The Evolution of Agrarian Landscapes in the Tropical Andes.

Changes in land-use practices have been a central element of human adaptation to Holocene climate change. Many practices that result in the short-term stabilization of socio-natural systems, however, have longer-term, unanticipated consequences that present cascading challenges for human subsistence strategies and opportunities for subsequent adaptations. Investigating complex sequences of interaction between climate change and human land-use in the past-rather than short-term causes and effects-is therefore essential for understanding processes of adaptation and change, but this approach has been stymied by a lack of suitably-scaled paleoecological data. Through a high-resolution paleoecological analysis, we provide a 7000-year history of changing climate and land management around Lake Acopia in the Andes of southern Peru. We identify evidence of the onset of pastoralism, maize cultivation, and possibly cultivation of quinoa and potatoes to form a complex agrarian landscape by c. 4300 years ago. Cumulative interactive climate-cultivation effects resulting in erosion ended abruptly c. 2300 years ago. After this time, reduced sedimentation rates are attributed to the construction and use of agricultural terraces within the catchment of the lake. These results provide new insights into the role of humans in the manufacture of Andean landscapes and the incremental, adaptive processes through which land-use practices take shape.

Phytolith assemblages reflect variability in human land use and the modern environment.

Phytoliths preserved in soils and sediments can be used to provide unique insights into past vegetation dynamics in response to human and climate change. Phytoliths can reconstruct local vegetation in terrestrial soils where pollen grains typically decay, providing a range of markers (or lack thereof) that document past human activities. The ca. 6 million km2 of Amazonian forests have relatively few baseline datasets documenting changes in phytolith representation across gradients of human disturbances. Here we show that phytolith assemblages vary on local scales across a gradient of (modern) human disturbance in tropical rainforests of Suriname. Detrended correspondence analysis showed that the phytolith assemblages found in managed landscapes (shifting cultivation and a garden), unmanaged forests, and abandoned reforesting sites were clearly distinguishable from intact forests and from each other. Our results highlight the sensitivity and potential of phytoliths to be used in reconstructing successional trajectories after site usage and abandonment. Percentages of specific phytolith morphotypes were also positively correlated with local palm abundances derived from UAV data, and with biomass estimated from MODIS satellite imagery. This baseline dataset provides an index of likely changes that can be observed at other sites that indicate past human activities and long-term forest recovery in Amazonia. Supplementary Information: The online version contains supplementary material available at 10.1007/s00334-023-00932-2.

Warming, drought, and disturbances lead to shifts in functional composition: A millennial-scale analysis for Amazonian and Andean sites.

Tropical forests are changing in composition and productivity, probably in response to changes in climate and disturbances. The responses to these multiple environmental drivers, and the mechanisms underlying the changes, remain largely unknown. Here, we use a functional trait approach on timescales of 10,000 years to assess how climate and disturbances influence the community-mean adult height, leaf area, seed mass, and wood density for eight lowland and highland forest landscapes. To do so, we combine data of eight fossil pollen records with functional traits and proxies for climate (temperature, precipitation, and El Niño frequency) and disturbances (fire and general disturbances). We found that temperature and disturbances were the most important drivers of changes in functional composition. Increased water availability (high precipitation and low El Niño frequency) generally led to more acquisitive trait composition (large leaves and soft wood). In lowland forests, warmer climates decreased community-mean height probably because of increased water stress, whereas in highland forests warmer climates increased height probably because of upslope migration of taller species. Disturbance increased the abundance of acquisitive, disturbance-adapted taxa with small seeds for quick colonization of disturbed sites, large leaves for light capture, and soft wood to attain fast height growth. Fire had weak effects on lowland forests but led to more stress-adapted taxa that are tall with fast life cycles and small seeds that can quickly colonize burned sites. Site-specific analyses were largely in line with cross-site analyses, except for varying site-level effects of El Niño frequency and fire activity, possibly because regional patterns in El Niño are not a good predictor of local changes, and charcoal abundances do not reflect fire intensity or severity. With future global changes, tropical Amazonian and Andean forests may transition toward shorter, drought- and disturbance-adapted forests in the lowlands but taller forests in the highlands.

Human-induced ecological cascades: Extinction, restoration, and rewilding in the Galápagos highlands.

Oceanic islands support unique biotas but often lack ecological redundancy, so that the removal of a species can have a large effect on the ecosystem. The larger islands of the Galápagos Archipelago once had one or two species of giant tortoise that were the dominant herbivore. Using paleoecological techniques, we investigate the ecological cascade on highland ecosystems that resulted from whalers removing many thousands of tortoises from the lowlands. We hypothesize that the seasonal migration of a now-extinct tortoise species to the highlands was curtailed by decreased intraspecific competition. We find the trajectory of plant community dynamics changed within a decade of the first whaling vessels visiting the islands. Novel communities established, with a previously uncommon shrub, Miconia, replacing other shrubs of the genera Alternanthera and Acalypha. It was, however, the introduction of cattle and horses that caused the local extirpation of plant species, with the most extreme impacts being evident after c. 1930. This modified ecology is considered the natural state of the islands and has shaped subsequent conservation policy and practice. Restoration of El Junco Crater should emphasize exclusion of livestock, rewilding with tortoises, and expanding the ongoing plantings of Miconia to also include Acalypha and Alternanthera.

