Quaternary International 400 (2016) 47e57
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Biochronological framework of Homo erectus horizons in China
Wei Dong
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, IVPP, CAS, Beijing 100044, China article info abstract
Article history: A total of 15 sites yielding Homo erectus were discovered in China. Numerical age dating results vary from Available online 27 October 2015 different authors. In an attempt to evaluate and possibly refine controversial dating results, the author proposes five stages as a biochronological framework for Homo erectus horizons in China by sequencing Keywords: all hominin horizons according to fauna antiquity coefficients, faunal binary similarity coefficients, faunal Biochron extinction rates, and ecological composition similarities of the faunas. The first stage is associated with Mammalian fauna the arrival of Homo erectus in East Asia, and the last stage is associated with the decline and final Homo erectus disappearance of Homo erectus as well as arrival of Homo sapiens in East Asia. The faunas experienced Horizon Pleistocene some kind of critical transition or turnover in the third stage. © China 2015 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction Well documented mammalian faunas associated with Homo erectus (e.g. Zdansky, 1928; Young, 1932; Li and Ji, 1981) tell us not It is well known that Homo erectus (s.l.) is the earliest prehistory only the probable diet components and paleoenvironments of the tool-maker human in East Asia. The systematic excavations during prehistoric humans, but also the relative ages of their horizons to the 1920s and 1930s at the Zhoukoudian site in Beijing started a calibrate the numerical age dating. The faunas were, however, series of research on Homo erectus and related prehistory in China mostly described alone and compared with a few related faunas (e.g. Black, 1927,1930; Zdansky, 1927,1928; Pei, 1929, 1931; Teilhard (e.g. Zdansky, 1927, 1928; Hu and Qi, 1978; Lin et al., 1978; Li and Ji, de Chardin and Young, 1929; Breuil, 1931, 1939; Black et al., 1933; 1981; Ji, 1999; Wang et al., 2007; Gong et al., 2014). Only a few Andersson, 1934; Weidenreich, 1935; Teilhard de Chardin, 1941). comparisons were at larger scale such as those of North China (Qi, More and more hominin localities were discovered and excavated 1989) or South China (Han and Xu, 1989), or even the whole since the systematic excavations at Zhoukoudian (Wu et al., 1999; country (Dong et al., 2000). Here, a biochronological framework of Liu et al., 2014). Nevertheless, the numerical age dating results of Homo erectus horizons in China is proposed, by sequencing all hominin horizons vary. For example, the lower layer yielding the faunas from the hominin horizons in the whole country, including first skull-cap of Homo erectus at Locality 1 of Zhoukoudian was those from Mohui and Hualongdong discovered and documented dated as 462 ka by Zhao et al. (1985) but 750 ka by Shen et al. in the recent decades, based on faunal evolution. An attempt is (2009); that of Hexian Man was 150e190 ka by Chen et al. (1987) made to evaluate and possibly refine the numerical ages with the but 300 ka by Huang et al. (1995); that of Nanjing Man was proposed biochrons. 350 ka by Chen et al. (1996) but 500 ka by Wang et al. (2002); and that of Lantian Man at Gongwangling was 1.15 Ma by An and Ho (1989) but1.7MabyZhu et al. (2015). The most spectacular dif- 2. Regional setting ference is that for Yuanmou Man, 700 ka by Hyodo et al. (2002) but 1.7MabyLi et al. (1976) and Zhu et al. (2008), with a difference of There are 15 sites yielding Homo erectus in China, since the fi 1 My. The latter variance is not very significant for Paleozoic or even discovery of the rst skull-cap of Peking Man at Zhoukoudian in Mesozoic paleontology, but it is important for that of the Cenozoic, 1929. Besides these, the purported 16th site of Homo erectus, especially for Quaternary and human evolution research. Xichuan Man site, remains unknown (Liu et al., 2014). Thirteen teeth of Xichuan Man were collected in 1973 from traditional Chi- nese medicine stores at Xichuan County in the same Nanyang Municipality as that of Nanzhao Man in Henan Province. The follow-up investigations at several potential localities were carried E-mail address: [email protected]. out but were fruitless (Wu and Wu, 1982). It is therefore impossible http://dx.doi.org/10.1016/j.quaint.2015.09.019 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved. 48 W. Dong / Quaternary International 400 (2016) 47e57 to include this unknown site in the present study. The geographic 2.4. Yunxian Man locality at Quyuanhekou and geological settings of the 15 sites are as follows. The Quyuanhekou (“Quyuan River mouth” in English) locality 0 00 0 00 2.1. Yuanmou Man locality at Danawu (GPS: 32 50 2.21 N, 110 35 4.20 E, 203 m) is about 21.3 km west (heading: 269.7 ) of Yunxian downtown and 56.4 km southwest The site (GPS: 25 40039.5400N, 101 54042.4200E, 1284 m) is about (heading: 250 ) of the Longgudong at Meipu of Yunxian County in 5 km southeast (heading: 128 ) of Yuanmou downtown, near the northwestern Hubei Province (Fig. 1E). It is located on the north middle part of eastern Yuanmou Basin in Yuanmou County, Yunnan bank of the Han River and on the west bank of Quyuan River mouth Province (Fig. 1A). The type section of the Late Cenozoic Yuanmou which is a tributary of the Han. Two skulls of Homo erectus and Formation ranges from the eastern slope of Dongshan in southern associated fauna were unearthed from