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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.

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.

For many amphibian species, reduced landscape connectivity results in reduced genetic connectivity among populations. However, large effective population sizes (Ne) slow the rate of genetic drift, causing subdivided populations to remain genetically similar despite little gene flow among them. Therefore, it is important to address the combined effects of Ne and matrix permeability to quantify the relative importance of gene flow and genetic drift on isolated amphibian populations. We applied a landscape genetic approach to investigate how patterns of gene flow (m), Ne (inferred via θ) and genetic differentiation differ among Eastern Red-backed Salamander (Plethodon cinereus) populations in a fragmented landscape (n = 4) compared to a continuous forest (n = 4). We assayed a panel of 10 microsatellite markers for population genetic analyses. Additionally, we constructed and validated a distribution model to generate resistance surfaces for examining the relationship between landscape connectivity, m, θ, and genetic differentiation (FST) using maximum-likelihood population-effects models (MLPE). Populations in continuous habitat were undifferentiated, whereas fragmented populations exhibited genetic structure driven by a single population. Results of the MLPE models in the fragmented landscape revealed spatial variation in θ as the best predictor of pairwise FST, followed by estimates of m, suggesting migration-drift interactions have a stronger influence on genetic differentiation than matrix permeability. Moreover, model coefficients for landscape resistance were comparable between landscapes. Overall, our results provide insight as to how the interaction of gene flow and genetic drift shapes population structure for a dispersal-limited species within a predominately anthropogenic landscape.

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.

Vulnerability to habitat fragmentation Habitat fragmentation caused by human activities has consequences for the distribution and movement of organisms. Betts et al. present a global analysis of how exposure to habitat fragmentation affects the composition of ecological communities (see the Perspective by Hargreaves). In a dataset consisting of 4489 animal species, regions that historically experienced little disturbance tended to harbor a higher proportion of species vulnerable to fragmentation. Species in more frequently disturbed regions were more resilient. High-latitude areas historically experienced more disturbance and harbor more resilient species, which suggests that extinction has removed fragmentation-sensitive species. Thus, conservation efforts to limit fragmentation are particularly important in the tropics. Science , this issue p. 1236 ; see also p. 1196

In a review of landscape-scale empirical studies, Fahrig (2017a) found that ecological responses to habitat fragmentation per se (fragmentation independent of habitat amount) were usually non-significant (>70% of responses) and that 76% of significant relationships were positive, with species abundance, occurrence, richness, and other response variables increasing with habitat fragmentation per se. Fahrig concluded that to date there is no empirical evidence supporting the widespread assumption that a group of small habitat patches generally has lower ecological value than large patches of the same total area. Fletcher et al. (2018) dispute this conclusion, arguing that the literature to date indicates generally negative ecological effects of habitat fragmentation per se. They base their argument largely on extrapolation from patch-scale patterns and mechanisms (effects of patch size and isolation, and edge effects) to landscape-scale effects of habitat fragmentation. We argue that such extrapolation is unreliable because: (1) it ignores other mechanisms, especially those acting at landscape scales (e.g., increased habitat diversity, spreading of risk, landscape complementation) that can counteract effects of the documented patch-scale mechanisms; and (2) extrapolation of a small-scale mechanism to a large-scale pattern is not evidence of that pattern but, rather a prediction that must be tested at the larger scale. Such tests were the subject of Fahrig's review. We find no support for Fletcher et al.'s claim that biases in Fahrig's review would alter its conclusions. We encourage further landscape-scale empirical studies of effects of habitat fragmentation per se, and research aimed at uncovering the mechanisms that underlie positive fragmentation effects.

