Presumed domestication? Evidence for wild rice cultivation and domestication in the fifth millennium BC of the Lower Yangtze region.

Fuller, Dorian Q. ; Harvey, Emma ; Qin, Ling 等

Presumed domestication

In some legal traditions, people are presumed innocent until proven guilty. In the study of agricultural origins it is perhaps prudent to presume plants are wild until evidence can be found to indicate domestication. This has not, however, been the convention in the archaeology of East Asia, where domestication is taken for granted, unproven and unquestioned. The recent research report by Jiang and Liu (Antiquity 80: 355-61), unfortunately continues this tradition. It seems a curious fact that little discussion has ever been devoted to wild rice foraging in Asia, which logically must have preceded agriculture, or how this might appear archaeologically. We believe that the rice chaff that was used to temper Shangshan pottery will most likely turn out to be a product of foraging, and we would like to take this opportunity to consider a larger body of available evidence from the Lower Yangtze area (Figure 1) that suggests the process of rice domestication came to an end (full domestication) closer to 4000 BC after an extended period, of a millennium or more, of pre-domestication cultivation and presumably a much longer period of wild rice use by foragers.

[FIGURE 1 OMITTED]

Writings on rice in the Lower Yangtze contain a paradox. It has been suggested that rice assemblages contain a mixture of indica and japonica cultivars, or just indica or just japonica, or intermediate 'ancient' forms of rice (e.g. You 1976; Zhou 1981; 2003; Li 1985; Zhao & Wu 1987; Oka 1988; Bellwood 1997: 206; Zhang 2002), but such claims are riddled with contradictions between different scholars looking at the same material, and all such claims are predicated on a now disproven theory of rice origins. In the 1980s prominent botanists, especially Oka (1988) but also T. Chang (1989), favoured a single origin for rice followed by differentiation into indica and japonica subspecies under cultivation. This scenario, however, is no longer tenable as data accumulated through newer genetic techniques indicates that indica and japonica are phylogenetically distinct, and represent separate domestication events from distinct progenitor species, Oryza nivara for indica and Oryza rufipogon (sensu stricto) for japonica (e.g. Sato et al. 1990; Sano & Morishima 1992; Chen et al. 1993; Wan & Ikehashi 1997; Cheng et al. 2003; Vaughan et al. 2003; Li et al. 2004; for nomenclature, see Vaughan 1994). Indeed, the most comprehensive study to date indicates two separate domestication events for japonica rice, in the South China region, and two domestications of indica rices in South Asia or western Southeast Asia (Londo et al. 2006). This evidence has only begun to be considered in the archaeological literature recently (e.g. Crawford & Shen 1998; Jones & Brown 2001; Fuller 2002: 297; Sato 2002). It should be noted that this genetic evidence overturns the assumption of a single Asian rice origin, which remains prominent in many textbooks (e.g. Bellwood 1997; Higham 2005). While this alone should call into question older archaeobotanical descriptions, since the dominance of 'indica' in early China should be impossible, in fact the attribution of these early finds to indica or japonica domesticated rice is not supported by available morphometric evidence.

Traditional identification ratios do not work unless the presence of wild species can be excluded. Attributions of ancient rice material has been made on the basis of grain or spikelet length-to-width ratios, with ratios of greater than 2.5 attributed to indica and ratios of less than 2.3 attributed to japonica. Recent morphometric data collected on modern rice species indicates that this does not work if wild species, including both the wild progenitors and other Oryza spp. are included (Figure 2). There is much variation in the size and proportions of domesticated and wild rice species today, making it difficult to assign one or a few grains to any given population (Thompson 1996: 176; Harvey 2006). Nevertheless, we would encourage the use of scatter-plots of actual measurements (Figure 3) as a more useful way to look at grain characters and suggest that some species distinctions can be made on an assemblage level. Also changes over time can be more easily tracked. A further problem is the issue of grain maturity, as immature grains will have exaggerated length-to-width ratios. We return to this problem below after considering what is expected in identifying the transition from wild to domesticated rice.

[FIGURES 2-3 OMITTED]

An evolutionary model of the rice domestication syndrome

There is an essential distinction between cultivation (human activity) and domestication (change in the plant). We should expect there to have been a phase, however brief, of pre-domestication cultivation (Helbaek 1960; Wilke et al. 1972; Hillman & Davies 1990; Harris 1996; Gepts 2004), and we should seek this archaeologically. In the Near East evidence suggests that pre-domestication cultivation was not brief, but lasted for one to two millennia (Tanno & Willcox 2006; Weiss et al. 2006). The domestication syndrome consists of traits that evolved under cultivation, but they are unlikely to have evolved entirely simultaneously. In rice we can consider three traits that should be recoverable through archaeology:

1. Relaxation of selection for natural dispersal aids, i.e. the reduction in awns and hairs which help the shed spikelet grip the soil. As humans start to plant seeds this should relax natural selection in favour of maintaining these characters. As others have remarked, many domesticated rices are awnless, and those with awns have many fewer awn hairs (Sato 2002). But there are also awnless wild rices such as Oryza meyeriana or O. granulata. (Vaughan 1994).

