Principles for area-based biodiversity conservation.

Recent international agreements have strengthened and expanded commitments to protect and restore native habitats for biodiversity protection ("area-based biodiversity conservation"). Nevertheless, biodiversity conservation is hindered because how such commitments should be implemented has been strongly debated, which can lead to suboptimal habitat protection decisions. We argue that, despite the debates, there are three essential principles for area-based biodiversity conservation. These principles are related to habitat geographic coverage, amount, and connectivity. They emerge from evidence that, while large areas of nature are important and must be protected, conservation or restoration of multiple small habitat patches is also critical for global conservation, particularly in regions with high land use. We contend that the many area-based conservation initiatives expected in the coming decades should follow the principles we identify, regardless of ongoing debates. Considering the importance of biodiversity for maintenance of ecosystem services, we suggest that this would bring widespread societal benefits.

Substituting space for time: Bird responses to forest loss in space provide a general picture of responses over time.

The practice of space-for-time substitution assumes that the responses of species or communities to land-use change over space represents how they will respond to that same change over time. Space-for-time substitution is commonly used in both ecology and conservation, but whether the assumption produces reliable insights remains inconclusive. Here, we tested space-for-time substitution using data from the North American Breeding Bird Survey (BBS) and Global Forest Change (GFC) to compare the effects of landscape-scale forest cover on bird richness and abundance over time and space, for 25 space-time comparisons. Each comparison consisted of a landscape that experienced at least 20% forest loss over 19 years (temporal site) and a set of 15-19 landscapes (spatial sites) that represented the same forest cover gradient over space in 2019 as experienced over time in their corresponding temporal site. Across the 25 comparisons, the observed responses of forest and open-habitat birds to forest cover over time generally aligned with their responses to forest cover over space, but with comparatively higher variability in the magnitude and direction of effect across the 25 temporal slopes than across the 25 spatial slopes. On average, the mean differences between the spatial and temporal slopes across the 25 space-time comparisons frequently overlapped with zero, suggesting that the spatial slopes are generally informative of the temporal slopes. However, we observed high variability around these mean differences, indicating that a single spatial slope is not strongly predictive of its corresponding temporal slope. We suggest that our results may be explained by annual variability in other relevant environmental factors that combine to produce complex effects on population abundances over time that are not easily captured by snapshots in space. While not being a 1:1 proxy, measuring bird responses to changes in habitat amount in space provides an idea on how birds might be expected to eventually equilibrate to similar changes in habitat amount over time. Further, analyses such as this could be potentially used to screen for cases of regional space-time mismatches where population-limiting factors other than habitat could be playing a more important role in the population trends observed there.

Landscape-scale habitat fragmentation is positively related to biodiversity, despite patch-scale ecosystem decay.

Positive effects of habitat patch size on biodiversity are often extrapolated to infer negative effects of habitat fragmentation on biodiversity at landscape scales. However, such cross-scale extrapolations typically fail. A recent, landmark, patch-scale analysis (Chase et al., 2020, Nature 584, 238-243) demonstrates positive patch size effects on biodiversity, that is, 'ecosystem decay' in small patches. Other authors have already extrapolated this result to infer negative fragmentation effects, that is, higher biodiversity in a few large than many small patches of the same cumulative habitat area. We test whether this extrapolation is valid. We find that landscape-scale patterns are opposite to their analogous patch-scale patterns: for sets of patches with equal total habitat area, species richness and evenness decrease with increasing mean size of the patches comprising that area, even when considering only species of conservation concern. Preserving small habitat patches will, therefore, be key to sustain biodiversity amidst ongoing environmental crises.

Are weak dispersers more vulnerable than strong dispersers to land use intensification?