Land-cover and climate change are both expected to alter species distributions and contribute to future biodiversity loss. However, the combined effects of land-cover and climate change on assemblages, especially at the landscape scale, remain understudied. Lowland tropical amphibians may be particularly susceptible to changes in land cover and climate warming because many species have narrow thermal safety margins resulting from air and body temperatures that are close to their critical thermal maxima (CTmax ). We examined how changing thermal landscapes may alter the area of thermally suitable habitat (TSH) for tropical amphibians. We measured microclimates in 6 land-cover types and CTmax of 16 frog species in lowland northeastern Costa Rica. We used a biophysical model to estimate core body temperatures of frogs exposed to habitat-specific microclimates while accounting for evaporative cooling and behavior. Thermally suitable habitat area was estimated as the portion of the landscape where species CTmax exceeded their habitat-specific maximum body temperatures. We projected changes in TSH area 80 years into the future as a function of land-cover change only, climate change only, and combinations of land-cover and climate-change scenarios representing low and moderate rates of change. Projected decreases in TSH area ranged from 16% under low emissions and reduced forest loss to 30% under moderate emissions and business-as-usual land-cover change. Under a moderate emissions scenario (A1B), climate change alone contributed to 1.7- to 4.5-fold greater losses in TSH area than land-cover change only, suggesting that future decreases in TSH from climate change may outpace structural habitat loss. Forest-restricted species had lower mean CTmax than species that occurred in altered habitats, indicating that thermal tolerances will likely shape assemblages in changing thermal landscapes. In the face of ongoing land-cover and climate change, it will be critical to consider changing thermal landscapes in strategies to conserve ectotherm species.

Human activities often replace native forests with warmer, modified habitats that represent novel thermal environments for biodiversity. Reducing biodiversity loss hinges upon identifying which species are most sensitive to the environmental conditions that result from habitat modification. Drawing on case studies and a meta-analysis, we examined whether observed and modelled thermal traits, including heat tolerances, variation in body temperatures, and evaporative water loss, explained variation in sensitivity of ectotherms to habitat modification. Low heat tolerances of lizards and amphibians and high evaporative water loss of amphibians were associated with increased sensitivity to habitat modification, often explaining more variation than non-thermal traits. Heat tolerances alone explained 24-66% (mean = 38%) of the variation in species responses, and these trends were largely consistent across geographic locations and spatial scales. As habitat modification alters local microclimates, the thermal biology of species will likely play a key role in the reassembly of terrestrial communities.

Climate change poses new challenges for natural resource managers. Predictive modeling of species–environment relationships using climate envelope models can enhance our understanding of climate change effects on biodiversity, assist in assessment of invasion risk by exotic organisms, and inform life-history understanding of individual species. While increasing interest has focused on the role of uncertainty in future conditions on model predictions, models also may be sensitive to the initial conditions on which they are trained. Although climate envelope models are usually trained using data on contemporary climate, we lack systematic comparisons of model performance and predictions across alternative climate data sets available for model training. Here, we seek to fill that gap by comparing variability in predictions between two contemporary climate data sets to variability in spatial predictions among three alternative projections of future climate. Overall, correlations between monthly temperature and precipitation variables were very high for both contemporary and future data. Model performance varied across algorithms, but not between two alternative contemporary climate data sets. Spatial predictions varied more among alternative general-circulation models describing future climate conditions than between contemporary climate data sets. However, we did find that climate envelope models with low Cohen's kappa scores made more discrepant spatial predictions between climate data sets for the contemporary period than did models with high Cohen's kappa scores. We suggest conservation planners evaluate multiple performance metrics and be aware of the importance of differences in initial conditions for spatial predictions from climate envelope models.

