Looking beyond history: tracing the dispersal of the Malaysian complex of crops to Africa.

In his 1959 book, Africa: Its Peoples and Their Culture History, George P. Murdock suggested that a Malaysian complex of crops dispersed to Africa in ancient times across the Indian Ocean along the Sabaean Lane. The Malaysian complex comprised bananas, sugarcane, taro, three yam species, rice, Polynesian arrowroot, breadfruit, coconut, areca palm, and betel leaf. Except for rice, arrowroot, and potentially taro, most of these crops were domesticated in the Island Southeast Asia-New Guinea region, from where they dispersed to Africa. Our reassessment of agronomic, archaeological, classical, genetic, and historical sources shows that we need to go beneath standard historical narratives to recover a much more ancient and complex history of crop introductions to Africa. Despite considerable uncertainty and fragmented research, we were able to conclude that the Malaysian complex of crops did not arrive in Africa as a complete assemblage at one time or along one route. Multiple lines of evidence suggest that these crops arrived in Africa at different times and followed different pathways of introduction to the continent.

Transition From Wild to Domesticated Pearl Millet (Pennisetum glaucum) Revealed in Ceramic Temper at Three Middle Holocene Sites in Northern Mali.

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.

Des empreintes d'épillets de mil domestiqué (Pennisetum glaucum (L.) R. Br.) observées dans des céramiques datées du 3e millénaire av. J.-C. provenant du nord du Mali constituent les plus anciens témoins de la mise en culture et de la domestication de cette céréale. Des tessons supplémentaires issus de la même région se rapportant aux 5e et 4e millénaires av. J.-C. ont été examinés et ont révélé des empreintes de balle de mil de morphologies sauvages. En plus de l'observation de leurs surfaces par stéréo-microscopie, et de l'observation des moulages d'empreintes au microscope à balayage, onze d'entre eux ont fait l'objet des micro-tomographies aux rayons X (microCT). Ces examens enrichissent considérablement l'ensemble des données archéologiques concernant l'utilisation du mil sauvage comme dégraissant végétal et son évolution morphologique à travers le temps. La taille des grains a aussi été estimée à partir des épillets conservés dans la céramique. En tenant compte des données enregistrées lors d'études antérieures, nous pouvons désormais retracer l'évolution du mil domestiqué en Afrique de l'Ouest à travers trois caractéristiques : l'évolution des involucres pédonculés à égrenage non-spontané; l'apparition d'involucres multiples par épillets, des épillets appariés le plus souvent; l'augmentation de la taille des grains au vu de leur largeur. Déjà au 4e millénaire avant J.-C., la largeur moyenne des grains a augmenté de 28% bien que les caractéristiques de l'épillet ressemblent au type sauvage. Au 3e millénaire avant J.-C., elle est 38% supérieure à celle du morphotype sauvage, tandis que des caractéristiques qualitatives de la domestication sont avérées par la présence d'épillets appariés et par celle d'involucres pédonculés à égrenage non-spontané. La non-déhiscence des épillets est un caractéristique de la domesticité qui s'est probablement fixé vers 2000 avant J.-C., tandis que l'augmentation de la taille moyenne des grains s'est poursuivie tout au long du 2e millénaire av. J.-C. Ces données fournissent désormais une séquence robuste concernant l'évolution morphologique du mil, la première céréale indigène domestiquée en Afrique de l'Ouest.

The domestication syndrome in vegetatively propagated field crops.

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'.

Ecological consequences of human niche construction: Examining long-term anthropogenic shaping of global species distributions.

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.

Evaluating the roles of directed breeding and gene flow in animal domestication.

For the last 150 y scholars have focused upon the roles of intentional breeding and genetic isolation as fundamental to understanding the process of animal domestication. This analysis of ethnoarchaeological, archaeological, and genetic data suggests that long-term gene flow between wild and domestic stocks was much more common than previously assumed, and that selective breeding of females was largely absent during the early phases of animal domestication. These findings challenge assumptions about severe genetic bottlenecks during domestication, expectations regarding monophyletic origins, and interpretations of multiple domestications. The findings also raise new questions regarding ways in which behavioral and phenotypic domestication traits were developed and maintained.

