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Managing agricultural landscapes to support biodiversity conservation requires profound structural changes worldwide. Often, discussions are centered on management at the field level. However, a wide and growing body of evidence calls for zooming out and targeting agricultural policies, research, and interventions at the landscape level to halt and reverse the decline in biodiversity, increase biodiversity-mediated ecosystem services in agricultural landscapes, and improve the resilience and adaptability of these ecosystems. We conducted the most comprehensive assessment to date on landscape complexity effects on nondomesticated terrestrial biodiversity through a meta-analysis of 1,134 effect sizes from 157 peer-reviewed articles. Increasing landscape complexity through changes in composition, configuration, or heterogeneity significatively and positively affects biodiversity. More complex landscapes host more biodiversity (richness, abundance, and evenness) with potential benefits to sustainable agricultural production and conservation, and effects are likely underestimated. The few articles that assessed the combined contribution of linear (e.g., hedgerows) and areal (e.g., woodlots) elements resulted in a near-doubling of the effect sizes (i.e., biodiversity level) compared to the dominant number of studies measuring these elements separately. Similarly, positive effects on biodiversity are stronger in articles monitoring biodiversity for at least 2 y compared to the dominant 1-y monitoring efforts. Besides, positive and stronger effects exist when monitoring occurs in nonoverlapping landscapes, highlighting the need for long-term and robustly designed monitoring efforts. Living in harmony with nature will require shifting paradigms toward valuing and promoting multifunctional agriculture at the farm and landscape levels with a research agenda that untangles complex agricultural landscapes’ contributions to people and nature under current and future conditions.

For this article, I reviewed empirical studies finding significant ecological responses to habitat fragmentation per se-in other words, significant responses to fragmentation independent of the effects of habitat amount (hereafter referred to as habitat fragmentation). I asked these two questions: Are most significant responses to habitat fragmentation negative or positive? And do particular attributes of species or landscapes lead to a predominance of negative or positive significant responses? I found 118 studies reporting 381 significant responses to habitat fragmentation independent of habitat amount . Of these responses, 76% were positive. Most significant fragmentation effects were positive, irrespective of how the authors controlled for habitat amount, the measure of fragmentation, the taxonomic group, the type of response variable, or the degree of specialization or conservation status of the species or species group. No support was found for predictions that most significant responses to fragmentation should be negative in the tropics, for species with larger movement ranges, or when habitat amount is low; most significant fragmentation effects were positive in all of these cases. Thus, although 24% of significant responses to habitat fragmentation were negative, I found no conditions in which most responses were negative. Authors suggest a wide range of possible explanations for significant positive responses to habitat fragmentation: increased functional connectivity, habitat diversity, positive edge effects, stability of predator-prey host-parasitoid systems, reduced competition, spreading of risk, and landscape complementation. A consistent preponderance of positive significant responses to fragmentation implies that there is no justification for assigning lower conservation value to a small patch than to an equivalent area within a large patch-instead, it implies just the opposite. This finding also suggests that land sharing will usually provide higher ecological value than land sparing.

This research was funded by the ERA-Net BiodivERsA, with the national funders French National Research Agency (ANR-11-EBID-0004), German Ministry of Research and Education (FKZ:01LC1104A) and Spanish Ministry of Economy and Competitiveness, part of the 2011 BiodivERsA call for research proposals. The contribution from the UK was funded by the UK Government Department of the Environment, Food and Rural Affairs (Defra), as project WC1034.

1. Increasing landscape heterogeneity of agroecosystems can enhance natural enemy populations and promote biological control. However, little is known about the multiscale effects of landscape heterogeneity on arthropod communities in rice agroecosystems, especially in combination with trophic interactions. 2. We examined for the first time how landscape heterogeneity, measured by four independent metrics of landscape composition and configuration at three spatial scales, affected species abundance and species richness of rice arthropods within four functional groups and the abundance of the most common species at 28 sites in the Philippines. We additionally examined the influence of trophic interactions among these functional groups. 3. We found that both the compositional and configurational landscape heterogeneity in combination with trophic interactions determined the structure of rice-arthropod communities. Herbivore abundance decreased with increasing landscape diversity. The abundance of parasitoids and species richness of both parasitoids and predators increased with the structural connectivity of rice bunds. Fragmentation of the rice landscape had a clear negative effect on most arthropod groups, except for highly mobile predatory arthropods. Abundance of common predators and detritivore species decreased with increasing complexity in the shape of rice patches. 4. Trophic interactions, measured as the abundance of prey, outweighed the importance of landscape heterogeneity for predators. In contrast, parasitoids responded positively to configurational landscape heterogeneity but were unaffected by prey abundance. 5. Synthesis and applications. Our research shows how landscape heterogeneity and trophic interactions have different effects on different functional groups. While predator abundance was solely driven by the availability of prey, all other functional groups in the rice-arthropod community were significantly affected by the composition and configuration of surrounding landscape features. Landscape management aiming to improve biodiversity and biological control in rice agroeco-systems should promote a diversity of land uses and habitat types within 100-300 m radii to reduce the presence of pests. Management practices should also focus on maintaining smaller rice patches and the structural connectivity of rice bunds to enhance populations of the natural enemies of rice pests. Future research should focus on the temporal and spatial manipulation of rice fields to maximize the effects of biological control.