Ecologists often state that weak dispersers are particularly at risk from land use intensification, and that they therefore should be prioritized for conservation. We reviewed the empirical evidence, to evaluate whether this idea should be used as a general rule in conservation. While 89% of authors predicted that weak dispersers are more vulnerable to land use intensification (80 out of 90 papers), only 56% of reported tests (235 out of 422) were consistent with this prediction. Thirty per cent of tests (128 out of 422) were consistent with the opposite prediction, that strong dispersers are more vulnerable to intensification, and 60% of articles (45 out of 75) had at least one test where strong dispersers were most vulnerable. The likelihood of finding that weak dispersers are more vulnerable to intensification than strong dispersers varied with latitude, taxonomic group and type of land use intensification. Notably, the odds of finding that weak dispersers are more vulnerable to intensification than strong dispersers was higher if the study was nearer to the equator. Taken together, our results show that the prediction that weak dispersers are more vulnerable than strong dispersers to intensification is not sufficiently supported to justify using weak dispersal as a general indicator of species risk in human-modified landscapes.

Support for an area-heterogeneity tradeoff for biodiversity in croplands.

Rapid expansion of the human population poses a challenge for wildlife conservation in agricultural landscapes. One proposal for addressing this challenge is to increase biodiversity in such landscapes by increasing crop diversity. However, studies report both positive and negative effects of crop diversity on biodiversity. One possible explanation, derived from the "area-heterogeneity tradeoff hypothesis," is that the effect of crop diversity on biodiversity depends on a tradeoff between increasing the number of crop types in a landscape and decreasing the amount of each single crop type. This should cause positive effects of increasing crop diversity at low to intermediate crop diversity and negative effects at intermediate to high crop diversity. We also propose two factors that could change the point at which the effect of increasing crop diversity shifts from positive to negative. First, we predicted that this shift would occur at a lower crop diversity when the surrounding landscape contains less semi-natural habitat and at a higher crop diversity when the landscape contains more semi-natural habitat. This should increase the likelihood of detecting negative effects of crop diversity when semi-natural cover is low and positive effects when it is high. Second, we predicted that the shift from a positive to negative effect would occur at a lower crop diversity when it is measured locally than when it is measured at greater distances from the site, making detection of negative crop diversity effects more likely when measurements are at local extents. We tested these predictions using data on the biodiversity of herbaceous plants, butterflies, syrphid flies, woody plants, bees, carabid beetles, spiders, and birds at 221 crop field edges in Eastern Ontario, Canada. We found support for an area-crop diversity tradeoff. Semi-natural cover and measurement extent influenced the biodiversity-crop diversity relationship, with positive effects when semi-natural cover was high and negative effects when semi-natural cover was low and when crop diversity was measured at local extents. The results suggest that policies/guidelines designed to increase crop diversity will not benefit biodiversity in the landscapes where conservation action is most urgently needed, that is, in landscapes with high agricultural use and low semi-natural cover.

Obstruction of biodiversity conservation by minimum patch size criteria.

Minimum patch size criteria for habitat protection reflect the conservation principle that a single large (SL) patch of habitat has higher biodiversity than several small (SS) patches of the same total area (SL > SS). Nonetheless, this principle is often incorrect, and biodiversity conservation requires placing more emphasis on protection of large numbers of small patches (SS > SL). We used a global database reporting the abundances of species across hundreds of patches to assess the SL > SS principle in systems where small patches are much smaller than the typical minimum patch size criteria applied for biodiversity conservation (i.e., ∼85% of patches <100 ha). The 76 metacommunities we examined included 4401 species in 1190 patches. From each metacommunity, we resampled species-area accumulation curves to evaluate how biodiversity responded to habitat existing as a few large patches or as many small patches. Counter to the SL > SS principle and consistent with previous syntheses, species richness accumulated more rapidly when adding several small patches (45.2% SS > SL vs. 19.9% SL > SS) to reach the same cumulative area, even for the very small patches in our data set. Responses of taxa to habitat fragmentation differed, which suggests that when a given total area of habitat is to be protected, overall biodiversity conservation will be most effective if that habitat is composed of as many small patches as possible, plus a few large ones. Because minimum patch size criteria often require larger patches than the small patches we examined, our results suggest that such criteria hinder efforts to protect biodiversity.