Early to mid-Holocene human activity exerted gradual influences on Amazonian forest vegetation.

Humans have been present in Amazonia throughout the Holocene, with the earliest archaeological sites dating to 12 000 years ago. The earliest inhabitants began managing landscapes through fire and plant domestication, but the total extent of vegetation modification remains relatively unknown. Here, we compile palaeoecological records from lake sediments containing charcoal and from pollen analyses to understand how human land-use affected vegetation during the early to mid-Holocene, and place our results in the context of previous archaeological work. We identified gradual, rather than abrupt changes in forest openness, disturbance and enrichment, with useful species at almost all sites. Early human occupations occurred in peripheral sites of Amazonia, where natural fires are part of the vegetation dynamics, so human-made fires did not exert a novel form of disturbance. Synchronicity between evidence of the onset of human occupation in lake records and archaeological sites was found for eastern Amazonia. For southwestern and western Amazonia and the Guiana Shield, the timing of the onset of human occupation differed by thousands of years between lake records and archaeological sites. Plant cultivation showed a different spatio-temporal pattern, appearing ca 2000 years earlier in western Amazonia than in other regions. Our findings highlight the spatial-temporal heterogeneity of Amazonia and indicate that the region cannot be treated as one entity when assessing ecological or cultural history. This article is part of the theme issue 'Tropical forests in the deep human past'.

Potential distributions of pre-Columbian people in Tropical Andean landscapes.

Much has yet to be learned of the spatial patterning of pre-Columbian people across the Tropical Andes. Using compiled archaeological data and a suite of environmental variables, we generate an ensemble species distribution model (SDM) that incorporates general additive models, random forest models and Maxent models to reconstruct spatial patterns of pre-Columbian people that inhabited the Tropical Andes east of the continental divide, within the modern countries of Bolivia, Peru and Ecuador. Within this region, here referred to as the eastern Andean flank, elevation, mean annual cloud frequency, distance to rivers and precipitation of the driest quarter are the environmental variables most closely related to human occupancy. Our model indicates that 11.04% of our study area (65 368 km2) was likely occupied by pre-Columbian people. Our model shows that 30 of 351 forest inventory plots, which are used to generate ecological understanding of Andean ecosystems, were likely occupied in the pre-Columbian period. In previously occupied sites, successional trajectories may still be shaping forest dynamics, and those forests may still be recovering from the ecological legacy of pre-Columbian impacts. Our ensemble SDM links palaeo- and neo-ecology and can also be used to guide both future archaeological and ecological studies. This article is part of the theme issue 'Tropical forests in the deep human past'.

How deregulation, drought and increasing fire impact Amazonian biodiversity.

Biodiversity contributes to the ecological and climatic stability of the Amazon Basin1,2, but is increasingly threatened by deforestation and fire3,4. Here we quantify these impacts over the past two decades using remote-sensing estimates of fire and deforestation and comprehensive range estimates of 11,514 plant species and 3,079 vertebrate species in the Amazon. Deforestation has led to large amounts of habitat loss, and fires further exacerbate this already substantial impact on Amazonian biodiversity. Since 2001, 103,079-189,755 km2 of Amazon rainforest has been impacted by fires, potentially impacting the ranges of 77.3-85.2% of species that are listed as threatened in this region5. The impacts of fire on the ranges of species in Amazonia could be as high as 64%, and greater impacts are typically associated with species that have restricted ranges. We find close associations between forest policy, fire-impacted forest area and their potential impacts on biodiversity. In Brazil, forest policies that were initiated in the mid-2000s corresponded to reduced rates of burning. However, relaxed enforcement of these policies in 2019 has seemingly begun to reverse this trend: approximately 4,253-10,343 km2 of forest has been impacted by fire, leading to some of the most severe potential impacts on biodiversity since 2009. These results highlight the critical role of policy enforcement in the preservation of biodiversity in the Amazon.

Andean drought and glacial retreat tied to Greenland warming during the last glacial period.

