Maintenance of different life stages of old-growth forest trees in deforested tropical landscapes.

Tropical tree species are increasingly being pushed to inhabit deforested landscapes. The habitat amount hypothesis posits that, in remaining forest patches, species diversity in equal-sized samples decreases with decreasing forest cover in the surrounding landscape. We tested this prediction by taking into account three important factors that can affect species responses to forest loss. First, forest loss effects can be linear (proportional) or nonlinear, as there can be threshold values of forest loss beyond which species extirpation may be accelerated. Second, such effects are usually scale dependent and may go unnoticed if assessed at suboptimal scales. Finally, species extirpation may take decades to become evident, so the effects of forest loss can be undetected when assessing long-lived organisms, like adult old-growth forest trees. Here, we evaluated the linear and nonlinear effects of landscape forest loss across different spatial scales on site-scale abundance and diversity of old-growth forest trees, separately for four plant-life stages (seeds, saplings, juveniles, and adults) in two rainforest regions with different levels of deforestation. We expected stronger (and negative) forest loss effects on early plant-life stages, especially in the region with the highest deforestation. Surprisingly, in 13 of 16 study cases (2 responses × 4 life stages × 2 regions), null models showed higher empirical support than linear and nonlinear models at any scale. Therefore, the species richness and abundance of local tree assemblages seem to be weakly affected by landscape-scale forest loss independently of the spatial scale, life stage, and region. Yet, as expected, the predictive power of forest cover was relatively lower in the least deforested region. Our findings suggest that landscape-scale forest loss is poorly related to site-scale processes, such as seed dispersal and seedling recruitment, or, at least, such effects are too small to shape the abundance and diversity of tree assemblages within forest patches. Therefore, our findings do not support the most important prediction of the habitat amount hypothesis but imply that, on a per-area basis, a unit of habitat (forest) in a highly deforested landscape has a conservation value similar to that of a more forested one, particularly in moderately deforested rainforests.

Underlying and proximate drivers of biodiversity changes in Mesoamerican biosphere reserves.

Protected areas are of paramount relevance to conserving wildlife and ecosystem contributions to people. Yet, their conservation success is increasingly threatened by human activities including habitat loss, climate change, pollution, and species overexploitation. Thus, understanding the underlying and proximate drivers of anthropogenic threats is urgently needed to improve protected areas' effectiveness, especially in the biodiversity-rich tropics. We addressed this issue by analyzing expert-provided data on long-term biodiversity change (last three decades) over 14 biosphere reserves from the Mesoamerican Biodiversity Hotspot. Using multivariate analyses and structural equation modeling, we tested the influence of major socioeconomic drivers (demographic, economic, and political factors), spatial indicators of human activities (agriculture expansion and road extension), and forest landscape modifications (forest loss and isolation) as drivers of biodiversity change. We uncovered a significant proliferation of disturbance-tolerant guilds and the loss or decline of disturbance-sensitive guilds within reserves causing a "winner and loser" species replacement over time. Guild change was directly related to forest spatial changes promoted by the expansion of agriculture and roads within reserves. High human population density and low nonfarming occupation were identified as the main underlying drivers of biodiversity change. Our findings suggest that to mitigate anthropogenic threats to biodiversity within biosphere reserves, fostering human population well-being via sustainable, nonfarming livelihood opportunities around reserves is imperative.

Landscape-scale forest cover drives the predictability of forest regeneration across the Neotropics.

Abandonment of agricultural lands promotes the global expansion of secondary forests, which are critical for preserving biodiversity and ecosystem functions and services. Such roles largely depend, however, on two essential successional attributes, trajectory and recovery rate, which are expected to depend on landscape-scale forest cover in nonlinear ways. Using a multi-scale approach and a large vegetation dataset (843 plots, 3511 tree species) from 22 secondary forest chronosequences distributed across the Neotropics, we show that successional trajectories of woody plant species richness, stem density and basal area are less predictable in landscapes (4 km radius) with intermediate (40-60%) forest cover than in landscapes with high (greater than 60%) forest cover. This supports theory suggesting that high spatial and environmental heterogeneity in intermediately deforested landscapes can increase the variation of key ecological factors for forest recovery (e.g. seed dispersal and seedling recruitment), increasing the uncertainty of successional trajectories. Regarding the recovery rate, only species richness is positively related to forest cover in relatively small (1 km radius) landscapes. These findings highlight the importance of using a spatially explicit landscape approach in restoration initiatives and suggest that these initiatives can be more effective in more forested landscapes, especially if implemented across spatial extents of 1-4 km radius.

