cerebral hemisphere, as well as grooves or vessels for meningeal arteries found in the outer portion of the skull, which supply it with blood.
A mean cranial capacity of about 1050 cc. in the Zhoukoudian skulls shows larger brain size compared to the somewhat earlier Java H. erectus, but there are many basic similarities. The low-slung, thick-walled skulls from both Java and China are widest at their base, and have large brow ridges, a sagittal keel on top, and a protruding ridge at the rear (occipital) bone (based on the reconstruction by Tattersall and Sawyer 1996).
Fig.7: Homo erectus skull and brain endocast of Zhoukoudian ZKD V (after Wu et al.2010 fig.2)
Subsistence modes and toolmaking of Homo erectus
In excavations which lasted (with wartime interruptions) from 1921-1982, the Longghushan cave site revealed over 100,000 artifacts of worked stone and cut or burned animal bone, the latter often found in the same levels producing Homo erectus fossils. The lithic artifacts, first systematically recorded in 1931, and dominated by quartz flakes, represent a Mode 1 or flake tool industry. Tool forms include retouched or sharpened flakes, scrapers, core tools or choppers, and pointed awls used for piercing. When discovered in the 1920s-30s, these were the first significant artifactual remains to be found in situ with hominid fossils. Since initial descriptions in the late 1930s, however, their interpretation has been rife with controversy (Boaz et al. 2000; Boaz and Ciochon 2004).
An abundance of animal bones found at Zhoukoudian included thousands of deer bones, some charred by fire. Given these apparent food remains, plus numerous stone tools, original interpretations of the cave site portrayed it as a home base for bands of Homo erectus who subsisted by hunting and gathering. Evidence of fire in levels containing burned bone suggested cooking, and the use of fire for warmth seemed consistent with this northernmost settlement area for Homo erectus. Bone artifacts including cut marks from stone tools were identified by Henri Breuil (1939), who visited China several times in the 1930s. Signs of possible cannibalism were also seen by Breuil (1939) and Weidenreich (1943) in breakage and widening of the foramen magna of several skull bases, possibly to remove the brain.
More recent reexamination of the evidence, however, has significantly altered views about a hunting subsistence for Homo erectus at Zhoukoudian. The overall sample of hominid bones itself provided the first important clues that the caves may have served other functions. Weidenreich (1943) first remarked on the fragmentary nature of the Homo erectus fossil assemblage at Zhoukoudian, which generally lacked long bones and extremities, and where skulls had been broken up apparently even before their deposition in cave sediments. This has been confirmed by recent, detailed examination of the remains at Zhoukoudian locality I by Boaz et al. (2000), revealing that hominid bones made up only half of one percent (0.5%) of the total bones at the site.
The high proportion of isolated teeth and skull parts, and the low incidence of long bones or hands and feet, is typical of the bone accumulation in a carnivore den. Some Homo erectus facial bones and crania (as first noted by Weidenreich) show punctures and other marks typical of carnivores, undoubtedly including the large, lion-sized Pleistocene cave hyena Pachycrocuta brevirostris, whose bone remains are the most frequent at the site. Hyenas also seem to have been responsible for the widening of the foramen magnum on H. erectus remains, evidence which could no longer pertain to cannibalism. Other evidence, nevertheless, of cut marks by stone tools on Homo erectus Skull V (first noted by Weidenreich) does show that cannibalism must have occurred at Zhoukoudian (Boaz and Ciochon 2004).
Postcranial bones also exhibit typical hyena tooth marks and breakage, as part of hyaenid scavaging or predation of Homo erectus. Many of the arm and leg bones would have been totally consumed by hyenas or other carnivores, thus accounting for their relative lack in the overall sample. The conclusion inevitably drawn by Boaz et al. (first advanced in 1929 by the archaeologist W. C. Pei [1938]) is that the Zhoukoudian caves, rather than being home bases for hominid hunters, were, for much of the time, dens for the giant cave hyena.