Abrupt warming events recorded in Greenland ice cores known as Dansgaard-Oeschger (DO) interstadials are linked to changes in tropical circulation during the last glacial cycle. Corresponding variations in South American summer monsoon (SASM) strength are documented, most commonly, in isotopic records from speleothems, but less is known about how these changes affected precipitation and Andean glacier mass balance. Here we present a sediment record spanning the last ~50 ka from Lake Junín (Peru) in the tropical Andes that has sufficient chronologic precision to document abrupt climatic events on a centennial-millennial time scale. DO events involved the near-complete disappearance of glaciers below 4700 masl in the eastern Andean cordillera and major reductions in the level of Peru's second largest lake. Our results reveal the magnitude of the hydroclimatic disruptions in the highest reaches of the Amazon Basin that were caused by a weakening of the SASM during abrupt arctic warming. Accentuated warming in the Arctic could lead to significant reductions in the precipitation-evaporation balance of the southern tropical Andes with deleterious effects on this densely populated region of South America.

On the vital role of enamel prism interfaces and graded properties in human tooth survival.

Teeth of omnivores face a formidable evolutionary challenge: how to protect against fracture and abrasive wear caused by the wide variety of foods they process. It is hypothesized that this challenge is met in part by adaptations in enamel microstructure. The low-crowned teeth of humans and some other omnivorous mammals exhibit multiple fissures running longitudinally along the outer enamel walls, yet remain intact. It is proposed that inter-prism weakness and enamel property gradation act together to avert entry of these fissures into vulnerable inner tooth regions and, at the same time, confer wear resistance at the occlusal surface. A simple indentation experiment is employed to quantify crack paths and energetics in human enamel, and an extended-finite-element model to evaluate longitudinal crack growth histories. Consideration is given as to how tooth microstructure may have played a vital role in human evolution, and by extension to other omnivorous mammals.

2,100 years of human adaptation to climate change in the High Andes.

Humid montane forests are challenging environments for human habitation. We used high-resolution fossil pollen, charcoal, diatom and sediment chemistry data from the iconic archaeological setting of Laguna de los Condores, Peru to reconstruct changing land uses and climates in a forested Andean valley. Forest clearance and maize cultivation were initiated during periods of drought, with periods of forest recovery occurring during wetter conditions. Between AD 800 and 1000 forest regrowth was evident, but this trend was reversed between AD 1000 and 1200 as drier conditions coincided with renewed land clearance, the establishment of a permanent village and the use of cliffs overlooking the lake as a burial site. By AD 1230 forests had regrown in the valley and maize cultivation was greatly reduced. An elevational transect investigating regional patterns showed a parallel, but earlier, history of reduced maize cultivation and forest regeneration at mid-elevation. However, a lowland site showed continuous maize agriculture until European conquest but very little subsequent change in forest cover. Divergent, climate-sensitive landscape histories do not support categorical assessments that forest regrowth and peak carbon sequestration coincided with European arrival.

A 7000-year history of changing plant trait composition in an Amazonian landscape; the role of humans and climate.

Tropical forests are shifting in species and trait composition, but the main underlying causes remain unclear because of the short temporal scales of most studies. Here, we develop a novel approach by linking functional trait data with 7000 years of forest dynamics from a fossil pollen record of Lake Sauce in the Peruvian Amazon. We evaluate how climate and human disturbances affect community trait composition. We found weak relationships between environmental conditions and traits at the taxon level, but strong effects for community-mean traits. Overall, community-mean traits were more responsive to human disturbances than to climate change; human-induced erosion increased the dominance of dense-wooded, non-zoochorous species with compound leaves, and human-induced fire increased the dominance of tall, zoochorous taxa with large seeds and simple leaves. This information can help to enhance our understanding of forest responses to past environmental changes, and improve predictions of future changes in tropical forest composition.

Past and future global transformation of terrestrial ecosystems under climate change.

Impacts of global climate change on terrestrial ecosystems are imperfectly constrained by ecosystem models and direct observations. Pervasive ecosystem transformations occurred in response to warming and associated climatic changes during the last glacial-to-interglacial transition, which was comparable in magnitude to warming projected for the next century under high-emission scenarios. We reviewed 594 published paleoecological records to examine compositional and structural changes in terrestrial vegetation since the last glacial period and to project the magnitudes of ecosystem transformations under alternative future emission scenarios. Our results indicate that terrestrial ecosystems are highly sensitive to temperature change and suggest that, without major reductions in greenhouse gas emissions to the atmosphere, terrestrial ecosystems worldwide are at risk of major transformation, with accompanying disruption of ecosystem services and impacts on biodiversity.

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.

Human disturbance amplifies Amazonian El Niño-Southern Oscillation signal.

