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Indigenous societies are known to have occupied the Amazon basin for more than 12,000 years, but the scale of their influence on Amazonian forests remains uncertain. We report the discovery, using LIDAR (light detection and ranging) information from across the basin, of 24 previously undetected pre-Columbian earthworks beneath the forest canopy. Modeled distribution and abundance of large-scale archaeological sites across Amazonia suggest that between 10,272 and 23,648 sites remain to be discovered and that most will be found in the southwest. We also identified 53 domesticated tree species significantly associated with earthwork occurrence probability, likely suggesting past management practices. Closed-canopy forests across Amazonia are likely to contain thousands of undiscovered archaeological sites around which pre-Columbian societies actively modified forests, a discovery that opens opportunities for better understanding the magnitude of ancient human influence on Amazonia and its current state.

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The southwestern Amazon Rainforest Ecotone (ARE) is the transitional landscape between the tropical forest and seasonally flooded savannahs of the Bolivian Llanos de Moxos. These heterogeneous landscapes harbour high levels of biodiversity and some of the earliest records of human occupation and plant domestication in Amazonia. While persistent Indigenous legacies have been demonstrated elsewhere in the Amazon, it is unclear how past human-environment interactions may have shaped vegetation composition and structure in the ARE. Here, we examine 6000 years of archaeological and palaeoecological data from Laguna Versalles (LV), Bolivia. LV was dominated by stable rainforest vegetation throughout the Holocene. Maize cultivation and cultural burning are present after ca 5700 cal yr BP. Polyculture cultivation of maize, manioc and leren after ca 3400 cal yr BP predates the formation of Amazonian Dark/Brown Earth (ADE/ABE) soils (approx. 2400 cal yr BP). ADE/ABE formation is associated with agroforestry indicated by increased edible palms, including Mauritia flexuosa and Attalea sp., and record levels of burning, suggesting that fire played an important role in agroforestry practices. The frequent use of fire altered ADE/ABD forest composition and structure by controlling ignitions, decreasing fuel loads and increasing the abundance of plants preferred by humans. Cultural burning and polyculture agroforestry provided a stable subsistence strategy that persisted despite pronounced climate change and cultural transformations and has an enduring legacy in ADE/ABE forests in the ARE. This article is part of the theme issue 'Tropical forests in the deep human past'.

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The expansion of forest farmers across tropical lowland South America during the Late Holocene has long been connected to climate change. The more humid conditions established during the Late Holocene are assumed to have driven the expansion of forests, which would have facilitated the dispersal of cultures that practised agroforestry. The Tupi, a language family of widespread distribution in South America, occupies a central place in the debate. Not only are they one of the largest families in the continent, but their expansion from an Amazonian homeland has long been hypothesized to have followed forested environments wherever they settled. Here, we assess that hypothesis using a simulation approach. We employ equation-based and cellular automaton models, simulating demic-diffusion processes under two different scenarios: a null model in which all land cells can be equally settled, and an alternative model in which non-forested cells cannot be settled or delay the expansion. We show that including land cover as a constraint to movement results in a better approximation of the Tupi expansion as reconstructed by archaeology and linguistics.

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Human expansions motivated by the spread of farming are one of the most important processes that shaped cultural geographies during the Holocene. The best known example of this phenomenon is the Neolithic expansion in Europe, but parallels in other parts of the globe have recently come into focus. Here, we examine the expansion of four archaeological cultures of widespread distribution in lowland South America, most of which originated in or around the Amazon basin and spread during the late Holocene with the practice of tropical forest agriculture. We analyze spatial gradients in radiocarbon dates of each culture through space-time regressions, allowing us to establish the most likely geographical origin, time and speed of expansion. To further assess the feasibility of demic diffusion as the process behind the archaeological expansions in question, we employ agent-based simulations with demographic parameters derived from the ethnography of tropical forest farmers. We find that, while some expansions can be realistically modeled as demographic processes, others are not easily explainable in the same manner, which is possibly due to different processes driving their dispersal (e.g. cultural diffusion) or problematic/incomplete archaeological data.

