Presumed domestication? Evidence for wild rice cultivation and domestication in the fifth millennium BC of the Lower region Dorian Q Fuller1, Emma Harvey1 & Ling Qin2

Prompted by a recent article by Jiang and Liu in Antiquity (80, 2006), Dorian Fuller and his co-authors return to the question of rice cultivation and consider some of the difficulties involved in identifying the transition from wild to domesticated rice. Using data from Eastern , they propose that, at least for the Lower Yangtze region, the advent of rice domestication around 4000 BC was preceded by a phase of pre-domestication cultivation that began around 5000 BC. This rice, together with other subsistence foods like nuts, acorns and waterchestnuts, was gathered by sedentary hunter-gatherer-foragers. The implications for sedentism and the spread of agriculture as a long term process are discussed. Keywords: East Asia, China, Yangtze region, sixth millennium BC, fifth millennium BC, rice, foraging, cultivation, origins of agriculture

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 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.

1 Institute of Archaeology, University College London, UK 2 School of Archaeology and Museology, Peking University, Beijing, China Received: 15 June 2006; Accepted: 7 September 2006; Revised: 15 September 2006 antiquity 81 (2007): 316–331 316 seilyOa(98 u loT hn 18) aordasnl rgnfrrc olwdby followed rice for origin botanists, single prominent a 1980s favoured into the (1989), differentiation In Chang T. origins. claims also rice such but of (1988) all Oka theory and especially disproven with material, now riddled same a are on the claims predicated at such are & looking but Zhao scholars 2002), 1985; different Zhang Li between 206; 2003; contradictions 1997: 1981; Bellwood Zhou 1988; 1976; You Oka (e.g. 1987; Wu rice of forms ‘ancient’ intermediate or of mixture a contain assemblages hn19;Jns&Bon20;Fle 02 9;St 02.I hudb oe that remains which noted origin, be rice should Asian It single 2002). a of Sato assumption & 297; the Crawford 2002: overturns (e.g. Fuller evidence recently 2001; genetic literature Brown this archaeological & the Jones in 1998; considered Shen be to begun only has Cheng 1997; Ikehashi e aga 94.Ide,tems opeesv td odt niae w separate two indicates date of to study comprehensive for events most domestication the Indeed, 1994). Vaughan see n uaqa;7 ahbn;8 aio .Snz;1.Xjaa;1.Cudn 2 edn 3 oga and Longnan 13. Weidun; 12. Chuodun; 11. Xujiawan; Luojiajiao Pengtoushan; 10. 20. Nanzhuangqiao, Bashidang; 19. Songze; including ; Chengtoushan. 18. Lingjiatan; Hemudu; 9. area, 21. 17. 1. Sanxingcun; Majiabang Maqiao; 16. article: Longqiuzhuang; this 15. 6. 8. Qiucheng; in 14. Caoxieshan; Liangzhu; mentioned Nanhebang; sites 5. 7. archaeological showing Pu’anqiao; Shangshan; region, Yangtze and 4. of inset Kuahuqiao; with 3. Asia East 2.Tianluoshan; of Map 1. Figure sn ogrtnbea aaacmltdtruhnwrgntctcnqe niae that indicates techniques genetic newer through accumulated data as indica tenable longer no is rmdsic rgntrspecies, progenitor distinct from for rtnso iei h oe agz oti aao.I a ensgetdta rice that suggested been has It paradox. a contain Yangtze Lower the in rice on Writings indica japonica and ie nSuhAi rwsenSuhatAi (Londo Asia Southeast western or Asia South in rices japonica eg Sato (e.g. r hlgntclydsic,adrpeetsprt oetcto events domestication separate represent and distinct, phylogenetically are indica tal. et tal et japonica oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian and 90 ao&Mrsia19;Chen 1992; Morishima & Sano 1990; . 03 Vaughan 2003; japonica indica rz nivara Oryza ie nteSuhCiargo,adtodomestications two and region, China South the in rice, useisudrcliain hsseai,however, scenario, This cultivation. under subspecies and 317 japonica tal. et for indica 03 Li 2003; utvr,o just or cultivars, and tal. et rz rufipogon Oryza tal. et 04 o nomenclature, for 2004; indica 06.Ti evidence This 2006). tal et 93 a & Wan 1993; . rjust or ( es stricto sensu japonica, )

Research Evidence for wild rice cultivation and domestication 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.