2. Increase in grain size, or grain weight. This character in cultivated cereals is related to increased efficiency and competitiveness in germination and early growth in open, heavily disturbed soils and with deeper burial of seeds, which is expected under tillage (see e.g. Harlan et al. 1973).

3. Finally, and perhaps most important, is selection against wild-type dispersal, i.e. the development of a non-shattering spikelet base, allowing plants to be efficiently harvested, as by uprooting or by sickle (Hillman & Davies 1990). It is this change which is most often taken as the key trait of domesticated cereals (see e.g. Zohary & Hopf 2000). This trait however, evolves in response to human harvesting practices and may evolve later than some of the other traits. The non-shedding trait occurs in rice by a toughening of the attachment of the spikelet base to the rachilla, and as shown by Thompson (1996; 1997), this is accompanied by a subtle change in the cross section of the rachilla attachment scar (also Sato 2002).

Of particular importance is that this last trait allows plants to retain all their grains at maturity, as opposed to wild plants that lose grain progressively as they come into maturity. This issue is particularly important because we expect foragers to have targeted rice for harvest when substantial number of grains were immature (as known amongst grass-using foragers, cf. Harris 1984). Immature spikelet bases may mimic those of domesticated types.

The challenge of rice foraging and grain immaturity

The challenge for hunter-gatherers targeting rice is that as grains mature they are shed into the water and mud where wild rice grows. Because rice panicles mature over an extended period, of 15-16 days, the number of grains available at one time on a plant changes. In order to maximise grain recovery it is necessary to target plants early in their grain production cycle, which means that a large proportion of immature spikelets will also be recovered (Figure 4). This has two important implications, first that it will reduce selection pressure for the evolution of domesticated (non-shattering) plants, and second it can be expected to produce assemblages with proportions of immature spikelets, including those without significant grain formation and those in which grains are long and skinny as a product of how rice grains fill out during maturation (Figure 5). This means that pre-domesticated assemblages will contain grains with exaggerated length:width ratios due to immaturity. The combination of increasingly mature harvesting, allowed by loss of wild shedding, and selection for larger grains under cultivation will both contribute to morphometric change in archaeobotanical assemblages.

[FIGURES 4-5 OMITTED]

The assemblages from Hemudu (5000-4000 BC, i.e. layers 4 & 3) and Kuahuqiao (6000-5400 BC) contained significant proportions of immature harvested spikelets, indicated both by spikelet/chaff remains and by grain morphometrics. At Hemudu finds included 'abundant empty husks of immature spikelets' (Zhou 2003:430 [Chinese original]). Similarly at Kuahuqiao, quantified rice remains included about 18 per cent grains, 47 per cent empty (dehusked) spikelets and 35 per cent intact spikelets (immature without full grain formation) (Zheng et al. 2004a). A reassessment of published grain measurements from the region, including Kuahuqiao and the Majiabang culture site of Longqiuzhuang (4800-4000 BC) indicate that grains fall into the immature and/or wild progenitor range, but that a marked shift in grain size occurred by the later levels at Longqiuzhuang (Songze culture, after 4000 BC), at which time grains appear longer, wider and presumably represent predominantly mature-harvested grains (Figure 6). In addition the site of Chuodun (late Majiabang period site, just before 4000 BC), shows a marked distinction from earlier sites in having wider grains. This suggests that the process of morphological domestication took place during the Majiabang/Hemudu period. Quantitative data from the Middle Yangtze region is more limited, but data from Bashidang (7000-6000 BC), a site often associated with early rice agriculture, has markedly thin rice, comparable to those from Kuahuqiao, which we interpret as immature and morphologically wild. Also of note are measurements from Jiahu (Henan Province Institute 1999), which are remarkably small, and more suggestive of a different wild rice, such as Oryza officinalis, which did not contribute to later domesticated populations but is a prolific wild seed producer.

[FIGURE 6 OMITTED]

An additional source of evidence that suggests a shift from immature to mature harvesting during the Majiabang period comes from bulliform phytoliths. While some of the variation in shape amongst bulliforms appears to be under genetic control and therefore reflects phylogenetics (Zheng et al. 2003; 2004a; 2004b; cf. Pearsall et al. 1995), there remains much to be understood about such phylogenetic variation in the past. However, some aspects of variation relate to plant maturity, especially horizontal length (HL) and vertical length (VL) show a strong correlation with plant maturity (Zheng et al. 2003: 1217). In other words more mature plants produce larger bulliforms. Recent metrical data on bulliforms from sites in the Lower Yangtze indicate a significant shift towards larger bulliforms through time (Figure 7). These data therefore agree with the evidence of grain morphometrics that earlier rice, e.g. of the Majiabang period, was being harvested substantially less mature than later, presumably domesticated rice of the Songze and Liangzhu phases.