Obstrucción de la conservación de la biodiversidad por el criterio del tamaño mínimo del fragmento Resumen Los criterios de tamaño mínimo de los fragmentos para la protección de los hábitats reflejan el principio de conservación según el cual un fragmento único grande (UG) de hábitat tiene mayor biodiversidad que varios fragmentos pequeños (VP) de la misma superficie total (UG > VP). Sin embargo, este principio a menudo es incorrecto; en su lugar, la conservación de la biodiversidad debería enfatizar más la protección de un gran número de pequeñas parcelas (VP > UG). Utilizamos una base de datos mundial que recopila la abundancia de especies en cientos de fragmentos para evaluar el principio UG > VP en sistemas donde los fragmentos pequeños son mucho menores que los criterios comunes de tamaño mínimo de fragmento aplicados para la conservación de la biodiversidad (es decir, ∼85% de parches <100 ha). Las 76 metacomunidades analizadas incluyeron 4,401 especies en 1,190 parcelas. Volvimos a muestrear las curvas de acumulación especie-área en cada metacomunidad para evaluar cómo respondía la biodiversidad al hábitat: como unas pocas manchas grandes o como fragmentos de muchas manchas pequeñas. Contrario al principio UG > VP y en congruencia con síntesis anteriores, la riqueza de especies se acumuló con mayor rapidez al añadir varios fragmentos pequeños (45.2% VP > UG frente a 19.9% UG > VP) para alcanzar la misma área acumulada, incluso para los fragmentos muy pequeños de nuestro conjunto de datos. Las respuestas de los taxones a la fragmentación del hábitat fueron diferentes, lo que sugiere que, cuando se trata de proteger una determinada superficie total de hábitat, la conservación global de la biodiversidad será más efectiva si ese hábitat está compuesto por el mayor número posible de pequeños fragmentos, más unos cuantos fragmentos grandes. Dado que los criterios de tamaño mínimo exigen a menudo fragmentos más grandes que los pequeños que examinamos, nuestros resultados sugieren que tales criterios dificultan los esfuerzos por proteger la biodiversidad.

Loss of the world's smallest forests.

A large number of small forests typically harbor higher biodiversity than a small number of large forests totaling the same area, suggesting that small patches are disproportionately valuable for biodiversity conservation. However, policies often favor protection of large forest patches. Here we demonstrate a global trend of higher deforestation in small than large forest patches: the likelihood that a randomly selected forest plot disappeared between 1992 and 2020 increased with decreasing size of the forest patch containing that plot. Our results imply a disproportionate impact of forest loss on biodiversity relative to the total forest area removed. Achieving recent commitments of the post-2020 Global Biodiversity Framework will require revision of current policies and increased societal awareness of the importance of small habitat patches for biodiversity protection.

Increasing crop heterogeneity enhances multitrophic diversity across agricultural regions.

Agricultural landscape homogenization has detrimental effects on biodiversity and key ecosystem services. Increasing agricultural landscape heterogeneity by increasing seminatural cover can help to mitigate biodiversity loss. However, the amount of seminatural cover is generally low and difficult to increase in many intensively managed agricultural landscapes. We hypothesized that increasing the heterogeneity of the crop mosaic itself (hereafter "crop heterogeneity") can also have positive effects on biodiversity. In 8 contrasting regions of Europe and North America, we selected 435 landscapes along independent gradients of crop diversity and mean field size. Within each landscape, we selected 3 sampling sites in 1, 2, or 3 crop types. We sampled 7 taxa (plants, bees, butterflies, hoverflies, carabids, spiders, and birds) and calculated a synthetic index of multitrophic diversity at the landscape level. Increasing crop heterogeneity was more beneficial for multitrophic diversity than increasing seminatural cover. For instance, the effect of decreasing mean field size from 5 to 2.8 ha was as strong as the effect of increasing seminatural cover from 0.5 to 11%. Decreasing mean field size benefited multitrophic diversity even in the absence of seminatural vegetation between fields. Increasing the number of crop types sampled had a positive effect on landscape-level multitrophic diversity. However, the effect of increasing crop diversity in the landscape surrounding fields sampled depended on the amount of seminatural cover. Our study provides large-scale, multitrophic, cross-regional evidence that increasing crop heterogeneity can be an effective way to increase biodiversity in agricultural landscapes without taking land out of agricultural production.