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

318 uhdt aetaiinlybe sdt riea h i of from mix ratios, the at length-to-width arrive on to based used ranges Provinicial been traditionally shape 1999; have given Archaeology data of within such Institute spikelets Provincial Zheng 2003; and Archaeology from of grains (data Institute Hemudu rice and Jiahu of (KHQ), proportions Kuahuqaio The 2. Figure aevldt ftepeec faywl ieseiscnb ue u,adacetgan r sue ob ul mature. fully be to assumed are grains ancient and out, ruled be can species rice wild any of presence the if validity have oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian tal. et 04) n rmmdr esrdrc ouain fo avy20) While 2006). Harvey (from populations rice measured modern from and 2004a), 319 indica and japonica riso elti ie,ti a only can this sites, on grains

Research Evidence for wild rice cultivation and domestication

Figure 3. A scatter plot of length and width of grains measured in modern populations (72 populations, 15 grains each, in the Institute of Archaeology, University College London reference collection: Harvey 2006). Ovoid boundaries emphasise the distribution of Oryza rufipogon the East Asian wild progenitor, and wild species that are not connected with domesticated rice.

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.

320 ih niae h oa ubro risrmiigo h ln tec tg n h rprin htaena aueand mature near are that lower proportions at the graph of and The each mature. stage at to each more). plant maturity at entire or plant into an days the come of six on grains to (by remaining the expected grains immature for are of days substantially we that 16 number this grains of total From total of the 1993). a indicates proportions indicates Hoshikawa i.e. left right the (after each, shows the plant days rice which on two approximately individual diagram right stages, an The top these on shedding. at initiation grain graph grain type the extension wild derived by assuming have and shed, yields pollen harvest of and stage maturation the panicle Rice 4. Figure ohkw 93,o hc ssae h eido ri mauiyi hc ri rprin ifrfo hs of those from grain differ immature proportions which during grain window to which 10-day closer approximate in thus an and immaturity is length, grain grey towards solid biased of in be Shown period will grain. proportions the mature the shaded on is based standards which on 1993), of Hoshikawa Chart 5. Figure japonica ieganmtia eeomn ntrso egh it n hcns esrs(after measures thickness and width length, of terms in development metrical grain rice oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian 321 indica or rufipogon ahrthan rather , japonica .

Research Evidence for wild rice cultivation and domestication

Figure 6. A scatter plot of grain measurements from selected Neolithic sites, showing maximum and minimum measured ranges with solid lines and statistical standard deviations with dashed lines (as reported). Notice that grains from Kuahuqiao, Bashidang and the lower (Majiabang period) levels (8-6) at Longqiuzhuang fall largely or entirely in the expected immature grain proportions (compare Figure 9), while the latest grains from Longqiuzhuang, Songze period (level 4) indicate a clear shift towards longer and fatter grains that can be regarded as fully mature, and thus domesticated. Chuoden (Late Majiabang) also indicates a shift towards mature japonica type grains, but suggests local population differences from the domesticated rice at Longqiuzhuang. The small grains from Jiahu are suggestive of wild rice not in the sativa complex, such as O. officinalis (compare Figure 5). Sources: Kuahuqiao (Zheng et al. 2004b), Longqiuzhuang (Huang & Zhang 2000), Jiahu (Henan Provincial Institute of Archaeology 1999), Chuodun (Tang 2003), and Middle Yangzte Bashidang (Pei 1998).

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. 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 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 predomin- antly 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