[FIGURE 7 OMITTED]

Other evidence argues for pre-domestication cultivation during the Hemudu/Majiabang period. As is well-known, Hemudu yielded a great many hafted or haftable bone scapula artefacts, which are regarded as spades or hoes, as well as some wooden spade blades (Chang 1986: 212; Zhejiang Provincial Institute 2003). This suggests manipulation of the soil through tillage. Coupled with the evidence that rice grains were largely immature we regard this as a strong case for wild plant food production. At the earlier Kuahuqiao, only four possible bone 'spades' were recovered, and these had poor hafting features suggesting they would not be suited to heavy tillage (Zhejiang Provincial Institute 2004: 176-7). This argues for the development of cultivation with systematic tillage sometime between Kuahuaqiao and Hemudu, or during Hemudu.

In addition, a study of a small sample of rice spikelets from Hemudu, indicated a mixture of domesticated, wild and intermediate characters (Sato 2002). Studies of awn remains had hair densities between those of modern wild and domesticated rices, thus suggesting that natural selection to maintain spikelet dispersal aids had been relaxed by cultivation. In addition spikelet bases included both smooth, wild types and torn types (although these may include immature wild as well as domesticated individuals).

From the later Majiabang period the first preserved field system, interpreted as small rice paddies, has been recovered from the sites of Caoxieshan (Jiangshu province) (Zou et al. 2000) and Choudun (Zhejiang Province) (Gu 2003). This development would have an important effect in terms of separating the cultivated wild rice from cross-pollination with free-growing populations, thus accelerating evolution. In the context of these small plots, experimentation with new harvesting methods, like uprooting, may also have played a role in domestication.

On the grounds of the above data, we suggest that wild plant food production began by or during the Hemudu phase (i.e. by c. 5000 BC) and that domesticated rice had evolved by c. 4000 BC (Figure 8). This implies that earlier finds of rice, such as Kuahuqiao and probably Shangshan represent wild rice gathered by foragers (with some cultivation perhaps beginning before the end of Kuahuqiao). It should be noted that this scenario implies an evolutionary process to evolve non-shattering rice plants

with a duration of the order 1000-1500 years. Data from the Near East indicate an equivalent period in the evolution from early cultivation to domesticated wheat and barley (Colledge 1998; Willcox 2004; Tanno & Willcox 2006; Weiss et al. 2006).

[FIGURE 8 OMITTED]

Wider implications for early East Asian agriculture

Our hypothesis has wider repercussions for understanding the origins and spread of agriculture in East Asia. Clearly, pottery developed in East Asia well in advance of agriculture, as is true in many other parts of the world (Rice 1999; Kuzmin 2006). It would also imply that sedentary hunter-gatherers preceded agriculture, rather than sedentism being driven by agriculture. In recent years, the orthodoxy has been that rice agriculture began early, perhaps at the start of the Holocene or late Pleistocene, in the Middle Yangtze, perhaps amongst seasonally inhabited cave-sites (e.g. Cohen 1998; Yasuda 2002; Higham 2005). The archaeobotanical basis for these inferences, however, has remained very limited (see Lu 1999; Crawford 2006). One site of the Pengtoushan culture, Bashidang, produced quantities of rice, but as indicated above this fits clearly with wild rice. Other criteria which have been used to suggest domestication include husk patterns (Zhang & Wang 1998) and husk phytoliths (Zhao 1998), although the relationship between these and evolution under cultivation (the domestication syndrome) remain unclear. Recent tests of these criteria on modern rice species fail to support their utility for inferring domestication (Harvey 2006). Nevertheless, some average grain 'types', reported from later Chengtoushan (Pei 1998), suggest the presence of both wild as well as mature/domestic types by the mid-fifth millennium BC and the Daxi Culture period.

The delayed domestication of rice in the Yangzte implies that millet domestication in northern China developed first. The earliest well-documented millets are from c. 6000 BC at Xinglonggou, in Eastern Inner Mongolia (Zhao 2005), while millet cultivation is regarded generally as established in the Yellow River basin by 5500 BC (the Beixin, Cishan, Peiligang and Dadiwan cultures). Rice from the south was added to this agricultural system only in the third millennium BC, with a few rice finds from Late Yangshao contexts (3000-2500 BC) and many more from the Longshan period (2500-2000 BC) (Crawford et al. 2005). This would suggest that within 1000-1500 years of fully domesticated rice in the South it had spread to Central China. The spread of rice southwards, to Taiwan and Vietnam, has a similar time flame with the earliest finds dating to c. 2500 BC (Higham 2005; Tsang 2005). The southward dispersal is often linked to migration and demographic expansion. Indeed the revised time frame makes this dispersal comparable in rate to the spread of crops from Greece through much of Europe, to the western Linearbandkeramik of Belgium and the Cardial Ware related pottery of Mediterranean France and Iberia. An earlier development of rice agriculture leaves unexplained a long latent period with little evidence for demographic growth and expansion.