the deposits of the fourth Yuanmou County to Laoyatang by the Longchuan River. It consists terrace of the Han River over the Sinian metamorphic series of four lithostratigraphic members. Member 4, the top member bedrock. The fourth terrace deposits were divided into three parts with a total thickness of 122.2 m, is composed of fluvial and alluvial based on sedimentary facies (Li and Feng, 2001). The lower part is silty clays interbedded with sandy conglomerates, and it is further mainly coarse sand and gravel bedload, the middle part is mainly fi divided into 5 lithostratigraphic layers with Layer 28 on the top and channel margin ne sand and silt, and the upper one is alluvial silt Layer 24 at the bottom (Qian and Zhou, 1991). A pair of upper first and clay. The upper part was further divided into three layers. Layer incisors of Homo erectus were collected from Layer 25 near Danawu 3, the lowermost one which yielded hominin skulls and associated (Hu, 1973), and a left tibia of Homo erectus was collected from the fauna, is fossiliferous light brown silt clay with a large amount of southern slope of Guojiabao (Zhou, 1998) which might be equiva- calcareous concretion, and its thickness is 1.3 m (Li and Feng, 2001). lent to Layer 27. Mammalian remains from Layers 25e27 were considered as the same fauna (Lin et al., 1978). 2.5. Yunxian Man locality at Meipu
Three teeth of Homo erectus were collected from the cave de- 2.2. Tiandong Man locality at Mohui cave posits in a karst cave (GPS: 33 006.4900N, 111 8057.8800E, 263 m) “ ” 0 00 0 00 called Longgudong ( Dragon Bone Cave ) in Yunxian County, Hubei The cave (GPS: 23 34 49.76 N,106 59 58.95 E, 215 m) is located Province (Fig. 1G). The cave was horizontally developed in a Sinian halfway up a PermianeTriassic limestone hill in Bubing Basin, limestone hill named Zhailiangzi. It is about 37 km northeast about 12.6 km west (heading: 258 ) of Tiandong downtown in (heading: 58 ) of Yunxian downtown and about 56.4 km northeast Guangxi Zhuang Autonomous Region (Fig. 1B). The cave deposits of (heading: 70 ) of the Yunxian Man locality at Quyuanhekou in the fi roughly 6.5 m thick were divided into ve lithostratigraphic layers same county. The cave deposits were divided into three layers (Xu, (Wang et al., 2005). A complete right lower second molar of Homo 1978). The upper one is a hard cemented layer with a thickness of erectus (s.l.) and abundant large mammal fossils were unearthed 30 cm; the middle one is mainly yellow sandy clay with occasional from Layer 3, which is composed of fossiliferous brown sandy clay breccia and gravels, its thickness ranges from 50 cm to 250 cm; and with breccia, and its thickness measures 120 cm. A broken upper the lower one is cemented yellow sediments with some small fi left rst or second molar was collected from disturbed deposits in limestone gravels. Only the middle layer yielded Homo erectus teeth the cave (Wang et al., 2005). and associated mammalian fossils (Xu, 1978).
2.3. Jianshi Man locality at Longgudong 2.6. Yunxi Man locality at Bailongdong
Longgudong (“Dragon-bone cave” in Chinese) is a karst cave The site (GPS: 32 59040.000N, 110 31033.600E, 550 m) is located in (GPS: 30 39054.2600N, 110 4035.1500E, 738 m) horizontally developed Shenwuling Village of Anjia Town of Yunxi County in northwestern in the Early Triassic limestone with a length of about 110 m. It is Hubei Province (Fig. 1I). It is only about 10 km east (heading: 89 )of about 34.5 km east (heading: 78 ) of Jianshi downtown in south- Yunxi downtown, and about 18.6 km northwest (heading: 344 )of western Hubei Province (Fig. 1C) and about 245 km south (heading: Yunxian Man locality at Quyuanhekou. Bailongdong (“White 191 ) of Yunxian Man locality at Quyuanhekou in northwestern Dragon cave” in Chinese) is developed in the Pliocene Sha-ping Hubei Province, but only about 45 km southeast (heading: 119 )of Formation composed of breccia interbedded with calsilutites and Longgupo Site of Chongqing Municipality adjacent to Hubei Prov- marls. The cave deposits were lithostratigraphically divided into 8 ince. The cave developed from the east entrance to west exit as an Layers (Wu et al., 2009; Liu et al., 2014). Layer 1 is loosely deposited irregularly shaped tunnel, and a secondary cave developed from the yellowish brown clay with fragments of marls and mammalian proximity of the west exit and extends northeastwards as a narrow bones and with a thickness of about 50 cm; its upper part was chamber. The cave deposits were formed in different parts of the considered as disturbed sediments. Layer 2 measures 54 cm and is cave. Seven sections along the length of the cave were excavated mainly brownish red fossiliferous clay with occasional calcareous from 1999 to 2000 (Zheng, 2004). The cave deposits with thickness concretions and gravels with a diameter of about 5 mm. An orange of 2e5 m in the seven sections were divided into 5e17 layers ac- yellow sandy clay lens with gravels and with a thickness of 7e9cm cording to lithostratigraphic character. The upper part of the de- is interbedded in the layer. Layers 3e8 are mainly clay with varied posits in all sections is characterized by calcareous flow-stone with scattered gravels and calcareous concretions and stratifications and varied thickness. The lower and main part of the deposits in all no fossils were found. Altogether, eight Homo erectus teeth and sections is composed of fossiliferous layers of clay, sub-clay and associated mammalian fossils were unearthed from Layer 2 (Wu sandy clay with occasional breccia on the bottoms. Three Homo et al., 2009; Liu et al., 2014). erectus (s.l.) teeth, an upper left third molar, an upper right third premolar and a lower right first molar were unearthed during the 2.7. Lantian Man locality at Gongwangling excavations from Layer 8 of Section AeA0 (with total 12 layers) near the east entrance, Layer 5 of Section GeG0 (with total 7 layers) in the Gongwangling locality (GPS: 34 11 03.3300N, 109 29023.9000E, west secondary cave and Layer 8 of Section CeC0 (with total 17 715 m) is located in southeastern proximity to Gongwang Village layers) in the center of the cave respectively (Zheng, 2004). and on the northern slope of a hill among Gongwangling ridges at W. Dong / Quaternary International 400 (2016) 47e57 49
