Search

The southward expansion of East Asian farmers profoundly influenced the social evolution of Southeast Asia by introducing cereal agriculture. However, the timing and routes of cereal expansion in key regions are unclear due to limited empirical evidence. Here we report macrofossil, microfossil, multiple isotopic (C/N/Sr/O) and paleoproteomic data directly from radiocarbon-dated human samples, which were unearthed from a site in Xingyi in central Yunnan and which date between 7000 and 3300 a BP. Dietary isotopes reveal the earliest arrival of millet ca. 4900 a BP, and greater reliance on plant and animal agriculture was indicated between 3800 and 3300 a BP. The dietary differences between hunter-gatherer and agricultural groups are also evident in the metabolic and immune system proteins analysed from their skeletal remains. The results of paleoproteomic analysis indicate that humans had divergent biological adaptations, with and without farming. The combined application of isotopes, archaeobotanical data and proteomics provides a new approach to documenting dietary and health changes across major subsistence transitions., Competing Interests: Conflict of interest The authors declare that they have no conflict of interest., (Copyright © 2023 Science China Press. Published by Elsevier B.V. All rights reserved.)

Cannabis grains are frequently reported from archaeological sites in Asia, and hypothesized centers of origins are China and Central Asia. Chinese early cannabis remains are often interpreted as evidence of hemp fabric production, in line with early textual evidence describing ritualistic hemp cloth use and hemp cultivation as a grain crop. Modern measurements on cannabis varieties show distinct sizes between fibre or oil/fibre and psychoactive varieties, the former having larger seeds on average than the latter. This paper reviews the current macro-botanical evidence for cannabis across East, Central and South Asia and builds a comparative framework based on modern cannabis seed measurements to help identify cannabis use in the past, through the metric analysis of archaeologically preserved seeds. Over 800 grains of cannabis were retrieved from the 2008 excavation of Haimenkou, Yunnan, Southwest China, dating to between 1650 and 400 bc. These are compared with other known archaeological cannabis and interpreted through the metric framework. This offers a basis for exploration of the seed morphometrics potential to infer cannabis cultivation and diversification in uses. At Haimenkou, cannabis seeds size mostly plot in the range of overlapping psychoactive/fibre types; we therefore suggest that the cannabis assemblage from Haimenkou is indicative of a crop beginning to undergo evolution from its early domesticated form towards a diversified crop specialized for alternative uses, including larger oilseed/fibre adapted varieties., Supplementary Information: The online version contains supplementary material available at 10.1007/s00334-023-00966-6., Competing Interests: Competing interestsThe authors declare no competing interests., (© The Author(s) 2023.)

Ancient Egyptian mummification was practiced for nearly 4000 years as a key feature of some of the most complex mortuary practices documented in the archaeological record. Embalming, the preservation of the body and organs of the deceased for the afterlife, was a central component of the Egyptian mummification process. Here, we combine GC-MS, HT-GC-MS, and LC-MS/MS analyses to examine mummification balms excavated more than a century ago by Howard Carter from Tomb KV42 in the Valley of the Kings. Balm residues were scraped from now empty canopic jars that once contained the mummified organs of the noble lady Senetnay, dating to the 18th dynasty, ca. 1450 BCE. Our analysis revealed balms consisting of beeswax, plant oil, fats, bitumen, Pinaceae resins, a balsamic substance, and dammar or Pistacia tree resin. These are the richest, most complex balms yet identified for this early time period and they shed light on balm ingredients for which there is limited information in Egyptian textual sources. They highlight both the exceptional status of Senetnay and the myriad trade connections of the Egyptians in the 2nd millennium BCE. They further illustrate the excellent preservation possible even for organic remains long removed from their original archaeological context., (© 2023. The Author(s).)

Plant life defines the environments to which animals adapt and provides the basis of food webs. This was equally true for hunter-gatherer economies of ancestral humans, yet through the domestication of plants and the creation of agricultural ecologies based around them, human societies transformed vegetation and transported plant taxa into new geographical regions. These human-plant interactions ultimately co-evolved, increasing human population densities, technologies of farming, and the diversification of landraces and crop complexes. Research in archaeology on preserved plant remains (archaeobotany) and on the genomes of crops, including ancient genomes, has transformed our scientific understanding of the complex relationships between humans and plants that are entailed by domestication. Key realizations of recent research include the recognition that: the co-evolution of domesticates and cultures was protracted, the adaptations of plant populations were unintended results of human economies rather than intentional breeding, domestication took place in dozens of world regions involving different crops and cultures, and convergent evolution can be recognized among cropping types - such as among seed crops, tuber crops, and fruit trees. Seven general domestication pathways can be defined for plants. Lessons for the present-day include: the importance of diversity in the past; genetic diversity within species has the potential to erode over time, but also to be rescued through processes of integration; similarly, diversification within agricultural ecosystems has undergone processes of decline, including marginalised, lost and 'forgotten' crops, as well as processes of renewal resulting from trade and human mobility that brought varied crops and varieties together., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

Asian rice ( Oryza sativa L.) is consumed by more than half of the world's population. Despite its global importance, the process of early rice domestication remains unclear. During domestication, wild rice ( Oryza rufipogon Griff.) acquired non-seed-shattering behavior, allowing humans to increase grain yield. Previous studies argued that a reduction in seed shattering triggered by the sh4 mutation led to increased yield during rice domestication, but our experiments using wild introgression lines show that the domesticated sh4 allele alone is insufficient for shattering loss in O . rufipogon . The interruption of abscission layer formation requires both sh4 and qSH3 mutations, demonstrating that the selection of shattering loss in wild rice was not as simple as previously suggested. Here we identified a causal single-nucleotide polymorphism at qSH3 within the seed-shattering gene OsSh1 , which is conserved in indica and japonica subspecies but absent in the circum -aus group of rice. Through harvest experiments, we further demonstrated that seed shattering alone did not significantly impact yield; rather, yield increases were observed with closed panicle formation controlled by SPR3 and further augmented by nonshattering, conferred by integration of sh4 and qSH3 alleles. Complementary manipulation of panicle shape and seed shattering results in a mechanically stable panicle structure. We propose a stepwise route for the earliest phase of rice domestication, wherein selection of visible SPR3 -controlled closed panicle morphology was instrumental in the sequential recruitment of sh4 and qSH3 , which together led to the loss of shattering.