322 nsaeaogtbliom per ob ne eei oto n hrfr reflects therefore and control genetic under be to appears (Zheng variation the bulliforms phylogenetics of some amongst While shape phytoliths. bulliform in from comes period Majiabang the during producer. seed wild prolific a is but populations HnnPoic nttt 99,wihaermral ml,admr ugsieo a of suggestive more and small, as remarkably such are rice, we Jiahu wild which from which different measurements 1999), Kuahuqiao, are Institute from note of Province those Also (Henan to wild. morphologically comparable and early rice, immature with as thin associated interpret often markedly site place a has BC), took agriculture, (7000-6000 region Bashidang domestication rice Yangtze from Middle data morphological but the limited, of from more data process is Quantitative the period. Majiabang/Hemudu that the suggests during This grains. wider lt,eprmnainwt e avsigmtos ieuroig a lohv played have small also these may domestication. of uprooting, in cross-pollination like context role from methods, the a harvesting rice In new wild evolution. with cultivated experimentation accelerating plots, the thus (Zou separating populations, province) of free-growing (Jiangshu terms with in Caoxieshan effect of important sites an the from recovered al. et been has these paddies, (although rice types individuals). torn domesticated and as well types as wild In wild cultivation. smooth, immature by suggesting include both relaxed thus may been included rices, had remains bases aids domesticated spikelet awn dispersal and spikelet addition of wild maintain modern to Studies selection of 2002). natural those that (Sato between characters densities intermediate hair had and wild domesticated, of Kuahuaqiao between sometime tillage Hemudu. during systematic or with four Hemudu, argues cultivation and only This they of 176-7). Kuahuqiao, suggesting 2004: development Institute earlier features Provincial the (Zhejiang hafting the for tillage poor heavy At to had production. suited these be food not and would plant recovered, soil wild were ‘spades’ regard for the we bone case immature of possible largely strong were manipulation a grains suggests rice as that this This evidence the (Chang 2003). with blades scapula Coupled spade Institute bone tillage. wooden Provincial through some haftable as Zhejiang or well as hafted 212; hoes, many or 1986: spades great as a regarded are yielded which Hemudu artefacts, well-known, is As period. of harvested evidence being the was larger with period, Liangzhu towards Majiabang and agree Songze the the shift phases. of therefore rice of domesticated significant presumably later, e.g. data than mature a rice, less These substantially on earlier indicate data 7). that Yangtze morphometrics metrical (Figure Lower grain Recent time bulliforms. the through larger in bulliforms produce sites plants from mature bulliforms more words other In egh(L hwasrn orlto ihpatmtrt (Zheng maturity plant with some vertical However, correlation and past. strong (HL) length the a horizontal in show especially maturity, variation (VL) plant phylogenetic length to relate such variation about of understood aspects be to much nadtoa oreo vdneta ugssasitfo mauet aueharvesting mature to immature from shift a suggests that evidence of source additional An rmteltrMjaagpro h rtpeevdfil ytm nepee ssmall as interpreted system, field preserved first the period Majiabang later the From mixture a indicated Hemudu, from spikelets rice of sample small a of study a addition, In Hemudu/Majiabang the during cultivation pre-domestication for argues evidence Other 00 n huu Zein rvne G 03.Ti eeomn ol have would development This 2003). (Gu Province) (Zhejiang Choudun and 2000) tal et oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian rz officinalis, Oryza 03 04;20b f Pearsall cf. 2004b; 2004a; 2003; . 323 hc i o otiuet ae domesticated later to contribute not did which tal. et 95,teeremains there 1995), tal. et 03 1217). 2003:

Research Evidence for wild rice cultivation and domestication

Figure 7. Size increase in Lower Yangtze rice phytoliths. The upper graph shows measured horizontal length (HL) and vertical length (VL) of rice bulliform phytoliths from Majiabang period samples (M), while the lower graph shows measurements from samples of the subsequent Songze (S) and Liangzhu (L) phases. The dashed oval represents the distribution of the Majiabang measurements. Data re-plotted, from Zheng et al. 1994; 2004a; 2004b; Wang & Ding 2000).

324 rdrn h euupae(..by (i.e. phase Hemudu the during or al utvto odmsiae ha n aly(olde19;Wlcx20;Tanno from 2004; evolution Willcox the 1998; in (Colledge Weiss period barley 2006; equivalent and Willcox an wheat & indicate domesticated an East to 1000- implies order Near cultivation the scenario the early of this from duration that a Data with noted years. plants be 1500 rice should non-shattering evolve It to Kuahuqiao). process of evolutionary perhaps cultivation end some the (with before foragers by beginning gathered rice wild represent Shangshan probably by aeo h hr h nerdpeec fteesae nteLwrYntervrae.As oe stergoa eln noaks in decline regional the the is at noted Also row area. the river Yangtze On Lower 1990). the in Harris (Tao stages data (after these the pollen cultivation of indicating in presence pre-domestication crops, indicated inferred domesticated and the on production chart based the food agriculture of to plant base foraging wild pure of from spectrum stages evolutionary significant idealised The 8. Figure yarclue nrcn er,teotooyhsbe htrc giutr ea early, began agriculture rice that been has orthodoxy driven imply the also being years, would sedentism It recent 2006). than of In Kuzmin rather 1999; agriculture. spread agriculture, (Rice by world preceded and the hunter-gatherers of origins sedentary parts other that the agriculture, many of in advance understanding in true well Asia is for East as in developed repercussions pottery Clearly, Asia. East wider in agriculture has hypothesis agriculture Our Asian East early for implications Wider ntegonso h bv aa esgetta idpatfo rdcinbgnby began production food plant wild that suggest we data, above the of grounds the On c 00B Fgr ) hsipista ale nso ie uha uhqa and Kuahuqiao as such rice, of finds earlier that implies This 8). (Figure BC 4000 . tal. et tal. et oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian 2006). 2006). c . 00B)adta oetctdrc a evolved had rice domesticated that and BC) 5000 325