Fig. 1. Geographic locations of Homo erectus horizons in China. A: Yuanmou Man locality at Danawu; B: Tiandong Man locality at Mohui Cave; C: Jianshi Man locality at Longgudong; D: Lantian Man locality at Gongwangling; E: Yunxian Man locality at Quyuanhekou; F: Locality 1 of Peking Man Site at Zhoukoudian; G: Yunxian Man locality at Meipu; H: Luonan Man locality at Donghe; I: Yunxi Man locality at Bailongdong; J: Hexian Man locality at Longtandong; K: Lantian Man locality at Chenjiawo; L: Yiyuan Man localities at Qizianshan; M: Nanzhao Man locality at Xinghuashan; N: Nanjing Man locality at Huludong; O: Dongzhi Man locality at Hualongdong. 50 W. Dong / Quaternary International 400 (2016) 47e57
Jiujianfang Town in Lantian County, Shaanxi Province (Fig. 1D). It is divided into 10 layers (Teilhard de Chardin and Young, 1929) and about 16.4 km east (heading: 78 ) of Lantian downtown, about the bottom layer, Layer 10, was later further divided into two as 50.7 km east (heading: 99 ) of Xi'an City and about 15 km southeast Layers 10 and 11 (Black et al., 1933). The soil micromorphology and (heading: 114.5 ) of Chenjiawo locality. Thick earth-like brownish Fourier Transform Infrared Spectrometry (FTIR) analyses of the red sandy clay (loess) was deposited above a very thick gravel bed sediments at Locality 1 (Goldberg et al., 2001) confirmed many on the upper terrace of the left bank of the Ba River, with total conclusions by Teilhard de Chardin and Young (1929). The Homo exposed strata 63 m thick. The earth-like deposits of 13 m thick in erectus materials and associated cultural remains and mammalian the excavated section of 60 m long were divided into 12 layers by faunas were unearthed from all layers except from the uppermost Wu et al. (1966). The skull-cap and other materials of Homo erectus Layers 1 and 2. Lithostratigraphical, paleontological and archaeo- as well as associated mammalian fossils were unearthed from the logical analyses of hominin layers 3e11 indicated that Layers 8e11, middle and upper parts of Layer 6, which is composed of light Layers 6e7 and Layers 3e5 represented three cycles of environ- grayish yellow earth-like sandy clay (loess) with some stratifica- mental change in accordance with the glacial-interglacial shifts (Xu tions and many concretions, and the thickness of the layer ranges et al., 1999). from 2.6 m in the west part to 0.4 m in the east (Wu et al., 1966). 2.11. Yiyuan Man localities at Qizianshan 2.8. Lantian Man locality at Chenjiawo The site (GPS: 36 14031.0800N, 118 7031.2000E, 369 m) is located at Chenjiawo locality (GPS: 34 14022.4400N, 109 20031.3000E, 691 m) the eastern foot of Qizianshan (“Riding Saddle Hill” in Chinese) is situated about 0.5 km south of Chenjiawo Village of Xiehu Town, about 1.6 km northwest (heading: 337 ) of Zhifang Village, Tumen Lantian County, Shaanxi Province (Fig. 1K). It is about 9.4 km Town, Yiyuan County, in the center of Shandong Province (Fig. 1L). northeast (heading: 18 ) of Lantian downtown, and about 15 km The site is composed of three localities in three different caves northwest (heading: 294.5 ) of Gongwanling locality. The exca- developed in Ordovician limestone within 80 m along the west side vated section is about 30 m thick and located about 9.3 km right of of a country road 4.5e6 m above the Ciyu River bed by the eastern the Ba River. The excavated deposits were divided by Zhang et al. foot of Qizianshan. Locality 1 is the most important and the cave (1964) into 18 layers. Layer 18, the top layer, is light yellow sandy deposits of 2.4e3.7 m thick were divided into five layers by Lu et al. earth interbedded with calcareous concretions and with a thick- (1989). The middle layer, Layer 3, is reddish brown silty clay with ness of 5.5 m. Layer 1, the bottom layer, is composed of gravels of considerable density and rigidity and yielded skull fragments and granite, quartzite, and quartz gneiss with diameters of 20e30 cm. two teeth of Homo erectus and rich mammalian fossils. Locality 2 is The other layers are mostly interbedding of sub-clay (loess) and 16 m south of Locality 1 and with the same deposits and paleosol. A mandible of Homo erectus and associated mammalian mammalian fossils. Locality 3 is 40 m south of Locality 2 and with fossils were unearthed from Layer 5, which is 3 m thick and the same deposits, and yielded five teeth of Homo erectus and many composed of light yellow sandy earth with calcareous concretions mammalian fossils from the middle layer as at Locality 1. The de- in the lower part. The layer is close to the bottom of the excavated posits from three localities were considered formed within the section (Zhang et al., 1964). same period based on their adjacency and the same composition of deposits, and they were correlated to the Middle Pleistocene Lishi 2.9. Luonan Man locality at Donghe Loess (Lu et al., 1989).