Though chickens are the most numerous and ubiquitous domestic bird, their origins, the circumstances of their initial association with people, and the routes along which they dispersed across the world remain controversial. In order to establish a robust spatial and temporal framework for their origins and dispersal, we assessed archaeological occurrences and the domestic status of chickens from ∼600 sites in 89 countries by combining zoogeographic, morphological, osteometric, stratigraphic, contextual, iconographic, and textual data. Our results suggest that the first unambiguous domestic chicken bones are found at Neolithic Ban Non Wat in central Thailand dated to ∼1650 to 1250 BCE, and that chickens were not domesticated in the Indian Subcontinent. Chickens did not arrive in Central China, South Asia, or Mesopotamia until the late second millennium BCE, and in Ethiopia and Mediterranean Europe by ∼800 BCE. To investigate the circumstances of their initial domestication, we correlated the temporal spread of rice and millet cultivation with the first appearance of chickens within the range of red junglefowl species. Our results suggest that agricultural practices focused on the production and storage of cereal staples served to draw arboreal red junglefowl into the human niche. Thus, the arrival of rice agriculture may have first facilitated the initiation of the chicken domestication process, and then, following their integration within human communities, allowed for their dispersal across the globe.

Most models of selection incorporate some notion of environmental degradation where the majority of the population becomes less fit concerning a character resulting in pressure to adapt. Such models have been variously associated with an adaptation cost, the substitution load. Conversely, adaptative mutations that represent an improvement in fitness in the absence of environmental change have generally been assumed to be associated with negligible cost. However, such adaptations could represent a competitive advantage that diminishes resource availability for others and so induces a cost. This type of adaptation in the form of seedling competition has been suggested as a mechanism for increases in seed sizes during domestication, a trait associated with the standard stabilizing selection model. We present a novel cost framework for competitive selection that demonstrates significant differences in behaviour to environmental-based selection in intensity, intensity over time and directly contrasts with the expectations of the standard model. Grain metrics of nine archaeological crops fit a mixed model in which episodes of competitive selection often emerge from shifting optimum episodes of stabilizing selection, highlighting the potential prevalence of the mechanism outlined here and providing fundamental insight into the factors driving domestication., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

The evidence from ancient crops over the past decade challenges some of our most basic assumptions about the process of domestication. The emergence of crops has been viewed as a technologically progressive process in which single or multiple localized populations adapt to human environments in response to cultivation. By contrast, new genetic and archaeological evidence reveals a slow process that involved large populations over wide areas with unexpectedly sustained cultural connections in deep time. We review evidence that calls for a new landscape framework of crop origins. Evolutionary processes operate across vast distances of landscape and time, and the origins of domesticates are complex. The domestication bottleneck is a redundant concept and the progressive nature of domestication is in doubt., Competing Interests: Declaration of interests No interests are declared., (Copyright © 2021 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Understanding the evolutionary history of crops, including identifying wild relatives, helps to provide insight for conservation and crop breeding efforts. Cultivated Brassica oleracea has intrigued researchers for centuries due to its wide diversity in forms, which include cabbage, broccoli, cauliflower, kale, kohlrabi, and Brussels sprouts. Yet, the evolutionary history of this species remains understudied. With such different vegetables produced from a single species, B. oleracea is a model organism for understanding the power of artificial selection. Persistent challenges in the study of B. oleracea include conflicting hypotheses regarding domestication and the identity of the closest living wild relative. Using newly generated RNA-seq data for a diversity panel of 224 accessions, which represents 14 different B. oleracea crop types and nine potential wild progenitor species, we integrate phylogenetic and population genetic techniques with ecological niche modeling, archaeological, and literary evidence to examine relationships among cultivars and wild relatives to clarify the origin of this horticulturally important species. Our analyses point to the Aegean endemic B. cretica as the closest living relative of cultivated B. oleracea, supporting an origin of cultivation in the Eastern Mediterranean region. Additionally, we identify several feral lineages, suggesting that cultivated plants of this species can revert to a wild-like state with relative ease. By expanding our understanding of the evolutionary history in B. oleracea, these results contribute to a growing body of knowledge on crop domestication that will facilitate continued breeding efforts including adaptation to changing environmental conditions., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)

Archaeological and paleoecological evidence shows that by 10,000 BCE, all human societies employed varying degrees of ecologically transformative land use practices, including burning, hunting, species propagation, domestication, cultivation, and others that have left long-term legacies across the terrestrial biosphere. Yet, a lingering paradigm among natural scientists, conservationists, and policymakers is that human transformation of terrestrial nature is mostly recent and inherently destructive. Here, we use the most up-to-date, spatially explicit global reconstruction of historical human populations and land use to show that this paradigm is likely wrong. Even 12,000 y ago, nearly three quarters of Earth's land was inhabited and therefore shaped by human societies, including more than 95% of temperate and 90% of tropical woodlands. Lands now characterized as "natural," "intact," and "wild" generally exhibit long histories of use, as do protected areas and Indigenous lands, and current global patterns of vertebrate species richness and key biodiversity areas are more strongly associated with past patterns of land use than with present ones in regional landscapes now characterized as natural. The current biodiversity crisis can seldom be explained by the loss of uninhabited wildlands, resulting instead from the appropriation, colonization, and intensifying use of the biodiverse cultural landscapes long shaped and sustained by prior societies. Recognizing this deep cultural connection with biodiversity will therefore be essential to resolve the crisis., Competing Interests: The authors declare no competing interest., (Copyright © 2021 the Author(s). Published by PNAS.)