Controversy has also persisted over the use of fire at the site. Ash in layers that had been taken as signs of hearth fires showed no evidence of phytoliths (silica particles) typically left by burning wood or other vegetation (Weiner et al. 1998). This has supported the interpretation that most fires at Zhoukoudian were not confined to hearth features, and thus could be due to natural causes (Binford and Stone 1986). Yet some of the main layers with Homo erectus fossils at Zhoukoudian cave (e.g., layer 10) also contain burned bone with stone tools, which would seemingly indicate hominid use of fire, very likely for cooking (Weiner et al. 1998).
The flake tool (Mode 1) assemblages contain lithic items useful for defleshing (scrapers and flakes) and for pounding bones to extract marrow (core choppers). Binford and Ho (1985) proposed that Homo erectus subsisted by scavenging, or foraging for meat left on bones in caves occupied primarily by cave hyenas, wolves, or bears. This is supported by close examination of animal bones at Zhoukoudian, which shows that cut marks made by Homo erectus with quartz flakes overlie carnivore bite marks. While the caves at Zhoukoudian often functioned primarily as dens for the cave hyena, they thus also must have served as foraging areas for Homo erectus, who may have camped nearby, possibly making shelters from tree branches (Boaz and Ciochon 2004).
Longuppo Cave
In the past two to three decades, several Lower Pleistocene hominid sites have been identified in China which predate the Homo erectus finds at Zhoukoudian. Among the most controversial is Longgupo Cave on the Yangtze River (fig.1), where a mandible fragment and teeth dated at 1.9 mya by ESR methods have been compared to early African Homo habilis and Homo erectus georgicus at Dmanisi (Wanpo et al. 1995). Associated with the teeth were pitted and abraided cobble tools of an exotic andesite-porphyrite.
Also found at Longgupo were teeth of Gigantopithecus, a huge Plio-Pleistocene ape also occurring with Homo erectus at another Chinese site, Jianshi Cave, and at Tham Khuyen Cave in Vietnam, dated at 475,000 BP (Ciochon et al. 1996). While debate continues on the hominid identification of the Longuppo teeth, the site is of potentially great significance in terms of initial dispersal from Africa.
Lantian
In Lantian county, a partial cranium from Gongwangling on the Yellow River, found in 1964, represents China’s earliest known Homo erectus fossil. Previously dated at about 1.15 million years ago (mya; Etler 1996), more recent testing
published in 2018 in the Journal of Human Evolution
has established a significantly earlier date of 1.63 mya,
making it the oldest fossil hominin cranium known in northeast Asia,
and the second oldest site with cranial remains outside Africa. Only
the Dmanisi crania from Georgia that, like Lantian, are relatively
small-brained, are older, at around 1.75 million years old. One skull of a female at Lantian, aged about
30, has a cranial capacity of 780 cc, comparable to individuals from
Koobi Fora and Sangiran, and the largest examples from Dmanisi.Fig.8: Skull of Lantian Homo erectus. A: view of skullcap from top. B: lateral view from right side (after Zhu et al. 2018).
Resarchers including Zhu Zhaoyu, Robin Dennell, and Huang Weiwen (2018) used various methods to redate the geological context of the Lantian skull. including loess stratigraphy, sedimentology and mineralogy, geochemistry, and paleomagnetism. The fossil horizon is situated in a paleosol unit located between the Gilsa Event (average age ca. 1.62 mya) and the Olduvai Subchron (top age 1.77 mya) . A similar subtropical faunal assemblage was found at both the Gongwangling sections and at Jiacun, in the same stratigraphic position identified with this paleosol unit. By extending the age of the site by half a million years, according to Dennell, the Gongwangling site has become a critical point of reference for understanding the migration and dispersal of early man in the Old World.
Nihewan Basin
Other early tool evidence comes from the Nihewan Basin in north China (40 N), a lacustrine region known for Villefranchian-age fauna (Zhu et al. 2001). While the site lacks hominid fossils, a Mode 1 flake tool assemblage of over 3000 items including side- and end-scrapers has been found in layers dated paleomagnetically to 1.36 mya, a relatively early date for hominids in a north-temperate zone.
By 800,000 years ago, bifacial stone tools were being made in the Bose Basin in south China, partly resembling an Acheulean industry but lacking handaxes (Hou Yamei et al. 2000).