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Flint corn processing method [coarse ground corn (CGC; 3.2 mm average particle size) or steam-flaked corn (SFC; 0.360 kg/L flake density)] was evaluated in conjunction with 4 levels of NDF from sugarcane bagasse (SCB) as roughage source (RNDF; 4%, 7%, 10%, and 13%; DM basis) to determine impact on growth performance, carcass characteristics, starch utilization, feeding behavior, and rumen morphometrics of Bos indicus beef cattle. Two hundred and forty Nellore bulls were blocked by initial BW (350 ± 37 kg), assigned to 32 feedlot pens and pens within weight block were randomly assigned, in a 2 × 4 factorial arrangement (2 corn processing and 4 levels of RNDF) to treatments. Effects of corn grain processing × RNDF level were not detected (P ≥ 0.14) for growth performance, dietary net energy concentration, carcass traits, rumen morphometrics, and feeding behavior, except for time spent ruminating and time spent resting (P ≤ 0.04), and a tendency for papillae width (P ≤ 0.09). Bulls fed SFC-based diets consumed 7% less (P = 0.001), had 10.6% greater carcass-adjusted ADG (P < 0.001) and 19% greater carcass-adjusted feed efficiency (P < 0.001) compared with bulls fed CGC-based diets. Observed net energy for maintenance and gain values were 14.9% and 19.4% greater (P < 0.001), respectively, for SFC than for CGC-based diets. Fecal starch concentration was less (P < 0.001) for bulls fed SFC compared with those fed CGC. No grain processing effects were detected (P = 0.51) for rumenitis score; however, cattle fed SFC presented smaller ruminal absorptive surface area (P = 0.03). Dry matter intake increased linearly (P = 0.02) and carcass-adjusted feed efficiency tended (P = 0.06) to decrease linearly as RNDF increased. Dietary RNDF concentration did not affect carcass characteristics (P ≥ 0.19), except for dressing percentage, which tended to decrease linearly (P = 0.06) as RNDF in finishing diets increased. Increasing RNDF in finishing diets had no effect (P = 0.26) on time spent eating, but time spent ruminating and resting increased linearly (min/d; P < 0.001) with increased dietary RNDF. Steam flaking markedly increased flint corn energy value, net energy of diets, and animal growth performance, and led to improvements on feed efficiency when compared with grinding, regardless of RNDF content of diets. Increasing dietary RNDF compromised feedlot cattle feed efficiency and carcass dressing.

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The long-term response of ancient societies to climate change has been a matter of global debate. Until recently, the lack of integrative studies using archaeological, palaeoecological and palaeoclimatological data prevented an evaluation of the relationship between climate change, distinct subsistence strategies and cultural transformations across the largest rainforest of the world, Amazonia. Here we review the most relevant cultural changes seen in the archaeological record of six different regions within Greater Amazonia during late pre-Columbian times. We compare the chronology of those cultural transitions with high-resolution regional palaeoclimate proxies, showing that, while some societies faced major reorganization during periods of climate change, others were unaffected and even flourished. We propose that societies with intensive, specialized land-use systems were vulnerable to transient climate change. In contrast, land-use systems that relied primarily on polyculture agroforestry, resulting in the formation of enriched forests and fertile Amazonian dark earth in the long term, were more resilient to climate change.

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The South American Monsoon System is responsible for the majority of precipitation in the continent, especially over the Amazon and the tropical savannah, known as 'Cerrado'. Compared to the extensively studied subtropical and temperate regions the effect of the Medieval Climate Anomaly (MCA) on the precipitation over the tropics is still poorly understood. Here, we present a multiproxy paleoprecipitation reconstruction showing a consistent change in the hydrologic regime during the MCA in the eastern Amazon and 'Cerrado', characterized by a substantial transition from humid to drier conditions during the Early (925-1150 C.E.) to Late-MCA (1150-1350 C.E.). We compare the timing of major changes in the monsoon precipitation with the expansion and abandonment of settlements reported in the archeological record. Our results show that important cultural successions in the pre-Columbian Central Amazon, the transition from Paredão to Guarita phase, are in agreement with major changes in the hydrologic regime. Phases of expansion and, subsequent abandonment, of large settlements from Paredão during the Early to Late-MCA are coherent with a reduction in water supply. In this context we argue that the sustained drier conditions during the latter period may have triggered territorial disputes with Guarita leading to the Paredão demise.

