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Reliable maps of species distributions are fundamental for biodiversity research and conservation. The International Union for Conservation of Nature (IUCN) range maps are widely recognized as authoritative representations of species' geographic limits, yet they might not always align with actual occurrence data. In recent area of habitat (AOH) maps, areas that are not habitat have been removed from IUCN ranges to reduce commission errors, but their concordance with actual species occurrence also remains untested. We tested concordance between occurrences recorded in camera trap surveys and predicted occurrences from the IUCN and AOH maps for 510 medium- to large-bodied mammalian species in 80 camera trap sampling areas. Across all areas, cameras detected only 39% of species expected to occur based on IUCN ranges and AOH maps; 85% of the IUCN only mismatches occurred within 200 km of range edges. Only 4% of species occurrences were detected by cameras outside IUCN ranges. The probability of mismatches between cameras and the IUCN range was significantly higher for smaller-bodied mammals and habitat specialists in the Neotropics and Indomalaya and in areas with shorter canopy forests. Our findings suggest that range and AOH maps rarely underrepresent areas where species occur, but they may more often overrepresent ranges by including areas where a species may be absent, particularly at range edges. We suggest that combining range maps with data from ground-based biodiversity sensors, such as camera traps, provides a richer knowledge base for conservation mapping and planning., (© 2024 The Authors. Conservation Biology published by Wiley Periodicals LLC on behalf of Society for Conservation Biology.)

Wildlife must adapt to human presence to survive in the Anthropocene, so it is critical to understand species responses to humans in different contexts. We used camera trapping as a lens to view mammal responses to changes in human activity during the COVID-19 pandemic. Across 163 species sampled in 102 projects around the world, changes in the amount and timing of animal activity varied widely. Under higher human activity, mammals were less active in undeveloped areas but unexpectedly more active in developed areas while exhibiting greater nocturnality. Carnivores were most sensitive, showing the strongest decreases in activity and greatest increases in nocturnality. Wildlife managers must consider how habituation and uneven sensitivity across species may cause fundamental differences in human-wildlife interactions along gradients of human influence., (© 2024. The Author(s).)

Biogeographic history can lead to variation in biodiversity across regions, but it remains unclear how the degree of biogeographic isolation among communities may lead to differences in biodiversity. Biogeographic analyses generally treat regions as discrete units, but species assemblages differ in how much biogeographic history they share, just as species differ in how much evolutionary history they share. Here, we use a continuous measure of biogeographic distance, phylobetadiversity, to analyze the influence of biogeographic isolation on the taxonomic and functional diversity of global mammal and bird assemblages. On average, biodiversity is better predicted by environment than by isolation, especially for birds. However, mammals in deeply isolated regions are strongly influenced by isolation; mammal assemblages in Australia and Madagascar, for example, are much less diverse than predicted by environment alone and contain unique combinations of functional traits compared to other regions. Neotropical bat assemblages are far more functionally diverse than Paleotropical assemblages, reflecting the different trajectories of bat communities that have developed in isolation over tens of millions of years. Our results elucidate how long-lasting biogeographic barriers can lead to divergent diversity patterns, against the backdrop of environmental determinism that predominantly structures diversity across most of the world., (© 2024. The Author(s).)

The United Nations recently agreed to major expansions of global protected areas (PAs) to slow biodiversity declines 1 . However, although reserves often reduce habitat loss, their efficacy at preserving animal diversity and their influence on biodiversity in surrounding unprotected areas remain unclear 2-5 . Unregulated hunting can empty PAs of large animals 6 , illegal tree felling can degrade habitat quality 7 , and parks can simply displace disturbances such as logging and hunting to unprotected areas of the landscape 8 (a phenomenon called leakage). Alternatively, well-functioning PAs could enhance animal diversity within reserves as well as in nearby unprotected sites 9 (an effect called spillover). Here we test whether PAs across mega-diverse Southeast Asia contribute to vertebrate conservation inside and outside their boundaries. Reserves increased all facets of bird diversity. Large reserves were also associated with substantially enhanced mammal diversity in the adjacent unprotected landscape. Rather than PAs generating leakage that deteriorated ecological conditions elsewhere, our results are consistent with PAs inducing spillover that benefits biodiversity in surrounding areas. These findings support the United Nations goal of achieving 30% PA coverage by 2030 by demonstrating that PAs are associated with higher vertebrate diversity both inside their boundaries and in the broader landscape., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

AbstractCommunity trait assembly, the formation of distributions of phenotypic characteristics across coexisting species, can occur via two main processes: filtering of trait distributions from the regional pool and in situ phenotypic evolution in local communities. But the relative importance of these processes remains unclear, largely because of the difficulty in determining the timing of evolutionary trait changes and biogeographic dispersal events in phylogenies. We assessed evolutionary and biogeographic transitions in woody plant species across the Indo-Malay archipelago, a series of island groups where the same plant lineages interact with different seed disperser and seed predator assemblages. Fruit size in 2,650 taxa spanning the angiosperm tree of life tended to be smaller in the Sulawesi and Maluku island groups, where frugivores are less diverse and smaller bodied, than in the regional source pool. While numerous plant lineages (not just small-fruited ones) reached the isolated islands, colonists tended to be the smaller-fruited members of each clade. Nearly all of the evolutionary transitions to smaller fruit size predated, often substantially, organismal dispersal to the islands. Our results suggest that filtering rather than within-island evolution largely determined the distribution of fruit sizes in these regions.