Tropical forest loss impoverishes arboreal mammal assemblages by increasing tree canopy openness.

Landscape-scale deforestation poses a major threat to global biodiversity, not only because it limits habitat availability, but also because it can drive the degradation of the remaining habitat. However, the multiple pathways by which deforestation directly and indirectly affects wildlife remain poorly understood, especially for elusive forest-dependent species such as arboreal mammals. Using structural equation models, we assessed the direct and indirect effects of landscape forest loss on arboreal mammal assemblages in the Lacandona rainforest, Mexico. We placed camera traps in 100 canopy trees, and assessed the direct effect of forest cover and their indirect effects via changes in tree basal area and canopy openness on the abundance and diversity (i.e., species richness and exponential of Shannon entropy) of arboreal mammals. We found that forest loss had negative indirect effects on mammal richness through the increase of tree canopy openness. This could be related to the fact that canopy openness is usually inversely related to resource availability and canopy connectivity for arboreal mammals. Furthermore, independently of forest loss, the abundance and richness of arboreal mammals was positively related to tree basal area, which is typically higher in old-growth forests. Thus, our findings suggest that arboreal mammals generally prefer old-growth vegetation with relatively low canopy openness and high tree basal area. However, unexpectedly, forest loss was directly and positively related to the abundance and richness of mammals, probably due to a crowding effect, a reasonable possibility given the relatively short history (~40 years) of deforestation in the study region. Conversely, the Shannon diversity was not affected by the predictors we evaluated, suggesting that rare mammals (not the common species) are the ones most affected by these changes. All in all, our findings emphasize that conservation measures ought to focus on increasing forest cover in the landscape, and preventing the loss of large trees in the remaining forest patches.

Homogenization of terrestrial mammals in fragmented rainforests: the loss of species turnover and its landscape drivers.

Understanding the factors and mechanisms shaping differences in species composition across space and time (ß-diversity) in human-modified landscapes has key ecological and applied implications. This topic is, however, challenging because landscape disturbance can promote either decreases (biotic homogenization) or increases (biotic differentiation) in ß-diversity. We assessed temporal differences in intersite ß-diversity of medium-bodied and large-bodied mammals in the fragmented Lacandona rainforest, Mexico. We hypothesized that, given the relatively short history of land-use changes in the region, and the gain and loss of some species caused by landscape spatial changes, ß-diversity would increase through time, especially its nestedness component. We estimated ß-diversity between 24 forest sites (22 forest patches and two continuous forest sites) in 2011 and 2017 to assess whether ß-diversity is decreasing or increasing in the region, and calculated its turnover and nestedness components to understand the mechanisms responsible for changes in ß-diversity, separately assessing mammal groups with different body mass, feeding guild, and habitat specialization. We then related such temporal changes in ß-diversity to temporal changes in five landscape variables (forest cover, matrix openness, number of patches, edge density and interpatch distance) to identify the landscape drivers of ß-diversity. In contrast with our expectations, ß-diversity decreased over time, suggesting an ongoing biotic homogenization process. This pattern was mostly driven by a decrease in species turnover in all mammal groups, especially in landscapes with decreasing forest cover and increasing forested matrices. Although the nestedness component showed a three-fold increase through time, species turnover was 22 and six times higher than nestedness in 2011 and 2017, respectively. The decreased turnover appears to be driven by an increase in dispersal (i.e., spillover) of native species among patches. The prevalence of species turnover over nestedness indicates that different forest sites have a fairly distinct subset of species (i.e., high complementarity in species composition). Therefore, conserving all remaining forest patches and increasing forest cover is of utmost importance to effectively maintain ß-diversity and conserve the total diversity (γ) of mammal assemblages in this Mesoamerican biodiversity hotspot.