An unresolved issue in the vegetation ecology of the Indian subcontinent is whether its savannas, characterized by relatively open formations of deciduous trees in C 4 -grass dominated understories, are natural or anthropogenic. Historically, these ecosystems have widely been regarded as anthropogenic-derived, degraded descendants of deciduous forests. Despite recent work showing that modern savannas in the subcontinent fall within established bioclimatic envelopes of extant savannas elsewhere, the debate persists, at least in part because the regions where savannas occur also have a long history of human presence and habitat modification. Here we show for the first time, using multiple proxies for vegetation, climate and disturbances from high-resolution, well-dated lake sediments from Lonar Crater in peninsular India, that neither anthropogenic impact nor fire regime shifts, but monsoon weakening during the past ~ 6.0 kyr cal. BP, drove the expansion of savanna at the expense of forests in peninsular India. Our results provide unambiguous evidence for a climate-induced origin and spread of the modern savannas of peninsular India at around the mid-Holocene. We further propose that this savannization preceded and drove the introduction of agriculture and development of sedentism in this region, rather than vice-versa as has often been assumed.

Historical sources describe irrigation and intensive agriculture being practiced in lowland Yunnan from at least the first century AD, but so far archaeobotanical remains allowing investigation of this issue have been scarce. Here, we present new archaeobotanical evidence, including macro-botanical and phytoliths results, from the Dian settlement site of Dayingzhuang, with direct AMS radiocarbon dates on two wheat grains falling between 750 and 390 BC. We compare these results with contemporary Dian sites and analyse the agricultural systems in Central Yunnan between the eight and fourth centuries BC. We propose that agriculture was intensified toward the end of the Dian through both multiple cropping seasons and increased evidence for irrigated rice fields., Supplementary Information: The online version contains supplementary material available at 10.1007/s12520-020-01268-y., Competing Interests: Conflict of interestThe authors declare that they have no conflict of interest., (© The Author(s) 2021.)

Imprints of domesticated pearl millet ( Pennisetum glaucum (L.) R. Br.) spikelets, observed as temper in ceramics dating to the third millennium BC, provide the earliest evidence for the cultivation and domestication process of this crop in northern Mali. Additional sherds from the same region dating to the fifth and fourth millennium BC were examined and found to have pearl millet chaff with wild morphologies. In addition to studying sherds by stereomicroscopy and subjecting surface casts to scanning electron microscopy (SEM), we also deployed X-ray microcomputed tomography (microCT) on eleven sherds. This significantly augmented the total dataset of archaeological pearl millet chaff remains from which to document the use of the wild pearl millet as ceramic temper and the evolution of its morphology over time. Grain sizes were also estimated from spikelets preserved in the ceramics. Altogether, we are now able to chart the evolution of domesticated pearl millet in western Africa using three characteristics: the evolution of nonshattering stalked involucres; the appearance of multiple spikelet involucres, usually paired spikelets; and the increase in grain size. By the fourth millennium BC, average grain breadth had increased by 28%, although spikelet features otherwise resemble the wild type. In the third millennium BC, the average width of seeds is 38% greater than that of wild seeds, while other qualitative features of domestication are indicated by the presence of paired spikelets and the appearance of nondehiscent, stalked involucres. Nonshattering spikelets had probably become fixed by around 2000 BC, while increases in average grain size continued into the second millennium BC. These data now provide a robust sequence for the morphological evolution of domesticated pearl millet, the first indigenous crop domesticated in western Africa., Supplementary Information: The online version contains supplementary material available at 10.1007/s10437-021-09428-8., Competing Interests: Conflict of interestThe authors declare no conflict of interest or ethical concerns with this paper., (© The Author(s) 2021.)

While narratives of the spread of agriculture are central to interpretation of African history, hard evidence of past crops and cultivation practices are still few. This research aims at filling this gap and better understanding the evolution of agriculture and foodways in West Africa. It reports evidence from systematic flotation samples taken at the settlement mounds of Sadia (Mali), dating from 4 phases (phase 0=before first-third century AD; phase 1=mid eighth-tenth c. AD; phase 2=tenth-eleventh c. AD; phase 3=twelfth-late thirteenth c. AD). Flotation of 2200 l of soil provided plant macro-remains from 146 archaeological samples. As on most West African sites, the most dominant plant is pearl millet ( Pennisetum glaucum ). But from the tenth century AD, sorghum ( Sorghum bicolor ) and African rice ( Oryza glaberrima ) appear in small quantities, and fonio ( Digitaria exilis ) and barnyard millet/hungry rice ( Echinochloa sp.), sometimes considered weeds rather than staple crops, are found in large quantities. Some samples also show remains of tree fruits from savannah parklands, such as baobab ( Adansonia digitata ), marula ( Sclerocarya birrea ), jujube ( Ziziphus sp.), shea butter ( Vittelaria paradoxa ) and African grapes ( Lannea microcarpa ). Fonio and Echinochloa sp. cultivation appears here to be a later addition that helped to diversify agriculture and buffer against failures that might affect the monoculture of pearl millet. This diversification at the end of the 1st millennium AD matches with other evidence found in West Africa., (© The Author(s) 2021.)

Rice (Oryza sativa) is one of the world's most important food crops, and is comprised largely of japonica and indica subspecies. Here, we reconstruct the history of rice dispersal in Asia using whole-genome sequences of more than 1,400 landraces, coupled with geographic, environmental, archaeobotanical and paleoclimate data. Originating around 9,000 yr ago in the Yangtze Valley, rice diversified into temperate and tropical japonica rice during a global cooling event about 4,200 yr ago. Soon after, tropical japonica rice reached Southeast Asia, where it rapidly diversified, starting about 2,500 yr BP. The history of indica rice dispersal appears more complicated, moving into China around 2,000 yr BP. We also identify extrinsic factors that influence genome diversity, with temperature being a leading abiotic factor. Reconstructing the dispersal history of rice and its climatic correlates may help identify genetic adaptations associated with the spread of a key domesticated species.