In this essay: I provide a brief history of habitat fragmentation research; I describe why its “non-questions” (‘Is habitat fragmentation a big problem for wildlife species?” and, “Are the effects of habitat fragmentation generally negative or positive?”) are important to conservation; I outline my role in tackling these questions; I discuss reasons why the culture of habitat fragmentation research is largely incapable of accepting the answers; and I speculate on the future of habitat fragmentation research.

Agricultural management intensity and landscape heterogeneity act as the main drivers of biodiversity loss in agricultural landscapes while also determining ecosystem services. The trait‐based functional diversity approach offers a way to assess changes in community functionality across agroecosystems. We focused on carabids and spiders, because they are an important component of crop field biodiversity and have significant biological control potential. We assessed the effect of small‐ vs. large‐scale agricultural landscapes, organic farming, and within‐field position on functional diversity of spiders and carabids. We sampled pairs of organic and conventional winter wheat fields in small‐scale agricultural landscapes (former West Germany) and in neighbouring large‐scale agricultural landscapes (former East Germany). We sampled arthropods with funnel traps in transects at field edges, field interiors (15 m from edge), and field centres. The gradient from field edges towards the centres played an important role: spider body size decreased; ballooning ability increased, and hunting strategy switched from active hunters to more web‐builders—presumably, due to higher microhabitat stability in the field centre. Higher trait diversity of spiders in field edges suggested higher biocontrol potential in small‐scale agriculture. In contrast, carabid feeding switched from herbivores to carnivores, presumably due to higher pest densities inside crop fields. Furthermore, small‐scale agricultural landscapes and organic management supported larger, i.e., less dispersive carabids. Synthesis and applications. In our research, spiders were more sensitive to edge effects and less sensitive to management and landscape composition than carabids. Smaller fields and longer edges, as well as organic management increase carabid functional diversity, which may increase resilience to environmental change. Since many spider species are confined to field edges, the effect of within‐field position on functional diversity is more important in small‐scale agricultural landscapes with more edge habitat than in large‐scale agricultural landscapes. Our findings suggest that European Union policy should acknowledge the high benefits of small‐scale agriculture for the functional role of major predators such as spiders and carabid beetles, as the benefits are equal to those from a conversion to organic agriculture. In our research, spiders were more sensitive to edge effects and less sensitive to management and landscape composition than carabids. Smaller fields and longer edges, as well as organic management increase carabid functional diversity, which may increase resilience to environmental change. Since many spider species are confined to field edges, the effect of within‐field position on functional diversity is more important in small‐scale agricultural landscapes with more edge habitat than in large‐scale agricultural landscapes. Our findings suggest that European Union policy should acknowledge the high benefits of small‐scale agriculture for the functional role of major predators such as spiders and carabid beetles, as the benefits are equal to those from a conversion to organic agriculture.

ContextThere is a long-standing quest in landscape ecology for holistic biodiversity metrics accounting for multi-taxa diversity in heterogeneous habitat mosaics. Passive acoustic monitoring of biodiversity may provide integrative indices allowing to investigate how soundscapes are shaped by compositional and configurational heterogeneity of mosaic landscapes.ObjectivesWe tested the effects of dominant habitat and landscape heterogeneity on acoustic diversity indices across a large range of mosaic landscapes from two long-term socio-ecological research areas in Occitanie, France and Arizona, USA.MethodsWe assessed acoustic diversity by automated recording for 44 landscapes distributed along gradients of compositional and configurational heterogeneity. We analyzed the responses of six acoustic indices and a composite multiacoustic index to habitat type and multi-scale landscape metrics for three time periods: 24 h-diel cycles, dawns and nights.ResultsLandscape mosaics dominated by permanent grasslands in Occitanie and woodlands in Arizona produced the highest values of acoustic diversity. Moreover, several indices including H, ADI, NDSI, NP and the multiacoustic index consistently responded to edge density in both study regions, but with contrasting patterns, increasing in Occitanie and decreasing in Arizona. Landscape configuration was a key driver of acoustic diversity for diel and nocturnal soundscapes, while dawn soundscapes depended more on landscape composition.ConclusionsAcoustic diversity was correlated more with configurational than compositional heterogeneity in both regions, with contrasting effects explained by the interplay between biogeography and land use history. We suggest that multiple acoustic indices are needed to properly account for complex responses of soundscapes to large-scale habitat heterogeneity in mosaic landscapes.