Resolving the SLOSS dilemma for biodiversity conservation: a research agenda.

The legacy of the 'SL > SS principle', that a single or a few large habitat patches (SL) conserve more species than several small patches (SS), is evident in decisions to protect large patches while down-weighting small ones. However, empirical support for this principle is lacking, and most studies find either no difference or the opposite pattern (SS > SL). To resolve this dilemma, we propose a research agenda by asking, 'are there consistent, empirically demonstrated conditions leading to SL > SS?' We first review and summarize 'single large or several small' (SLOSS) theory and predictions. We found that most predictions of SL > SS assume that between-patch variation in extinction rate dominates the outcome of the extinction-colonization dynamic. This is predicted to occur when populations in separate patches are largely independent of each other due to low between-patch movements, and when species differ in minimum patch size requirements, leading to strong nestedness in species composition along the patch size gradient. However, even when between-patch variation in extinction rate dominates the outcome of the extinction-colonization dynamic, theory can predict SS > SL. This occurs if extinctions are caused by antagonistic species interactions or disturbances, leading to spreading-of-risk of landscape-scale extinction across SS. SS > SL is also predicted when variation in colonization dominates the outcome of the extinction-colonization dynamic, due to higher immigration rates for SS than SL, and larger species pools in proximity to SS than SL. Theory that considers change in species composition among patches also predicts SS > SL because of higher beta diversity across SS than SL. This results mainly from greater environmental heterogeneity in SS due to greater variation in micro-habitats within and across SS habitat patches ('across-habitat heterogeneity'), and/or more heterogeneous successional trajectories across SS than SL. Based on our review of the relevant theory, we develop the 'SLOSS cube hypothesis', where the combination of three variables - between-patch movement, the role of spreading-of-risk in landscape-scale population persistence, and across-habitat heterogeneity - predict the SLOSS outcome. We use the SLOSS cube hypothesis and existing SLOSS empirical evidence, to predict SL > SS only when all of the following are true: low between-patch movement, low importance of spreading-of-risk for landscape-scale population persistence, and low across-habitat heterogeneity. Testing this prediction will be challenging, as it will require many studies of species groups and regions where these conditions hold. Each such study would compare gamma diversity across multiple landscapes varying in number and sizes of patches. If the prediction is not generally supported across such tests, then the mechanisms leading to SL > SS are extremely rare in nature and the SL > SS principle should be abandoned.

Preserving 40% forest cover is a valuable and well-supported conservation guideline: reply to Banks-Leite et al.

Banks-Leite et al. (2021) claim that our suggestion of preserving ≥ 40% forest cover lacks evidence and can be problematic. We find these claims unfounded, and discuss why conservation planning urgently requires valuable, well-supported and feasible general guidelines like the 40% criterion. Using region-specific thresholds worldwide is unfeasible and potentially harmful.

Corrigendum: A Severe Lack of Evidence Limits Effective Conservation of the World's Primates.

[This corrects the article DOI: 10.1093/biosci/biaa082.].

A Severe Lack of Evidence Limits Effective Conservation of the World's Primates.

Threats to biodiversity are well documented. However, to effectively conserve species and their habitats, we need to know which conservation interventions do (or do not) work. Evidence-based conservation evaluates interventions within a scientific framework. The Conservation Evidence project has summarized thousands of studies testing conservation interventions and compiled these as synopses for various habitats and taxa. In the present article, we analyzed the interventions assessed in the primate synopsis and compared these with other taxa. We found that despite intensive efforts to study primates and the extensive threats they face, less than 1% of primate studies evaluated conservation effectiveness. The studies often lacked quantitative data, failed to undertake postimplementation monitoring of populations or individuals, or implemented several interventions at once. Furthermore, the studies were biased toward specific taxa, geographic regions, and interventions. We describe barriers for testing primate conservation interventions and propose actions to improve the conservation evidence base to protect this endangered and globally important taxon.