Concluding remarks: nut foragers and rice domestication

At Hemudu the presence of rice has been emphasised whereas the quantities of nuts, especially acorns, has been mainly ignored. Both Hemudu and Kuahuqiao produced substantial quantities of waterlogged plant remains. Rice was a small component of a broader subsistence base with a focus on nuts. Table 1 summarises the species present at these sites (Zhejiang Provincial Institute 2003; 2004). What is striking about this list of taxa is the wide range of nuts, in particular acorns, which were found in large quantities in storage pits. The authors have recently begun a collaboration with the Zhejiang Provincial Institute of Archaeology and Chinese Academy of Social Sciences on the archaeobotany of Tian Luo Shan, a site of the Hemudu culture. Here, large quantities of acorns and waterchestnuts are preserved in water-logged samples and storage pits, in addition to rice spikelet bases and other wild (weed?) seeds. Nut remains substantially outnumber those of rice, which was probably also true at Hemudu although the remains were never quantified and the acorns were ignored in publications on the site. The ongoing investigation of new material will provide an opportunity to check the model for a late evolution of domesticated rice. Jiahu also produced large quantities of acorns (Henan Provincial Institute of Archaeology 1999; Zhao Zhijun, personal communication), suggesting that these were a significant, if not the staple, resource.

Our hypothesis of a late rice domestication not only provides a coherent integration of archaeobotanical evidence with expectations from botany and genetics, but suggests an explanatory framework for the origins of rice agriculture. Regional pollen evidence indicates a marked decline in oaks at the end of the sixth millennium BC (Tao et al. 2006), which could suggest an important 'push' factor that encouraged the development of rice cultivation. Models of agricultural origins drawn from evolutionary ecology (the 'diet breadth model'), suggest that many early crops were secondary resources, but were reliable sources of subsistence risk-buffeting which became increasingly important when primary resources (such as nuts) declined, whether due to over exploitation or climatic factors (e.g. Winterhalder & Goland 1993; Keeley 1995; Piperno & Pearsall 1998). A social pull factor can also be suggested. Specialised craft production, and potential status artefacts (such as jades and fine ground axes), began to be produced during the Kuahuqiao through Majiabang horizon, and emerged as key status symbols in the Songze (4000-3300 BC) and subsequent Liangzhu periods (3300-2200 BC). The cultivation of wild rice in small wetland plots could also have provided a source of wealth production, as its surplus could have been more reliably produced and controlled than foraged resources. Our evolutionary model for rice also makes sense in terms of what is known from archaeology of settlement patterns and inferred social organisation (Qin 2000; 2003; Fuller et al. in press), as it is only after the Majiabang period (during the Songze and subsequent Liangzhu periods) that evidence suggests a demographic filling in of the landscape, social differentiation and specialised craft production. A post-4000 BC demographic explosion therefore becomes explicable.

The available data clearly indicate the need for archaeologists to cease and desist in presuming that all rice finds are domesticated and equate to agriculture. Rather we need to look for the long-term processes, by which nut collectors became rice farmers, and one of many wild marsh grasses in the genus Oryza became the world's most productive crop.

Acknowledgements

The authors would like to thank our colleagues at Zhejiang Provincial Institute of Cultural Relics and Archaeology, especially Sun Guoping and Zheng Yunfei, who first introduced Dorian to the material at the Hemudu museum and other sites under research by the provincial institute. This occurred while he was on an AHRB Research Leave grant. We have also benefited from discussions with Zhao Zhijun. Our current archaeobotanical collaboration at Tianluoshan has been made possible by Sun, Zheng, Zhao and the provincial director, Cao Jin Yan. The authors' involvement in this project is supported by grants from the Sino-British Trust of the British Academy and China National Education Ministry Project Grant for the Center for the Study of Chinese Archaeology at Peking University. Research on the morphometrics of modern rice species was conducted by E. Harvey as a research student supported by an Art and Humanities Research Board studentship. We thank two anonymous peer-reviewers for their suggestions.

Received: 15 June 2006. Accepted: 7 September 2006. Revised: 15 September 2006

References

BELLWOOD, P. 1997. Prehistory of the Indo-Malaysian archipelago, second edition. Honolulu: University of Hawaii Press.

CHANG, K.-C. 1986. The Archaeology of Ancient China. New Haven: Yale University Press.

CHANG, T.T. 1989. Domestication and the spread of cultivated rices, in D.R. Harris & G.C. Hillman (ed.) Foraging and Farming. The Evolution of Plant Exploitation: 408-17. London: Unwin Hyman.