Background: Vegetatively propagated crops are globally significant in terms of current agricultural production, as well as for understanding the long-term history of early agriculture and plant domestication. Today, significant field crops include sugarcane (Saccharum officinarum), potato (Solanum tuberosum), manioc (Manihot esculenta), bananas and plantains (Musa cvs), sweet potato (Ipomoea batatas), yams (Dioscorea spp.) and taro (Colocasia esculenta). In comparison with sexually reproduced crops, especially cereals and legumes, the domestication syndrome in vegetatively propagated field crops is poorly defined., Aims and Scope: Here, a range of phenotypic traits potentially comprising a syndrome associated with early domestication of vegetatively propagated field crops is proposed, including: mode of reproduction, yield of edible portion, ease of harvesting, defensive adaptations, timing of production and plant architecture. The archaeobotanical visibility of these syndrome traits is considered with a view to the reconstruction of the geographical and historical pathways of domestication for vegetatively propagated field crops in the past., Conclusions: Although convergent phenotypic traits are identified, none of them are ubiquitous and some are divergent. In contrast to cereals and legumes, several traits seem to represent varying degrees of plastic response to growth environment and practices of cultivation, as opposed to solely morphogenetic 'fixation'., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.)

The introduction of wheat into central China is thought to have been one of the significant contributions of interactions between China and Central Asia which began in the 3rd millennium bc. However, only a limited number of Neolithic wheat grains have been found in central China and even fewer have been directly radiocarbon dated, making the date when wheat was adopted in the region and its role in subsistence farming uncertain. Based on systematic archaeobotanical data and direct dating of wheat remains from the Xiazhai site in central China, as well as a critical review of all reported discoveries of Neolithic and Bronze Age wheat from this region, we conclude that many wheat finds are intrusive in Neolithic contexts. We argue that the role of wheat in the subsistence of the late Neolithic and early Bronze Age of central China was minimal, and that wheat only began to increase in its subsistence role in the later Bronze Age during the Zhou dynasty after ca. 1000 bc., (© The Author(s) 2019.)

The present paper reports the first systematic archaeobotanical evidence from Bangladesh together with direct AMS radiocarbon dates on crop remains. Macro-botanical remains were collected by flotation from two sites, Wari-Bateshwar (WB), an Early Historic archaeological site, dating mainly between 400 and 100 BC, with a later seventh century AD temple complex, and Raghurampura Vikrampura (RV), a Buddhist Monastery ( vihara ) located within the Vikrampura city site complex and dating to the eleventh and sixteenth centuries AD. Despite being a tropical country, with high rainfall and intensive soil processes, our work demonstrates that conventional archaeobotany, the collection of macro-remains through flotation, has much potential towards putting together a history of crops and agricultural systems in Bangladesh. The archaeobotanical assemblage collected from both sites indicates the predominance of rice agriculture, which would have been practiced in summer. Spikelet bases are of domesticated type rice, while grain metrics suggest the majority of rice was probably subspecies japonica. The presence of some wetland weeds suggests at least some of the rice was grown in wet (flooded) systems, but much of it may have been rainfed as inferred from the Southeast Asian weed Acmella paniculata . Other crops include winter cereals, barley and possible oat, and small numbers of summer millets ( Pennisetum glaucum , Sorghum bicolor , Setaria italica ), a wide diversity of summer and winter pulses (14 spp.), cotton, sesame and mustard seed. Pulse crops included many known from India. Thus, while most crops indicate diffusion of crops from India eastwards, the absence of indica rice could also indicate some diffusion from Southeast Asia. The later site RV also produced evidence of the rice bean ( Vigna umbellata ), a domesticate of mainland Southeast Asia. These data provide the first empirical evidence for reconstructing past agriculture in Bangladesh and for the role of connections to both India and mainland Southeast Asia in the development of crop diversity in the Ganges delta region., (© The Author(s) 2020.)

This paper presents new excavation data and new radiometric dates for Jebel Moya, south-central Sudan. These data suggest revisions to previous chronological understandings of the site. New excavations, initiated in 2017, show a longer, more continuous occupation of the site than has been previously recognised. Archaeozoological and archaeobotanical analyses provide evidence for domesticated taxa. Archaeobotanical evidence is dominated by domesticated sorghum ( Sorghum bicolor ), radiocarbon dated to c . 2550-2210 BC. Faunal remains include cattle and goat/sheep. A late third-millennium BC date on the human skeleton excavated in the 2017 season also shows that mortuary activity began early in the site's history, contemporary with domesticated faunal and botanical remains. These initial results indicate the long-term association of the site with pastoralism and agriculture and with environmental change. Jebel Moya's continued potential to serve as a chronological and cultural reference point for future studies in south-central Sudan and the eastern Sahel is reinforced.

Tetraploid emmer wheat (Triticum turgidum ssp. dicoccon) is a progenitor of the world's most widely grown crop, hexaploid bread wheat (Triticum aestivum), as well as the direct ancestor of tetraploid durum wheat (T. turgidum subsp. turgidum). Emmer was one of the first cereals to be domesticated in the old world; it was cultivated from around 9700 BC in the Levant 1,2 and subsequently in south-western Asia, northern Africa and Europe with the spread of Neolithic agriculture 3,4 . Here, we report a whole-genome sequence from a museum specimen of Egyptian emmer wheat chaff, 14 C dated to the New Kingdom, 1130-1000 BC. Its genome shares haplotypes with modern domesticated emmer at loci that are associated with shattering, seed size and germination, as well as within other putative domestication loci, suggesting that these traits share a common origin before the introduction of emmer to Egypt. Its genome is otherwise unusual, carrying haplotypes that are absent from modern emmer. Genetic similarity with modern Arabian and Indian emmer landraces connects ancient Egyptian emmer with early south-eastern dispersals, whereas inferred gene flow with wild emmer from the Southern Levant signals a later connection. Our results show the importance of museum collections as sources of genetic data to uncover the history and diversity of ancient cereals.