Designing optimal human-modified landscapes for forest biodiversity conservation.

Agriculture and development transform forest ecosystems to human-modified landscapes. Decades of research in ecology have generated myriad concepts for the appropriate management of these landscapes. Yet, these concepts are often contradictory and apply at different spatial scales, making the design of biodiversity-friendly landscapes challenging. Here, we combine concepts with empirical support to design optimal landscape scenarios for forest-dwelling species. The supported concepts indicate that appropriately sized landscapes should contain ≥ 40% forest cover, although higher percentages are likely needed in the tropics. Forest cover should be configured with c. 10% in a very large forest patch, and the remaining 30% in many evenly dispersed smaller patches and semi-natural treed elements (e.g. vegetation corridors). Importantly, the patches should be embedded in a high-quality matrix. The proposed landscape scenarios represent an optimal compromise between delivery of goods and services to humans and preserving most forest wildlife, and can therefore guide forest preservation and restoration strategies.

Support for the habitat amount hypothesis from a global synthesis of species density studies.

Decades of research suggest that species richness depends on spatial characteristics of habitat patches, especially their size and isolation. In contrast, the habitat amount hypothesis predicts that (1) species richness in plots of fixed size (species density) is more strongly and positively related to the amount of habitat around the plot than to patch size or isolation; (2) habitat amount better predicts species density than patch size and isolation combined, (3) there is no effect of habitat fragmentation per se on species density and (4) patch size and isolation effects do not become stronger with declining habitat amount. Data on eight taxonomic groups from 35 studies around the world support these predictions. Conserving species density requires minimising habitat loss, irrespective of the configuration of the patches in which that habitat is contained.

Regional context and dispersal mode drive the impact of landscape structure on seed dispersal.

Land-use change modifies the spatial structure of tropical landscapes, shaping global biodiversity patterns. Yet, it remains unknown how key ecological processes, such as seed dispersal, can be affected by changes in landscape patterns, and whether such effects differ among regions with different climate and disturbance intensity. We assessed the effect of five landscape metrics (forest cover, matrix openness, forest edge density, forest fragmentation, and interpatch distance) on the seed rain recorded in two Mexican fragmented regions (20 forest sites per region): the more deforested, defaunated, and windy Los Tuxtlas rainforest (LTX), and the better-preserved Lacandona rainforest (LAC). We quantified the proportions of dispersed tree species and their seeds, separately evaluating wind- and animal-dispersed species. Our findings support the hypothesis that forest loss is more important than fragmentation per se, negatively impacting the seed rain in both regions. As expected, landscape patterns were comparatively more important for wind-dispersed seeds in LTX, probably because of stronger wind events in this region. Specifically, proportions of wind-dispersed seeds and species decreased with increasing edge density, suggesting that forest edges prevent dispersal of wind-dispersed species, which may occur if edges create physical barriers that limit wind flow. This pattern can also be caused by source limitation in landscapes with more forest edges, as tree mortality rates usually increase at forest edges. The wind-dispersed seed rain was also positively related to matrix openness, especially in LTX, where wind flow can be favored by the dominance of treeless anthropogenic matrices. Surprisingly, the proportion of animal-dispersed seeds in LTX was positively related to matrix openness and patch isolation, suggesting that seed dispersers in more deforested regions may be forced to concentrate in isolated patches and use the available habitat more intensively. Yet, as expected, patch isolation limited wind-dispersed seeds in LAC. Therefore, dispersal (and potentially regeneration) of wind-dispersed trees is favored in regions exposed to stronger wind events, especially in landscapes dominated by regularly shaped patches surrounded by open areas. Conversely, animal-dispersed seeds are primarily favored by increasing forest cover. Preventing forest loss is therefore critical to promote animal seed dispersal and forest recovery in human-modified rainforests.

Extinction filters mediate the global effects of habitat fragmentation on animals.