The loss of large animals due to overhunting and habitat loss potentially affects tropical tree populations and carbon cycling. Trees reliant on large-bodied seed dispersers are thought to be particularly negatively affected by defaunation. But besides seed dispersal, defaunation can also increase or decrease seed predation. It remains unclear how these different defaunation effects on early life stages ultimately affect tree population dynamics. We reviewed the literature on how tropical animal loss affects different plant life stages, and we conducted a meta-analysis of how defaunation affects seed predation. We used this information to parameterize models that altered matrix projection models from a suite of tree species to simulate defaunation-caused changes in seed dispersal and predation. We assessed how applying these defaunation effects affected population growth rates. On average, population-level effects of defaunation were negligible, suggesting that defaunation may not cause the massive reductions in forest carbon storage that have been predicted. In contrast to previous hypotheses, we did not detect an effect of seed size on changes in seed predation rates. The change in seed predation did not differ significantly between exclosure experiments and observational studies, although the results of observational studies were far more variable. Although defaunation surely affects certain tree taxa, species that benefit or are harmed by it and net changes in forest carbon storage cannot currently be predicted based on available data. Further research on how factors such as seed predation vary across tree species and defaunation scenarios is necessary for understanding cascading changes in species composition and diversity., (© 2022 Society for Conservation Biology.)

The numerous explanations for why Earth's biodiversity is concentrated at low latitudes fail to explain variation in the strength and even direction of the gradient through deep time. Consequently, we do not know if today's gradient is representative of what might be expected on other planets or is merely an idiosyncrasy of Earth's history. We propose a hierarchy of factors driving the latitudinal distribution of diversity: (i) over geologically long time spans, diversity is largely predicted by climate; (ii) when climatic gradients are shallow, diversity tracks habitat area; and (iii) historical contingencies linked to niche conservatism have geologically short-term, transient influence at most. Thus, latitudinal diversity gradients, although variable in strength and direction, are largely predictable on our planet and possibly others., Competing Interests: Declaration of interests The authors declare no conflicts of interest., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Patterns of co-occurrence among species can help reveal the structure and assembly of ecological communities. However, studies have been limited by measuring co-occurrence in either space or time but not both simultaneously. This is especially problematic in systems such as masting forests where resources are highly variable, meaning that spatial use and co-occurrence patterns can change on fine spatiotemporal scales. We develop an analytical framework for assessing species co-occurrence at fine spatial and temporal scales simultaneously and apply these models to a camera trapping dataset from Borneo. We sought to determine how substantial variation in food availability across space and time affects co-occurrence among terrestrial vertebrates. We detect many significant, mostly positive, co-occurrence patterns among species, but almost entirely in unlogged forest and during dipterocarp mast years. The most strongly co-occurring pair of species, bearded pig (Sus barbatus) and sambar (Rusa unicolor), only positively co-occur in areas and years when fruit is locally abundant. Species occurrences in logged forest and non-mast years are mostly random with respect to other species. This suggests that frugivore-granivore species positively co-occur when resources are plentiful (i.e., large trees are present and fruiting), likely because they use the same resources; these patterns disappear when food availability is lower. Our approach demonstrates the utility of measuring co-occurrence in space and time together and highlights the importance of resource abundance for driving the co-occurrence structure of communities. Furthermore, our method could be broadly applied to other systems to assess fine-scale spatiotemporal patterns across a range of taxa., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Conservation outcomes could be greatly enhanced if strategies addressing anthropogenic land-use change considered the impacts of these changes on entire communities as well as on individual species. Examining how species interactions change across gradients of habitat disturbance allows us to predict the cascading consequences of species extinctions and the response of ecological networks to environmental change. We conducted the first detailed study of changes in a commensalist network of mammals and dung beetles across an environmental disturbance gradient, from primary tropical forest to plantations, which varied in above-ground carbon density (ACD) and mammal communities. Mammal diversity changed only slightly across the gradient, remaining high even in oil palm plantations and fragmented forest. Dung beetle species richness, however, declined in response to lower ACD and was particularly low in plantations and the most disturbed forest sites. Three of the five network metrics (nestedness, network specialization and functionality) were significantly affected by changes in dung beetle species richness and ACD, but mammal diversity was not an important predictor of network structure. Overall, the interaction networks remained structurally and functionally similar across the gradient, only becoming simplified (i.e. with fewer dung beetle species and fewer interactions) in the most disturbed sites. We suggest that the high diversity of mammals, even in disturbed forests, combined with the generalist feeding patterns of dung beetles, confer resilience to the commensalist dung beetle-mammal networks. This study highlights the importance of protecting logged and fragmented forests to maintain interaction networks and potentially prevent extinction cascades in human-modified systems., (© 2021 British Ecological Society.)

Mitigating the massive impacts of defaunation on natural ecosystems requires understanding and predicting hunting effort across the landscape. But such understanding has been hindered by the difficulty of assessing the movement patterns of hunters in thick forests and across complex terrain. We statistically tested hypotheses about the spatial distribution of hunting with circuit theory and structural equation models. We used a data set of >7000 known kill locations in Guyana and hunter movement models to test these methods. Comparing models with different resistance layers (i.e., different estimates of how terrain and land cover influence human movement speed) showed that rivers, on average, limited movement rather than serving as transport arteries. Moreover, far more kills occurred close to villages than in remote areas. This, combined with the lack of support for structural equation models that included latent terms for prey depletion driven by past overhunting, suggests that kill locations in this system tended to be driven by where hunters were currently foraging rather than by influences of historical harvest. These analyses are generalizable to a variety of ecosystems, species, and data types, providing a powerful way of enhancing maps and predictions of hunting effort across complex landscapes., (© 2020 Society for Conservation Biology.)

Overhunting reduces important plant-animal interactions such as vertebrate seed dispersal and seed predation, thereby altering plant regeneration and even above-ground biomass. It remains unclear, however, if non-hunted species can compensate for lost vertebrates in defaunated ecosystems. We use a nested exclusion experiment to isolate the effects of different seed enemies in a Bornean rainforest. In four of five tree species, vertebrates kill many seeds (13-66%). Nonetheless, when large mammals are excluded, seed mortality from insects and fungi fully compensates for the lost vertebrate predation, such that defaunation has no effect on seedling establishment. The switch from seed predation by generalist vertebrates to specialist insects and fungi in defaunated systems may alter Janzen-Connell effects and density-dependence in plants. Previous work using simulation models to explore how lost seed dispersal will affect tree species composition and carbon storage may require reevaluation in the context of functional redundancy within complex species interactions networks.