Management of communities in fragmented systems requires application of models for predicting and understanding patterns of diversity at relevant scales. Metacommunity models may help explain patterns of beta-diversity, but more empirical investigations are needed to determine the generality of these models and the importance of matrix identity as a mediator of metacommunity processes. We studied patterns of beta diversity among cypress domes in a landscape composed of two different matrix types within Big Cypress National Preserve to determine whether community composition differed by matrix type and to evaluate predictions of metacommunity models. We sampled fully-aquatic vertebrates in 16 cypress domes. A causal modeling framework was used to assess the relative importance of space and environmental variables measured in the domes and in the nearby matrix in explaining variation in community similarities. Our results show that community composition was influenced by matrix type, which violates the common metacommunity assumption that matrix identity has negligible effect on patch patterns. We found different drivers of beta-diversity patterns between matrix types; similarities among dome communities within cypress-prairie matrix were influenced by local environmental conditions and matrix characteristics, while communities in pine-rockland domes were most influenced by conditions in the matrix. Our results have implications for use of metacommunity models in conservation planning and we suggest that future research should focus on the importance of matrix identity and complexity in mediating community patterns in patchy landscapes.

Aim Connectivity is a key determinant of the distribution and abundance of organisms and is greatly influenced by anthropogenic landscape modification, yet we lack a synthetic perspective on the magnitude and extent of matrix effects on connectivity. We synthesize results from published studies to understand the importance of matrix effects on fragmented animal populations. Location Global. Methods We conduct a meta-analysis of 283 fragmented populations representing 184 terrestrial animal taxa to determine the strength of matrix composition effects on the occurrence and abundance of animals in fragmented habitat. Results Studies that use data on matrix composition report greater effects on abundance and occupancy of fragmented populations than studies that define connectivity without regard to the surrounding matrix (i.e. ‘binary’ studies that describe only characteristics of patch habitat). Main conclusions Our findings underscore that conservation strategies must consider the importance of matrix habitat, have important implications for metapopulation and metacommunity paradigms, and provide direct large-scale, multi-taxa evidence that matrix habitat is an important driver of ecological dynamics in heterogeneous landscapes.

Exotic ecosystem engineers induce structural and qualitative habitat changes in invaded landscapes, yet studies rarely examine the effects of both of these changes on native taxa. We used a factorial experiment in natural, predator-containing environments to determine whether performance of amphibian larvae was affected by predators and/or changes in habitat structure or chemistry associated with the invasive shrub Lonicera maackii. Invertebrate predators significantly reduced survival of American toad Anaxyrus americanus larvae, whereas tadpole development was accelerated in pools inoculated with the chemical signature of L. maackii. The significant effect of L. maackii chemistry on A. americanus larvae suggests that invasive species may have non-intuitive effects even on native taxa with which they share no trophic connection, and may represent cryptic components of the multiple, interactive drivers of biodiversity change.

We quantified patterns of species richness and species composition of frogs and reptiles (lizards and snakes) among three habitats (continuous forest, forest islands, and a seasonally flooded savannah) and between forest island size and isolation classes in a floristic transition zone in northeastern Santa Cruz Department, Bolivia. Species richness was similar across macrohabitats, as was faunal composition of forested habitats, although savannah harbored a distinct herpetofauna. On forest islands, richness and composition of forest frogs was largely related to isolation, whereas reptiles were affected by both isolation and habitat. The observation that isolation rather than area was the primary driver of distribution patterns on forest islands stands in contrast to many studies, and may be a function of (1) the greater range in forest island isolation values compared to area or (2) the long history of isolation in this landscape.

Habitat loss and fragmentation threaten a large proportion of terrestrial biodiversity, and identifying the ecological traits associated with extinction proneness is of widespread interest. We used a multivariate statistical approach to identify combinations of ecological traits that best allowed us to identify extinction-prone amphibians and reptiles in a fragmented landscape in north-eastern Bolivia. Extinction-prone amphibians were rare and did not utilize the savannah matrix separating forest islands, whereas extinction-prone reptiles were trophically specialized. Rarity and matrix aversion are among the most widely reported correlates to extinction proneness, and we argue that an increased understanding of their role as drivers of extinction processes is necessary. We suggest that the absence of reptilian vertebrate predators may exacerbate trophic cascades in habitat patches.

