Pre-contact and post-colonial ecological legacies shape Surinamese rainforests.

Disturbances in tropical forests can have long-lasting ecological impacts, but their manifestations (ecological legacies) in modern forests are uncertain. Many Amazonian forests bear the mark of past soil modifications, species enrichments, and fire events, but the trajectories of ecological legacies from the pre-contact or post-colonial period remain relatively unexplored. We assessed the fire and vegetation history from 15 soil cores ranging from 0 to 10 km from a post-colonial Surinamese archaeological site. We show that (1) fires occurred from 96 bc to recent times and induced significant vegetation change, (2) persistent ecological legacies from pre-contact and post-colonial fire and deforestation practices were mainly within 1 km of the archaeological site, and (3) palm enrichment of Attalea, Oenocarpus and Astrocaryum occurred within 0, 1, and 8 km of the archaeological site, respectively. Our results challenge the notion of spatially extensive and persistent ecological legacies. Instead, our data indicate that the persistence and extent of ecological legacies are dependent on their timing, frequency, type, and intensity. Examining the mechanisms and manifestations of ecological legacies is crucial in assessing forest resilience and Indigenous and local land rights in the highly threatened Amazonian forests.

Long-term fire and vegetation change in northwestern Amazonia.

Amazonian forest plots are used to quantify biodiversity and carbon sequestration, and provide the foundation for much of what is known about tropical ecology. Many plots are assumed to be undisturbed, but recent work suggests that past fire, forest openings, and cultivation created vegetation changes that have persisted for decades to centuries (ecological legacies). The Yasuní Forest Dynamics plot is one of the most biodiverse places on earth, yet its human history remains unknown. Here, we use charcoal and phytolith analysis to investigate the fire and vegetation history of the Yasuní forest plot, and compare results with nearby forest plots in Colombia (Amacayacu) and Peru (Medio Putumayo-Algodón [MPA]) to explore the spatial variability of past disturbances and ecological legacies in northwestern Amazonia. Three 14C dated charcoal fragments provided evidence for a modern (1956 CE) and a past fire event ca. 750 years ago at Yasuní, compared with fire ages of 1000-1600 years ago documented at Amacayacu and MPA. Small-scale disturbances and localized canopy openings also occurred in the Yasuní plot. Phytolith assemblages from Yasuní and Amacayacu showed more variability in past vegetation change than MPA. Low-intensity, non-continuous disturbances occurred at all three plots in the past, and our results highlight the variability of past human activities both in space and time in northwestern Amazonia. Our data also suggest that post-Columbian human disturbances from the Rubber Boom (AD 1850-1920) and subsequent oil exploration have likely left stronger ecological legacies than those left by pre-Columbian peoples in our studied regions.

A 5,000-year vegetation and fire history for tierra firme forests in the Medio Putumayo-Algodón watersheds, northeastern Peru.

This paper addresses an important debate in Amazonian studies; namely, the scale, intensity, and nature of human modification of the forests in prehistory. Phytolith and charcoal analysis of terrestrial soils underneath mature tierra firme (nonflooded, nonriverine) forests in the remote Medio Putumayo-Algodón watersheds, northeastern Peru, provide a vegetation and fire history spanning at least the past 5,000 y. A tree inventory carried out in the region enables calibration of ancient phytolith records with standing vegetation and estimates of palm species densities on the landscape through time. Phytolith records show no evidence for forest clearing or agriculture with major annual seed and root crops. Frequencies of important economic palms such as Oenocarpus, Euterpe, Bactris, and Astrocaryum spp., some of which contain hyperdominant species in the modern flora, do not increase through prehistoric time. This indicates pre-Columbian occupations, if documented in the region with future research, did not significantly increase the abundance of those species through management or cultivation. Phytoliths from other arboreal and woody species similarly reflect a stable forest structure and diversity throughout the records. Charcoal 14C dates evidence local forest burning between ca. 2,800 and 1,400 y ago. Our data support previous research indicating that considerable areas of some Amazonian tierra firme forests were not significantly impacted by human activities during the prehistoric era. Rather, it appears that over the last 5,000 y, indigenous populations in this region coexisted with, and helped maintain, large expanses of relatively unmodified forest, as they continue to do today.

Comment on "Persistent effects of pre-Columbian plant domestication on Amazonian forest composition".