Loss of habitats and native species, introduction of invasive species, and changing climate regimes lead to the homogenization of landscapes and communities, affecting the availability of habitats and resources for economically important guilds, such as pollinators. Understanding how pollinators and their interactions vary along resource diversity gradients at different scales may help to determine their adaptability to the current diversity loss related to global change. We used data on 20 plant–pollinator communities along gradients of flower richness (local diversity) and landscape heterogeneity (landscape diversity) to understand how the diversity of resources at local and landscape scales affected (1) wild pollinator abundance and richness (accounting also for honey bee abundance), (2) the structure of plant–pollinator networks, (3) the proportion of actively selected interactions (those not occurring by neutral processes), and (4) pollinator diet breadth and species' specialization in networks. Wild pollinator abundance was higher overall in flower-rich and heterogeneous habitats, while wild pollinator richness increased with flower richness (more strongly for beetles and wild bees) and decreased with honeybee abundance. Network specialization (𝐻₂ʹ), modularity, and functional complementarity were all positively related to floral richness and landscape heterogeneity, indicating niche segregation as the diversity of resources increases at both scales. Flower richness also increased the proportion of actively selected interactions (especially for wild bees and flies), whereas landscape heterogeneity had a weak negative effect on this variable. Overall, network-level metrics responded to larger landscape scales than pollinator-level metrics did. Higher floral richness resulted in a wider taxonomic and functional diet for all the study guilds, while functional diet increased mainly for beetles. Despite this, specialization in networks (𝑑ʹ) increased with flower richness for all the study guilds, because pollinator species fed on a narrower subset of plants as communities became richer in species. Our study indicates that pollinators are able to adapt their diet to resource changes at local and landscape scales. However, resource homogenization might lead to poor and generalist pollinator communities, where functionally specialized interactions are lost. This study highlights the importance of including different scales to understand the effects of global change on pollination service through changes in resource diversity.

1. Individual differences in contact rate can arise from host, group and landscape heterogeneity and can result in different patterns of spatial spread for diseases in wildlife populations with concomitant implications for disease control in wildlife of conservation concern, livestock and humans. While dynamic disease models can provide a better understanding of the drivers of spatial spread, the effects of landscape heterogeneity have only been modelled in a few well-studied wildlife systems such as rabies and bovine tuberculosis. Such spatial models tend to be either purely theoretical with intrinsic limiting assumptions or individual-based models that are often highly species- and system-specific, limiting the breadth of their utility. 2. Our goal was to review studies that have utilized dynamic, spatial models to answer questions about pathogen transmission in wildlife and identify key gaps in the literature. We begin by providing an overview of the main types of dynamic, spatial models (e.g., metapopulation, network, lattice, cellular automata, individual-based and continuous-space) and their relation to each other. We investigate different types of ecological questions that these models have been used to explore: pathogen invasion dynamics and range expansion, spatial heterogeneity and pathogen persistence, the implications of management and intervention strategies and the role of evolution in host-pathogen dynamics. 3. We reviewed 168 studies that consider pathogen transmission in free-ranging wildlife and classify them by the model type employed, the focal host-pathogen system, and their overall research themes and motivation. We observed a significant focus on mammalian hosts, a few well-studied or purely theoretical pathogen systems, and a lack of studies occurring at the wildlife-public health or wildlife-livestock interfaces. 4. Finally, we discuss challenges and future directions in the context of unprecedented human-mediated environmental change. Spatial models may provide new insights into understanding, for example, how global warming and habitat disturbance contribute to disease maintenance and emergence. Moving forward, better integration of dynamic, spatial disease models with approaches from movement ecology, landscape genetics/genomics and ecoimmunology may provide new avenues for investigation and aid in the control of zoonotic and emerging infectious diseases.

Assessments of ecosystem services (ES) and biodiversity (hereafter ecological parameters) provide a comprehensive view of the links between landscapes, ecosystem functioning and human well-being. The investigation of consistent associations between ecological parameters, called bundles, and of their links to landscape composition and structure is essential to inform management and policy, yet it is still in its infancy. We mapped over the French Alps an unprecedented array of 18 ecological parameters (16 ES and two biodiversity parameters) and explored their co-occurrence patterns underpinning the supply of multiple ecosystem services in landscapes. We followed a three-step analytical framework to i) detect the ES and biodiversity associations relevant at regional scale, ii) identify the clusters supplying consistent bundles of ES at subregional scale and iii) explore the links between landscape heterogeneity and ecological parameter associations at landscape scale. We used successive correlation coefficients, overlap values and self-organizing maps to characterize ecological bundles specific to given land cover types and geographical areas of varying biophysical characteristics and human uses at nested scales from regional to local. The joint analysis of land cover richness and ES gamma diversity demonstrated that local landscape heterogeneity alone did not imply compatibility across multiple ecosystem services, as some homogeneous landscape could supply multiple ecosystem services.Synthesis and applications. Bundles of ecosystem services and biodiversity parameters are shaped by the joint effects of biophysical characteristics and of human history. Due to spatial congruence and to underlying functional interdependencies, ecological parameters should be managed as bundles even when management targets specific objectives. Moreover, depending on the abiotic context, the supply of multiple ecosystem services can arise either from deliberate management in homogeneous landscapes or from spatial heterogeneity. Bundles of ecosystem services and biodiversity parameters are shaped by the joint effects of biophysical characteristics and of human history. Due to spatial congruence and to underlying functional interdependencies, ecological parameters should be managed as bundles even when management targets specific objectives. Moreover, depending on the abiotic context, the supply of multiple ecosystem services can arise either from deliberate management in homogeneous landscapes or from spatial heterogeneity.