Habitat loss is the primary driver of biodiversity decline worldwide, but the effects of fragmentation (the spatial arrangement of remaining habitat) are debated. We tested the hypothesis that forest fragmentation sensitivity-affected by avoidance of habitat edges-should be driven by historical exposure to, and therefore species' evolutionary responses to disturbance. Using a database containing 73 datasets collected worldwide (encompassing 4489 animal species), we found that the proportion of fragmentation-sensitive species was nearly three times as high in regions with low rates of historical disturbance compared with regions with high rates of disturbance (i.e., fires, glaciation, hurricanes, and deforestation). These disturbances coincide with a latitudinal gradient in which sensitivity increases sixfold at low versus high latitudes. We conclude that conservation efforts to limit edges created by fragmentation will be most important in the world's tropical forests.

A global assessment of primate responses to landscape structure.

Land-use change modifies the spatial structure of terrestrial landscapes, potentially shaping the distribution, abundance and diversity of remaining species assemblages. Non-human primates can be particularly vulnerable to landscape disturbances, but our understanding of this topic is far from complete. Here we reviewed all available studies on primates' responses to landscape structure. We found 34 studies of 71 primate species (24 genera and 10 families) that used a landscape approach. Most studies (82%) were from Neotropical forests, with howler monkeys being the most frequently studied taxon (56% of studies). All studies but one used a site-landscape or a patch-landscape study design, and frequently (34% of studies) measured landscape variables within a given radius from the edge of focal patches. Altogether, the 34 studies reported 188 responses to 17 landscape-scale metrics. However, the majority of the studies (62%) quantified landscape predictors within a single spatial scale, potentially missing significant primate-landscape responses. To assess such responses accurately, landscape metrics need to be measured at the optimal scale, i.e. the spatial extent at which the primate-landscape relationship is strongest (so-called 'scale of effect'). Only 21% of studies calculated the scale of effect through multiscale approaches. Interestingly, the vast majority of studies that do not assess the scale of effect mainly reported null effects of landscape structure on primates, while most of the studies based on optimal scales found significant responses. These significant responses were primarily to landscape composition variables rather than landscape configuration variables. In particular, primates generally show positive responses to increasing forest cover, landscape quality indices and matrix permeability. By contrast, primates show weak responses to landscape configuration. In addition, half of the studies showing significant responses to landscape configuration metrics did not control for the effect of forest cover. As configuration metrics are often correlated with forest cover, this means that documented configuration effects may simply be driven by landscape-scale forest loss. Our findings suggest that forest loss (not fragmentation) is a major threat to primates, and thus, preventing deforestation (e.g. through creation of reserves) and increasing forest cover through restoration is critically needed to mitigate the impact of land-use change on our closest relatives. Increasing matrix functionality can also be critical, for instance by promoting anthropogenic land covers that are similar to primates' habitat.

Species sorting and mass effect along forest succession: Evidence from taxonomic, functional, and phylogenetic diversity of amphibian communities.

Species recovery after forest disturbance is a highly studied topic in the tropics, but considerable debate remains on the role of secondary forests as biodiversity repositories, especially regarding the functional and phylogenetic dimensions of biodiversity. Also, studies generally overlook how alpha and beta diversities interact to produce gamma diversity along successional gradients.We used a metacommunity approach to assess how species sorting (i.e., environmental filtering) and mass effect (i.e., source-sink dynamics) affect 14 complementary metrics of amphibian taxonomic, functional, and phylogenetic diversity along a successional gradient in southern Mexico. As amphibians have narrow environmental tolerances and low dispersal capabilities, we expected that species sorting may be relatively more important than mass effect in structuring amphibian communities.Between 2010 and 2012, we sampled frogs, salamanders, and caecilians in 23 communities distributed in four successional stages: young (2-5 years old) and intermediate (13-28 years old) secondary forests, old-growth forest fragments, and old-growth continuous forest. We assessed 15 ecologically relevant functional traits per species and used a time-calibrated molecular phylogeny.We recorded 1,672 individuals belonging to 30 species and 11 families. Supporting our expectations from the species sorting perspective, from the poorest (younger forests) to the best quality (continuous forest) scenarios, we observed (a) an increase in alpha diversity regardless of species abundances; (b) a clear taxonomic segregation across successional stages; (c) an increase in functional richness and dispersion; (d) an increase in mean phylogenetic distance and nearest taxon index; and (e) a reduction in mean nearest taxon distance. However, 10 species occurred in all successional stages, resulting in relatively low beta diversity. This supports a mass effect, where interpatch migrations contribute to prevent local extinctions and increase compositional similarity at the regional scale.Our findings indicate that amphibian metacommunities along forest successional gradients are mainly structured by species sorting, but mass effects may also play a role if high levels of forest cover are conserved in the region. In fact, secondary forests and forest fragments can potentially safeguard different aspects of amphibian diversity, but their long-term conservation value requires preventing additional deforestation.