Biodiversity is declining worldwide. Because species interact with one another and with their environment, losses of particular organisms alter the function of ecosystems. Our understanding of the global rates and specific causes of functional decline remains limited, however. Species losses also reduce the cumulative amount of extant evolutionary history ("phylogenetic diversity" [PD]) in communities-our biodiversity heritage. Here we provide a global assessment of how each known anthropogenic threat is driving declines in functional diversity (FD) and PD, using terrestrial mammals as a case study. We find that habitat loss and harvest (e.g., legal hunting, poaching, snaring) are by far the biggest drivers of ongoing FD and PD loss. Declines in FD in high-biodiversity countries, particularly in Southeast Asia and South America, are greater than would be expected if species losses were random with respect to ecological function. Among functional guilds, herbivores are disproportionately likely to be declining from harvest, with important implications for plant communities and nutrient cycling. Frugivores are particularly likely to be declining from both harvest and habitat loss, with potential ramifications for seed dispersal and even forest carbon storage. Globally, phylogenetically unique species do not have an elevated risk of decline, but in areas such as Australia and parts of Southeast Asia, both habitat loss and harvest are biased toward phylogenetically unique species. Enhanced conservation efforts, including a renewed focus on harvest sustainability, are urgently needed to prevent the deterioration of ecosystem function, especially in the South American and equatorial Asian tropics., Competing Interests: The authors declare no competing interest.

Conservation planning tends to focus on protecting species' ranges or landscape connectivity but seldom both-particularly in the case of diverse taxonomic assemblages and multiple planning goals. Therefore, information on potential trade-offs between maintaining landscape connectivity and achieving other conservation objectives is lacking. We developed an optimization approach to prioritize the maximal protection of species' ranges, ecosystem types, and forest carbon stocks, while also including habitat connectivity for range-shifting species and dispersal corridors to link protected area. We applied our approach to Sabah, Malaysia, where the state government mandated an increase in protected-area coverage of approximately 305,000 ha but did not specify where new protected areas should be. Compared with a conservation planning approach that did not incorporate the 2 connectivity features, our approach increased the protection of dispersal corridors and elevational connectivity by 13% and 21%, respectively. Coverage of vertebrate and plant species' ranges and forest types were the same whether connectivity was included or excluded. Our approach protected 2% less forest carbon and 3% less butterfly range than when connectivity features were not included. Hence, the inclusion of connectivity into conservation planning can generate large increases in the protection of landscape connectivity with minimal loss of representation of other conservation targets., (© 2019 Society for Conservation Biology.)

Unsustainable hunting is emptying forests of large animals around the world, but current understanding of how human foraging spreads across landscapes has been stymied by data deficiencies and cryptic hunter behaviour. Unlike other global threats to biodiversity like deforestation, climate change and overfishing, maps of wild meat hunters' movements-often based on forest accessibility-typically cover small scales and are rarely validated with real-world observations. Using camera trapping data from rainforests across Malaysian Borneo, we show that while hunter movements are strongly correlated with the accessibility of different parts of the landscape, accessibility measures are most informative when they integrate fine-scale habitat features like topography and land cover. Measures of accessibility naive to fine-scale habitat complexity, like distance to the nearest road or settlement, generate poor approximations of hunters' movements. In comparison, accessibility as measured by high-resolution movement models based on circuit theory provides vastly better reflections of real-world foraging movements. Our results highlight that simple models incorporating fine-scale landscape heterogeneity can be powerful tools for understanding and predicting widespread threats to biodiversity.

Although dispersal is generally viewed as a crucial determinant for the fitness of any organism, our understanding of its role in the persistence and spread of plant populations remains incomplete. Generalizing and predicting dispersal processes are challenging due to context dependence of seed dispersal, environmental heterogeneity and interdependent processes occurring over multiple spatial and temporal scales. Current population models often use simple phenomenological descriptions of dispersal processes, limiting their ability to examine the role of population persistence and spread, especially under global change. To move seed dispersal ecology forward, we need to evaluate the impact of any single seed dispersal event within the full spatial and temporal context of a plant's life history and environmental variability that ultimately influences a population's ability to persist and spread. In this perspective, we provide guidance on integrating empirical and theoretical approaches that account for the context dependency of seed dispersal to improve our ability to generalize and predict the consequences of dispersal, and its anthropogenic alteration, across systems. We synthesize suitable theoretical frameworks for this work and discuss concepts, approaches and available data from diverse subdisciplines to help operationalize concepts, highlight recent breakthroughs across research areas and discuss ongoing challenges and open questions. We address knowledge gaps in the movement ecology of seeds and the integration of dispersal and demography that could benefit from such a synthesis. With an interdisciplinary perspective, we will be able to better understand how global change will impact seed dispersal processes, and potential cascading effects on plant population persistence, spread and biodiversity., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Wild meat (or 'bushmeat') hunting is nearly ubiquitous across the tropics and is very often unsustainable-driving declines and extirpation of numerous mammal populations. Loss of these animals can alter the transport of nutrients within and between ecosystems. But whether the physical removal of vertebrate carcasses and the nutrients that they store can reduce overall nutrient availability in ecosystems has been little explored. At 32 sites on three continents, we show that annual phosphorus (P) loss via mammal exploitation was low relative to the rate of atmospheric P deposition. But at four sites in Africa and Southeast Asia, removal of P in the skeletons of hunted mammals exceeded the atmospheric input of this nutrient by 10-fold or more. Because P is the growth-limiting nutrient for many tropical terrestrial ecosystems and certain large mammals, the imbalance created by the removal of mammal biomass under very high hunting scenarios could reduce ecosystem carrying capacity if no compensatory P additions occur in the system. This biogeochemical perspective on bushmeat exploitation raises further concerns about harvest sustainability and human food security in areas where hunting rates are high and ecosystem P inputs low.