Levis et al (Research Articles, 3 March 2017, p. 925) concluded that pre-Columbian tree domestication has shaped present-day Amazonian forest composition. The study, however, downplays five centuries of human influence following European arrival to the Americas. We show that the effects of post-Columbian activities in Amazonia are likely to have played a larger role than pre-Columbian ones in shaping the observed floristic patterns.

Assessing elements of an extended evolutionary synthesis for plant domestication and agricultural origin research.

The development of agricultural societies, one of the most transformative events in human and ecological history, was made possible by plant and animal domestication. Plant domestication began 12,000-10,000 y ago in a number of major world areas, including the New World tropics, Southwest Asia, and China, during a period of profound global environmental perturbations as the Pleistocene epoch ended and transitioned into the Holocene. Domestication is at its heart an evolutionary process, and for many prehistorians evolutionary theory has been foundational in investigating agricultural origins. Similarly, geneticists working largely with modern crops and their living wild progenitors have documented some of the mechanisms that underwrote phenotypic transformations from wild to domesticated species. Ever-improving analytic methods for retrieval of empirical data from archaeological sites, together with advances in genetic, genomic, epigenetic, and experimental research on living crop plants and wild progenitors, suggest that three fields of study currently little applied to plant domestication processes may be necessary to understand these transformations across a range of species important in early prehistoric agriculture. These fields are phenotypic (developmental) plasticity, niche construction theory, and epigenetics with transgenerational epigenetic inheritance. All are central in a controversy about whether an Extended Evolutionary Synthesis is needed to reconceptualize how evolutionary change occurs. An exploration of their present and potential utility in domestication study shows that all three fields have considerable promise in elucidating important issues in plant domestication and in agricultural origin and dispersal research and should be increasingly applied to these issues.

Particularism and the retreat from theory in the archaeology of agricultural origins.

The introduction of new analytic methods and expansion of research into previously untapped regions have greatly increased the scale and resolution of data relevant to the origins of agriculture (OA). As a result, the recognition of varied historical pathways to agriculture and the continuum of management strategies have complicated the search for general explanations for the transition to food production. In this environment, higher-level theoretical frameworks are sometimes rejected on the grounds that they force conclusions that are incompatible with real-world variability. Some of those who take this position argue instead that OA should be explained in terms of local and historically contingent factors. This retreat from theory in favor of particularism is based on the faulty beliefs that complex phenomena such as agricultural origins demand equally complex explanations and that explanation is possible in the absence of theoretically based assumptions. The same scholars who are suspicious of generalization are reluctant to embrace evolutionary approaches to human behavior on the grounds that they are ahistorical, overly simplistic, and dismissive of agency and intent. We argue that these criticisms are misplaced and explain why a coherent theory of human behavior that acknowledges its evolutionary history is essential to advancing understanding of OA. Continued progress depends on the integration of human behavior and culture into the emerging synthesis of evolutionary developmental biology that informs contemporary research into plant and animal domestication.

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.

Plant foods and the dietary ecology of Neanderthals and early modern humans.

One of the most important challenges in anthropology is understanding the disappearance of Neanderthals. Previous research suggests that Neanderthals had a narrower diet than early modern humans, in part because they lacked various social and technological advances that lead to greater dietary variety, such as a sexual division of labor and the use of complex projectile weapons. The wider diet of early modern humans would have provided more calories and nutrients, increasing fertility, decreasing mortality and supporting large population sizes, allowing them to out-compete Neanderthals. However, this model for Neanderthal dietary behavior is based on analysis of animal remains, stable isotopes, and other methods that provide evidence only of animal food in the diet. This model does not take into account the potential role of plant food. Here we present results from the first broad comparison of plant foods in the diets of Neanderthals and early modern humans from several populations in Europe, the Near East, and Africa. Our data comes from the analysis of plant microremains (starch grains and phytoliths) in dental calculus and on stone tools. Our results suggest that both species consumed a similarly wide array of plant foods, including foods that are often considered low-ranked, like underground storage organs and grass seeds. Plants were consumed across the entire range of individuals and sites we examined, and none of the expected predictors of variation (species, geographic region, or associated stone tool technology) had a strong influence on the number of plant species consumed. Our data suggest that Neanderthal dietary ecology was more complex than previously thought. This implies that the relationship between Neanderthal technology, social behavior, and food acquisition strategies must be better explored.