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Domesticated maize evolved from wild teosinte under human influences in Mexico beginning around 9000 years before the present (yr B.P.), traversed Central America by ~7500 yr B.P., and spread into South America by ~6500 yr B.P. Landrace and archaeological maize genomes from South America suggest that the ancestral population to South American maize was brought out of the domestication center in Mexico and became isolated from the wild teosinte gene pool before traits of domesticated maize were fixed. Deeply structured lineages then evolved within South America out of this partially domesticated progenitor population. Genomic, linguistic, archaeological, and paleoecological data suggest that the southwestern Amazon was a secondary improvement center for partially domesticated maize. Multiple waves of human-mediated dispersal are responsible for the diversity and biogeography of modern South American maize.

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The legacy of pre-Columbian land use in the Amazonian rainforest is one of the most controversial topics in the social1-10 and natural sciences11,12. Until now, the debate has been limited to discipline-specific studies, based purely on archaeological data8, modern vegetation13, modern ethnographic data3 or a limited integration of archaeological and palaeoecological data12. The lack of integrated studies to connect past land use with modern vegetation has left questions about the legacy of pre-Columbian land use on the modern vegetation composition in the Amazon, unanswered11. Here, we show that persistent anthropogenic landscapes for the past 4,500 years have had an enduring legacy on the hyperdominance of edible plants in modern forests in the eastern Amazon. We found an abrupt enrichment of edible plant species in fossil lake and terrestrial records associated with pre-Columbian occupation. Our results demonstrate that, through closed-canopy forest enrichment, limited clearing for crop cultivation and low-severity fire management, long-term food security was attained despite climate and social changes. Our results suggest that, in the eastern Amazon, the subsistence basis for the development of complex societies began ~4,500 years ago with the adoption of polyculture agroforestry, combining the cultivation of multiple annual crops with the progressive enrichment of edible forest species and the exploitation of aquatic resources. This subsistence strategy intensified with the later development of Amazonian dark earths, enabling the expansion of maize cultivation to the Belterra Plateau, providing a food production system that sustained growing human populations in the eastern Amazon. Furthermore, these millennial-scale polyculture agroforestry systems have an enduring legacy on the hyperdominance of edible plants in modern forests in the eastern Amazon. Together, our data provide a long-term example of past anthropogenic land use that can inform management and conservation efforts in modern Amazonian ecosystems.

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In the highlands of southern Brazil an anthropogenitcally driven expansion of forest occurred at the expense of grasslands between 1410 and 900 cal BP, coincident with a period of demographic and cultural change in the region. Previous studies have debated the relative contributions of increasing wetter and warmer climate conditions and human landscape modifications to forest expansion, but generally lacked high resoltiuon proxies to measure these effects, or have relied on single proxies to reconstruct both climate and vegetation. Here, we develop and test a model of natural ecosystem distribution against vegetation histories, paleoclimate proxies, and the archaeological record to distinguish human from temperature and precipitation impacts on the distribution and expansion of Araucaria forests during the late Holocene. Carbon isotopes from soil profiles confirm that in spite of climatic fluctuations, vegetation was stable and forests were spatially limited to south-facing slopes in the absence of human inputs. In contrast, forest management strategies for the past 1400 years expanded this economically important forest beyond its natural geographic boundaries in areas of dense pre-Columbian occupation, suggesting that landscape modifications were linked to demographic changes, the effects of which are still visible today.

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