Management activities such as law enforcement and community outreach are thought to affect conservation outcomes in protected areas, but their importance relative to intrinsic environmental characteristics of the parks and extrinsic human pressures surrounding the parks have not been explored. Furthermore, it is not clear which is more related to conservation outcomes-the management itself or local people's perceptions of the management. We measured objective (reports by park staff) and subjective (reports by local people) levels of community outreach and law enforcement based on responses to 374 questionnaires. We estimated mammal abundance and diversity of 6 protected areas based on data from 115 camera traps in Xishuangbanna, southwest China, a biodiversity hotspot with high hunting and land-conversion pressures. We then examined correlations among them and found that local people's perception of law enforcement was positively related to the local abundance of 2 large, hunted species, wild boar (Sus scrofa) (β = 15.22) and muntjac (Muntiacus vaginalis) (β = 14.82), but not related to the abundance of smaller mammals or to objective levels of enforcement. The subjective frequency of outreach by park staff to local communities (β = 3.42) and park size (β = 3.28) were significantly and positively related to mammal species richness, whereas elevation, human population density, and subjective frequency of law enforcement were not. We could not conclude that community outreach and law enforcement were directly causing increased mammal abundance and diversity. Nevertheless, the patterns we detected are some of the first empirical evidence consistent with the idea that biodiversity in protected areas may be more positively and strongly related to local perceptions of the intensity of park management than to either intrinsic (e.g., elevation, park size) or extrinsic (e.g., human population density) environmental factors., (© 2018 Society for Conservation Biology.)

Periods of extreme food abundance, such as irregular masting events, can dramatically affect animal populations and communities, but the extent to which anthropogenic disturbances alter animal responses to mast events is not clear. In South-East Asia, dipterocarp trees reproduce in mast fruiting events every 2-10 years in some of the largest masting events on the planet. These trees, however, are targeted for selective logging, reducing the intensity of fruit production and potentially affecting multiple trophic levels. Moreover, animal responses to resource pulse events have largely been studied in systems where the major mast consumers have been extirpated. We sought to evaluate the influence of human-induced habitat disturbance on animal responses to masting in a system where key mast consumers remain extant. We used motion-triggered camera traps to quantify terrestrial mammal and bird occurrences in Sabah, Malaysian Borneo, relative to variation in fruit biomass from 69 plant families during a major (2014) and minor (2015) masting event and a non-mast year (2013), in both logged and unlogged forests. Bearded pigs (Sus barbatus) showed the clearest responses to masting and occurrence rates were highest in unlogged forest in the year following the major mast, suggesting that the pulse in fruit availability increased immigration or reproduction. We also detected local-scale spatial tracking of dipterocarp fruits in bearded pigs in unlogged forest, while this was equivocal in other species. In contrast, pigs and other vertebrate taxa in our study showed limited response to spatial or temporal variation in fruit availability in logged forest. Our findings suggest that vertebrates, namely bearded pigs, may respond to masting via movement and increased reproduction, but that these responses may be attenuated by habitat disturbance., (© 2019 The Authors. Journal of Animal Ecology © 2019 British Ecological Society.)

A fundamental tenet of modern ecology and conservation science is the fact that species occurrence in habitat patches can be determined by patch area and isolation. But such island biogeographic models often poorly predict actual species occurrences in structurally complex landscapes that typify most ecosystems. Recent advances in circuit theory have enhanced estimates of species dispersal and can provide powerful ways to predict landscape-scale distribution of species assemblages through integration with island biogeography. Applying such an integrative analytical framework to 43 bird species in Tanzania improved model fit by an average of 2.2-fold over models where patch isolation was estimated without accounting for landscape matrix heterogeneity. This approach also allowed us to assess species-specific dispersal rates and quantify differences among land cover types in their permeability to animal movement. These results reaffirm the utility of foundational island biogeographic principles, yet with an important caveat. Two-thirds of the variance in species occurrence in habitat fragments can be explained simply by patch area and isolation, conditional on isolation explicitly accounting for the spatial configuration of different land cover types in the landscape matrix.

The expansion of shrubs and trees across high-latitude ecosystems is one of the most dramatic ecological manifestations of climate change. Most of the work quantifying these changes has been done in small areas and over relatively recent time scales. These land-cover transitions are highly spatially variable, and we have limited understanding of the factors underlying this variation. We use repeat photography to generate a data set of land-cover changes in Denali National Park and Preserve, Alaska, stretching back a century and spanning a range of edaphic, topographic, and climatic conditions. Most land-cover classes were quite stable, with low probabilities of transitioning to other land-cover types. The advance of woody vegetation into low-stature tundra, and the spread of conifer trees into shrub-dominated areas, were both more likely at low elevations and in areas without permafrost. Permafrost also reduced the likelihood of herbaceous vegetation transitioning to woody cover. Exceptions to the general trend of relative stability included nearly all (96%) of the broadleaf forest-dominated areas being invaded by conifers, an expected successional trajectory, and many open gravel river bars (17.8%) transitioning to thick shrubs. These floodplain areas were distinctly not at equilibrium, as only 0.1% of shrub-dominated areas converted to gravel. Warming temperatures in coming decades and concomitant declines in the extent of permafrost are predicted to enhance the spread of woody vegetation in Denali further, but only by ~3%. Land-cover transitions, notably the rapid advance of trees and shrubs observed in other studies, could be less likely and more spatially heterogeneous here than in other high-latitude systems., (© 2019 by the Ecological Society of America.)

Species interactions have long been predicted to increase in intensity toward the tropics and low elevations because of gradients in climate, productivity, or biodiversity. Despite their importance for understanding global ecological and evolutionary processes, plant-animal interaction gradients are particularly difficult to test systematically across large geographic gradients, and evidence from smaller, disparate studies is inconclusive. By systematically measuring postdispersal seed predation using 6995 standardized seed depots along 18 mountains in the Pacific cordillera, we found that seed predation increases by 17% from the Arctic to the Equator and by 17% from 4000 meters above sea level to sea level. Clines in total predation, likely driven by invertebrates, were consistent across treeline ecotones and within continuous forest and were better explained by climate seasonality than by productivity, biodiversity, or latitude. These results suggest that species interactions play predictably greater ecological and evolutionary roles in tropical, lowland, and other less seasonal ecosystems.