Preceramic maize from Paredones and Huaca Prieta, Peru.

Maize (Zea mays ssp. mays) is among the world's most important and ancient domesticated crops. Although the chronology of its domestication and initial dispersals out of Mexico into Central and South America has become more clear due to molecular and multiproxy archaeobotanical research, important problems remain. Among them is the paucity of information on maize's early morphological evolution and racial diversification brought about in part by the poor preservation of macrofossils dating to the pre-5000 calibrated years before the present period from obligate dispersal routes located in the tropical forest. We report newly discovered macrobotanical and microbotanical remains of maize that shed significant light on the chronology, land race evolution, and cultural contexts associated with the crop's early movements into South America and adaptation to new environments. The evidence comes from the coastal Peruvian sites of Paredones and Huaca Prieta, Peru; dates from the middle and late preceramic and early ceramic periods (between ca. 6700 and 3000 calibrated years before the present); and constitutes some of the earliest known cobs, husks, stalks, and tassels. The macrobotanical record indicates that a diversity of racial complexes characteristic of the Andean region emerged during the preceramic era. In addition, accelerator mass spectrometry radiocarbon determinations carried out directly on different structures of preserved maize plants strongly suggest that assays on burned cobs are more reliable than those on unburned cobs. Our findings contribute to knowledge of the early diffusion of maize and agriculture and have broader implications for understanding the development of early preindustrial human societies.

Microfossils in calculus demonstrate consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium).

The nature and causes of the disappearance of Neanderthals and their apparent replacement by modern humans are subjects of considerable debate. Many researchers have proposed biologically or technologically mediated dietary differences between the two groups as one of the fundamental causes of Neanderthal disappearance. Some scenarios have focused on the apparent lack of plant foods in Neanderthal diets. Here we report direct evidence for Neanderthal consumption of a variety of plant foods, in the form of phytoliths and starch grains recovered from dental calculus of Neanderthal skeletons from Shanidar Cave, Iraq, and Spy Cave, Belgium. Some of the plants are typical of recent modern human diets, including date palms (Phoenix spp.), legumes, and grass seeds (Triticeae), whereas others are known to be edible but are not heavily used today. Many of the grass seed starches showed damage that is a distinctive marker of cooking. Our results indicate that in both warm eastern Mediterranean and cold northwestern European climates, and across their latitudinal range, Neanderthals made use of the diverse plant foods available in their local environment and transformed them into more easily digestible foodstuffs in part through cooking them, suggesting an overall sophistication in Neanderthal dietary regimes.

Starch grain and phytolith evidence for early ninth millennium B.P. maize from the Central Balsas River Valley, Mexico.

Questions that still surround the origin and early dispersals of maize (Zea mays L.) result in large part from the absence of information on its early history from the Balsas River Valley of tropical southwestern Mexico, where its wild ancestor is native. We report starch grain and phytolith data from the Xihuatoxtla shelter, located in the Central Balsas Valley, that indicate that maize was present by 8,700 calendrical years ago (cal. B.P.). Phytolith data also indicate an early preceramic presence of a domesticated species of squash, possibly Cucurbita argyrosperma. The starch and phytolith data also allow an evaluation of current hypotheses about how early maize was used, and provide evidence as to the tempo and timing of human selection pressure on 2 major domestication genes in Zea and Cucurbita. Our data confirm an early Holocene chronology for maize domestication that has been previously indicated by archaeological and paleoecological phytolith, starch grain, and pollen data from south of Mexico, and reshift the focus back to an origin in the seasonal tropical forest rather than in the semiarid highlands.

The cultural and chronological context of early Holocene maize and squash domestication in the Central Balsas River Valley, Mexico.