CHEN, W.-B., I. NAKAMURA, Y.-I. SATO & H. NAKAI. 1993. Indica and Japonica differentiation in Chinese landraces. Euphytica 74: 195-201.

CHENG, C., R. MOTOHASHI, S. TCHUCHIMOTO, Y. FUKUTA, H. OHTSUBO & E. OHTSUBO. 2003. Polyphyletic Origin of Cultivated Rice: Based on the Interspersion Patterns of SINEs. Molecular Biology and Evolution 20: 67-75.

COHEN, D.J. 1998. The Origins of Domesticated Cereals and the Pleistocene-Holocene Transition in East Asia. The Review of Archaeology 19: 22-9.

COLLEDGE, S. 1998. Identifying pre-domestication cultivation using multivariate analysis, in A.B. Damania, J. Valkoun, G. Willcox & C.O. Qualset (ed.) The origins of agriculture and crop domestication: 121-31. Aleppo, Syria: ICARDA.

CRAWFORD, G. 2006. East Asian Plant Domestication, in M.T. Stark (ed.) Archaeology of Asia: 77-95. Oxford: Blackwell.

CRAWFORD, G. & C. SHEN. 1998. The origins of rice agriculture: recent progress in East Asia. Antiquity 72: 858-66.

CRAWFORD, G., A. UNDERHILL, Z. ZHAO, G. LEE, G. FEINMAN, L. NICHOLAS, F. LUAN, H. YU, H. FANG & F. CAI. 2005. Late Neolithic Plant Remains from Northern China: Preliminary Results from Liangchengzhen, Shandong. Current Anthropology 46(2): 309-17.

FULLER, D.Q. 2002. Fifty Years of Archaeobotanical Studies in India: Laying a Solid Foundation, in S. Settar & R. Korisettar (ed.) Indian Archaeology in Retrospect, Volume III. Archaeology and Interactive Disciplines: 247-363. New Delhi: Manohar.

FULLER, D.Q., L. QIN & E.L. HARVEY. In press. Evidence for a late onset of agriculture in the Lower Yangtze region and challenges for an archaeobotany of rice, in A. Sanchez-Mazas, R. Blench, M. Ross, M. Lin & I. Pejros (ed.) Human migrations in continental East Asia and Taiwan: genetic, linguistic and archaeological evidence. London: Taylor & Francis.

GEPTS, P. 2004. Crop Domestication as a long-term selection experiment. Plant Breeding Reviews 24: 1-44.

GU, J. 2003. The Rice Paddy Land from Chuodun Site of Majiabang Period, in Chuodun Shah Site: collected papers, Dongnan Wenhua [southeast Culture] 2003 (special issue 1): 42-5 [in Chinese].

HARLAN, J.R., J.M.J. DE WET & E.G. PRICE. 1973. Comparative evolution of cereals. Evolution 27: 311-25.

HARRIS, D.R. 1984. Ethnohistorical evidence for the exploitation of wild grasses and forbs: its scope and archaeological implications, in W.A. van Zeist & W.C. Casparie (ed.) Plants and Ancient Man: Studies in Palaeoethnobotany: 63-9. Rotterdam: A.A. Balkema.

--1990. Settling Down and Breaking Ground: Rethinking the Neolithic Revolution (Twelfth Kroon-Voordact Lecture). Amsterdam: Netherlands Museum of Anthropology and Prehistory.

--1996. Introduction: themes and concepts in the study of early agriculture, in D.R. Harris (ed.) The Origins and Spread of Agriculture and Pastoralism in Eurasia: 1-9. London: UCL Press.

HARVEY, E.L. 2006. Early agricultural communities in Northern and Eastern India: an archaeobotanical investigation. Unpublished PhD thesis, University College London.

HELBAEK, H. 1960. The palaeoethnobotany of the Near East and Europe, in R.J. Braidwood & B. Howe (ed.) Prehistoric Investigations in Iraqi Kurdistan: 99-108. Chicago: University of Chicago Press.

HENAN PROVINCIAL INSTITUTE OF ARCHAEOLOGY. 1999. Wuyang jiahu (Jiahu Site in Wuyang). Beijing: Science Press [in Chinese].

HIGHAM, C. 2005. East Asian Agriculture and its Impact, in C. Scarre (ed.) The Human Past. World Prehistory and the Development of Human Societies: 234-63. London: Thames & Hudson.

HILLMAN, G. & M.S. DAVIES. 1990. Measured Domestication Rates in Wild Wheats and Barley Under Primitive Cultivation and their Archaeological Implications. Journal of World Prehistory 4:157-222.

HOSHIKAWA, K. 1993. Anthesis, Fertilization and Development of Caryopsis, in T. Matsuo & K. Hoshikawa (ed.) Science of the Rice Plant, Volume One. Morphology: 339-76. Tokyo: Food and Agriculture Policy Research Center.