Research Evidence for wild rice cultivation and domestication 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 , 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 , 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 frame 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 Pronvincial Institute of Archaeology and Chinese Academy of Social Sciences on the archaeobotany of Tian Luo Shan, a site of the . 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

326 hoZiu,proa omncto) ugsigta hs eeasgicn,i o the not if significant, a 1999; were Archaeology these resource. Jiahu that of staple, rice. suggesting Institute domesticated Provincial communication), of personal (Henan evolution Zhijun, acorns late Zhao of a quantities for large model produced the also check to opportunity an provide n.Rc,Ch. Rice, Eng. n.Ok,aon,Ch. acorns, Oaks, Eng. Polygonaceae, rnsmume Prunus uyl ferox Euryale n.Fxu,“ognses;Ch. seeds”; “gorgon Foxnut, Eng. rnsarmeniaca Prunus dnie nterpr ftebsso efrmis .O h pcfi dniyaddomestication and identity specific the On text. 3. in was remains; discussion species leaf see this of rice, museum; basis of Hemudu the status the of at report authors the the in by identified examined seeds, fruit unidentified amongst aa omnnms(nls,Ciee rbbeuse Probable axillaris Choerospondias siceraria Lagenaria Chinese) (English, names Common Taxa; Asian East the receptacles acorn examined Acorn and 1. illustrated oaks’ Notes: by ‘qinggang 1977; judging Menninger but 2003; 1974; difficult, Archaeology their Usher are of and on identifications Institute remains, based Provinicial fruit uses from (Zhejiang documented (K) reports Ethnographically Kuahuqiao final 2004). and the (H) from Hemudu taken from uses, identified possible species Plant 1. Table n.“otenSu uue,Ch. Jujube”, Sour “Southern Eng. Ch. gourd, Bottle Eng. Sophora shi Quercus rz rufipogon/sativa Oryza mgau davidiana Amygdalus rp bispinosa Trapa oxlachryma-jobi Coix mgau Puu)persica (Prunus) Amygdalus n.Kowe fml) Ch. (family), Knotweed Eng. n.Mm pio;Ch. apricot; Mume Eng. n.Arct Ch. Apricot, Eng. n.Spoa Ch. Sophora, Eng. n.Wtrcetu,Ch. chestnut, Water Eng. n.Cieemuti ec,Ch. peach, mountain Chinese Eng. n.Jbstears, Job’s Eng. n.Tu ec,Ch. peach, True Eng. hntao shan Cyclobalanopsis poal mainly (probably a unzao suan Nan sp. p.( spp. es lato sensu Cyclobalanopsis oaks) Dao Lithocarpus (syn 3 Xing iyi Yi [Poaceae], Huai ) 1 rnsdavidiana Prunus [Rosaceae], a tao Mao oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian ulu Hu in zi Xiang unmei Suan p. pert oiae .Ntrcre nrpr,btincluded but report, in recorded Not 2. dominate. to appear spp., and igjiao Ling qian ) 2 327 HK HK HK HK HK HK H H H K K K K sabove. As dbesaoa ris ed a eeaten be can seeds fruits, seasonal Edible dbesaoa ris ed a eeaten be can seeds fruits, seasonal Edible dbewt osig pce nti family this in species roasting; with Edible riseil scra (storable) cereal as edible Grains evso ot sdmdcnly( medicinally used roots or Leaves dbent trbe oeta staple(?) potential storable, nut, Edible dbesaoa ris ed a eeaten be can seeds fruits, seasonal Edible oeta abhdaesal (storable) staple carbohydrate Potential ed re omk trh or(storable); flour starchy make to dried Seeds oties signtflas ed a be can seeds floats, net fishing Containers, oeta abhdaesal (storable) staple carbohydrate Potential dbefut,rc nvtmnC also C; vitamin in rich fruits, Edible ose lk lod) n trdi stone in stored and almonds), (like roasted ose lk lod) n trdi stone in stored and almonds), (like roasted ot mrc n oo Japan Jomon and aboriginal America in North used been have to known p.;pd sda elwdetf ( dyestuff yellow japonica a as used pods spp.); ose lk lod) n trdi stone in stored and almonds), (like roasted tm n ot ae svegetable as eaten roots and stems rcse o iykernal oily for processed medicinal ) Sophora S.