The reasons and processes that led hunter-gatherers to transition into a sedentary and agricultural way of life are a fundamental unresolved question of human history. Here we present results of excavations of two single-occupation early Neolithic sites (dated to 7.9 and 7.4 ka) and two high-resolution archaeological surveys in northeast China, which capture the earliest stages of sedentism and millet cultivation in the second oldest center of domestication in the Old World. The transition to sedentism coincided with a significant transition to wetter conditions in north China, at 8.1-7.9 ka. We suggest that these wetter conditions were an empirical precondition that facilitated the complex transitional process to sedentism and eventually millet domestication in north China. Interestingly, sedentism and plant domestication followed different trajectories. The sedentary way of life and cultural norms evolved rapidly, within a few hundred years, we find complex sedentary villages inhabiting the landscape. However, the process of plant domestication, progressed slowly over several millennia. Our earliest evidence for the beginning of the domestication process appear in the context of an already complex sedentary village (late Xinglongwa culture), a half millennia after the onset of cultivation, and even in this phase domesticated plants and animals were rare, suggesting that the transition to domesticated (sensu stricto) plants in affluent areas might have not played a substantial role in the transition to sedentary societies., Competing Interests: The authors have declared that no competing interests exist.

The evolution of domesticated cereals was a complex interaction of shifting selection pressures and repeated episodes of introgression. Genomes of archaeological crops have the potential to reveal these dynamics without being obscured by recent breeding or introgression. We report a temporal series of archaeogenomes of the crop sorghum (Sorghum bicolor) from a single locality in Egyptian Nubia. These data indicate no evidence for the effects of a domestication bottleneck, but instead reveal a steady decline in genetic diversity over time coupled with an accumulating mutation load. Dynamic selection pressures acted sequentially to shape architectural and nutritional domestication traits and to facilitate adaptation to the local environment. Later introgression between sorghum races allowed the exchange of adaptive traits and achieved mutual genomic rescue through an ameliorated mutation load. These results reveal a model of domestication in which genomic adaptation and deterioration were not focused on the initial stages of domestication but occurred throughout the history of cultivation.

Date palm ( Phoenix dactylifera L.) is a major fruit crop of arid regions that were domesticated ∼7,000 y ago in the Near or Middle East. This species is cultivated widely in the Middle East and North Africa, and previous population genetic studies have shown genetic differentiation between these regions. We investigated the evolutionary history of P. dactylifera and its wild relatives by resequencing the genomes of date palm varieties and five of its closest relatives. Our results indicate that the North African population has mixed ancestry with components from Middle Eastern P. dactylifera and Phoenix theophrasti , a wild relative endemic to the Eastern Mediterranean. Introgressive hybridization is supported by tests of admixture, reduced subdivision between North African date palm and P. theophrasti , sharing of haplotypes in introgressed regions, and a population model that incorporates gene flow between these populations. Analysis of ancestry proportions indicates that as much as 18% of the genome of North African varieties can be traced to P. theophrasti and a large percentage of loci in this population are segregating for single-nucleotide polymorphisms (SNPs) that are fixed in P. theophrasti and absent from date palm in the Middle East. We present a survey of Phoenix remains in the archaeobotanical record which supports a late arrival of date palm to North Africa. Our results suggest that hybridization with P. theophrasti was of central importance in the diversification history of the cultivated date palm., Competing Interests: The authors declare no conflict of interest., (Copyright © 2019 the Author(s). Published by PNAS.)

The transition to urbanism has long focused on annual staple crops (cereals and legumes), perhaps at the expense of understanding other changes within agricultural practices that occurred between the end of the initial domestication period and urbanisation. This paper examines the domestication and role of fruit tree crops within urbanisation in both Western Asia and China, using a combination of evidence for morphological change and a database that documents both the earliest occurrence of tree fruit crops and their spread beyond their wild range. In Western Asia the domestication of perennial fruit crops likely occurs between 6500 bc and 3500 bc, although it accompanies a shift in location from that of the earliest domestications within the Fertile Crescent to Mesopotamia, where the earliest urban societies arose. For China, fruit-tree domestication dates between ca 4000 and 2500 bc, commencing after millet domestication and rice domestication in Northern and Southern China, respectively, but within the period that led up to the urban societies that characterised the Longshan period in the Yellow River basin and the Liangzhu Culture in the Lower Yangtze. These results place the domestication of major fruit trees between the end of the domestication of staple annual crops and the rise of urbanism. On this basis it is argued that arboriculture played a fundamental role within the re-organisation of existing land use, shifting the emphasis from short-term returns of cereal crops into longer term investment in the developing agricultural landscape in both Western and East Asia. In this respect perennial tree crops can be placed alongside craft specialisation, such as metallurgy and textiles, in the formation of urban centres and the shaping the organisational administration that accompanied the rise of urbanism.

The origins of bread have long been associated with the emergence of agriculture and cereal domestication during the Neolithic in southwest Asia. In this study we analyze a total of 24 charred food remains from Shubayqa 1, a Natufian hunter-gatherer site located in northeastern Jordan and dated to 14.6-11.6 ka cal BP. Our finds provide empirical data to demonstrate that the preparation and consumption of bread-like products predated the emergence of agriculture by at least 4,000 years. The interdisciplinary analyses indicate the use of some of the "founder crops" of southwest Asian agriculture (e.g., Triticum boeoticum , wild einkorn) and root foods (e.g., Bolboschoenus glaucus , club-rush tubers) to produce flat bread-like products. The available archaeobotanical evidence for the Natufian period indicates that cereal exploitation was not common during this time, and it is most likely that cereal-based meals like bread become staples only when agriculture was firmly established., Competing Interests: The authors declare no conflict of interest., (Copyright © 2018 the Author(s). Published by PNAS.)

