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Open for Competition: Domesticates, Parasitic Domesticoids and the Agricultural Niche

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Open for Competition: Domesticates, Parasitic Domesticoids and the Agricultural Niche Fuller, D Q and Stevens, C J 2017 Open for Competition: Domesticates, Parasitic Domesticoids and the Agricultural Niche. Archaeology International, No. 20: pp. 110–121, DOI: https://doi.org/10.5334/ai-359 RESEARCH ARTICLE Open for Competition: Domesticates, Parasitic Domesticoids and the Agricultural Niche Dorian Q. Fuller and Chris J. Stevens This paper explores the relationship of weeds and crop parasites in the domestication of crop-plants within the Old World, drawing predominately on China and the Near East. This relationship is explored using the concept of niche construction in which the act of cultivation sets about chains of feedback in which the ecological worlds of plants and humans became increasingly intertwined resulting in ever increasing spheres of interdependence. Into this domestication entanglement a number of peripheral organisms (termed parasitic domesticoids) were drawn, from the weeds which came to inhabit arable fields, to the insect pests and rodents that came to settle in the grain stores of the first farmers. The evolution and spread of these organisms is then outlined against that of the crop itself. Introduction There is no doubt that the suite of human and population increase, whereas domesti- activities involved in cultivation created coids are parasitic in that they prosper at the environments that would not otherwise expense of domesticates and humans. These exist, and non-human organisms have categories are not necessarily fixed for all adapted to these environments. The adapted, time, as crops can become weeds and vice- co-evolved organisms include domesticated versa some weeds have become crops. These plants (true domesticates) as well as what we categories can be applied beyond the plants will refer to as domesticoids, comprising taxa in the field, in that other organisms, such that tagged along becoming also increas- as rodents and insects can be considered ingly part of the cultivated environment. in these terms. While terms such as synan- These domesticoids include weeds and crop thrope or commensal are often applied to pests. In terms of biological coevolution, such taxa, we suggest that parasitic domesti- domestication creates a symbiotic relation- coid is more fitting, as these taxa have under- ship, in which both organisms, crops and gone adaptions that parallel domestication, people benefit in terms of reproduction making them increasingly dependent on the human created niche of agriculture and sedentism, and parasitic in that they derive UCL Institute of Archaeology, London WC1H 0PY, UK their nutrients at the expense of the cultural Corresponding author: Dorian Q Fuller food production. In other words the culturally ([email protected]) maintained and transmitted agricultural niche Fuller and Stevens: Open for Competition 111 is integral to their survival, and they are more dormancy, increase in seed size and reduction than mere fellow travellers. of seed dispersal aids likes barbs and awns Niche Construction has become widely (Harlan, de Wet and Price 1973). While these accepted as a useful conceptual framework changes ‘improve’ crop plants, they come for the discussion of plant and animal domes- with costs, in that they entrap people in new tication (e.g. Fuller and Lucas 2017; Scott forms of labour, including maintenance of soil 2017; Watkins 2017; Zeder 2016). Cultural fertility and new steps in crop-processing niche construction implicates processes (Fuller, Allaby and Stevens 2010; Fuller and of change via feedback between culturally Lucas 2017; Fuller et al. 2016). Growth in transmitted practices, long-term modifica- archaeobotanical evidence and refinements tions to the environment (clearance, arable in methods have made it possible to docu- soils, etc.), and genetic evolution in various ment not just the presence but also the organisms, not just the human associated quantitative increase in these traits over time, taxa that were the targets of domestication, allowing rates of change to be estimated but fellow travellers, opportunists, and even (Fuller et al. 2014). Such information in turn unseen micro-organisms. can then be related to archaeological context The present paper explores the entangle- and innovations in technology, techniques ments between the evolution of cultivation and labour organization of subsistence. and storage practices with the adaptations Cultivation comes in a wide range of forms in crops, weeds and pests. First, how the but clearance of pre-existing vegetation and domestication syndrome of traits in seed disturbance of the soil is largely universal. crops adapted them to cultivation and pro- More intensive forms of