Effective conservation strategies must ensure that species remain not just extant, but able to maintain key roles in species interactions and in the maintenance of communities and ecosystems. Such ecological functions, however, have not been well incorporated into management or policy. We present a framework for quantifying ecological function that is complementary to population viability analysis (PVA) and that allows function to be integrated into strategic planning processes. Ecological function analysis (EFA) focuses on preventing secondary extinctions and maintaining ecosystem structure, biogeochemical processes, and resiliency. EFA can use a range of modeling approaches and, because most species interactions are relatively weak, EFA needs to be performed for relatively few species or functions, making it a realistic way to improve conservation management., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Protected areas form the cornerstone of global efforts to conserve biodiversity. Most current methods for designing reserve networks focus on maximizing the representation of species, but with no assurance that those species will persist in the protected landscapes into the future. We present a new strategy for reserve design that combines metapopulation theory with spatial conservation prioritization to estimate conservation solutions that minimize extinction risk across numerous species simultaneously. Our framework optimizes the spatial configuration of reserves to maximize metapopulation persistence for an entire assemblage of species by accounting for both species representation and landscape connectivity. As a case study, we design a reserve network for 114 terrestrial mammal species in Indonesian New Guinea. Our approach builds on Marxan, the flagship representation-based reserve design tool, improving estimated persistence (metapopulation capacity) by an average of 4.6-fold across species, without increasing the socioeconomic cost. We suggest that enhancing species persistence, rather than protecting arbitrary proportions of species' ranges, should be the ultimate objective of conservation planning., (© 2018 by the Ecological Society of America.)

Animals can have both positive (e.g. via seed dispersal) and negative (e.g. via herbivory) impacts on plants. The net effects of these interactions remain difficult to predict and may be affected by overhunting and habitat disturbance, two widespread threats to tropical forests. Recent studies have documented their separate effects on plant recruitment but our understanding of how defaunation and logging interact to influence tropical tree communities is limited. From 2013 to 2016, we followed the fate of marked tree seedlings ( n = 1489) from 81 genera in and outside experimental plots. Our plots differentially excluded small, medium and large-bodied mammal herbivores in logged and unlogged forest in Malaysian Borneo. We assessed the effects of experimental defaunation and logging on taxonomic diversity and plant trait (wood density, specific leaf area, fruit size) composition of seedling communities. Although seedling mortality was highest in the presence of all mammal herbivores (44%), defaunation alone did not alter taxonomic diversity nor plant trait composition. However, herbivores (across all body sizes) significantly reduced mean fruit size across the seedling community over time (95% confidence interval (CI): -0.09 to -0.01), particularly in logged forest (95% CI: -0.12 to -0.003). Our findings suggest that impacts of mammal herbivores on plant communities may be greater in forests with a history of disturbance and could subsequently affect plant functional traits and ecological processes associated with forest regeneration., (© 2018 The Author(s).)

Background: A crucial step in conserving biodiversity is to identify the distributions of threatened species and the factors associated with species threat status. In the biodiversity hotspot of the Himalaya, very little is known about which locations harbour the highest diversity of threatened species and whether diversity of such species is related to area, mid-domain effects (MDE), range size, or human density. In this study, we assessed the drivers of variation in richness of threatened birds, mammals, reptiles, actinopterygii, and amphibians along an elevational gradient in Nepal Himalaya., Results: Although geometric constraints (MDE), species range size, and human population density were significantly related to threatened species richness, the interaction between range size and human population density was of greater importance. Threatened species richness was positively associated with human population density and negatively associated with range size., Conclusions: In areas with high richness of threatened species, species ranges tend to be small. The preponderance of species at risk of extinction at low elevations in the subtropical biodiversity hotspot could be due to the double impact of smaller range sizes and higher human density.

Large, fruit-eating vertebrates have been lost from many of the world's ecosystems. The ecological consequences of this defaunation can be severe, but the evolutionary consequences are nearly unknown because it remains unclear whether frugivores exert strong selection on fruit traits. I assessed the macroevolution of fruit traits in response to variation in the diversity and size of seed-dispersing vertebrates. Across the Indo-Malay Archipelago, many of the same plant lineages have been exposed to very different assemblages of seed-dispersing vertebrates. Phylogenetic analysis of >400 plant species in 41 genera and five families revealed that average fruit size tracks the taxonomic and functional diversity of frugivorous birds and mammals. Fruit size was 40.2-46.5% smaller in the Moluccas and Sulawesi (respectively), with relatively depauperate assemblages of mostly small-bodied animals, than in the Sunda Region (Borneo, Sumatra, and Peninsular Malaysia), with a highly diverse suite of large and small animals. Fruit color, however, was unrelated to vertebrate diversity or to the representation of birds versus mammals in the frugivore assemblage. Overhunting of large animals, nearly ubiquitous in tropical forests, could strongly alter selection pressures on plants, resulting in widespread, although trait-specific, morphologic changes., Competing Interests: The author declares no conflict of interest., (Published under the PNAS license.)

Vertebrate granivores destroy plant seeds, but whether animal-induced seed mortality alters plant recruitment varies with habitat context, seed traits, and among granivore species. An incomplete understanding of seed predation makes it difficult to predict how widespread extirpations of vertebrate granivores in tropical forests might affect tree communities, especially in the face of habitat disturbance. Many tropical forests are simultaneously affected by animal loss as well as habitat disturbance, but the consequences of each for forest regeneration are often studied separately or additively, and usually on a single plant demographic stage. The combined impacts of these threats could affect plant recruitment in ways that are not apparent when studied in isolation. We used wire cages to exclude large (elephants), medium, (sambar deer, bearded pigs, muntjac deer), and small (porcupines, chevrotains) ground-dwelling mammalian granivores and herbivores in logged and unlogged forests in Malaysian Borneo. We assessed the interaction between habitat disturbance (selective logging) and experimental defaunation on seed survival, germination, and seedling establishment in five dominant dipterocarp tree species spanning a 21-fold gradient in seed size. Granivore-induced seed mortality was consistently higher in logged forest. Germination of unpredated seeds was reduced in logged forest and in the absence of small to large-bodied mammals. Experimental defaunation increased germination and reduced seed removal but had little effect on seed survival. Seedling recruitment however, was more likely where logging and animal loss occurred together. The interacting effects of logging and hunting could therefore, actually increase seedling establishment, suggesting that the loss of mammals in disturbed forest could have important consequences for forest regeneration and composition., (© 2017 by the Ecological Society of America.)