Research Evidence for wild rice cultivation and domestication

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-buffering 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 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.

References Chang, T.T. 1989. Domestication and the spread of cultivated rices, in D.R. Harris & G.C. Hillman Bellwood, P. 1997. Prehistory of the Indo-Malaysian (ed.) Foraging and Farming. The Evolution of Plant archipelago, second edition. Honolulu: University of Exploitation: 408-17. London: Unwin Hyman. Hawaii Press. Chen, W.-B., I. Nakamura, Y.-I. Sato & H. Nakai. Chang, K.-C. 1986. The Archaeology of Ancient China. 1993. Indica and Japonica differentiation in New Haven: Yale University Press. Chinese landraces. Euphytica 74: 195-201.

328 ars D.R. Harris, Wet De J.M.J. J.R., Harlan, Gu et,P Gepts, Qin L. D.Q., Fuller, G. Lee, G. Zhao, Z. Underhill, A. G., Crawford, olde S. Colledge, ulr D.Q Fuller, D.J Cohen, Y. Tchuchimoto, S. Motohashi, R. C., Cheng, rwod G. Crawford, rwod G Crawford, .20.TeRc ad adfo hou Site Chuodun from Land Paddy Rice The 2003. J. , ..Balkema. A.A. nPalaeoethnobotany in (ed.) & Casparie Zeist W.C. and van W.A. scope in its implications, forbs: archaeological and grasses wild of exploitation 311-25. cereals. of evolution Comparative Chinese]. [in Culture] papers collected in Period, Majiabang of 1-44. giutr:rcn rgesi atAsia. East in progress recent agriculture: eeto experiment. selection Francis. evidence archaeological linguistic and genetic, Taiwan: and Asia East (ed.) continental Pejros Ross, I. M. & Blench, Lin R. M. Sanchez-Mazas, A. in archaeobotany rice, an of Lower for the challenges in and agriculture region of Yangtze onset late a for Evidence Disciplines Volume Retrospect, Fang H. Yu, H. & Luan, F. Nicholas, L. Feinman, 858-66. 72: etr&R oietr(ed.) S. Korisettar in R. Foundation, & Solid Settar a Laying India: in Studies Shandong. Liangchengzhen, from Results Preliminary China: Northern 22-9. Asia. East in Transition Pleistocene-Holocene the and Cereals 62:309-17. 46(2): Evolution and Biology SINEs. of Patterns on Interspersion Based the Rice: Cultivated of Origin Polyphyletic Ohtsubo H. Fukuta, xod Blackwell. Oxford: (ed.) Stark M.T. in ICARDA. domestication Qualset C.O. & (ed.) Willcox G. Valkoun, J. A.B. Damania, in analysis, multivariate using cultivation .Cai. F. 04 rpDmsiaina long-term a as Domestication Crop 2004. . h rgn farclueadcrop and agriculture of origins The 98 h rgn fDomesticated of Origins The 1998. . 03(pca su ) 42-5 1): issue (special 2003 02 it er fArchaeobotanical of Years Fifty 2002. . 4-6.NwDli Manohar. Delhi: New 247-363. : 94 tnhsoia vdnefrthe for evidence Ethnohistorical 1984. 98 dniyn pre-domestication Identifying 1998. 05 aeNoihcPatRmisfrom Remains Plant Neolithic Late 2005. 06 atAinPatDomestication, Plant Asian East 2006. . & h eiwo Archaeology of Review The 2-1 lpo Syria: Aleppo, 121-31. : , .Shen C. oga eha[southeast Wenhua Dongnan III. & rhelg fAsia of Archaeology lnsadAcetMn Studies Man: Ancient and Plants 39 Rotterdam: 63-9. : ..Harvey E.L. ln reigReviews Breeding Plant 0 67-75. 