HUANG, F. & M. ZHANG. 2000. Pollen and phytolith evidence for rice cultivation during the Neolithic at Longquizhuang, eastern Jainghuai, China. Vegetation History and Archaeobotany 9: 161-8.

JIANG, L. & L. LIU. 2006. New evidence for the origins of sedentism and rice domestication in the Lower Yangzi River, China. Antiquiy 80: 355-61.

JONES, M.K. & T. BROWN. 2001. Agricultural origins: the evidence from modern and ancient DNA. The Holocene 10: 769-76.

KEELEY, L.H. 1995. Protoagricultural practices among Hunter-Gatherers: a cross-cultural survey, in T.D. Price & A.B. Gebauer (ed.) Last Hunters, First Farmers. New Perspectives on the Prehistoric Transition to Agriculture: 243-72. Santa Fe: School of American Research Press.

KUZMIN, Y.V. 2006. Chronology of the earliest pottery in East Asia: progress and pitfalls. Antiquity 80: 362-71.

LI, C., Y. ZHANG, K. YING, X. DANG & B. HAN. 2004. Sequence variation of simple sequence repeats on chromosome-4 in two subspecies of Asian cultivated rice. Theoretical and Applied Genetics 108: 392-400.

LIU, J. 1985. Some observations on the archaeological site of Hemudu, Zhejiang Province, China. Bulletin of the Indo-Pacific Prehistory Association 6: 40-5.

LONDO, J.P., Y.-C. CHIANG, K.-H. HUNG, T.-Y. CHIANG & B.A. SCHAAL. 2006. Phylogeography of Asian wild rice, Oryza rufipogon, reveals multiple independent domestications of cultivated rice, Oryza sativa. Proceedings of the National Academy of Sciences (USA) 103: 9578-83.

LU, T.L.D. 1999. The Transition from Foraging to Farming and the Origin of Agriculture in China (British Archaeological Reports International Series 774). Oxford: John & Erica Hedges.

MENNINGER, E.A. 1977. Edible nuts of the world. Stuart (FL): Horticultural Books.

OKA, H.I. 1988. Origin of Cultivated Rice. Tokyo: Japanese Scientific Societies Press.

PEARSALL, D.M., D.R. PIPERNO, E.H. DINAN, M. UMLAUF, Z. ZHAO & R.A. BENFER JR. 1995. Distinguishing rice (Oryza sativa Poaceae) from wild Oryza species through phytolith analysis: results of preliminary research. Economic Botany 49(2): 183-96.

PEI, A. 1998. Notes on new advancements and revelations in the agricultural archaeology of early rice domestication in the Dongting Lake region. Antiquity 72: 878-85.

PIPERNO, D.R. & D.M. PEARSALL. 1998. The Origins of Agriculture in the Lowland Neotropics. New York: Academic Press.

QIN, L. 2000. Review of the history and issues in the study of Liangzhu culture, Kaoguxueyanjiu [Archaelogical Studies] 2000 (4): 77-100 [in Chinese].

--2003. A Study of Prehistoric Social Structure in Taihu Area. Unpublished PhD thesis, Peking University [in Chinese].

RICE, P. 1999. On the origins of pottery. Journal of Archaeological Method and Theory 6: 1-54.

SANO, R. & H. MORISHIMA. 1992. Indica-Japonica differentiation of rice cultivars viewed from variations in key characters of isozyme, with special reference to Himalayan hilly areas. Theoretical and Applied Genetics, 84: 266-74.

SATO, Y.-I. 2002. Origin of Rice Cultivation in the Yangtze River Basin, in Y. Yasuda (ed.) The Origins of Pottery and Agriculture: 143-50. New Delhi: Lustre Press/Roll Books.

SATO, Y.-I., R. ISHIKAWA & H. MORISHIMA. 1990. Nonrandom association of genes and characters found in indica x japonica hybrids of rice. Heredity 65: 75-9.

TANG, L. 2003. The Primitive Rice Remains from Chuodun Site, in Chuodun Shan Site: collected papers, Dongnan Wenhua [southeast Culture], 2003 (special issue 1): 46-9 [in Chinese].

TANNO, K.-I. & G. WILLCOX. 2006. How Fast was Wild Wheat Domesticated? Science 311: 1886.

TAO, J., M.-T. CHEN & S. XU. 2006. A Holocene environmental record from the southern Yangtze River delta, eastern China. Palaeogeography, Palaeoclimatology, Palaeoecology 230: 204-29.

THOMPSON, G.B. 1996. The Excavations of Khok Phanom Di, a prehistoric site in Central Thailand. Volume IV. Subsistence and environment: the botanical evidence. The Biological Remains Part III. London: The Society of Antiquaries of London.

--1997. Archaeobotanical indicators of rice domestication--a critical evaluation of diagnostic criteria, in R. Ciarla & F. Rispoli (ed.) South-East Asian Archaeology 1992:159-74. Rome: Istituto Italiano per l'Africa e l'Oriente.