This paper asks whether we can identify a recurrent domestication syndrome for tree crops (fruits, nuts) and track archaeologically the evolution of domestication of fruits from woody perennials. While archaeobotany has made major contributions to documenting the domestication process in cereals and other annual grains, long-lived perennials have received less comparative attention. Drawing on examples from across Eurasia, comparisons suggest a tendency for the larger domesticated fruits to contain seeds that are proportionally longer, thinner and with more pointed (acute to attenuated) apices. Therefore, although changes in flavour, such as increased sweetness, are not recoverable, seed metrics and shape provide an archaeological basis for tracking domestication episodes in fruits from woody perennials. Where available, metrical data suggest length increases, as well as size diversification over time, with examples drawn from the Jomon of Japan ( Castanea crenata ), Neolithic China ( Prunus persica ) and the later Neolithic of the Near East ( Olea europaea, Phoenix dactylifera ) to estimate rates of change. More limited data allow us to also compare Mesoamerica avocado ( Persea americana ) and western Pacific Canarium sp. nuts and Spondias sp. fruits. Data from modern Indian jujube ( Ziziphus mauritiana ) are also considered in relation to seed length:width trends in relation to fruit contents (flesh proportion, sugar content). Despite the long generation time in tree fruits, rates of change in their seeds are generally comparable to rates of phenotypic evolution in annual grain crops, suggesting that gradual evolution via unconscious selection played a key role in initial processes of tree domestication, and that this had begun in the later Neolithic once annual crops had been domesticated, in both west and east Asia., (© The Author(s) 2017.)

Four decades have passed since Harlan and Stemler (1976) proposed the eastern Sahelian zone as the most likely center of Sorghum bicolor domestication. Recently, new data on seed impressions on Butana Group pottery, from the fourth millennium BC in the southern Atbai region of the far eastern Sahelian Belt in Africa, show evidence for cultivation activities of sorghum displaying some domestication traits. Pennisetum glaucum may have been undergoing domestication shortly thereafter in the western Sahel, as finds of fully domesticated pearl millet are present in southeastern Mali by the second half of the third millennium BC, and present in eastern Sudan by the early second millennium BC. The dispersal of the latter to India took less than 1000 years according to present data. Here, we review the middle Holocene Sudanese archaeological data for the first time, to situate the origins and spread of these two native summer rainfall cereals in what is proposed to be their eastern Sahelian Sudan gateway to the Red Sea and the Indian Ocean trade., Competing Interests: Compliance with Ethical StandardsThe authors declare that they have no conflict of interest.

To what degree is cultural multi-level selection responsible for the rise of environmentally transformative human behaviors? And vice versa? From the clearing of vegetation using fire to the emergence of agriculture and beyond, human societies have increasingly sustained themselves through practices that enhance environmental productivity through ecosystem engineering. At the same time, human societies have increased in scale and complexity from mobile bands of hunter-gatherers to telecoupled world systems. We propose that these long-term changes are coupled through positive feedbacks among social and environmental changes, coevolved primarily through selection acting at the group level and above, and that this can be tested by combining archeological evidence with mechanistic experiments using an agent-based virtual laboratory (ABVL) approach. A more robust understanding of whether and how cultural multi-level selection couples human social change with environmental transformation may help in addressing the long-term sustainability challenges of the Anthropocene.

Domestication is the process by which plants or animals evolved to fit a human-managed environment, and it is marked by innovations in plant morphology and anatomy that are in turn correlated with new human behaviours and technologies for harvesting, storage and field preparation. Archaeobotanical evidence has revealed that domestication was a protracted process taking thousands of plant generations. Within this protracted process there were changes in the selection pressures for domestication traits as well as variation across a geographic mosaic of wild and cultivated populations. Quantitative data allow us to estimate the changing selection coefficients for the evolution of non-shattering (domestic-type seed dispersal) in Asian rice ( Oryza sativa L.), barley ( Hordeum vulgare L.), emmer wheat ( Triticum dicoccon (Shrank) Schübl.) and einkorn wheat ( Triticum monococcum L.). These data indicate that selection coefficients tended to be low, but also that there were inflection points at which selection increased considerably. For rice, selection coefficients of the order of 0.001 prior to 5500 BC shifted to greater than 0.003 between 5000 and 4500 BC, before falling again as the domestication process ended 4000-3500 BC. In barley and the two wheats selection was strongest between 8500 and 7500 BC. The slow start of domestication may indicate that initial selection began in the Pleistocene glacial era.This article is part of the themed issue 'Process and pattern in innovations from cells to societies'., (© 2017 The Author(s).)

Reduction of seed dormancy mechanisms, allowing for rapid germination after planting, is a recurrent trait in domesticated plants, and can often be linked to changes in seed coat structure, in particular thinning. We report evidence for seed coat thinning between 2,000 BC and 1,200 BC, in southern Indian archaeological horsegram (Macrotyloma uniflorum), which it has been possible to document with high precision and non-destructively, through high resolution x-ray computed tomography using a synchrotron. We find that this trait underwent stepped change, from thick to semi-thin to thin seed coats, and that the rate of change was gradual. This is the first time that the rate of evolution of seed coat thinning in a legume crop has been directly documented from archaeological remains, and it contradicts previous predictions that legume domestication occurred through selection of pre-adapted low dormancy phenotypes from the wild.