The site (GPS: 34 7040.3700N, 110 9036.4400E, 950 m) is located in 2.12. Nanzhao Man locality at Xinghuashan Longyanandong (“Dragon teeth's south cave” in Chinese) on Hua- shilang Hill east of Donghe Village and on the left bank of the About 24.5 km east (heading: 94.5 ) of Nanzhao downtown, the mouth of a very small river called Shimen, which is a left tributary site (GPS: 33 28016.6100N, 112 41020.9300E, 228 m) is located on the of the Luo River, Chengguang Town, Luonan County, Shaanxi second terrace on the right or west bank of the Jihe (“Chicken River” Province (Fig. 1H). The cave, or rather a vertical fissure of 1 m wide, in Chinese) and at the eastern foot of Xinghuashan (“Apricot Flower was developed about 50 m above the bed of the Shimen River along Hill” in Chinese) at Ruanzhuang Village, Yunyang Town, Nanzhao a joint in Cambrian dolomitic limestone of the Sanchuan Formation. County, southwestern Henan Province (Fig. 1M). The excavated The cave deposits are mainly grayish yellow calcareous clay (Xue deposits were divided into five layers, and the right mandibular et al., 1999). A right upper first molar of Homo erectus and some fourth premolar of Homo erectus and associated mammalian fossils teeth of mammals were collected from the upper part of the cave were collected from Layer 5, the bottom layer with excavated deposits (Xue,1987). A parallel cave called Longyabeidong (“Dragon thickness of 1.8 m, which is composed of brownish yellow sandy teeth's north cave”) also about 50 m above the bed of the Shimen clay with limestone breccia and gravels (Qiu et al., 1982). River was developed 30 m north of Longyanandong in the same limestone hill along two joints, and the cave is larger. It was exca- 2.13. Nanjing Man locality at Huludong vated from 1995 to 1996. A large amount of stone artifacts and mammalian fossils was unearthed, and the mammalian fossils were About 20.9 km east (heading: 85.3 ) of Nanjing Provincial considered to be from the same fauna as that of the parallel cave Museum in Nanjing downtown, the site (GPS: 32 3028.5400N, (Xue et al., 1999). 119 2027.2300E, 73 m) is located in Huludong (“Calabash Cave” in Chinese) on the hillside of Leigong Hill, Tanshan Town, Jiangning 2.10. Locality 1 (Peking Man Site) at Zhoukoudian District of Nanjing Municipality, Jiansu Province (Fig. 1N). The karst cave was developed in the limestone formed from the Late The locality (GPS: 39 41018.5300N,115 55031.6300E,115 m) is about Cambrian to Ordovician and is composed of a major chamber, a 47 km southwest (heading: 238 ) of the Forbidden City and located minor chamber, and a tunnel between the two chambers. The de- in a collapsed karst cave developed in Ordovician limestone on the posits in the minor chamber were divided into four layers. Skull I of eastern slope of Dragon-Bone Hill in the western margin of Zhou- Nanjing Homo erectus and associated mammalian fossils were koudian downtown, Fangshan District, Beijing Municipality collected from Layer 2, the second lowermost layer of 37 cm thick (Fig. 1F). The cave deposits of more than 30 m thickness were which is composed of light reddish brown clay (Wu et al., 2002). W. Dong / Quaternary International 400 (2016) 47e57 51
The deposits in the tunnel were higher and divided into six layers. 3.2. Methods Skull II of Nanjing Homo erectus and a few mammalian materials were collected from Layer 5, the second uppermost layer of 40 cm All extinct taxa, including regional extinct ones, from all Homo thick which is composed of calcareous concreted gravels (Wu et al., erectus horizons from China were considered as a presumed archaic 2002). fauna and those of extant taxa as a presumed extant fauna. The presence or absence of a taxon in a fauna is a binary variable. Simple matching coefficients are therefore used for comparing the simi- 2.14. Hexian Man locality at Longtandong larity of fauna lists (see Appendix III for detailed description). A fi fi 0 00 0 00 simple matching coef cient of two compared fauna is de ned in The site (GPS: 31 53 22.90 N, 118 12 10.54 E, 23 m) is located the present study as a binary similarity coefficient of the paired on the northern slope of Wangjia Hill and 23.7 km northwest faunas. The faunas from each Homo erectus horizon as well as the (heading: 322.5 ) of Hexian downtown in eastern Anhui Province presumed archaic and extant faunas were compared with each (Fig. 1J). It is 231 km northeast (heading: 30.7 ) of Dongzhi Man site fi fi other in pairs to nd all binary similarity coef cients (Dong et al., and 92.6 km east (heading: 84.9 ) of Hefei downtown. Long- 2010). A binary similarity coefficient of the presumed archaic “ ” tandong ( Dragon Pool Cave in Chinese) was developed in fauna and a compared fauna is defined as the antiquity coefficient e Cambrian dolomite. The excavated cave deposits of 4.5 5.6 m thick of the compared fauna (Dong et al., 2013, 2014). All binary similarity fi were divided into ve layers. A skull-cap and other materials of coefficients were then ranked according to BrainerdeRobinson's Homo erectus and associated mammalian fossils were unearthed (B.eR.) rule (Chen, 1983, 2005; Appendix III), and the faunas side by from Layer 2, the second lowermost layer which is composed of side in the resulted list are those taxonomically most similar. The e yellowish brown clay with a thickness of 0.7 1.4 m (Huang et al., fauna extinction rate is classically defined as the percentage of 1982). extinct taxa number over total taxa number of the fauna. The ecological composition of the faunas was based mainly on diet 2.15. Dongzhi Man locality at Hualongdong preferences of the fauna members. The diet and habitat preferences of taxa were based on information from Nowak and Paradiso (1983) The site (GPS: 30 6027.3300N, 116 56055.1000E, 93 m) is located in and Wang (2003). The ratios of different diet preferences were also Hualongdong (“Elegant Dragon Cave” in Chinese) at Wang Village, compared in percentage, and the ratios are indications of ecological Raodu Town, Dongzhi County, southwestern Anhui Province similarity of compared faunas. (Fig. 1O). It is 231 km southwest (heading: 211.3 ) of Hexian Man site and 191.8 km south (heading: 188 ) of Hefei downtown. The excavated deposits of 3.1e4.8 m thick were divided into four layers 4. Results and discussion and two skull fragments and two teeth of Homo erectus and asso- ciated mammalian fossils and artifacts were collected from Layer 3, 4.1. Biochronological sequence the second lowermost layer of 1.25e1.75 m thick, which is composed of firm brownish red clay (Gong et al., 2014). The sequence of Homo erectus horizons according to faunal antiquity coefficients are listed in Table 1 with calculated extinction rates, sediment facies and available dating results for comparison. 3. Material and methods The locations of Lower, Middle and Upper Zhoukoudian hominin horizons sequenced by antiquity coefficients are in accordance with 3.1. Materials the stratigraphic sequence, but the Middle Zhoukoudian horizon would be lower than the Lower Zhoukoudian horizon if sequenced The faunas from all Homo erectus localities from China by extinction rates. This test supports the reliability of antiquity described above, e.g. those of Danawu (Lin et al., 1978), Mohui coefficients over extinction rates. (Wang et al., 2007), Gongwangling (Hu and Qi, 1978), Longgudong Among the sequenced horizons in Table 1, Yuanmou Man ho- (Zheng, 2004), Quyuanhekou (Ji, 1999), Locality 1 of Zhoukoudian rizon at Danawu has the highest antiquity coefficient and extinc- (Li and Ji, 1981), Meipu (Xu, 1978), Donghe (Xue et al., 1999), tion rate, and remains as the oldest horizon. This supports the Bailongdong (Wu et al., 2009), Longtandong (Huang et al., 2012), paleomagnetic dating results of 1.7 Ma by Zhu et al. (2008), over Chenjiawo (Qi, 1989), Qizianshan (Lu et al., 1989), Xinghuashan other dating results such as that of 700 ka by Hyodo et al. (2002). (Qiu et al., 1982), Huludong (Wu et al., 2002), Hualongdong (Gong The second oldest horizon is that of Mohui cave, by both antiquity et al., 2014), were selected as materials for the present study. The coefficients and extinction rates. Longgudong horizon is just later taxa selected from each locality were from the layer, or layers, than that of Mohui by antiquity coefficients. This result supports the yielding Homo erectus. They indicate the corresponding biochron biochronological sequence of Gigantopithecus faunas in East Asia by of the Homo erectus horizon of the locality, except for Locality 1 Jin et al. (2009), but is in contradiction with the paleomagnetic (Peking Man site) of Zhoukoudian, where there are 9 layers dating of the hominin horizon at Longgudong of 2.14 Ma (Gao and yielding Homo erectus materials and the hominin horizons were Cheng, 2004). The horizons from Mohui cave and Longgudong defined in the present study according to three cycles of sedi- share many Early Pleistocene taxa such as Gigantopithecus blacki, mentation related to environmental changes. Layers 8e11, Layers Hystrix kiangsenensis, Cuon dubius, Pachycrocuta brevirostris licenti, 6e7 and Layers 3e5 were considered as lower (L), middle (M) and Rhinoceros sinensis, Tapirus sanyuanensis, Sus peii, Sus xiaozhu, Cervus upper (U) Homo erectus horizons respectively. Because the screen- (Rusa) yunnanensis, but that of Mohui cave has more archaic forms wash technique was not used in the excavations at certain local- such as Ailuropoda microta, Stegodon preorientalis, Dorcabune liu- ities, small mammals are absent from such localities. The taxa chengensis (see Supplementary Table 1). The extinction rate of selected for the present study are therefore large mammals or Gongwangling follows that of Mohui as the third highly rated. those collected without screen wash. All taxa from all Homo Gongwangling horizon is the fourth oldest one by antiquity erectus horizons from China are listed in online Supplementary coefficients, supporting the dating of 1.15 Ma by An and Ho (1989). Table S1. Gongwangling horizon shares many Early Pleistocene taxa with 52 W. Dong / Quaternary International 400 (2016) 47e57
Table 1 Sequenced Homo erectus horizons according to faunal antiquity coefficients.
