The biocultural origins and dispersal of domestic chickens.

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.

A stepwise route to domesticate rice by controlling seed shattering and panicle shape.

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.

People have shaped most of terrestrial nature for at least 12,000 years.

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.

The Evolutionary History of Wild, Domesticated, and Feral Brassica oleracea (Brassicaceae).

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.

A novel cost framework reveals evidence for competitive selection in the evolution of complex traits during plant domestication.

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.

Cross-species hybridization and the origin of North African date palms.

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.

Archaeobotanical evidence reveals the origins of bread 14,400 years ago in northeastern Jordan.

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.

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.

Ancient crops provide first archaeological signature of the westward Austronesian expansion.

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 Rice Paradox: Multiple Origins but Single Domestication in Asian Rice.

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.

On the Origins and Dissemination of Domesticated Sorghum and Pearl Millet across Africa and into India: a View from the Butana Group of the Far Eastern Sahel.

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.

Quatre décennies se sont écoulées depuis que Harlan et Stemler ont proposé la zone sahélienne orientale comme le centre le plus probable de la domestication du sorgho bicolore. Récemment, de nouvelles données sur les impressions de semences sur les poteries du groupe Butana du IVe millénaire avant JC dans la région sud d'Atbai dans la ceinture sahélienne d'Afrique orientale montrent des preuves d'activités de culture du sorgho présentant certains traits de domestication. Pennisetum glaucum pourrait être en cours de domestication peu après dans l'ouest du Sahel, puisque le millet perlé entièrement domestiqué est. présent dans le sud-est du Mali vers la seconde moitié du troisième millénaire avant J.-C. et présent dans l'est du Soudan au début du deuxième millénaire avant notre ère. La dispersion de ce dernier en Inde a pris moins de mille ans selon les données actuelles. Nous examinons ici pour la première fois les données archéologiques soudanaises de l'Holocène moyen pour situer les origines et la propagation de ces deux céréales de pluie estivales indigènes dans ce qui est. proposé comme leur porte d'entrée soudanienne du Sahel oriental à la mer Rouge et à l'océan Indien.

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.

Storytelling and story testing in domestication.

The domestication of plants and animals marks one of the most significant transitions in human, and indeed global, history. Traditionally, study of the domestication process was the exclusive domain of archaeologists and agricultural scientists; today it is an increasingly multidisciplinary enterprise that has come to involve the skills of evolutionary biologists and geneticists. Although the application of new information sources and methodologies has dramatically transformed our ability to study and understand domestication, it has also generated increasingly large and complex datasets, the interpretation of which is not straightforward. In particular, challenges of equifinality, evolutionary variance, and emergence of unexpected or counter-intuitive patterns all face researchers attempting to infer past processes directly from patterns in data. We argue that explicit modeling approaches, drawing upon emerging methodologies in statistics and population genetics, provide a powerful means of addressing these limitations. Modeling also offers an approach to analyzing datasets that avoids conclusions steered by implicit biases, and makes possible the formal integration of different data types. Here we outline some of the modeling approaches most relevant to current problems in domestication research, and demonstrate the ways in which simulation modeling is beginning to reshape our understanding of the domestication process.

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.

Used planet: a global history.

Human use of land has transformed ecosystem pattern and process across most of the terrestrial biosphere, a global change often described as historically recent and potentially catastrophic for both humanity and the biosphere. Interdisciplinary paleoecological, archaeological, and historical studies challenge this view, indicating that land use has been extensive and sustained for millennia in some regions and that recent trends may represent as much a recovery as an acceleration. Here we synthesize recent scientific evidence and theory on the emergence, history, and future of land use as a process transforming the Earth System and use this to explain why relatively small human populations likely caused widespread and profound ecological changes more than 3,000 y ago, whereas the largest and wealthiest human populations in history are using less arable land per person every decade. Contrasting two spatially explicit global reconstructions of land-use history shows that reconstructions incorporating adaptive changes in land-use systems over time, including land-use intensification, offer a more spatially detailed and plausible assessment of our planet's history, with a biosphere and perhaps even climate long ago affected by humans. Although land-use processes are now shifting rapidly from historical patterns in both type and scale, integrative global land-use models that incorporate dynamic adaptations in human-environment relationships help to advance our understanding of both past and future land-use changes, including their sustainability and potential global effects.

The nature of selection during plant domestication.

Plant domestication is an outstanding example of plant-animal co-evolution and is a far richer model for studying evolution than is generally appreciated. There have been numerous studies to identify genes associated with domestication, and archaeological work has provided a clear understanding of the dynamics of human cultivation practices during the Neolithic period. Together, these have provided a better understanding of the selective pressures that accompany crop domestication, and they demonstrate that a synthesis from the twin vantage points of genetics and archaeology can expand our understanding of the nature of evolutionary selection that accompanies domestication.

Fluvial landscapes of the Harappan civilization.

The collapse of the Bronze Age Harappan, one of the earliest urban civilizations, remains an enigma. Urbanism flourished in the western region of the Indo-Gangetic Plain for approximately 600 y, but since approximately 3,900 y ago, the total settled area and settlement sizes declined, many sites were abandoned, and a significant shift in site numbers and density towards the east is recorded. We report morphologic and chronologic evidence indicating that fluvial landscapes in Harappan territory became remarkably stable during the late Holocene as aridification intensified in the region after approximately 5,000 BP. Upstream on the alluvial plain, the large Himalayan rivers in Punjab stopped incising, while downstream, sedimentation slowed on the distinctive mega-fluvial ridge, which the Indus built in Sindh. This fluvial quiescence suggests a gradual decrease in flood intensity that probably stimulated intensive agriculture initially and encouraged urbanization around 4,500 BP. However, further decline in monsoon precipitation led to conditions adverse to both inundation- and rain-based farming. Contrary to earlier assumptions that a large glacier-fed Himalayan river, identified by some with the mythical Sarasvati, watered the Harappan heartland on the interfluve between the Indus and Ganges basins, we show that only monsoonal-fed rivers were active there during the Holocene. As the monsoon weakened, monsoonal rivers gradually dried or became seasonal, affecting habitability along their courses. Hydroclimatic stress increased the vulnerability of agricultural production supporting Harappan urbanism, leading to settlement downsizing, diversification of crops, and a drastic increase in settlements in the moister monsoon regions of the upper Punjab, Haryana, and Uttar Pradesh.