Convergent evolution and parallelism in plant domestication revealed by an expanding archaeological record.

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.

Using ancient DNA to study the origins and dispersal of ancestral Polynesian chickens across the Pacific.

The human colonization of Remote Oceania remains one of the great feats of exploration in history, proceeding east from Asia across the vast expanse of the Pacific Ocean. Human commensal and domesticated species were widely transported as part of this diaspora, possibly as far as South America. We sequenced mitochondrial control region DNA from 122 modern and 22 ancient chicken specimens from Polynesia and Island Southeast Asia and used these together with Bayesian modeling methods to examine the human dispersal of chickens across this area. We show that specific techniques are essential to remove contaminating modern DNA from experiments, which appear to have impacted previous studies of Pacific chickens. In contrast to previous reports, we find that all ancient specimens and a high proportion of the modern chickens possess a group of unique, closely related haplotypes found only in the Pacific. This group of haplotypes appears to represent the authentic founding mitochondrial DNA chicken lineages transported across the Pacific, and allows the early dispersal of chickens across Micronesia and Polynesia to be modeled. Importantly, chickens carrying this genetic signature persist on several Pacific islands at high frequencies, suggesting that the original Polynesian chicken lineages may still survive. No early South American chicken samples have been detected with the diagnostic Polynesian mtDNA haplotypes, arguing against reports that chickens provide evidence of Polynesian contact with pre-European South America. Two modern specimens from the Philippines carry haplotypes similar to the ancient Pacific samples, providing clues about a potential homeland for the Polynesian chicken.

Current perspectives and the future of domestication studies.

It is difficult to overstate the cultural and biological impacts that the domestication of plants and animals has had on our species. Fundamental questions regarding where, when, and how many times domestication took place have been of primary interest within a wide range of academic disciplines. Within the last two decades, the advent of new archaeological and genetic techniques has revolutionized our understanding of the pattern and process of domestication and agricultural origins that led to our modern way of life. In the spring of 2011, 25 scholars with a central interest in domestication representing the fields of genetics, archaeobotany, zooarchaeology, geoarchaeology, and archaeology met at the National Evolutionary Synthesis Center to discuss recent domestication research progress and identify challenges for the future. In this introduction to the resulting Special Feature, we present the state of the art in the field by discussing what is known about the spatial and temporal patterns of domestication, and controversies surrounding the speed, intentionality, and evolutionary aspects of the domestication process. We then highlight three key challenges for future research. We conclude by arguing that although recent progress has been impressive, the next decade will yield even more substantial insights not only into how domestication took place, but also when and where it did, and where and why it did not.

Ancient voyaging and Polynesian origins.

The "Polynesian motif" defines a lineage of human mtDNA that is restricted to Austronesian-speaking populations and is almost fixed in Polynesians. It is widely thought to support a rapid dispersal of maternal lineages from Taiwan ~4000 years ago (4 ka), but the chronological resolution of existing control-region data is poor, and an East Indonesian origin has also been proposed. By analyzing 157 complete mtDNA genomes, we show that the motif itself most likely originated >6 ka in the vicinity of the Bismarck Archipelago, and its immediate ancestor is >8 ka old and virtually restricted to Near Oceania. This indicates that Polynesian maternal lineages from Island Southeast Asia gained a foothold in Near Oceania much earlier than dispersal from either Taiwan or Indonesia 3-4 ka would predict. However, we find evidence in minor lineages for more recent two-way maternal gene flow between Island Southeast Asia and Near Oceania, likely reflecting movements along a "voyaging corridor" between them, as previously proposed on archaeological grounds. Small-scale mid-Holocene movements from Island Southeast Asia likely transmitted Austronesian languages to the long-established Southeast Asian colonies in the Bismarcks carrying the Polynesian motif, perhaps also providing the impetus for the expansion into Polynesia.

Multidisciplinary perspectives on banana (Musa spp.) domestication.