Homo erectus horizons Antiquity coefficients Facies Extinction rate % Absolute age dating
Layer 3 81 cave 36.36 Hualongdong, Dongzhi Layer 2 87 cave 43.75 350 ka (Chen et al., 1996) Huludong, Nanjing 500 ka (Wang et al., 2002) Layer 5 88 terrace 40.00 Xinghuashan, Nanzhao Layers 3e5 89 cave 47.06 230e310 ka (Zhao et al., 1985) Zhoukoudian (Upper), Beijing >400 ka (Shen et al., 2001) Middle layers 90 cave 53.85 Qizianshan, Yiyuan Layer 5 92 loess 50.00 650 ka (An and Ho, 1989) Chenjiawo, Lantian Layers 6e7 94 cave 59.09 380 ka (Zhao et al., 1985) Zhoukoudian (Middle), Beijing Layer 2 95 cave 58.06 150e190 ka (Chen et al., 1987) Longtandong, Hexian 300 ka (Huang et al., 1995) Layer 2 97 cave 46.43 Bailongdong, Yunxi Upper layers 98 cave 57.89 Donghe, Luonan Layer 2 99 cave 33.33 Meipu, Yunxian Layers 8e11 102 cave 55.26 462 ka (Zhao et al., 1985) Zhoukoudian (Lower), Beijing 750 ka (Shen et al., 2009) Layer 3 104 terrace 65.22 936 ka (de Lumley et al., 2008) Quyuanhekou, Yunxian Layer 6 106 loess 75.00 1.15 Ma (An and Ho, 1989) Gongwangling, Lantian 1.7 Ma (Zhu et al., 2015) Middle Layers 114 cave 72.50 2.14 Ma (Gao and Cheng, 2004) Longgudong, Jianshi Layer 3 117 cave 80.00 Mohui, Tiandong Layers 25, 27 120 terrace 80.77 700 ka (Hyodo et al., 2002) Danawu,Yuanmou 1.7 Ma (Zhu et al., 2008)
horizons of Yuanmou Man, Mohui cave and Longgudong, but it has Middle Zhoukoudian (Layers 6e7). This is contrary to the later forms such as Canis lupus, Elaphodus cephalophus, Cervus (Sika) traditional conclusion that the Longtandong horizon was grayi, Capricornis sumatraensis. If we accept the dating of 1.7 Ma of equivalent to that of Upper Zhoukoudian (Xu and You, 1984; Gongwangling by Zhu et al. (2015), the horizons of Yuanmou Man, Chen et al., 1987; Huang et al., 1995, 2012). The extinction rate Mohui cave and Longgudong should be earlier than 1.7 Ma based on of the Longtandong horizon is higher than that of Lower Zhou- the present biochronological analyses. koudian but lower than that of Middle Zhoukoudian (Table 1), a The sequence of Quyuanhekou is later than Gongwangling by confusing result. The numerical age dating results show that the antiquity coefficients and extinction rates tallies with the dating of horizon of Longtandong is younger than that of Middle 936 ka by de Lumley et al. (2008) for Quyuanhekou and that of 1.15 Ma Zhoukoudian. by An and Ho (1989) or 1.7 Ma by Zhu et al. (2015) for Gongwangling. The hominin horizon of Middle Zhoukoudian is lower than that The lower hominin horizon (Layers 8e11) of Zhoukoudian is of Chenjiawo by both antiquity coefficients and extinction rates. younger than that of Quyuanhekou by antiquity coefficients and However, this does not agree with the numerical age dating results extinction rates, in accordance with the dating of 936 ka by de of 380 ka for that of Middle Zhoukoudian (Zhao et al., 1985) and Lumley et al. (2008) for Quyuanhekou and that of 750 ka by Shen 650 ka for that of Chenjiawo (An and Ho, 1989). This might result et al. (2009) or 462 ka by Zhao et al. (1985) for the Lower from the much smaller number of identified taxa from Chenjiawo Zhoukoudian. compared to Middle Zhoukoudian. The hominin horizons of Meipu of Yunxian, Donghe of Luonan The horizon of Yiyuan Man at Qizianshan is higher than that of and Bailongdong of Yunxi have no available numerical dating data Chenjiawo and lower than that of Upper Zhoukoudian (Layers for comparison. Their antiquity coefficient sequence from oldest to 3e5) according to antiquity coefficients. Its extinction rate is youngest is Meipu, Donghe, Bailongdong. They are all younger than higher than those of both Chenjiawo and Upper Zhoukoudian. the Lower Zhoukoudian but older than Longtandong by antiquity The difference is likely due to the smaller taxa numbers of both coefficients. Although their extinction rates, especially those of Qizianshan and Chenjiawo horizons compared to Upper Zhou- Meipu and Bailongdong, are all lower than that of Longtandong, the koudian. No numerical age dating results were available for the presence of archaic forms such as Pachycrocuta brevirostris licenti Qizianshan horizon. and Gomphotherium sp. in Meipu horizon, Ailuropoda wuling- The Upper Zhoukoudian horizon is older than that of Xinghua- shanensis, Pachycrocuta brevirostris licenti, Sus peii and Cervavitus shan by both antiquity coefficients and extinction rates. Xinghua- fenqii in Bailongdong horizon, and Ursus cf. etruscus and Gom- shan is older than Huludong by antiquity coefficients only. The photheriidae indet. in Donghe horizon supports the antiquity co- numerical age dating data show that Huludong hominin horizon is efficients more than extinction rates. older than the Upper Zhoukoudian. The Hualongdong horizon is The Longtandong horizon of Hexian Man site is sequenced younger than those of Huludong and Xinghuashan by both antiq- just later than that of Bailongdong and just earlier than that of uity coefficients and extinction rates. W. Dong / Quaternary International 400 (2016) 47e57 53
4.2. Fauna similarity sequence coefficient between Yuanmou Man horizon and that of Hua- longdong is not large (199). The sequenced faunal binary similarity coefficients of hominin horizons according to BrainerdeRobinson's rule are listed in 4.3. Paleoecological composition and paleoenvironment Table 2, i.e. the horizons with maximum binary similarity co- efficients are ranked side by side. The largest B.eR. binary similarity The taxa based percentages of browsers, grazers, omnivores, coefficient is found with the pair of Huludong at Tangshan and scavengers and carnivores of every hominin horizon are illustrated Qizianshan horizons (235), followed by that of Qizianshan and in Fig. 2. The horizons most similar are those of Quyuanhekou and Chenjiawo (232), and then that of Chenjiawo and Hualongdong Meipu, and these horizons are also closely related by B.eR. binary (231). The smallest binary similarity coefficient is with the pair of similarity coefficient. Those of Xinghuashan, Bailongdong and Mohui and Longtandong horizons (169), followed by that of Longgudong are also similar to each other, as well as those of Longtandong and Gongwangling (178), then that of Bailongdong Longtandong and Huludong at Tangshan. However, they are not and Longgudong (193). closely related by B.eR. binary similarity coefficients. The Upper,
Table 2 Binary faunal similarity coefficients of Homo erectus horizons e sequenced according to BrainerdeRobinson's rule.