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.

Bridging research and practice in conservation.

Calls for biodiversity conservation practice to be more evidence based are growing, and we agree evidence use in conservation practice needs improvement. However, evidence-based conservation will not be realized without improved access to evidence. In medicine, unlike in conservation, a well-established and well-funded layer of intermediary individuals and organizations engage with medical practitioners, synthesize primary research relevant to decision making, and make evidence easily accessible. These intermediaries prepare targeted evidence summaries and distribute them to practitioners faced with time-sensitive and value-laden decisions. To be effective, these intermediaries, who we refer to as evidence bridges, should identify research topics based on the priorities of practitioners; synthesize evidence; prepare and distribute easy-to-find and easy-to-use evidence summaries; and develop and maintain networks of connections with researchers and practitioners. Based on a review of the literature regarding evidence intermediaries in conservation and environmental management, as well as an anonymous questionnaire searching for such organizations, we found few intermediaries that met all these criteria. Few evidence bridges that do exist are unable to reach most conservation practitioners, which include resource managers in government and industry, conservation organizations, and farmers and other private landowners. We argue that the lack of evidence bridges from research to practitioners contributes to evidence complacency and limits the use of evidence in conservation action. Nevertheless, several existing organizations help reduce the gap between evidence and practice and could serve as a foundation for building additional components of evidence bridges in conservation. Although evidence bridges need expertise in research and evidence synthesis, they also require expertise in identifying and communicating with the community of practitioners most in need of clear and concise syntheses of evidence. Article Impact Statement: Evidence-based conservation will not be realized without improved access to evidence. We call for intermediary evidence bridges.

Vinculación entre la Investigación y la Práctica en la Conservación Resumen Cada vez existen más peticiones para que las prácticas de conservación de la biodiversidad estén más basadas en evidencias, además de que apoyamos la idea de que el uso de evidencias en la práctica de la conservación necesita mejorar. Sin embargo, la conservación basada en la evidencia no se logrará sin un acceso mejorado a las evidencias. En la medicina, no como en la conservación, un estrato bien establecido y financiado de individuos y organizaciones intermediarias interactúan con los médicos, sintetizan las investigaciones primarias relevantes para la toma de decisiones y hacen que las evidencias sean de fácil acceso. Estos intermediarios preparan resúmenes de evidencias específicas y los distribuyen a los médicos que enfrentan decisiones urgentes y muy valiosas. Para que sean efectivos, estos intermediarios, a quienes nos referimos como puentes de evidencias, deben poder identificar los temas de estudio con base en las prioridades de los practicantes, sintetizar evidencias, preparar y distribuir resúmenes fáciles de encontrar y fáciles de usar, y desarrollar y mantener redes de conexiones con los investigadores y los practicantes. Con base en una revisión de la literatura correspondiente a los intermediarios de evidencias en la conservación y el manejo ambiental, así como en un cuestionario anónimo que busca a dichas organizaciones, encontramos a pocos intermediarios que cumplieran con estos criterios. Los pocos puentes de evidencias que existen no son capaces de llegar a la mayoría de los practicantes de la conservación, los cuales incluyen a los gestores de recursos en el gobierno y en la industria, a las organizaciones de conservación y a los agricultores y otros terratenientes privados. Argumentamos que la falta de puentes de evidencia entre los investigadores y los practicantes contribuye a la indulgencia de evidencias y limita el uso de evidencias en las acciones de conservación. Sin embargo, varias organizaciones existentes ayudan a reducir la brecha entre la evidencia y la práctica y podrían funcionar como base para la construcción de componentes adicionales para los puentes de evidencia en la conservación. Aunque los puentes de evidencias necesitan experiencia con la investigación y con la síntesis de evidencias, también requieren experiencia con la identificación de y comunicación con la comunidad de practicantes que más necesitan una síntesis clara y concisa de la evidencia.