Ecological traits of the world's primates.

Ecosystems largely depend, for both their functioning and their ecological integrity, on the ecological traits of the species that inhabit them. Non-human primates have a wide geographic distribution and play vital roles in ecosystem structure, function, and resilience. However, there is no comprehensive and updated compilation of information on ecological traits of all the world's primate species to accurately assess such roles at a global scale. Here we present a database on some important ecological traits of the world's primates (504 species), including home range size, locomotion type, diel activity, trophic guild, body mass, habitat type, current conservation status, population trend, and geographic realm. We compiled this information through a careful review of 1,216 studies published between 1941 and 2018, resulting in a comprehensive, easily accessible and user-friendly database. This database has broad applicability in primatological studies, and can potentially be used to address many research questions at all spatial scales, from local to global.

The conservation value of human-modified landscapes for the world's primates.

Land-use change pushes biodiversity into human-modified landscapes, where native ecosystems are surrounded by anthropic land covers (ALCs). Yet, the ability of species to use these emerging covers remains poorly understood. We quantified the use of ALCs by primates worldwide, and analyzed species' attributes that predict such use. Most species use secondary forests and tree plantations, while only few use human settlements. ALCs are used for foraging by at least 86 species with an important conservation outcome: those that tolerate heavily modified ALCs are 26% more likely to have stable or increasing populations than the global average for all primates. There is no phylogenetic signal in ALCs use. Compared to all primates on Earth, species using ALCs are less often threatened with extinction, but more often diurnal, medium or large-bodied, not strictly arboreal, and habitat generalists. These findings provide valuable quantitative information for improving management practices for primate conservation worldwide.

Direct and cascading effects of landscape structure on tropical forest and non-forest frugivorous birds.

Global biodiversity is increasingly threatened by land-use change, but the direct and indirect drivers of species diversity in human-modified tropical landscapes are poorly known. Forest-dependent species are expected to be particularly sensitive to changes in landscape composition (e.g., forest loss) and configuration (e.g., increase of forest edges), both directly and indirectly through cascading landscape effects on local patterns of forest structure and resource availability. In contrast, non-forest-dependent species are probably more strongly related to landscape changes than to local forest patterns, as these species are able to use resources not only from the forest, but also from other landscape elements over larger spatial scales. We tested these hypotheses using structural equation modeling. In particular, we sampled 20 landscapes (115 ha each) from the Brazilian Atlantic rainforest to assess the effect of landscape-scale forest cover and amount of forest edges on the diversity of frugivorous birds, both directly and indirectly through the effect that these landscape variables may have on vegetation complexity and fruit biomass. We separately assessed the response of forest-dependent and non-forest-dependent frugivores to infer potential mechanisms underlying bird assemblages in fragmented landscapes. The diversity of forest-dependent birds mainly decreased with the simplification of vegetation complexity in more deforested landscapes, but increased with increasing fruit biomass in more forested landscapes (indirect effects). Both patterns were significant, thus supporting a strong bottom-up control, i.e., local habitat simplification and resource scarcity in highly deforested landscapes limits the maintenance of forest-dependent birds. Conversely, but as expected, non-forest-dependent birds were more strongly and directly related to landscape-scale patterns. In particular, landscapes with higher forest edge amount showed higher bird species diversity, probably because the increasing length of ecotones and interspersion/juxtaposition of different habitat types in landscapes with more forest edges can increase resource availability and foraging efficiency of non-forest-dependent birds. As the seed dispersal services offered by forest-dependent species cannot be ecologically compensated for by the proliferation of non-forest-dependent species, preventing forest loss is imperative to maintain forest-dependent birds and forest regeneration in this vanishing biodiversity hotspot.