cultivation can be vided increasing returns to yield, but in expected to remove all pre-existing visible doing so forced increased human effort vegetation and, through digging and turning which manifested in the evolution of cultural the soil, influence below ground biomass in practices and technologies. Second, how the terms of tubers and the seedbank. The axe, arable ecosystem constructed new commu- simple digging sticks, and fire, together with nities of parasitic domesticoids, a category human effort, can achieve this (Steensberg applied to storage pests and weeds, with 1986). Cultivation creates a habitat in which some of the latter eventually converging annual disturbance is both uniform and on and becoming true domesticates. Taken highly predictable, sunlight penetrates to together these entail that arable systems, the soil, becoming available for growth and and the social systems that maintained germination, and evapotranspiration often them, acquired an increasing number of increases. This creates a scenario of altered- innovations that were transmitted through competition, in which plants adapted to a combination of genetics, environment and rapid germination are favoured and faster culture. growing plants from more rigorous seedlings can quickly shade out their neighbours, per- Transformation to the Agricultural haps related to increased seed size (Harlan, Niche: Hook and Pull de Wet and Price 1973). Tillage also creates Domestication transforms crops and species deeper cracks which may bury seeds more so that they are better adapted to being deeply than they would be if they had fallen planted and harvested by people, and a set on natural soil surfaces, for which larger of recurrent parallel adaptations is found seeds with more nutrient reserves may also across species. In grain crops, planted from have an advantage. seed and harvested for their seeds, there is a Comparisons across taxa, including those well-documented domestication syndrome: cultivated as vegetables or tubers, indicates including loss of automatic seed dispersal, that increased seed size is a recurrent trait of loss of germination inhibition or seed domesticated species (Kluyver et al. 2017). 112 Fuller and Stevens: Open for Competition Figure 1: Showing size increase plotted on the Y-axis against time on the X-axis for the major crops from the Near East (top) and China (bottom). Average grain and seed size data for given sites and time periods from archaeobotancial data based on that in Fuller et al. 2014. Archaeobotanical data for several species cultivation by providing a predictable clearly documents this increase in average resource base for plants promotes a shift seed size over the millennia of early cultivation to nutrient-acquisition strategies that can (Fig. 1). While increased seed size may some- be contrasted with nutrient conservation times be linked to genetic changes favouring strategies seen in more stable environments other organ enlargements, it appears rather (Milla et al. 2015). as a selected trait, part of a general strategy of In turn increased seed size pulls farmers nutrient acquisition, including higher growth into a greater commitment to cultivars. rate, highly adapted to the nutrient-rich, Larger seeds have a higher caloric value, bare soils and open habitats of competi- augmented by an increased number of tion found in early cultivated fields. Thus grains per ear or panicle harvested. However, Fuller and Stevens: Open for Competition 113 to support this increased yield, each plant crops in cultivated fields, weeds have simi- requires more resources: soil nutrients and larly been selected for such environments. water, hence human efforts towards supplying In particular this has favoured the spread of these must also increase. Some of the taxa with transient seedbanks, which either earliest farming ecologies took advantage of germinate shortly after sowing, characteristic soils naturally replenished by annual flooding. of larger grained grasses and forbs of season- These include many early Pre-Pottery ally dry grassland (Type I), or after winter, Neolithic sites in the Levant, inferred from often characteristic of seasonally disturbed site locations (Scott 2017: 66; Sherratt 2007), coastal and mountain habitats (Type II) and the decrue rice agricultural systems of (Thompson and Grime 1979). In contrast China in use prior to evolution of domesti- are those weeds with persistent seedbanks cated, non-shattering rice (Fuller, Weisskopf (Thompson and Grime 1979) in which a large and Castillo 2016; Weisskopf et al. 2015). But dormant population is maintained in the soil as cultivation expanded, and crops became (Type IV). morphologically closer to domesticated These two contrasting ecologies, along forms, thus more demanding, intensive with intermediate seedbank form (Type III), management of soil conditions became are important
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