20: rhelg n Interactive and Archaeology 98 h rgn frice of origins The 1998. . & ChuodunShanSite: ua irtosin migrations Human .Ohtsubo E. odn alr& Taylor London: . & ninAcaooyin Archaeology Indian urn Anthropology Current oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian ..Price E.G. Evolution npress. In . Molecular 77-95. : 2003. . 19: Antiquity 1973. . 27: 24: 329 –1990. oe,M.K. Jones, L. Jiang, F. Huang, i . .Zag .Yn,X Liang X. Ying, K. Zhang, Y. C., Li, Y.V Kuzmin, L.H. Keeley, ohkw,K. Hoshikawa, G. Hillman, C Higham, ea rvnilIsiueo Archaeology of Institute Provincial Henan ebe,H. Helbaek, E.L Harvey, study the in concepts and themes Introduction: –1996. fsdnimadrc oetcto nteLower the China. River, in Yangzi domestication rice and sedentism of 0:392-400. 108: utvtdrice. Asian cultivated of subspecies two on in repeats chromosome-4 sequence simple of variation Sequence h vdnefo oenadacetDNA. ancient Holocene and modern from evidence the Archaeobotany and History Vegetation China. at Jainghuai, Neolithic eastern the Longquizhuang, during cultivation rice for evidence giutr oiyRsac Center. Research Policy Agriculture Morphology One. K. (ed.) & Matsuo Hoshikawa T. in Caryopsis, of Development nEs sa rgesadpitfalls. 362-71. and progress Asia: East in Press. Research American of Agriculture to PrehistoricTransition the on Perspectives New Farmers. (ed.) Gebauer A.B. & T.D.Price in survey, cross-cultural a Hunter-Gatherers: Prehistory Implications. Archaeological their and Cultivation Barley Primitive and Under Wheats Wild in Rates Domestication 3-3 odn hms&Hudson. & Thames Societies London: Human 234-63. of Development the and Prehistory (ed.) Scarre C. in Impact, Chinese]. [in Press Science 1999. 918 hcg:Uiest fCiaoPress. Chicago of University Chicago: 99-108. atadErp,i ..Badod&B Howe B. & Braidwood (ed.) R.J. in Europe, and East London. College University thesis, PhD Unpublished archaeobotanical investigation. an India: Eastern and Northern in Pastoralism and Eurasia Agriculture of Spread (ed.) and Harris Origins D.R. in agriculture, early of Prehistory. and Anthropology of Museum Netherlands Amsterdam: Lecture). Revolution Neolithic the etigDw n raigGon:Rethinking Ground: Breaking and Down Settling rhsoi netgtosi rq Kurdistan Iraqi in Investigations Prehistoric & uagjau(ih iei Wuyang). in Site (Jiahu jiahu Wuyang -.Lno:ULPress. UCL London: 1-9. : & 05 atAinArclueadits and Agriculture Asian East 2005. . .Liu L. 06 al giutrlcmuiisin communities agricultural Early 2006. . & 0 769-76. 10: 06 hoooyo h aletpottery earliest the of Chronology 2006. . 90 h aaotnbtn fteNear the of palaeoethnobotany The 1960. & :157-222. 4: 95 rtarclua rcie among practices Protoagricultural 1995. .Zhang M. .Brown. T. 93 nhss etlzto and Fertilization Anthesis, 1993. ..Davies M.S. 06 e vdnefrteorigins the for evidence New 2006. . hoeia n ple Genetics Applied and Theoretical 3-6 oy:Fo and Food Tokyo: 339-76. : cec fteRc ln,Volume Plant, Rice the of Science Antiquity 00 olnadphytolith and Pollen 2000. . 4-2 at e School Fe: Santa 243-72. : TefhKroon-Voordact (Twelfth 01 giutrlorigins: Agricultural 2001. 90 Measured 1990. . h ua at World Past. Human The atHnes First Hunters, Last ora fWorld of Journal 0 355-61. 80: & Antiquity .Han B. :161-8. 9: The 80: Beijing: 2004. . . The : :

