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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.)

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.

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.

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.

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.)

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).)

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.)