Yunxian
Two crania (EV 9001 and EV 9002) were found in 1989 and 1990 near Mitousi Village, Yunxian County, Hubei Province. These are described by Li and Etler (1992) as Chinese Homo erectus, distinct from "archaic" Homo sapiens such as known in Europe. A mean ESR date from mammal tooth enamel at Yunxian is
581,000 BP with a range from 800,000 to 455,000 over ten dates (Chen et al.1997) The date of 581,000 BP falls near the middle of the Chinese Homo erectus sequence, and is signifianctly older than the earliest dated examples of archaic Homo sapiens in China (Jinniushan dated at 280,000 BP, Dali at 200,000 BP, and Mapa at 132,000 BP). Recovered lithic artefacts at Yunxian include a small number of bifaces (Li et al. 1998).Fig.9: Skull EV 9002 from Yunxian (after Li and Etler 1992, fig.4b)
Of the two crania, EV 9002 (fig.9) is better preserved but still extensively fractured and distorted. While few cranial dimensions can be certain, it is clear that both crania are relatively massive. Secondly, if facial distortion is taken into account there is a close similarity in both size and morphology between the facial skeletons of Sangiran 17 and Yunxian EV 9002. Neither resemble Jinniushan (Wu, R. 1988) or Dali (Wu, X. 1981; Brown xxxx)
.
Nanjing
Chinese Homo erectus occupations from 670,000-400,000 occurred not only at Zhoukoudian, but also at Tangshan Hill near Nanjing, in Yunxian at Quyuanhekou (both about 620,000-580,000 BP), and the Hexian site at Anhi (over 400,000 BP). The Nanjing and Zhoukoudian Homo erectus specimens seem close in morphology, and Middle Pleistocene faunal assemblages from the two sites are very similar.
Jinniushan
Later Chinese hominids: Based on these revised dates, fossil evidence of Homo erectus is missing in China between about 400,000 and 280,000 BP. Later hominids then appear with more rounded skulls and (often) larger brain cases, including a young male from Jinniushan dated at 280,000 years ago, with a modern-sized cranium of 1400 cc.
Hexian
At Hexian, a well-preserved Homo cranium (1025 cc) with reduced brow ridges has been dated to about 200,000 BP, contemporary with the Dali cranium (fig.10), which shows large brow ridges and a flat, high face. The still later Mapa cranium (about 132,000 BP) may be contemporary with Homo soloensis populations from Java’s Ngandong deposits. Controversy persists on whether Homo heidelbergensis populations migrated into China and overlapped the latest Homo erectus groups as a separate species, or there was a more gradual, in situ transition.
Dali
The Dali cranium (fig.10) was discovered in 1978 in a loess terrace near Jiefang Village, Dali County, Shaanxi Province. The skull is largely present and relatively well preserved, with some postdepositional crushing and displacement of the palate and left maxilla. A large section of the right parietal is missing, as are the maxillary teeth and left zygomatic arch. Uranium-series dating of ox teeth from the site obtained a date of 209,000 BP (Chen et al 1994).
Stone tools, primarily scrapers, were also recovered from the site. (Brown xxxx)Wu (1981, 1989) found that most of the cranial dimensions and morphological features of Dali were intermediate between Homo erectus and Homo sapiens, with Dali assigned to "archaic" Homo sapiens. Craniofacial anatomy and vault shape are distinct from European Neanderthals and earlier European hominids like those at Petralona and Atapuerca. Unlike Homo erectus, maximum cranial breadth is located on the posterior-superior temporal, rather than the cranial base. Most of the vault superstructures in the temporal, occipital and frontal regions are robust. The mastoid process is small and the occipital and nuchal plane form a sharp angle similar to Homo erectus, with an endocranial volume of 1100–1200 m3, comparable to some Zhoukoudian skulls (Brown xxxx).
While Dali’s cranial vault is relatively robust, with a mixture of Homo erectus and Homo sapiens traits, the facial skeleton contains some gracile features such as the zygomatic arch, which is verry similar to those in modern Homo sapiens.
Fig.10: Skull from Dali, China (American Museum of Natural History cast; photo: Athena Review)
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Zhu Zhaoyu, Robin Dennell, and Huang Weiwen (2018)
Glossary