The responses of lowland tropical communities to climate change will critically influence global biodiversity but remain poorly understood. If species in these systems are unable to tolerate warming, the communities-currently the most diverse on Earth-may become depauperate ('biotic attrition'). In response to temperature changes, animals can adjust their distribution in space or their activity in time, but these two components of the niche are seldom considered together. We assessed the spatio-temporal niches of rainforest mammal species in Borneo across gradients in elevation and temperature. Most species are not predicted to experience changes in spatio-temporal niche availability, even under pessimistic warming scenarios. Responses to temperature are not predictable by phylogeny but do appear to be trait-based, being much more variable in smaller-bodied taxa. General circulation models and weather station data suggest unprecedentedly high midday temperatures later in the century; predicted responses to this warming among small-bodied species range from 9% losses to 6% gains in spatio-temporal niche availability, while larger species have close to 0% predicted change. Body mass may therefore be a key ecological trait influencing the identity of climate change winners and losers. Mammal species composition will probably change in some areas as temperatures rise, but full-scale biotic attrition this century appears unlikely., (© 2017 The Author(s).)

Although deforestation and forest degradation have long been considered the most significant threats to tropical biodiversity, across Southeast Asia (Northeast India, Indochina, Sundaland, Philippines) substantial areas of natural habitat have few wild animals (>1 kg), bar a few hunting-tolerant species. To document hunting impacts on vertebrate populations regionally, we conducted an extensive literature review, including papers in local journals and reports of governmental and nongovernmental agencies. Evidence from multiple sites indicated animal populations declined precipitously across the region since approximately 1980, and many species are now extirpated from substantial portions of their former ranges. Hunting is by far the greatest immediate threat to the survival of most of the region's endangered vertebrates. Causes of recent overhunting include improved access to forests and markets, improved hunting technology, and escalating demand for wild meat, wildlife-derived medicinal products, and wild animals as pets. Although hunters often take common species, such as pigs or rats, for their own consumption, they take rarer species opportunistically and sell surplus meat and commercially valuable products. There is also widespread targeted hunting of high-value species. Consequently, as currently practiced, hunting cannot be considered sustainable anywhere in the region, and in most places enforcement of protected-area and protected-species legislation is weak. The international community's focus on cross-border trade fails to address overexploitation of wildlife because hunting and the sale of wild meat is largely a local issue and most of the harvest is consumed in villages, rural towns, and nearby cities. In addition to improved enforcement, efforts to engage hunters and manage wildlife populations through sustainable hunting practices are urgently needed. Unless there is a step change in efforts to reduce wildlife exploitation to sustainable levels, the region will likely lose most of its iconic species, and many others besides, within the next few years., (© 2016 Society for Conservation Biology.)

We examined the links between the science and policy of habitat corridors to better understand how corridors can be implemented effectively. As a case study, we focused on a suite of landscape-scale connectivity plans in tropical and subtropical Asia (Malaysia, Singapore, and Bhutan). The process of corridor designation may be more efficient if the scientific determination of optimal corridor locations and arrangement is synchronized in time with political buy-in and establishment of policies to create corridors. Land tenure and the intactness of existing habitat in the region are also important to consider because optimal connectivity strategies may be very different if there are few, versus many, political jurisdictions (including commercial and traditional land tenures) and intact versus degraded habitat between patches. Novel financing mechanisms for corridors include bed taxes, payments for ecosystem services, and strategic forest certifications. Gaps in knowledge of effective corridor design include an understanding of how corridors, particularly those managed by local communities, can be protected from degradation and unsustainable hunting. There is a critical need for quantitative, data-driven models that can be used to prioritize potential corridors or multicorridor networks based on their relative contributions to long-term metacommunity persistence., (© 2016 Society for Conservation Biology.)

Elucidating how dispersal and landscape connectivity influence metacommunity stability will shed light on natural processes structuring ecosystems and help prioritize conservation actions in an increasingly fragmented world. Much of the theoretical and mathematical development of the metacommunity concept has been based on simplified experimental systems or simulated data. We still have limited understanding of how variation in the habitat matrix and species-specific differences in dispersal ability contribute to metacommunity dynamics in heterogeneous landscapes. We model a metacommunity of rainforest mammals in Borneo, a tropical biodiversity hotspot, where protected areas are increasingly isolated by ongoing habitat disturbance and loss. We employ a combination of hierarchical models of local abundance, circuit-theory-based dispersal analysis, and metapopulation models. Our goal was to understand which landscape links were the most important to metapopulation persistence and metacommunity stability. Links were particularly important if they were short and connected two large patches. This was partly because only the very shortest links could be traversed by poorly dispersing species, including small herbivores such as chevrotains (Tragulus spp.) and porcupines. Links that join large patches into a "super-patch" may also promote island-mainland rather than Levins-type metapopulation dynamics for good dispersers, particularly large carnivores such as clouded leopards (Neofelis diardi) and sun bears (Helarctos malayanus), reducing metapopulation extinction risk and thereby enhancing metacommunity stability. Link importance to metacommunity stability was highly correlated between heterogeneous and homogeneous landscapes. But link importance to metapopulation capacity varied strongly across species, and the correlation between heterogeneous and homogeneous landscape matrix scenarios was low for poorly dispersing taxa. This suggests that the environmental conditions in the area between habitat patches, the landscape matrix, is important for assessing certain individual species but less so for understanding the stability of the entire metacommunity., (© 2016 by the Ecological Society of America.)