TSANG, C.-H. 2005. Recent discoveries at the Tapenkeng culture sites in Taiwan: implications for the problem of Austronesian origins, in L. Sagart, R. Blench & A. Sanchez-Mazas (ed.) The Peopling of East Asia. Putting together archaeology, linguistics and genetics: 63-74. London: Routledge-Curzon.

USHER, G. 1974. Dictionary of plants used by man. New York: Hafner.

VAUGHAN, D.A. 1994. The wild relatives of rice: a genetic resources handbook. Los Banos, Phillipines: International Rice Research Institute.

VAUGHAN, D.A., H. MORISHIMA & K. KADOWAKI. 2003. Diversity in the Oryza genus. Current Opinion in Plant Biology 6: 139-46.

WAN, J. & H. IKEHASHI. 1997. Identification of two types of differentiation in cultivated rice (Oryza sativa L.) detected by polymorphism of isozymes and hybrid sterility. Euphytica 94: 151-61.

WANG, C. & J. DING. 2000. The Plant Opal Analysis of Shaoqingshan Site, Kunshan City, Jiangsu Province, Kaogu (Archaeology) 2000(4): 87-93 [in Chinese].

WEISS, E., M.E. KISLEV & A. HARTMANN. 2006. Autonomous cultivation before domestication. Science 312: 1608-10.

WILKE, P.J., R. BETTINGER, T.F. KING & J.F. O'CONNELL. 1972. Harvest selection and domestication in seed plants. Antiquity 46: 203-9.

WILLCOX, G. 2004. Measuring grain size and identifying Near Eastern cereal domestication: evidence from the Euphrates valley. Journal of Archaeological Science 31: 145-50.

WINTERHALDER, B. & C. GOLAND. 1993. On population, foraging efficiency and plant domestication. Current Anthropology 34:710-15.

YASUDA, Y. 2002. Origins of pottery and agriculture in East Asia, in Y. Yasuda (ed.) The Origins of Pottery and Agriculture: 119-42. New Delhi: Lustre Press/ Roli Books.

YOU, X. 1976. Some Issues on Rice Grains and Bone-Spade Si Unearthed from Layer 4 of Hemudu Site. Wenwu (Cultural Relics) 8:20-23 [in Chinese].

ZHANG, J. & X. WANG. 1998. Notes on the recent discovery of ancient cultivated rice at Jiahu, Henan Province: a new theory concerning the origin of Oryza japonica in China. Antiquity 72: 897-901.

ZHANG, W. 2002. The bi-peak tubercle of rice, the character of ancient rice and the origin of cultivated rice, in Y. Yasuda (ed.) The Origins of Pottery and Agriculture: 205-16. New Delhi: Lustre Press/Roli Books.

ZHAO, S. & W.-T. Wu. 1987. Early Neolithic Hemudu Culture along the Hangzhou estuary and the Origin of Domestic Paddy Rice in China. Asian Perspectives 27: 29-34.

ZHAO, Z. 1998. The Middle Yangtze region in China is one place where rice was domesticated: phytolith evidence from the Diaotonghuan Cave, Northern Jaingxi. Antiquity 72: 885-97.

--2005. Palaeoethnobotany and its new achievements in China. Kaogu (Archaeology) 2005(7):42-9 [in Chinese].

Zhejiang Provincial Institute of Cultural Relics and Archaeology. 2003. Hemudu: xinshiqi shidai yizhi kaogu fajue baogao (Hemudu: an Excavation Report of the Neolithic Site). Beijing: Cultural Relics Publishing House [in Chinese].

--2004. Kuahuqiao. Beijing: Cultural Relics Publishing House [in Chinese].

ZHENG, Y., X. You, J. Xu, Q. BIAN & W. YU. 1994. Rice Phytolithic Analysis of Hemudu Site. Journal of Zhejiang Agriculture University 1994(1): 81-5 [in Chinese].

ZHENG, Y., Y. DONG, A. MATSUI, T. UDATSU & H. FUJIWARA. 2003. Molecular genetic basis of determining subspecies of ancient rice using shape of phytoliths, Journal of Archaeological Science 30: 1215-21.

ZHENG, Y., L.-P. JIANG & J.-M. ZHENG. 2004a. Study on the Remains of Ancient Rice from Kuahuqiao Site in Zhejiang Province. Chinese Journal of Rice Science 18:119-24 [in Chinese].

ZHENG, Y., A. MATSUI & H. FUJIWARA. 2004b. Phytoliths of rice detected in the Neolithic sites in the Valley of the Taihu Lake in China. Environmental Archaeology 8: 177-83.

ZHOU, J. 1981. Investigation of Ancient Rice from Mid and Lower Yangtze River. Yunnan nongye keji (Yunnan Agriculture Science and Technique) 6:1-6 [in Chinese].