Research Evidence for wild rice cultivation and domestication

Liu, J. 1985. Some observations on the archaeological Tanno, K.-I. & G. Willcox. 2006. How Fast was site of Hemudu, Zhejiang Province, China. Bulletin Wild Wheat Domesticated? Science 311: 1886. of the Indo-Pacific Prehistory Association 6: 40-5. Tao, J., M.-T. Chen & S. Xu. 2006. A Holocene Londo, J.P., Y.-C. Chiang, K.-H. Hung, T.-Y. environmental record from the southern Yangtze Chiang & B.A. Schaal. 2006. Phylogeography of River delta, eastern China. Palaeogeography, Asian wild rice, Oryza rufipogon, reveals multiple Palaeoclimatology, Palaeoecology 230: 204-29. independent domestications of cultivated rice, Thompson, G.B. 1996. The Excavations of Khok Oryza sativa. Proceedings of the National Academy of Phanom Di, a prehistoric site in Central Thailand. Sciences (USA) 103: 9578-83. Volume IV. Subsistence and environment: the Lu, T.L.D. 1999. The Transition from Foraging to botanical evidence. The Biological Remains Part III. Farming and the Origin of London: The Society of Antiquaries of London. (British Archaeological Reports International Series –1997. Archaeobotanical indicators of rice 774). Oxford: John & Erica Hedges. domestication – a critical evaluation of diagnostic Menninger, E.A. 1977. Edible nuts of the world. criteria, in R. Ciarla & F. Rispoli (ed.) South-East Stuart (FL): Horticultural Books. Asian Archaeology 1992: 159-74. Rome: Istituto Oka, H.I. 1988. Origin of Cultivated Rice.Tokyo: Italiano per l’Africa e l’Oriente. Japanese Scientific Societies Press. Tsang, C.-H. 2005. Recent discoveries at the Pearsall, D.M., D.R. Piperno, E.H. Dinan, Tapenkeng culture sites in Taiwan: implications for M. Umlauf, Z. Zhao & R.A. Benfer Jr. 1995. the problem of Austronesian origins, in L. Sagart, Distinguishing rice (Oryza sativa Poaceae) from R. Blench & A. Sanchez-Mazas (ed.) The Peopling wild Oryza species through phytolith analysis: of East Asia. Putting together archaeology, linguistics results of preliminary research. Economic Botany and genetics: 63-74. London: Routledge-Curzon. 49(2): 183-96. Usher, G. 1974. Dictionary of plants used by man. Pei, A. 1998. Notes on new advancements and New York: Hafner. revelations in the agricultural archaeology of early Vaughan, D.A. 1994. The wild relatives of rice: a genetic rice domestication in the Dongting Lake region. resources handbook. Los Banos, Phillipines: Antiquity 72: 878-85. International Rice Research Institute. Piperno, D.R. & D.M. Pearsall. 1998. The Origins of Vaughan, D.A., H. Morishima & K. Kadowaki. Agriculture in the Lowland Neotropics. New York: 2003. Diversity in the Oryza genus. Current Academic Press. Opinion in Plant Biology 6: 139-46. Qin, L. 2000. Review of the history and issues in the Wan, J. & H. Ikehashi. 1997. Identification of two study of , Kaoguxue yanjiu types of differentiation in cultivated rice (Oryza [Archaelogical Studies] 2000(4): 77-100 sativa L.) detected by polymorphism of isozymes [in Chinese]. and hybrid sterility. Euphytica 94: 151-61. –2003. A Study of Prehistoric Social Structure in Taihu Wang, C. & J. Ding. 2000. The Plant Opal Analysis Area. Unpublished PhD thesis, Peking University of Shaoqingshan Site, Kunshan City, [in Chinese]. Province, Kaogu (Archaeology) 2000(4): 87-93 [in Rice, P. 1999. On the origins of pottery. Journal of Chinese]. Archaeological Method and Theory 6: 1-54. Weiss, E., M.E. Kislev & A. Hartmann. 2006. Sano, R. & H. Morishima. 1992. Indica-Japonica Autonomous cultivation before domestication. differentiation of rice cultivars viewed from Science 312: 1608-10. variations in key characters of isozyme, with special Wilke, P.J., R. Bettinger, T.F. King & reference to Himalayan hilly areas. Theoretical and J.F. O’Connell. 1972. Harvest selection and Applied Genetics, 84: 266-74. domestication in seed plants. Antiquity 46: Sato, Y.-I. 2002. Origin of Rice Cultivation in the 203-9. Yangtze River Basin, in Y. Yasuda (ed.) The Origins Willcox, G. 2004. Measuring grain size and of Pottery and Agriculture: 143-50. New Delhi: identifying Near Eastern cereal domestication: Lustre Press/Roli Books. evidence from the Euphrates valley. Journal of Sato, Y.-I., R. Ishikawa & H. Morishima. 1990. Archaeological Science 31: 145-50. Nonrandom association of genes and characters Winterhalder, B. & C. Goland. 1993. On found in indica x japonica hybrids of rice. Heredity population, foraging efficiency and plant 65: 75-9. domestication. Current Anthropology 34: 710-15. Tang, L. 2003. The Primitive Rice Remains from Yasuda, Y. 2002. Origins of pottery and agriculture in Chuodun Site, in Chuodun Shan Site: collected East Asia, in Y. Yasuda (ed.) The Origins of Pottery papers, Dongnan Wenhua [southeast Culture], 2003 and Agriculture: 119-42. New Delhi: Lustre Press/ (special issue 1): 46-9 [in Chinese]. Roli Books.