Unsustainable hunting does more than threaten wildlife populations and human livelihood - Carlos Peres and colleagues recently showed that it can also reduce how much carbon the forest stores. Maintaining primates and other important seed dispersing animals in tropical forests could be an important piece of the climate change mitigation puzzle., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Background: Plasmodium knowlesi is a zoonotic pathogen, transmitted among macaques and to humans by anopheline mosquitoes. Information on P. knowlesi malaria is lacking in most regions so the first step to understand the geographical distribution of disease risk is to define the distributions of the reservoir and vector species., Methods: We used macaque and mosquito species presence data, background data that captured sampling bias in the presence data, a boosted regression tree model and environmental datasets, including annual data for land classes, to predict the distributions of each vector and host species. We then compared the predicted distribution of each species with cover of each land class., Results: Fine-scale distribution maps were generated for three macaque host species (Macaca fascicularis, M. nemestrina and M. leonina) and two mosquito vector complexes (the Dirus Complex and the Leucosphyrus Complex). The Leucosphyrus Complex was predicted to occur in areas with disturbed, but not intact, forest cover (> 60% tree cover) whereas the Dirus Complex was predicted to occur in areas with 10-100% tree cover as well as vegetation mosaics and cropland. Of the macaque species, M. nemestrina was mainly predicted to occur in forested areas whereas M. fascicularis was predicted to occur in vegetation mosaics, cropland, wetland and urban areas in addition to forested areas., Conclusions: The predicted M. fascicularis distribution encompassed a wide range of habitats where humans are found. This is of most significance in the northern part of its range where members of the Dirus Complex are the main P. knowlesi vectors because these mosquitoes were also predicted to occur in a wider range of habitats. Our results support the hypothesis that conversion of intact forest into disturbed forest (for example plantations or timber concessions), or the creation of vegetation mosaics, will increase the probability that members of the Leucosphyrus Complex occur at these locations, as well as bringing humans into these areas. An explicit analysis of disease risk itself using infection data is required to explore this further. The species distributions generated here can now be included in future analyses of P. knowlesi infection risk.

Agricultural expansion is the largest threat to global biodiversity. In particular, the rapid spread of tree plantations is a primary driver of deforestation in hyperdiverse tropical regions. Plantations tend to support considerably lower biodiversity than native forest, but it remains unclear whether plantation traits affect their ability to sustain native wildlife populations, particularly for threatened taxa. If animal diversity varies across plantations with different characteristics, these traits could be manipulated to make plantations more "wildlife friendly." The degree to which plantations create edge effects that degrade habitat quality in adjacent forest also remains unclear, limiting our ability to predict wildlife persistence in mixed-use landscapes. We used systematic camera trapping to investigate mammal occurrence and diversity in oil palm plantations and adjacent forest in Sabah, Malaysian Borneo. Mammals within plantations were largely constrained to locations near native forest; the occurrence of most species and overall species richness declined abruptly with decreasing forest proximity from an estimated 14 species at the forest ecotone to -1 species 2 km into the plantation. Neither tree height nor canopy cover within plantations strongly affected mammal diversity or occurrence, suggesting that manipulating tree spacing or planting cycles might not make plantations more wildlife friendly. Plantations did not appear to generate strong edge effects; mammal richness within forest remained high and consistent up to the plantation ecotone. Our results suggest that land-sparing strategies, as opposed to efforts to make plantations more wildlife-friendly, are required for regional wildlife conservation in biodiverse tropical ecosystems.

Habitat corridors are important tools for maintaining connectivity in increasingly fragmented landscapes, but generally they have been considered in single-species approaches. Corridors intended to facilitate the movement of multiple species could increase persistence of entire communities, but at the likely cost of being less efficient for any given species than a corridor intended specifically for that species. There have been few tests of the trade-offs between single- and multispecies corridor approaches. We assessed single-species and multispecies habitat corridors for 5 threatened mammal species in tropical forests of Borneo. We generated maps of the cost of movement across the landscape for each species based on the species' local abundance as estimated through hierarchical modeling of camera-trap data with biophysical and anthropogenic covariates. Elevation influenced local abundance of banded civets (Hemigalus derbyanus) and sun bears (Helarctos malayanus). Increased road density was associated with lower local abundance of Sunda clouded leopards (Neofelis diardi) and higher local abundance of sambar deer (Rusa unicolor). Pig-tailed macaque (Macaca nemestrina) local abundance was lower in recently logged areas. An all-species-combined connectivity scenario with least-cost paths and 1 km buffers generated total movement costs that were 27% and 23% higher for banded civets and clouded leopards, respectively, than the connectivity scenarios for those species individually. A carnivore multispecies connectivity scenario, however, increased movement cost by 2% for banded civets and clouded leopards. Likewise, an herbivore multispecies scenario provided more effective connectivity than the all-species-combined scenario for sambar and macaques. We suggest that multispecies habitat connectivity plans be tailored to groups of ecologically similar, disturbance-sensitive species to maximize their effectiveness., (© 2014 Society for Conservation Biology.)

Humans influence tropical rainforest animals directly via exploitation and indirectly via habitat disturbance. Bushmeat hunting and logging occur extensively in tropical forests and have large effects on particular species. But how they alter animal diversity across landscape scales and whether their impacts are correlated across species remain less known. We used spatially widespread measurements of mammal occurrence across Malaysian Borneo and recently developed multispecies hierarchical models to assess the species richness of medium- to large-bodied terrestrial mammals while accounting for imperfect detection of all species. Hunting was associated with 31% lower species richness. Moreover, hunting remained high even where richness was very low, highlighting that hunting pressure persisted even in chronically overhunted areas. Newly logged sites had 11% lower species richness than unlogged sites, but sites logged >10 years previously had richness levels similar to those in old-growth forest. Hunting was a more serious long-term threat than logging for 91% of primate and ungulate species. Hunting and logging impacts across species were not correlated across taxa. Negative impacts of hunting were the greatest for common mammalian species, but commonness versus rarity was not related to species-specific impacts of logging. Direct human impacts appeared highly persistent and lead to defaunation of certain areas. These impacts were particularly severe for species of ecological importance as seed dispersers and herbivores. Indirect impacts were also strong but appeared to attenuate more rapidly than previously thought. The lack of correlation between direct and indirect impacts across species highlights that multifaceted conservation strategies may be needed for mammal conservation in tropical rainforests, Earth's most biodiverse ecosystems., (© 2014 Society for Conservation Biology.)