The origin of domesticated Asian rice (Oryza sativa) has been a contentious topic, with conflicting evidence for either single or multiple domestication of this key crop species. We examined the evolutionary history of domesticated rice by analyzing de novo assembled genomes from domesticated rice and its wild progenitors. Our results indicate multiple origins, where each domesticated rice subpopulation (japonica, indica, and aus) arose separately from progenitor O. rufipogon and/or O. nivara. Coalescence-based modeling of demographic parameters estimate that the first domesticated rice population to split off from O. rufipogon was O. sativa ssp. japonica, occurring at ∼13.1-24.1 ka, which is an order of magnitude older then the earliest archeological date of domestication. This date is consistent, however, with the expansion of O. rufipogon populations after the Last Glacial Maximum ∼18 ka and archeological evidence for early wild rice management in China. We also show that there is significant gene flow from japonica to both indica (∼17%) and aus (∼15%), which led to the transfer of domestication alleles from early-domesticated japonica to proto-indica and proto-aus populations. Our results provide support for a model in which different rice subspecies had separate origins, but that de novo domestication occurred only once, in O. sativa ssp. japonica, and introgressive hybridization from early japonica to proto-indica and proto-aus led to domesticated indica and aus rice., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)

This paper presents an integrated methodology for the analysis of archaeological remains of cereal meals, based on scanning electronic microscopic analyses of microstructures of charred food fragments from Neolithic Çatalhöyük (Turkey). The remains of cereal foods as 'bread-like' or 'porridge-like' small charred lumps of various amalgamated plant materials are frequently recovered from Neolithic and later archaeological sites in southwest Asia and Europe. Cereal food remains have recently attracted interest because the identification of their plant contents, the forms of food that they represent and the methods used in their creation can provide unique information about ancient culinary traditions and routine food processing, preparation and cooking techniques. Here, we focus on three methodological aspects: (1) the analysis of their composition; (2) the analysis of their microstructure to determine preparation and cooking processes; (3) the comparison with experimental reference materials. Preliminary results are presented on the botanical composition and cooking processes represented by the charred cereal preparations found at Neolithic Çatalhöyük (Turkey), for example cereals processed into bread, dough and/or porridge.

The period from the late third millennium BC to the start of the first millennium AD witnesses the first steps towards food globalization in which a significant number of important crops and animals, independently domesticated within China, India, Africa and West Asia, traversed Central Asia greatly increasing Eurasian agricultural diversity. This paper utilizes an archaeobotanical database (AsCAD), to explore evidence for these crop translocations along southern and northern routes of interaction between east and west. To begin, crop translocations from the Near East across India and Central Asia are examined for wheat ( Triticum aestivum ) and barley ( Hordeum vulgare ) from the eighth to the second millennia BC when they reach China. The case of pulses and flax ( Linum usitatissimum ) that only complete this journey in Han times (206 BC-AD 220), often never fully adopted, is also addressed. The discussion then turns to the Chinese millets, Panicum miliaceum and Setaria italica , peaches ( Amygdalus persica ) and apricots ( Armeniaca vulgaris ), tracing their movement from the fifth millennium to the second millennium BC when the Panicum miliaceum reaches Europe and Setaria italica Northern India, with peaches and apricots present in Kashmir and Swat. Finally, the translocation of japonica rice from China to India that gave rise to indica rice is considered, possibly dating to the second millennium BC. The routes these crops travelled include those to the north via the Inner Asia Mountain Corridor, across Middle Asia, where there is good evidence for wheat, barley and the Chinese millets. The case for japonica rice, apricots and peaches is less clear, and the northern route is contrasted with that through northeast India, Tibet and west China. Not all these journeys were synchronous, and this paper highlights the selective long-distance transport of crops as an alternative to demic-diffusion of farmers with a defined crop package.

African rice (Oryza glaberrima Steud.) is a cereal crop species closely related to Asian rice (Oryza sativa L.) but was independently domesticated in West Africa ∼3,000 years ago. African rice is rarely grown outside sub-Saharan Africa but is of global interest because of its tolerance to abiotic stresses. Here we describe a map of 2.32 million SNPs of African rice from whole-genome resequencing of 93 landraces. Population genomic analysis shows a population bottleneck in this species that began ∼13,000-15,000 years ago with effective population size reaching its minimum value ∼3,500 years ago, suggesting a protracted period of population size reduction likely commencing with predomestication management and/or cultivation. Genome-wide association studies (GWAS) for six salt tolerance traits identify 11 significant loci, 4 of which are within ∼300 kb of genomic regions that possess signatures of positive selection, suggesting adaptive geographical divergence for salt tolerance in this species.

The Austronesian settlement of the remote island of Madagascar remains one of the great puzzles of Indo-Pacific prehistory. Although linguistic, ethnographic, and genetic evidence points clearly to a colonization of Madagascar by Austronesian language-speaking people from Island Southeast Asia, decades of archaeological research have failed to locate evidence for a Southeast Asian signature in the island's early material record. Here, we present new archaeobotanical data that show that Southeast Asian settlers brought Asian crops with them when they settled in Africa. These crops provide the first, to our knowledge, reliable archaeological window into the Southeast Asian colonization of Madagascar. They additionally suggest that initial Southeast Asian settlement in Africa was not limited to Madagascar, but also extended to the Comoros. Archaeobotanical data may support a model of indirect Austronesian colonization of Madagascar from the Comoros and/or elsewhere in eastern Africa.

The exhibition of increasingly intensive and complex niche construction behaviors through time is a key feature of human evolution, culminating in the advanced capacity for ecosystem engineering exhibited by Homo sapiens A crucial outcome of such behaviors has been the dramatic reshaping of the global biosphere, a transformation whose early origins are increasingly apparent from cumulative archaeological and paleoecological datasets. Such data suggest that, by the Late Pleistocene, humans had begun to engage in activities that have led to alterations in the distributions of a vast array of species across most, if not all, taxonomic groups. Changes to biodiversity have included extinctions, extirpations, and shifts in species composition, diversity, and community structure. We outline key examples of these changes, highlighting findings from the study of new datasets, like ancient DNA (aDNA), stable isotopes, and microfossils, as well as the application of new statistical and computational methods to datasets that have accumulated significantly in recent decades. We focus on four major phases that witnessed broad anthropogenic alterations to biodiversity-the Late Pleistocene global human expansion, the Neolithic spread of agriculture, the era of island colonization, and the emergence of early urbanized societies and commercial networks. Archaeological evidence documents millennia of anthropogenic transformations that have created novel ecosystems around the world. This record has implications for ecological and evolutionary research, conservation strategies, and the maintenance of ecosystem services, pointing to a significant need for broader cross-disciplinary engagement between archaeology and the biological and environmental sciences.

