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.
Analysis of a global set of 35 studies suggests that habitat amount, rather than patch area, isolation or fragmentation per se, determined species richness in sample plots at scales ranging from 13 to 11 000 ha. Minimising species losses requires protecting and restoring as much habitat as possible, irrespective of the configuration of that habitat.
1. Land-use change is the main driver of global biodiversity loss, but its relative impact on species turnover (β-diversity) across multiple spatial scales remains unclear. Plant communities in ...fragmented rain forests can undergo declines (floristic homogenization) or increases (floristic differentiation) in β-diversity. 2. We tested these alternative hypotheses analysing a large vegetation data base from a hierarchically nested sampling design (450 plots in 45 forest patches in 3 landscapes with different deforestation levels) at Los Tuxtlas rain forest, Mexico. Differences in β-diversity across spatial scales (i.e. among plots, among patches, and among landscapes) were analysed using multiplicative diversity decompositions of Hill numbers. 3. Plant β-diversity among plots within forest patches decreased in landscapes with higher deforestation levels, leading to floristic homogenization within patches. This homogenization process can be explained by the loss of rare and shade-tolerant plant species, and the recruitment and dominance of disturbance-adapted species, and can limit the accumulation of species (γ-diversity) in landscapes with higher deforestation. 4. Nevertheless, the landscape with the highest deforestation level showed the highest floristic differentiation among patches. This landscape showed the greatest isolation distances between patches; a landscape spatial pattern that can limit the interchange of seeds (and species) between patches. Because the study patches are undergoing secondary succession following disturbances (e.g. logging, edge effects), different disturbance regimes and increased distance among patches could lead to higher β-diversity. 5. Synthesis. These findings indicate that patterns of floristic homogenization and differentiation depend on the landscape configuration and on the spatial scale of analysis. At the landscape scale, our results suggest that, in accordance with non-equilibrium dynamics and the landscape-divergence hypothesis, patches located in landscapes with different forest cover and different connectivity can experience contrasting successional pathways due to increasing levels of compositional differentiation between patches. These novel findings add further uncertainties to the maintenance of biodiversity in severely deforested tropical landscapes and have key ecological implications for biodiversity conservation planning.
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.
ABSTRACT
Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified ...tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.
1. Seasonally dry tropical forests (SDTFs) are one of the most threatened forests world-wide. These species-rich forests not only cope with several acute (e.g. forest loss) and chronic (e.g. ...overgrazing and firewood extraction) human disturbances but also with climate change (e.g. longer and more severe droughts); yet, the isolated and combined effects of climate and acute and chronic human disturbances on SDTF vegetation are poorly known. 2. Given the environmental filter imposed by drought in SDTFs, the composition and structure of vegetation is expected to be strongly associated with annual precipitation, and thus the effects of human disturbances on vegetation may also depend on precipitation (i.e. interacting effect). 3. We tested these hypotheses in the Brazilian Caatinga – a SDTF threatened by climate change and human disturbances. We evaluated the isolated and combined (both additive and multiplicative) effect of precipitation, a chronic disturbance index and acute disturbance (landscape forest cover) on the diversity, stem density, evenness, taxonomic composition and above-ground biomass of adult trees and shrubs across 19 0·1-ha plots distributed along a disturbance and precipitation gradients. 4. We recorded 5541 stems from 129 species. Precipitation showed a stronger (positive) effect on species diversity than acute and chronic disturbances and, as expected, the effect of disturbance depended on precipitation (interacting effect): that is, species diversity (especially the number of rare species) was negatively related to forest loss but positively related to chronic disturbance in wetter sites, whereas in drier sites, species diversity was weakly related to forest cover, but strongly and negatively related to chronic disturbance. Contrary to species diversity, community evenness, stem density and biomass were weakly related to all predictors. 5. Synthesis. Precipitation appears to be a strong environmental filter determining the distribution of water-demanding plant species. Chronic disturbance in wetter (high-productive) forests may favour species diversity by increasing ecosystem heterogeneity (intermediate disturbance hypothesis). Yet, the biodiversity costs of chronic disturbance are higher in drier (low-productive) forests; that is, there is a co-limitation imposed by drought and disturbance in drier forests. Overall, our findings indicate that rapid climatic changes in the region will probably have strong negative effects on this seasonally dry tropical forest.
1. Secondary forests are increasingly dominant in human-modified tropical landscapes, but the drivers of forest recovery remain poorly understood. Soil conditions influence plant community ...composition, and are expected to change over a gradient of succession. However, the role of soil conditions as an environmental filter driving community assembly during forest succession has rarely been explicitly assessed. 2. We evaluated the role of stand basal area and soil conditions on community assembly and its consequences for community functional properties along a chronosequence of Atlantic forest regeneration following sugar cane cultivation. Specifically, we tested whether community functional properties are related to stand basal area, soil fertility and soil moisture. Our expectations were that edaphic environmental filters play an increasingly important role along secondary succession by increasing functional trait convergence towards more conservative attributes. 3. We sampled soil and woody vegetation features across 15 second-growth (3-30 years) and 11 old-growth forest plots (300 m² each). We recorded tree functional traits related to resource-use strategies (specific leaf area, SLA; leaf dry matter content, LDMC; leaf area, LA; leaf thickness, LT; and leaf succulence, LS) and calculated community functional properties using the community-weighted mean (CWM) of each trait and the functional dispersion (FDis) of each trait separately and all traits together. 4. With exception of LA, all leaf traits were strongly associated with stand basal area; LDMC and SLA increased, while LT and LS decreased with forest development. Such changes in LDMC, LT and LS were also related to the decrease in soil nutrient availability and pH along succession, while soil moisture was weakly related to community functional properties. Considering all traits, as well as leaf thickness and succulence separately, FDis strongly decreased with increasing basal area and decreasing soil fertility along forest succession, presenting the lowest values in oldgrowth forests. 5. Synthesis. Our findings suggest that tropical forest regeneration may be a deterministic process shaped by soil conditions. Soil fertility operates as a key filter causing functional convergence towards more conservative resource-use strategies, such as leaves with higher leaf dry matter content.
ABSTRACT
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.
With the decreasing affordability of protecting large blocks of pristine tropical forests, ecologists have staked their hopes on the management of human-modified landscapes (HMLs) to conserve ...tropical biodiversity. Here, we examine key forces affecting the dynamics of HMLs, and propose a framework connecting human disturbances, land use, and prospects for both tropical biodiversity and ecosystem services. We question the forest transition as a worldwide source of new secondary forest; the role played by regenerating (secondary) forest for biodiversity conservation, and the resilience of HMLs. We then offer a conceptual model describing potential successional trajectories among four major landscape types (natural, conservation, functional, and degraded) and highlight the potential implications of our model in terms of research agendas and conservation planning.
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.