Scientific interest in the impact of habitat fragmentation on biodiversity is increasing, but our un- derstanding of fragmentation is clouded by a lack of appreciation for fundamental similarities and differences across studies representing a wide range of taxa and landscape types. In an effort to synthesize data describ- ing ecological responses of animals to fragmentation across two classes of independent variables (taxonomic group and landscape), we sampled 148 studies of five major faunal groups from the primary literature and an- alyzed data on 13 variables extracted from those studies. We focused our analyses on three classes of dependent variables (effects of area and isolation on species richness, z values, and nestedness and species composition). Area ranged over more orders of magnitude than isolation and tended to explain more variation in species richness than isolation. There were few matrix or taxon effects on the patterns we investigated, although we did find that sky islands tended to manifest isolation effects on both species richness and nestedness more frequently than other patch types. Sky islands may offer insight into the future of habitat patches fragmented by contemporary habitat loss, and because they show a stronger effect of isolation than other patch types, we suggest that isolation will play an increasing role in the biology of habitat fragments. We use multiple lines of evidence to suggest that our understanding of the role of isolation on community assembly in fragmented landscapes is inadequate. Finally, our observation that consistent taxonomic differences in community pat- terns were minimal suggests that conservation actions intended to mitigate the negative effects of extinction may have far-reaching effects across taxonomic groups.

Scientific interest in the impact of habitat fragmentation on biodiversity is increasing, but our understanding of fragmentation is clouded by a lack of appreciation for fundamental similarities and differences across studies representing a wide range of taxa and landscape types. In an effort to synthesize data describing ecological responses of animals to fragmentation across two classes of independent variables (taxonomic group and landscape), we sampled 148 studies offive major faunal groups from the primary literature and analyzed data on 13 variables extracted from those studies. We focused our analyses on three classes of dependent variables (effects of area and isolation on species richness, z values, and nestedness and species composition). Area ranged over more orders of magnitude than isolation and tended to explain more variation in species richness than isolation. There were few matrix or taxon effects on the patterns we investigated, although we did find that sky islands tended to manifest isolation effects on both species richness and nestedness more frequently than other patch types. Sky islands may offer insight into the future of habitat patches fragmented by contemporary habitat loss, and because they show a stronger effect of isolation than other patch types, we suggest that isolation will play an increasing role in the biology of habitat fragments. We use multiple lines of evidence to suggest that our understanding of the role of isolation on community assembly in fragmented landscapes is inadequate. Finally, our observation that consistent taxonomic differences in community patterns were minimal suggests that conservation actions intended to mitigate the negative effects of extinction may have far-reaching effects across taxonomic groups.

The presence of multiple interacting threats to biodiversity and the increasing rate of species extinction make it critical to prioritize management efforts on species and communities that maximize conservation success. We implemented a multi-step approach that coupled vulnerability assessments evaluating threats to Florida taxa such as climate change, sea-level rise, and habitat fragmentation with in-depth literature surveys of taxon-specific ecological traits. The vulnerability, adaptive capacity, and ecological traits of 12 threatened and endangered subspecies were compared to non-listed subspecies of the same parent species. Overall, the threatened and endangered subspecies showed high vulnerability and low adaptive capacity, in particular to sea level rise and habitat fragmentation. They also exhibited larger home ranges and greater dispersal limitation compared to non-endangered subspecies, which may inhibit their ability to track changing climate in fragmented landscapes. There was evidence for lower reproductive capacity in some of the threatened or endangered taxa, but not for most. Taxa located in the Florida Keys or in other low coastal areas were most vulnerable to sea level rise, and also showed low levels of adaptive capacity, indicating they may have a lower probability of conservation success. Our analysis of at-risk subspecies and closely related non-endangered subspecies demonstrates that ecological traits help to explain observed differences in vulnerability and adaptive capacity. This study points to the importance of assessing the relative contributions of multiple threats and evaluating conservation value at the species (or subspecies) level when resources are limited and several factors affect conservation success.