Extinctions beget further extinctions when species lose obligate mutualists, predators, prey, or hosts. Here, we develop a conceptual model of species and community attributes affecting secondary extinction likelihood, incorporating mechanisms that buffer organisms against partner loss. Specialized interactors, including 'cryptic specialists' with diverse but nonredundant partner assemblages, incur elevated risk. Risk is also higher for species that cannot either evolve new traits following partner loss or obtain novel partners in communities reorganizing under changing environmental conditions. Partner loss occurs alongside other anthropogenic impacts; multiple stressors can circumvent ecological buffers, enhancing secondary extinction risk. Stressors can also offset each other, reducing secondary extinction risk, a hitherto unappreciated phenomenon. This synthesis suggests improved conservation planning tactics and critical directions for research on secondary extinctions., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

After a decade with nine of the lowest arctic sea-ice minima on record, including the historically low minimum in 2012, we synthesize recent developments in the study of ecological responses to sea-ice decline. Sea-ice loss emerges as an important driver of marine and terrestrial ecological dynamics, influencing productivity, species interactions, population mixing, gene flow, and pathogen and disease transmission. Major challenges in the near future include assigning clearer attribution to sea ice as a primary driver of such dynamics, especially in terrestrial systems, and addressing pressures arising from human use of arctic coastal and near-shore areas as sea ice diminishes.

Behaviorally mediated trophic cascades (BMTCs) occur when the fear of predation among herbivores enhances plant productivity. Based primarily on systems involving small-bodied predators, BMTCs have been proposed as both strong and ubiquitous in natural ecosystems. Recently, however, synthetic work has suggested that the existence of BMTCs may be mediated by predator hunting mode, whereby passive (sit-and-wait) predators have much stronger effects than active (coursing) predators. One BMTC that has been proposed for a wide-ranging active predator system involves the reintroduction of wolves (Canis lupus) to Yellowstone National Park, USA, which is thought to be leading to a recovery of trembling aspen (Populus tremuloides) by causing elk (Cervus elaphus) to avoid foraging in risky areas. Although this BMTC has been generally accepted and highly popularized, it has never been adequately tested. We assessed whether wolves influence aspen by obtaining detailed demographic data on aspen Stands using tree rings and by monitoring browsing levels in experimental elk exclosures arrayed across a gradient of predation risk for three years. Our study demonstrates that the historical failure of aspen to regenerate varied widely among stands (last recruitment year ranged from 1892 to 1956), and our data do not indicate an abrupt cessation of recruitment. This pattern of recruitment failure appears more consistent with a gradual increase in elk numbers rather than a rapid behavioral shift in elk foraging following wolf extirpation. In addition, our estimates of relative survivorship of young browsable aspen indicate that aspen are not currently recovering in Yellowstone, even in the presence of a large wolf population. Finally, in an experimental test of the BMTC hypothesis we found that the impacts of elk browsing on aspen demography are not diminished in sites where elk are at higher risk of predation by wolves. These findings suggest the need to further evaluate how trophic cascades are mediated by predator-prey life history and ecological context.

Myriad tropical vertebrates are threatened by overharvest. Whether this harvest has indirect effects on nonhunted organisms that interact with the game species is a critical question. Many tropical birds and mammals disperse seeds. Their overhunting in forests can cause zoochorous trees to suffer from reduced seed dispersal. Yet how these reductions in seed dispersal influence tree abundance and population dynamics remains unclear. Reproductive parameters in long-lived organisms often have very low elasticities; indeed the demographic importance of seed dispersal is an open question. We asked how variation in hunting pressure across four national parks with seasonal forest in northern Thailand influenced the relative abundance of gibbons, muntjac deer, and sambar deer, the sole dispersers of seeds of the canopy tree Choerospondias axillaris. We quantified how variation in disperser numbers affected C. axillaris seed dispersal and seedling abundance across the four parks. We then used these data in a structured population model based on vital rates measured in Khao Yai National Park (where poaching pressure is minimal) to explore how variation in illegal hunting pressure might influence C. axillaris population growth and persistence. Densities of the mammals varied strongly across the parks, from relatively high in Khao Yai to essentially zero in Doi Suthep-Pui. Levels of C. axillaris seed dispersal and seedling abundance positively tracked mammal density. If hunting in Khao Yai were to increase to the levels seen in the other parks, C. axillaris population growth rate would decline, but only slightly. Extinction of C. axillaris is a real possibility, but may take many decades. Recent and ongoing extirpations of vertebrates in many tropical forests could be creating an extinction debt for zoochorous trees whose vulnerability is belied by their current abundance.

Many plants interact with groups of mutualist pollinators and seed dispersers. A key issue for both basic ecology and conservation is whether the different species within these guilds of mutualist animals are functionally equivalent. Comparing the relative effects of sympatric mutualists is important for understanding the evolution of multispecies mutualisms and for predicting mutualism stability in the face of anthropogenic change. However, empirical comparisons of the population-level impacts of mutualist animals on their host plant are rare, particularly for seed dispersal mutualisms in species-rich ecosystems. We compared the influence of three seed-dispersing tropical mammals, lar gibbons (Hylobates lar), sambar deer (Rusa unicolor), and red muntjac deer (Muntiacus muntjak), on the demography of a shared host tree in Thailand, Choerospondias axillaris (Anacardiaceae). Sambar and muntjac dispersed far more C. axillaris seeds than did gibbons. While sambar deposited many seeds under female tree canopies, muntjac were the only disperser to move seeds to open microhabitats, where C. axillaris seed germination, seedling survival, and initial growth are enhanced. Using stage-based population models, we assessed how disperser-specific seed dispersal, variation in the frequency of canopy gap formation, and their interaction influenced the potential population growth of C. axillaris. Large differences in dispersal quantity and small differences in dispersal quality among sambar and gibbons resulted in similar and negligible impacts on the tree's population dynamics. Muntjac, by taking some of the seeds to open microhabitats, are projected to have a greater positive impact on C. axillaris demography than either sambar or gibbons. Model comparisons of population-level species impacts may allow us to predict which ecological interactions are at risk from loss of critical species.