330 ho S. Zhao, X. You, Zheng,Y.,X.You,J.Xu,Q.Bian–2004. and Relics Cultural of Institute Provincial Zhejiang in achievements new its and Palaeoethnobotany –2005. Z Zhao, W Zhang, hn,J. Zhang, rgno oetcPdyRc nChina. in Perspectives Rice Paddy Domestic the of and Origin estuary Hangzhou the along Culture Books. Agriculture cultivated (ed.) Yasuda of Y. origin in the rice, and rice ancient of character os i Chinese]. [in House Chinese]. [in House Publishing Site) Neolithic Report the Excavation of an (Hemudu: baogao fajue kaogu 2003. Archaeology. Chinese]. [in China. Northern Jaingxi. Cave, Diaotonghuan the phytolith from domesticated: evidence was rice where place one rvne e hoycnenn h rgnof origin japonica the Oryza concerning theory Henan new Jiahu, a at Province: rice cultivated ancient of discovery Site. Bone-Spade Chinese]. University Agriculture Zhejiang Site. of Hemudu of Analysis Phytolithic Rice Kuahuqiao. 96 oeIse nRc risand Grains Rice on Issues Some 1976. 98 h ideYntergo nCiais China in region Yangtze Middle The 1998. . ew Clua Relics) (Cultural Wenwu & & 02 h ipa ueceo ie the rice, of tubercle bi-peak The 2002. . ag (Archaeology) Kaogu Antiquity .T Wu W.-T. .Wang. X. 0-6 e eh:Lsr Press/Roli Lustre Delhi: New 205-16. : 7 29-34. 27: Si nate rmLyr4o Hemudu of 4 Layer from Unearthed nChina. in ejn:Clua eisPublishing Relics Cultural Beijing: 2 885-97. 72: 97 al elti Hemudu Neolithic Early 1987. . ejn:Clua Relics Cultural Beijing: . euu isiisia yizhi shidai xinshiqi Hemudu: 98 oe nterecent the on Notes 1998. h rgn fPteyand Pottery of Origins The Antiquity 057:42-9 2005(7): oinQFle,Em avy&Ln Qin Ling & Harvey Emma Fuller, Q Dorian :2-3[nChinese]. [in 20-23 8: & 941:8- [in 81-5 1994(1): .Yu W. 2 897-901. 72: 1994. . Asian Journal 331 hn,Y,A Matsui A. Y., Zheng, Jiang L.-P. Y., Zheng, o,H,J u .L,L ag .Ding J. Tang, L. Li, M. Gu, J. H., Zou, 20.TeMrhlgclIvsiainand Investigation Morphological The –2003. J Zhou, hn,Y,Y og .Mtu,T Udatsu T. Matsui, A. Dong, Y. Y., Zheng, oay D. Zohary, ulsigHue[nChinese]. [in House Publishing Cities and Y. (ed.) & Yasuda Yan W. in at Province, Culture Jiangsu Majiabang Caoxieshan, of paddies of Findings 2000. h alyo h ah aei China. Archaeology in Environmental Lake Taihu in the sites of Neolithic Valley the the in detected rice of Phytoliths Science Province. Kuahuqiao Zhejiang from in Rice Site Ancient of Remains the on 1215-21. shape phytoliths, using of rice ancient of subspecies determining Fujiwara. nvriyPress. University World Old the in hjagPoica nttt fClua eisand Relics Archaeology Cultural of in Institute Site, Provincial Hemudu Zhejiang from Grain Rice of Identification Chinese]. [in Technique) and Science Agriculture (Yunnan River. Yangtze Lower and ulsigHue[nChinese]. [in House Publishing the of Site) Report Neolithic Excavation an (Hemudu: baogao fajue 91 netgto fAcetRc rmMid from Rice Ancient of Investigation 1981. . 8192 i Chinese]. [in 18:119-24 & 714 ejn:Clua Relics Cultural Beijing: 97-114. : 03 oeua eei ai of basis genetic Molecular 2003. .Hopf. M. h rgn fRc giutr,Pottery Agriculture, Rice of Origins The euu isiisia ih kaogu yizhi shidai xinshiqi Hemudu: 2-0 ejn:Clua Relics Cultural Beijing: 429-30. : ora fAcaooia Science Archaeological of Journal hr dto.Ofr:Oxford Oxford: edition. third , & & .Fujiwara. H. 2000. .M Zheng. J.-M. hns ora fRice of Journal Chinese unnnny keji nongye Yunnan :177-83. 8: oetcto fPlants of Domestication 2004b. 04.Study 2004a. & .Yao Q. & :1-6 6: H. 30: .

Research