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.

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.

Inference in road ecology research: what we know versus what we think we know.

Roads and traffic impacts on wildlife populations are well documented. Three major mechanisms can cause them: reduced connectivity, increased mortality and reduced habitat quality. Researchers commonly recommend mitigation based on the mechanism they deem responsible. We reviewed the 2012-2016 literature to evaluate authors' inferences, to determine whether they explicitly acknowledge all possible mechanisms that are consistent with their results. We found 327 negative responses of wildlife to roads, from 307 studies. While most (84%) of these responses were consistent with multiple mechanisms, 60% of authors invoked a single mechanism. This indicates that many authors are over-confident in their inferences, and that the literature does not allow estimation of the relative importance of the mechanisms. We found preferences in authors' discussion of mechanisms. When all three mechanisms were consistent with the response measured, authors were 2.4 and 2.9 times as likely to infer reduced habitat quality compared to reduced connectivity or increased mortality, respectively. When both reduced connectivity and increased mortality were consistent with the response measured, authors were 5.2 times as likely to infer reduced connectivity compared to increased mortality. Given these results, road ecologists and managers are likely over-recommending mitigation for improving habitat quality and connectivity, and under-recommending measures to reduce road-kill.

Landscape configurational heterogeneity by small-scale agriculture, not crop diversity, maintains pollinators and plant reproduction in western Europe.

Agricultural intensification is one of the main causes for the current biodiversity crisis. While reversing habitat loss on agricultural land is challenging, increasing the farmland configurational heterogeneity (higher field border density) and farmland compositional heterogeneity (higher crop diversity) has been proposed to counteract some habitat loss. Here, we tested whether increased farmland configurational and compositional heterogeneity promote wild pollinators and plant reproduction in 229 landscapes located in four major western European agricultural regions. High-field border density consistently increased wild bee abundance and seed set of radish (Raphanus sativus), probably through enhanced connectivity. In particular, we demonstrate the importance of crop-crop borders for pollinator movement as an additional experiment showed higher transfer of a pollen analogue along crop-crop borders than across fields or along semi-natural crop borders. By contrast, high crop diversity reduced bee abundance, probably due to an increase of crop types with particularly intensive management. This highlights the importance of crop identity when higher crop diversity is promoted. Our results show that small-scale agricultural systems can boost pollinators and plant reproduction. Agri-environmental policies should therefore aim to halt and reverse the current trend of increasing field sizes and to reduce the amount of crop types with particularly intensive management.

Habitat specialist birds disperse farther and are more migratory than habitat generalist birds.

Some theories predict habitat specialists should be less dispersive and migratory than generalists, while other theories predict the opposite. We evaluated the cross-species relationship between the degree of habitat specialization and dispersal and migration status in 101 bird species breeding in North America and the United Kingdom, using empirical estimates of the degree of habitat specialization from breeding bird surveys and mean dispersal distance estimates from large-scale mark-recapture studies. We found that habitat specialists dispersed farther than habitat generalists, and full migrants had more specialized habitat than partial migrants or resident species. To our knowledge this is the first large-scale, multi-species study to demonstrate a positive relationship between the degree of habitat specialization and dispersal, and it is opposite to the pattern found for invertebrates. This finding is particularly interesting because it suggests that trade-offs between the degree of habitat specialization and dispersal ability are not conserved across taxonomic groups. This cautions against extrapolation of trait co-occurrence from one species group to another. In particular, it suggests that efforts aimed at conserving the most habitat-specialist temperate-breeding birds will not lead to conservation of the most dispersal-limited species.

An experimental test of the habitat-amount hypothesis for saproxylic beetles in a forested region.