The long-term interaction between human activity and climate is subject to increasing scrutiny. Humans homogenize landscapes through deforestation, agriculture, and burning and thereby might reduce the capacity of landscapes to provide archives of climate change. Alternatively, land-use change might overwhelm natural buffering and amplify latent climate signals, rendering them detectable. Here we examine a sub-annually resolved sedimentary record from Lake Sauce in the western Amazonian lowlands that spans 6900 years. Finely-laminated sediments were deposited from ca. 5000 years ago until the present, and human activity in the watershed was revealed through the presence of charcoal and maize agriculture. The laminations, analyzed for color content and bandwidth, showed distinctive changes that were coupled to more frequent occurrence of fossil maize pollen. As agricultural activity intensified ca. 2200 cal. BP, the 2- to 8-year periodicity characteristic of El Niño-Southern Oscillation became evident in the record. These agricultural activities appeared to have amplified an existing, but subtle climatic signal that was previously absorbed by natural vegetation. When agricultural activity slowed, or land use around Lake Sauce changed at ca. 800 cal. BP, the signal of El Niño-Southern Oscillation (ENSO) activity became erratic.

The functional extinction of Andean megafauna.

Controversy exists over the cause and timing of the extinction of the Pleistocene megafauna. In the tropical Andes, deglaciation and associated rapid climate change began ~8,000 years before human arrival, providing an opportunity to separate the effects of climate change from human hunting on megafaunal extinction. We present a paleoecological record spanning the last 25,000 years from Lake Pacucha, Peru (3,100 m elevation). Fossil pollen, charcoal, diatoms, and the dung fungus Sporormiella, chronicle a two-stage megaherbivore population collapse. Sporormiella abundance, the proxy for megafaunal presence, fell sharply at ~21,000 years ago, but rebounded prior to a permanent decline between ~16,800 and 15,800 years ago. This two-stage decline in megaherbivores resulted in a functional extinction by ~15,800 years ago, 3,000 years earlier than known human occupation of the high Andes. Declining megaherbivore populations coincided with warm, wet intervals. Climatic instability and megafaunal population collapse triggered an ecological cascade that resulted in novel floral assemblages, and increases in woody species, fire frequency, and plant species that were sensitive to trampling. Our data revealed that Andean megafaunal populations collapsed due to positive feedbacks between habitat quality and climate change rather than human activity.

On the evolutionary advantage of multi-cusped teeth.

A hallmark of mammalian evolution is a progressive complexity in postcanine tooth morphology. However, the driving force for this complexity remains unclear: whether to expand the versatility in diet source, or to bolster tooth structural integrity. In this study, we take a quantitative approach to this question by examining the roles of number, position and height of multiple cusps in determining sustainable bite forces. Our approach is to use an extended finite-element methodology with due provision for step-by-step growth of an embedded crack to determine how fracture progresses with increasing occlusal load. We argue that multi-cusp postcanine teeth are well configured to withstand high bite forces provided that multiple cusps are contacted simultaneously to share the load. However, contact on a single near-wall cusp diminishes the strength. Location of the load points and cusp height, rather than cusp number or radius, are principal governing factors. Given these findings, we conclude that while complex tooth structures can enhance durability, increases in cusp number are more likely to be driven by the demands of food manipulation. Structural integrity of complex teeth is maintained when individual cusps remain sufficiently distant from the side walls and do not become excessively tall relative to tooth width.

Sporormiella as a tool for detecting the presence of large herbivores in the Neotropics / Sporormiella como ferramenta para detectar a presença de grandes herbívoros na região Neotropical

The reliability of using the abundance of Sporormiella spores as a proxy for the presence and abundance of megaherbivores was tested in southern Brazil. Mud-water interface samples from nine lakes, in which cattle-use was categorized as high, medium, or low, were assayed for Sporormiella representation. The sampling design allowed an analysis of both the influence of the number of animals using the shoreline and the distance of the sampling site from the nearest shoreline. Sporormiella was found to be a reliable proxy for the presence of large livestock. The concentration and abundance of spores declined from the edge of the lake toward the center, with the strongest response being in sites with high livestock use. Consistent with prior studies in temperate regions, we find that Sporormiella spores are a useful proxy to study the extinction of Pleistocene megafauna or the arrival of European livestock in Neotropical landscapes.

A confiabilidade dos valores de Sporormiella como um proxy para estimar a presença e abundância de megaherbívoros foi testada na região sudeste do Brasil. Amostras superficiais de nove lagos, categorizados quanto a presença de gados em alto, médio e baixo uso do seu entorno foram coletadas para a análise de abundância de Sporormiella. O modelo amostral aplicado permitiu a interpretação tanto da influência do número de animais que usam a margem do lago quanto a distância da margem do lago sobre a quantidade de esporos encontrados. As análises indicam que esporos de Sporormiella é um excelente proxy para detectar a presença de grandes herbívoros. A concentração e abundância de esporos reduz em direção ao centro do lago, o que fica mais evidente em locais com alto uso do entorno do lago por esses animais. Consistente com estudos realizados em regiões temperadas, nós concluímos que o uso de Sporormiella se mostra de grande valia para entender a extinção da megafauna do Pleistoceno como também a introdução de animais nas paisagens neotropicais.