Arc YmM Hld Cjw Qzas Ts ZkdM ZkdU ZkdL Xhs Dh Mp Qyhk Bld Lgd Mh Ltd Gwl exta
Arc 258 YmM 120 258 Hld 81 199 258 Cjw 92 196 231 258 Qzas 90 188 227 232 258 Ts 87 181 224 227 235 258 ZkdM 94 168 209 216 234 229 258 ZkdU 89 150 193 202 218 211 226 258 ZkdL 102 142 181 190 204 205 214 220 258 Xhs 88 186 221 224 226 225 214 202 194 258 Dh 98 187 218 217 215 220 205 189 177 217 258 Mp 99 189 216 211 217 210 199 179 171 211 218 258 Qyhk 104 184 207 204 206 205 190 178 170 202 209 225 258 Bld 97 179 198 191 189 194 173 161 147 189 192 204 219 258 Lgd 114 168 181 176 174 171 154 142 128 164 177 183 188 193 258 Mh 117 189 200 195 197 192 179 165 153 187 184 198 189 192 209 258 Ltd 95 154 201 208 206 215 206 188 180 206 197 193 186 171 154 169 258 Gwl 106 170 197 202 198 199 186 178 170 196 203 195 208 191 166 165 178 258 exta 0 132 175 168 168 171 164 162 158 170 157 159 154 161 144 141 160 152 258
Abbreviations: Arc. Presumed archaic fauna composed of all extinct taxa from all Homo erectus horizons from China; Bld. Bailongdong at Yunxi; Cjw. Chenjiawo at Lantian; Dh. Donghe at Luonan; Exta. Presumed extant fauna composed of all extant taxa from all Homo erectus horizons from China; Gwl. Gongwangling at Lantian; Hld. Hualongdong at Dongzhi; Lgd. Longgudong at Jianshi; Ltd. Longtandong at Hexian; Mh. Mohui Cave at Tiandong; Mp. Meipu at Yunxian; Qyhk. Quyuanhekou (i.e. Quyuan River mouth) at Yunxian; Qzas. Qizi-anshan at Yiyuan; Ts. Huludong at Tanshan in Nanjing; Xhs. Xinghuashan at Nanzhao; YmM. Yuanmou Man Site; ZkdU. Layers 3e5 of Locality 1 of Zhoukoudian Site in Beijing; ZkdM. Layers 6e7 of Locality 1 of Zhoukoudian Site in Beijing; ZkdL. Layers 8e11 of Locality 1 of Zhoukoudian Site in Beijing. See materials and methods for definition and data source of every Homo erectus horizon.
A few closely related horizons by B.eR. binary similarity co- Middle and Lower Zhoukoudian horizons are roughly similar to efficients, such as those of Upper, Middle and Lower Zhoukoudian, each other, and are closely related by B.eR. binary similarity are geographically in the same location. Similarly, those of coefficients. Quyuanhekou, Meipu and Bailongdong closely related by B.eR. The percentage of browsers in most horizons is larger than that binary similarity coefficients are in the same area and geographi- of grazers except for Chenjiawo, Upper, Middle and Lower Zhou- cally adjacent. koudian. Those of browsers and grazers are equal in Qizianshan. In Certain closely related pairs by B.eR. binary similarity co- most horizons, the hominin environment was relatively humid and efficients such as Chenjiawo and Qizianshan, Meipu and Donghe, temperate. Longgudong and Mohui, are also closely related by binary antiquity The taxa based percentages of browsers, grazers, omnivores, coefficients. Some closely related pairs by B.eR. binary similarity scavengers and carnivores of every hominin horizon are compared coefficients such as Huludong at Tangshan (350 ka or 500 ka) and in Fig. 3, where the sequence of the horizons follows the antiquity Middle Zhoukoudian (380 ka), Longtandong (150e190 ka or coefficients from the oldest on the left to the youngest on the right. 300 ka) and Upper Zhoukoudian (230e310 ka), are in accordance Browsers experienced four undulations, two relatively strong and with their similar numerical age dating results (see Table 1). two relatively mild. The grazers experienced six undulations, and The Yuanmou Man horizon is the closest to presumed archaic one of them is relatively strong. The scavengers experienced six fauna when all horizons are compared with the archaic one as undulations and two relatively sharp downturns. They were rela- expected, but also unexpectedly the closest to the youngest horizon tively abundant at Qizianshan and Huludong. The latter was at Hualongdong when all horizons are compared with that of considered as having been a hyena den based on the presence of Yuanmou Man. This is because many taxa from Hualongdong are hyena fossils and absence of lithic industry associated with homi- identified at generic level only and Yuanmou Man and Hua- nin remains (Vialet et al., 2010). The carnivores experienced four longdong horizons have more common genera. The B.eR. similarity undulations, with one relatively sharp rise which is associated with 54 W. Dong / Quaternary International 400 (2016) 47e57
100%
90%
80%
70%
60% Grazer Browser 50% Scavenger Carnivore
40% Omnivore
30%
20%
10%
0% YmM Dh Gwl Mh Qyhk Mp Bld Xhs Lgd Ltd Ts ZkdA ZkdB ZkdC Qzas Cjw Hld
Fig. 2. Comparison of ecological composition of Homo erectus horizons in China. The abbreviations for horizons are the same as for Table 2.