The habitat-amount hypothesis challenges traditional concepts that explain species richness within habitats, such as the habitat-patch hypothesis, where species number is a function of patch size and patch isolation. It posits that effects of patch size and patch isolation are driven by effects of sample area, and thus that the number of species at a site is basically a function of the total habitat amount surrounding this site. We tested the habitat-amount hypothesis for saproxylic beetles and their habitat of dead wood by using an experiment comprising 190 plots with manipulated patch sizes situated in a forested region with a high variation in habitat amount (i.e., density of dead trees in the surrounding landscape). Although dead wood is a spatio-temporally dynamic habitat, saproxylic insects have life cycles shorter than the time needed for habitat turnover and they closely track their resource. Patch size was manipulated by adding various amounts of downed dead wood to the plots (~800 m³ in total); dead trees in the surrounding landscape (~240 km2 ) were identified using airborne laser scanning (light detection and ranging). Over 3 yr, 477 saproxylic species (101,416 individuals) were recorded. Considering 20-1,000 m radii around the patches, local landscapes were identified as having a radius of 40-120 m. Both patch size and habitat amount in the local landscapes independently affected species numbers without a significant interaction effect, hence refuting the island effect. Species accumulation curves relative to cumulative patch size were not consistent with either the habitat-patch hypothesis or the habitat-amount hypothesis: several small dead-wood patches held more species than a single large patch with an amount of dead wood equal to the sum of that of the small patches. Our results indicate that conservation of saproxylic beetles in forested regions should primarily focus on increasing the overall amount of dead wood without considering its spatial arrangement. This means dead wood should be added wherever possible including in local landscapes with low or high dead-wood amounts. For species that have disappeared from most forests owing to anthropogenic habitat degradation, this should, however, be complemented by specific conservation measures pursued within their extant distributional ranges.

Experimental study designs to improve the evaluation of road mitigation measures for wildlife.

An experimental approach to road mitigation that maximizes inferential power is essential to ensure that mitigation is both ecologically-effective and cost-effective. Here, we set out the need for and standards of using an experimental approach to road mitigation, in order to improve knowledge of the influence of mitigation measures on wildlife populations. We point out two key areas that need to be considered when conducting mitigation experiments. First, researchers need to get involved at the earliest stage of the road or mitigation project to ensure the necessary planning and funds are available for conducting a high quality experiment. Second, experimentation will generate new knowledge about the parameters that influence mitigation effectiveness, which ultimately allows better prediction for future road mitigation projects. We identify seven key questions about mitigation structures (i.e., wildlife crossing structures and fencing) that remain largely or entirely unanswered at the population-level: (1) Does a given crossing structure work? What type and size of crossing structures should we use? (2) How many crossing structures should we build? (3) Is it more effective to install a small number of large-sized crossing structures or a large number of small-sized crossing structures? (4) How much barrier fencing is needed for a given length of road? (5) Do we need funnel fencing to lead animals to crossing structures, and how long does such fencing have to be? (6) How should we manage/manipulate the environment in the area around the crossing structures and fencing? (7) Where should we place crossing structures and barrier fencing? We provide experimental approaches to answering each of them using example Before-After-Control-Impact (BACI) study designs for two stages in the road/mitigation project where researchers may become involved: (1) at the beginning of a road/mitigation project, and (2) after the mitigation has been constructed; highlighting real case studies when available.

Landscape context affects genetic diversity at a much larger spatial extent than population abundance.