Original multidisciplinary research hereby clarifies the complex geodomestication pathways that generated the vast range of banana cultivars (cvs). Genetic analyses identify the wild ancestors of modern-day cvs and elucidate several key stages of domestication for different cv groups. Archaeology and linguistics shed light on the historical roles of people in the movement and cultivation of bananas from New Guinea to West Africa during the Holocene. The historical reconstruction of domestication processes is essential for breeding programs seeking to diversify and improve banana cvs for the future.

Plant domestication and agricultural ecologies.

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.

Multidisciplinary evidence for early banana (Musa cvs.) cultivation on Mabuyag Island, Torres Strait.

Multiproxy archaeobotanical analyses (starch granule, phytolith and microcharcoal) of an abandoned agricultural terrace at Wagadagam on Mabuyag Island, Torres Strait, Australia, document extensive, low-intensity forms of plant management from at least 2,145-1,930 cal yr BP and intensive forms of cultivation at 1,376-1,293 cal yr BP. The agricultural activities at 1,376-1,293 cal yr BP are evidenced from terrace construction, banana (Musa cultivars) cultivation and dramatic transformations to the local palaeoenvironment. The robust evidence for the antiquity of horticulture in western Torres Strait provides an historical basis for understanding the diffusion of cultivation practices and cultivars, most likely from New Guinea. This study also provides a methodological template for the investigation of plant management, potentially including forms of cultivation that were practiced in northern Australia before European colonization.

Contemporaneity of Australopithecus, Paranthropus, and early Homo erectus in South Africa.

Understanding the extinction of Australopithecus and origins of Paranthropus and Homo in South Africa has been hampered by the perceived complex geological context of hominin fossils, poor chronological resolution, and a lack of well-preserved early Homo specimens. We describe, date, and contextualize the discovery of two hominin crania from Drimolen Main Quarry in South Africa. At ~2.04 million to 1.95 million years old, DNH 152 represents the earliest definitive occurrence of Paranthropus robustus, and DNH 134 represents the earliest occurrence of a cranium with clear affinities to Homo erectus These crania also show that Homo, Paranthropus, and Australopithecus were contemporaneous at ~2 million years ago. This high taxonomic diversity is also reflected in non-hominin species and provides evidence of endemic evolution and dispersal during a period of climatic variability.

Archaeological assessment reveals Earth's early transformation through land use.

Environmentally transformative human use of land accelerated with the emergence of agriculture, but the extent, trajectory, and implications of these early changes are not well understood. An empirical global assessment of land use from 10,000 years before the present (yr B.P.) to 1850 CE reveals a planet largely transformed by hunter-gatherers, farmers, and pastoralists by 3000 years ago, considerably earlier than the dates in the land-use reconstructions commonly used by Earth scientists. Synthesis of knowledge contributed by more than 250 archaeologists highlighted gaps in archaeological expertise and data quality, which peaked for 2000 yr B.P. and in traditionally studied and wealthier regions. Archaeological reconstruction of global land-use history illuminates the deep roots of Earth's transformation and challenges the emerging Anthropocene paradigm that large-scale anthropogenic global environmental change is mostly a recent phenomenon.

MicroCT reveals domesticated rice (Oryza sativa) within pottery sherds from early Neolithic sites (4150-3265 cal BP) in Southeast Asia.

Rice (Oryza sativa) was domesticated in the Yangtze Valley region at least 6000-8000 years ago, yet the timing of dispersal of domesticated rice to Southeast Asia is contentious. Often rice is not well-preserved in archaeobotanical assemblages at early Neolithic sites in the wet tropics of Southeast Asia and consequently rice impressions in pottery have been used as a proxy for rice cultivation despite their uncertain taxonomic and domestication status. In this research, we use microCT technology to determine the 3D microscale morphology of rice husk and spikelet base inclusions within pottery sherds from early Neolithic sites in Vietnam. In contrast to surface impressions, microCT provides images of the entire husk and spikelet base preserved within the pottery, including the abscission scar characteristic of domesticated rice. This research demonstrates the potential of microCT to be a new, non-destructive method for the identification of domesticated plant remains within pottery sherds, especially in contexts where archaeobotanical preservation is poor and chaff-tempered sherds are rare and unavailable for destructive analysis. The method has the potential to greatly advance the understanding of crop domestication and agricultural dispersal for ceramic cultures in different parts of the world.