60
50
40
Omnivore 30 Carnivore
Scavenger
20 Browser
Grazer
10
0 YmM Mh Lgd Gwl Qyhk ZkdL Mp Dh Bld Ltd ZkdM Cjw Qzas ZkdU Xhs Ts Hld
Fig. 3. Evolution of ecological composition of Homo erectus horizons in China. The abbreviations for horizons are the same as for Table 2. W. Dong / Quaternary International 400 (2016) 47e57 55 a relatively sharp decrease of browsers. The omnivores also expe- span of 350 ky. The fourth stage is represented by Chenjiawo, rienced six undulations, with one relatively sharp rise associated Middle Zhoukoudian, Qizianshan and Longtandong horizons. with a relatively sharp fall of carnivores. Chenjiawo is the oldest, and the estimated numerical age ranges from 600 ka to 300 ka with a time span of 300 ky. The fifth and last stage is represented by Huludong, Xinghuashan, Upper Zhou- 4.4. Conclusions koudian and Hualongdong horizons, with Huludong the oldest and Hualongdong the youngest. The estimated numerical age ranges Based on the antiquity coefficients and with consideration of from 400 ka to 150 ka, with a time span of 250 ky. The last stage is B.eR. binary similarity coefficients, extinction rates and numerical associated with the decline and final disappearance of Homo erectus age dating results, five biochronological stages are proposed as the as well as arrival of Homo sapiens in East Asia. framework for Homo erectus horizons in China as illustrated in Besides the significant taxa of each stage listed in Table 3, Axis Table 3. Some age overlaps between two adjacent stages are kept shansius and Ailuropoda microta of the first stage disappeared since for estimated numerical ages to compare different dating the second stage. Rhinopithecus lantianensis, Ailuropoda wuling- results.
Table 3 Biochronological framework of Homo erectus horizons in China.
Biochrons Homo erectus horizons Significant taxa Estimated age
Stage 5 Hualongdong, Dongzhi Macaca robustus 150e400 ka Zhoukoudian (Upper), Beijing Trogontherium cuvieri Xinghuashan, Nanzhao Nyctereutes sinensis Huludong, Nanjing Ursus arctos Ailuropoda baconi Pachycrocuta sinensis Megaloceros pachyosteus Cervus (Sika) grayi Stage 4 Longtandong, Hexian Macaca robustus 300e600 ka Qizianshan, Yiyuan Trogontherium cuvieri Zhoukoudian (Middle), Beijing Canis lupus variabilis Chenjiawo, Lantian Nyctereutes sinensis Ursus arctos Ailuropoda baconi Pachycrocuta sinensis Lutra melina Stegodon orientalis Megaloceros pachyosteus Cervus (Sika) grayi Stage 3 Donghe, Luonan Macaca robustus 500e850 ka Zhoukoudian (Lower), Beijing Trogontherium cuvieri Bailongdong, Yunxi Nyctereutes sinensis Meipu, Yunxian Canis lupus variabilis Ailuropoda wulingshanensis Pachycrocuta sinensis Pachycrocuta brevirostris licenti Megaloceros pachyosteus Cervus (Sika) grayi Stage 2 Quyuanhekou, Yunxian Rhinopithecus lantianensis 0.8e1.3 Ma Gongwangling, Lantian Hystrix kiangsenensis Longgudong, Jianshi Ailuropoda wulingshanensis Pachycrocuta brevirostris licenti Stegodon orientalis Cervus (Rusa) yunnanensis Stage 1 Mohui, Tiandong Hystrix kiangsenensis 1.2e1.8 Ma Danawu, Yuanmou Ailuropoda microta Pachycrocuta brevirostris licenti Cervus (Rusa) yunnanensis Axis shansius
N.B.: The bold taxa are the first appeared ones in the stage, i.e. absent from the previous stage(s) and the underlined taxa will disappear in the next stage.
The first stage is represented by Yuanmou Man and Tiandong shanensis and Stegodon orientalis appeared in horizons at the sec- Man horizons. The latter is relatively younger; the estimated nu- ond stage. Rhinopithecus lantianensis of the second stage and Cervus merical age ranges from 1.8 Ma to 1.2 Ma with a time span of 600 ky. (Rusa) yunnanensis from the previous stages disappeared after the The first stage is associated with the arrival of Homo erectus in East third stage. Many significant taxa such as Macaca robustus, Tro- Asia. The second stage is represented by the Longgudong, Gong- gontherium cuvieri, Nyctereutes sinensis, Pachycrocuta sinensis, wangling and Quyuanhekou horizons, and the latter is relatively Megaloceros pachyosteus and Cervus (Sika) grayi appeared in the younger; the estimated numerical age ranges from 1.3 Ma to 800 ka third stage. The Ailuropoda wulingshanensis of the second and third with a time span of 500 ky. 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