Regional landscape context influences the fate of local populations, yet the spatial extent of this influence (called the "scale of effect") is difficult to predict. Thus, a major problem for conservation management is to understand the factors governing the scale of effect such that landscape structure surrounding a focal area is measured and managed at the biologically relevant spatial scale. One unresolved question is whether and how scale of effect may depend on the population response measured (e.g., abundance vs. presence/absence). If scales of effect differ across population outcomes of a given species, management based on one outcome may compromise another, further complicating conservation decision making. Here we used an individual-based simulation model to investigate how scales of effect of landscapes that vary in the amount and fragmentation of habitat differ among three population responses (local abundance, presence/absence, and genetic diversity). We also explored how the population response measured affects the relative importance of habitat amount and fragmentation in shaping local populations, and how dispersal distance mediates the magnitude and spatial scale of these effects. We found that the spatial scale most strongly influencing local populations depended on the outcome measured and was predicted to be small for abundance, medium-sized for presence/absence, and large for genetic diversity. Increasing spatial scales likely resulted from increasing temporal scales over which outcomes were regulated (with local genetic diversity being regulated over the largest number of generations). Thus, multiple generations of dispersal and gene flow linked local population patterns to regional population size. The effects of habitat amount dominated the effects of fragmentation for all three outcomes. Increased dispersal distance strongly reduced abundance, but not presence/absence or genetic diversity. Our results suggest that managing protected species at spatial scales based on population abundance data may ignore broader landscape effects on population genetic diversity and persistence, lending support to the importance of managing large buffers surrounding areas of conservation concern.

Why is a landscape perspective important in studies of primates?

With accelerated deforestation and fragmentation through the tropics, assessing the impact that landscape spatial changes may have on biodiversity is paramount, as this information is required to design and implement effective management and conservation plans. Primates are expected to be particularly dependent on the landscape context; yet, our understanding on this topic is limited as the majority of primate studies are at the local scale, meaning that landscape-scale inferences are not possible. To encourage primatologists to assess the impact of landscape changes on primates, and help future studies on the topic, we describe the meaning of a "landscape perspective" and evaluate important assumptions of using such a methodological approach. We also summarize a number of important, but unanswered, questions that can be addressed using a landscape-scale study design. For example, it is still unclear if habitat loss has larger consistent negative effects on primates than habitat fragmentation per se. Furthermore, interaction effects between habitat area and other landscape effects (e.g., fragmentation) are unknown for primates. We also do not know if primates are affected by synergistic interactions among factors at the landscape scale (e.g., habitat loss and diseases, habitat loss and climate change, hunting, and land-use change), or whether landscape complexity (or landscape heterogeneity) is important for primate conservation. Testing for patterns in the responses of primates to landscape change will facilitate the development of new guidelines and principles for improving primate conservation.

Overcoming confusion and stigma in habitat fragmentation research.

Anthropogenic habitat loss is widely recognized as a primary environmental concern. By contrast, debates on the effects of habitat fragmentation persist. To facilitate overcoming these debates, here we: (i) review the state of the literature on habitat fragmentation, finding widespread confusion and stigma; (ii) identify consequences of this for biodiversity conservation and ecosystem management; and (iii) suggest ways in which research can move forward to resolve these problems. Confusion is evident from the 25 most-cited fragmentation articles published between 2017 and 2021. These articles use five distinct concepts of habitat fragmentation, only one of which clearly distinguishes habitat fragmentation from habitat area and other factors ('fragmentation per se'). Stigmatization is evident from our new findings that fragmentation papers are more charged with negative sentiments when compared to papers from other subfields in the environmental sciences, and that fragmentation papers with more negative sentiments are cited more. While most empirical studies of habitat fragmentation per se find neutral or positive effects on species and biodiversity outcomes, which implies that small habitat patches have a high cumulative value, confusion and stigma in reporting and discussing such results have led to suboptimal habitat protection policy. For example, government agencies, conservation organizations, and land trusts impose minimum habitat patch sizes on habitat protection. Given the high cumulative value of small patches, such policies mean that many opportunities for conservation are being missed. Our review highlights the importance of reducing confusion and stigma in habitat fragmentation research. To this end, we propose implementing study designs in which multiple sample landscapes are selected across independent gradients of habitat amount and fragmentation, measured as patch density. We show that such designs are possible for forest habitat across Earth's biomes. As such study designs are adopted, and as language becomes more precise, we expect that confusion and stigma in habitat fragmentation research will dissipate. We also expect important breakthroughs in understanding the situations where effects of habitat fragmentation per se are neutral, positive, or negative, and the reasons for these differences. Ultimately this will improve efficacy of area-based conservation policies, to the benefit of biodiversity and people.