Molecular evidence indicates that the wild ancestor of maize is presently native to the seasonally dry tropical forest of the Central Balsas watershed in southwestern Mexico. We report here on archaeological investigations in a region of the Central Balsas located near the Iguala Valley in Guerrero state that show for the first time a long sequence of human occupation and plant exploitation reaching back to the early Holocene. One of the sites excavated, the Xihuatoxtla Shelter, contains well-stratified deposits and a stone tool assemblage of bifacially flaked points, simple flake tools, and numerous handstones and milling stone bases radiocarbon dated to at least 8700 calendrical years B.P. As reported in a companion paper (Piperno DR, et al., in this issue of PNAS), starch grain and phytolith residues from the ground and chipped stone tools, plus phytoliths from directly associated sediments, provide evidence for maize (Zea mays L.) and domesticated squash (Cucurbita spp.) in contexts contemporaneous with and stratigraphically below the 8700 calendrical years B.P. date. The radiocarbon determinations, stratigraphic integrity of Xihuatoxtla's deposits, and characteristics of the stone tool assemblages associated with the maize and squash remains all indicate that these plants were early Holocene domesticates. Early agriculture in this region of Mexico appears to have involved small groups of cultivators who were shifting their settlements seasonally and engaging in a variety of subsistence pursuits.

Human behavioral ecology, phenotypic (developmental) plasticity, and agricultural origins: insights from the emerging evolutionary synthesis.

The fields of human behavioral ecology (HBE) and evolutionary developmental biology (evo-devo) both stand to make significant contributions to our understanding of agricultural origins. These two approaches share a concern with phenotypic-plasticity and its evolutionary significance. HBE considers the adaptive plasticity of the human phenotype in response to resource distribution in time and space and has helped to advance understanding of the economic costs and benefits of food production. However, evo-devo and the associated subject of phenotypic (developmental) plasticity have so far been largely neglected as sources of insight into the domestication of plants, despite growing evidence for their evolutionary importance in nature and their roles in the origins of novel traits. We argue that it is important to consider environmentally induced phenotypic variation resulting directly from both natural- and human-induced ecological change as a source of the distinctive morphologies of domesticated plants.

Starch grains on human teeth reveal early broad crop diet in northern Peru.

Previous research indicates that the Nanchoc Valley in northern Peru was an important locus of early and middle Holocene human settlement, and that between 9200 and 5500 (14)C yr B.P. the valley inhabitants adopted major crop plants such as squash (Cucurbita moschata), peanuts (Arachis sp.), and cotton (Gossypium barbadense). We report here an examination of starch grains preserved in the calculus of human teeth from these sites that provides direct evidence for the early consumption of cultivated squash and peanuts along with two other major food plants not previously detected. Starch from the seeds of Phaseolus and Inga feuillei, the flesh of Cucurbita moschata fruits, and the nuts of Arachis was routinely present on numerous teeth that date to between 8210 and 6970 (14)C yr B.P. Early plant diets appear to have been diverse and stable through time and were rich in cultivated foods typical of later Andean agriculture. Our data provide early archaeological evidence for Phaseolus beans and I. feuillei, an important tree crop, and indicate that effective food production systems that contributed significant dietary inputs were present in the Nanchoc region by 8000 (14)C yr B.P. Starch grain studies of dental remains document plants and edible parts of them not normally preserved in archaeological records and can assume primary roles as direct indicators of ancient human diets and agriculture.

Identification of teosinte, maize, and Tripsacum in Mesoamerica by using pollen, starch grains, and phytoliths.

We examined pollen grains and starch granules from a large number of modern populations of teosinte (wild Zea spp.), maize (Zea mays L.), and closely related grasses in the genus Tripsacum to assess their strengths and weaknesses in studying the origins and early dispersals of maize in its Mesoamerican cradle of origin. We report new diagnostic criteria and question the accuracy of others used previously by investigators to identify ancient maize where its wild ancestor, teosinte, is native. Pollen grains from teosinte overlap in size with those of maize to a much greater degree than previously reported, making the differentiation of wild and domesticated maize in palynological studies difficult. There is presently no valid method for separating maize and teosinte pollen on a morphological basis. Starch grain analysis, a recently developed tool of archaeobotany, appears to be of significant utility in distinguishing the seeds of teosinte from maize. We propose that the differences in starch grain morphology and size between wild and domesticated maize defined in this study may be associated with domestication genes in Zea that have been documented in the starch biosynthesis pathway. As previously reported, phytoliths effectively discriminate the female reproductive structures of Tripsacum, teosinte, and maize. Multiproxy microfossil studies of archaeological and paleoecological contexts appear to be effective tools for investigating the earliest stages of maize domestication and dispersals.