--2003. The Morphological Investigation and Identification of Rice Grain from Hemudu Site, in Zhejiang Provincial Institute of Cultural Relics and Archaeology Hemudu: xinshiqi shidai yizhi kaogu fajue baogao (Hemudu: an Excavation Report of the Neolithic Site): 429-30. Beijing: Cultural Relics Publishing House [in Chinese].

ZOHARY, D. & M. HOPF. 2000. Domestication of Plants in the Old World, third edition. Oxford: Oxford University Press.

ZOU, H., J. GU, M. LI, L. TANG, J. DING & Q. YAO. 2000. Findings of paddies of Majiabang Culture at Caoxieshan, Jiangsu Province, in W. Yan & Y. Yasuda (ed.) The Origins of Rice Agriculture, Pottery and Cities: 97-114. Beijing: Cultural Relics Publishing House [in Chinese].

Dorian Q Fuller (1), Emma Harvey (1) & Ling Qin (2)

(1) Institute of Archaeology, University College London, UK

(2) School of Archaeology and Museology, Peking University, Beijing, China

Table 1. Plant species identified from Hemudu (H) and Kuahuqiao (K)
from fruit remains, and their possible uses, taken from the final
reports (Zhejiang Provincial Institute of Archaeology 2003;
2004). Ethnographically documented uses based on Usher 1974;
Menninger 1977; Notes: 1. Acorn identifications are difficult, but
judging by illustrated and examined acorn receptacles the East Asian
'qinggang oaks' Cyclobalanopsis spp., appear to dominate. 2. Not
recorded in report, but included amongst unidentified fruit seeds,
examined by the authors at the Hemudu museum; this species was
identified in the report of the basis of leaf remains; 3. On the
specific identity and domestication status of rice, see discussion
in text.

 Taxa; Common names
 (English, Chinese) Probable use

Lagenaria siceraria HK Containers, fishing
 net floats, seeds can
Eng. Bottle gourd, Ch. be processed for
 [TEXT NOT REPRODUCIBLE oily kernal
 IN ASCII] Hu lu

Quercus spp. (sensu lato) HK Potential carbohydrate
 staple (storable)
Eng. Oaks, acorns, Ch.
 [TEXT NOT REPRODUCIBLE
 IN ASCII] Xiang zi
 (probably mainly
 Lithocarpus and Cyclo-
 balanopsis oaks) (1)

Choerospondias axillaris HK Edible fruits, rich
 in vitamin C; also
Eng. "Southern Sour Jujube", medicinal
 Ch. [TEXT NOT REPRODUCIBLE
 IN ASCII] Nan suan zao

Amygdalus davidiana H Edible seasonal fruits,
 (syn Prunus davidiana) (2) seeds can be eaten
 roasted (like almonds),
Eng. Chinese mountain peach, and stored in stone
 Ch. [TEXT NOT REPRODUCIBLE
 IN ASCII] shan tao

Amygdalus (Prunus) K As above.
persica [Rosaceae],

Eng. True peach, Ch.
[TEXT NOT REPRODUCIBLE
IN ASCII] Mao tao

Prunus mume K Edible seasonal fruits,
 seeds can be eaten
Eng. Mume apricot; Ch. roasted (like almonds),
[TEXT NOT REPRODUCIBLE and stored in stone
IN ASCII] Suan mei

Prunus armeniaca K Edible seasonal fruits,
 seeds can be eaten
Eng. Apricot, Ch. [TEXT roasted (like almonds),
NOT REPRODUCIBLE IN and stored in stone
ASCII] Xing

Euryale ferox HK Seeds dried to make
 starchy flour
Eng. Foxnut, "gorgon (storable); stems and
seeds"; Ch. [TEXT roots eaten as
NOT REPRODUCIBLE IN vegetable
ASCII] gian shi

Sophora sp. H Leaves or roots used
 medicinally (Sophora
Eng. Sophora, Ch. [TEXT spp.); pods used as a
NOT REPRODUCIBLE IN yellow dyestuff (S.
ASCII] Huai japonica)

Coix lachryma jobi H Grains edible as
 cereal (storable)
Eng. Job's tears, [TEXT
NOT REPRODUCIBLE IN
ASCII] Yi yi

Trapa bispinosa HK Edible nut, storable,
 potential staple(?)
Eng. Water chestnut, Ch.
[TEXT NOT REPRODUCIBLE
IN ASCII] Ling jiao

Polygonaceae, K Edible with roasting;
 species in this family
Eng. Knotweed (family), known to have been
Ch. [TEXT NOT REPRO- used in aboriginal
DUCIBLE IN ASCII] North America and
 Jomon Japan

Oryza rufipogonlsativa HK Potential carbohydrate
(3) [Poaceae], staple (storable)
Eng. Rice, Ch. [TEXT
NOT REPRODUCIBLE IN
ASCII] Dao