Integrating expert knowledge and ecological niche models to estimate Mexican primates' distribution.

Ecological niche modeling is used to estimate species distributions based on occurrence records and environmental variables, but it seldom includes explicit biotic or historical factors that are important in determining the distribution of species. Expert knowledge can provide additional valuable information regarding ecological or historical attributes of species, but the influence of integrating this information in the modeling process has been poorly explored. Here, we integrated expert knowledge in different stages of the niche modeling process to improve the representation of the actual geographic distributions of Mexican primates (Ateles geoffroyi, Alouatta pigra, and A. palliata mexicana). We designed an elicitation process to acquire information from experts and such information was integrated by an iterative process that consisted of reviews of input data by experts, production of ecological niche models (ENMs), and evaluation of model outputs to provide feedback. We built ENMs using the maximum entropy algorithm along with a dataset of occurrence records gathered from a public source and records provided by the experts. Models without expert knowledge were also built for comparison, and both models, with and without expert knowledge, were evaluated using four validation metrics that provide a measure of accuracy for presence-absence predictions (specificity, sensitivity, kappa, true skill statistic). Integrating expert knowledge to build ENMs produced better results for potential distributions than models without expert knowledge, but a much greater improvement in the transition from potential to realized geographic distributions by reducing overprediction, resulting in better representations of the actual geographic distributions of species. Furthermore, with the combination of niche models and expert knowledge we were able to identify an area of sympatry between A. palliata mexicana and A. pigra. We argue that the inclusion of expert knowledge at different stages in the construction of niche models in an explicit and systematic fashion is a recommended practice as it produces overall positive results for representing realized species distributions.

Fragmentation and matrix contrast favor understory plants through negative cascading effects on a strong competitor palm.

Understanding the patterns and processes driving biodiversity maintenance in fragmented tropical forests is urgently needed for conservation planning, especially in species-rich forest reserves. Of particular concern are the effects that habitat modifications at the landscape scale may have on forest regeneration and ecosystem functioning: a topic that has received limited attention. Here, we assessed the effects of landscape structure (i.e., forest cover, open area matrices, forest fragmentation, and mean inter-patch isolation distance) on understory plant assemblages in the Los Tuxtlas Biosphere Reserve, Mexico. Previous studies suggest that the demographic burst of the strong competitor palm Astrocaryum mexicanum in the core area of this reserve limits plant recruitment and imperils biodiversity conservation within this protected area. Yet, the local and landscape predictors of this palm, and its impact on tree recruitment at a regional scale are unknown. Thus, we used structural equation modeling to assess the direct and cascading effects of landscape structure on stem and species density in the understory of 20 forest sites distributed across this biodiversity hotspot. Indirect paths included the effect of landscape structure on tree basal area (a proxy of local disturbance), and the effects of these variables on A. mexicanum. Density of A. mexicanum mainly increased with decreasing both fragmentation and open areas in the matrix (matrix contrast, hereafter), and such an increase in palm density negatively affected stem and species density in the understory. The negative direct effect of matrix contrast on stem density was overridden by the indirect positive effects (i.e., through negative cascading effects on A. mexicanum), resulting in a weak effect of matrix contrast on stem density. These findings suggest that dispersal limitation and negative edge effects in more fragmented landscapes dominated by open areas prevent the proliferation of this palm species, enhancing the diversity and abundance of understory trees. This "positive" news adds to an increasing line of evidence suggesting that fragmentation may have some positive effects on biodiversity, in this case by preventing the proliferation of species that can jeopardize biodiversity conservation within tropical reserves.