Current debate concerns the pace at which domesticated plants emerged from cultivated wild populations and how many genes were involved. Using an individual-based model, based on the assumptions of Haldane and Maynard Smith, respectively, we estimate that a surprisingly low number of 50-100 loci are the most that could be under selection in a cultivation regime at the selection strengths observed in the archaeological record. This finding is robust to attempts to rescue populations from extinction through selection from high standing genetic variation, gene flow, and the Maynard Smith-based model of threshold selection. Selective sweeps come at a cost, reducing the capacity of plants to adapt to new environments, which may contribute to the explanation of why selective sweeps have not been detected more frequently and why expansion of the agrarian package during the Neolithic was so frequently associated with collapse.

Phytoliths and biomolecular components extracted from ancient plant remains from Chang'an (Xi'an, the city where the Silk Road begins) and Ngari (Ali) in western Tibet, China, show that the tea was grown 2100 years ago to cater for the drinking habits of the Western Han Dynasty (207BCE-9CE), and then carried toward central Asia by ca.200CE, several hundred years earlier than previously recorded. The earliest physical evidence of tea from both the Chang'an and Ngari regions suggests that a branch of the Silk Road across the Tibetan Plateau, was established by the second to third century CE.

Rice (Oryza sativa) is regarded as the only grass that was selected for cultivation and eventual domestication in the Yangtze basin of China. Although both macro-fossils and micro-fossils of rice have been recovered from the Early Neolithic site of Shangshan, dating to more than 10,000 years before present (BP), we report evidence of phytolith and starch microfossils taken from stone tools, both for grinding and cutting, and cultural layers, that indicating barnyard grass (Echinochloa spp.) was a major subsistence resource, alongside smaller quantities of acorn starches (Lithocarpus/Quercus sensu lato) and water chestnuts (Trapa). This evidence suggests that early managed wetland environments were initially harvested for multiple grain species including barnyard grasses as well as rice, and indicate that the emergence of rice as the favoured cultivated grass and ultimately the key domesticate of the Yangtze basin was a protracted process.

Baligang is a Neolithic site on a northern tributary of the middle Yangtze and provides a long archaeobotanical sequence from the Seventh Millennium BC upto the First Millennium BC. It provides evidence for developments in rice and millet agriculture influenced by shifting cultural affiliation with the north (Yangshao and Longshan) and south (Qujialing and Shijiahe) between 4300 and 1800 BC. This paper reports on plant macro-remains (seeds), from systematic flotation of 123 samples (1700 litres), producing more than 10,000 identifiable remains. The earliest Pre-Yangshao occupation of the sites provide evidence for cultivation of rice (Oryza sativa) between 6300-6700 BC. This rice appears already domesticated in on the basis of a dominance of non-shattering spikelet bases. However, in terms of grain size changes has not yet finished, as grains are still thinner than more recent domesaticated rice and are closer in grain shape to wild rices. This early rice was cultivated alongside collection of wild staple foods, especially acorns (Quercus/Lithicarpus sensu lato). In later periods the sites has evidence for mixed farming of both rice and millets (Setaria italica and Panicum miliaceum). Soybean appears on the site in the Shijiahe period (ca.2500 BC) and wheat (Triticum cf. aestivum) in the Late Longshan levels (2200-1800 BC). Weed flora suggests an intensification of rice agriculture over time with increasing evidence of wetland weeds. We interpret these data as indicating early opportunistic cultivation of alluvial floodplains and some rainfed rice, developing into more systematic and probably irrigated cultivation starting in the Yangshao period, which intensified in the Qujialing and Shijiahe period, before a shift back to an emphasis on millets with the Late Longshan cultural influence from the north.

We have compiled an extensive database of archaeological evidence for rice across Asia, including 400 sites from mainland East Asia, Southeast Asia and South Asia. This dataset is used to compare several models for the geographical origins of rice cultivation and infer the most likely region(s) for its origins and subsequent outward diffusion. The approach is based on regression modelling wherein goodness of fit is obtained from power law quantile regressions of the archaeologically inferred age versus a least-cost distance from the putative origin(s). The Fast Marching method is used to estimate the least-cost distances based on simple geographical features. The origin region that best fits the archaeobotanical data is also compared to other hypothetical geographical origins derived from the literature, including from genetics, archaeology and historical linguistics. The model that best fits all available archaeological evidence is a dual origin model with two centres for the cultivation and dispersal of rice focused on the Middle Yangtze and the Lower Yangtze valleys.

Recent increases in archaeobotanical evidence offer insights into the processes of plant domestication and agricultural origins, which evolved in parallel in several world regions. Many different crop species underwent convergent evolution and acquired domestication syndrome traits. For a growing number of seed crop species, these traits can be quantified by proxy from archaeological evidence, providing measures of the rates of change during domestication. Among domestication traits, nonshattering cereal ears evolved more quickly in general than seed size. Nevertheless, most domestication traits show similarly slow rates of phenotypic change over several centuries to millennia, and these rates were similar across different regions of origin. Crops reproduced vegetatively, including tubers and many fruit trees, are less easily documented in terms of morphological domestication, but multiple lines of evidence outline some patterns in the development of vegecultural systems across the New World and Old World tropics. Pathways to plant domestication can also be compared in terms of the cultural and economic factors occurring at the start of the process. Whereas agricultural societies have tended to converge on higher population densities and sedentism, in some instances cultivation began among sedentary hunter-gatherers whereas more often it was initiated by mobile societies of hunter-gatherers or herder-gatherers.