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ABSTRACT Animal behavior is an important component of individual, population, and community responses to anthropogenic habitat alteration. For example, antipredator behavior (e.g., vigilance) and animal movement behavior may both be important behavioral responses to the increased density of habitat edges and changes in patch connectivity that characterize highly modified habitats. Importantly, edge density and connectivity might interact, and this interaction is likely to mediate animal behavior: linear, edge‐rich landscape features often provide structural connectivity between patches, but the functional connectedness of patches for animal use could depend upon how edge density modifies animal vigilance and movement. Using remote cameras in large‐scale experimental landscapes that manipulate edge density (high‐ vs. low‐density edges) and patch connectivity (isolated or connected patches), we examined the effects of edge density and connectivity on the antipredator behavior and movement behavior of white‐tailed deer (Odocoileus virginianus). Deer vigilance was 1.38 times greater near high‐density edges compared to low‐density edges, regardless of whether patches were connected or isolated. Deer were also more likely to move parallel to connected high‐density edges than all other edge types, suggesting that connectivity promotes movement along high‐density edges. These results suggest that increases in edge density that accompany human fragmentation of existing habitats may give rise to large‐scale changes in the antipredator behavior of deer. These results also suggest that conservation strategies that simultaneously manipulate edge density and connectivity (i.e., habitat corridors) may have multiple effects on different aspects of deer behavior: linear habitat corridors were areas of high vigilance, but also areas where deer movement behavior implied increased movement along the habitat edge. 

Abstract Soil microorganisms play outsized roles in nutrient cycling, plant health, and climate regulation. Despite their importance, we have a limited understanding of how soil microbes are affected by habitat fragmentation, including their responses to conditions at fragment edges, or “edge effects.” To understand the responses of soil communities to edge effects, we analyzed the distributions of soil bacteria, archaea, and fungi in an experimentally fragmented system of open patches embedded within a forest matrix. In addition, we identified taxa that consistently differed among patch, edge, or matrix habitats (“specialists”) and taxa that showed no habitat preference (“nonspecialists”). We hypothesized that microbial community turnover would be most pronounced at the edge between habitats. We also hypothesized that specialist fungi would be more likely to be mycorrhizal than nonspecialist fungi because mycorrhizae should be affected more by different plant hosts among habitats, whereas specialist prokaryotes would have smaller genomes (indicating reduced metabolic versatility) and be less likely to be able to sporulate than nonspecialist prokaryotes. Across all replicate sites, the matrix and patch soils harbored distinct microbial communities. However, sites where the contrasts in vegetation and pH between the patch and matrix were most pronounced exhibited larger differences between patch and matrix communities and tended to have edge communities that differed from those in the patch and forest. There were similar numbers of patch and matrix specialists, but very few edge specialist taxa. Acidobacteria and ectomycorrhizae were more likely to be forest specialists, while Chloroflexi, Ascomycota, and Glomeromycota (i.e., arbuscular mycorrhizae) were more likely to be patch specialists. Contrary to our hypotheses, nonspecialist bacteria were not more likely than specialist bacteria to have larger genomes or to be spore‐formers. We found partial support for our mycorrhizal hypothesis: arbuscular mycorrhizae, but not ectomycorrhizae, were more likely to be specialists. Overall, our results indicate that soil microbial communities are sensitive to edges, but not all taxa are equally affected, with arbuscular mycorrhizae in particular showing a strong response to habitat edges. In the context of increasing habitat fragmentation worldwide, our results can help inform efforts to maintain the structure and functioning of the soil microbiome. 

Abstract 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. 

Abstract Habitat loss is a major threat to biodiversity, but the effects of habitat fragmentation are less clear. Examining drivers of key demographic processes, such as reproduction, will clarify species‐level responses to fragmentation and broader effects on biodiversity. Yet, understanding how fragmentation affects demography has been challenging due to the many ways landscapes are altered by co‐occurring habitat loss and fragmentation, coupled with the rarity of experiments to disentangle these effects.In a large, replicated fragmentation experiment with open savanna habitats surrounded by pine plantation forests, we tested the effects of inter‐patch connectivity, patch edge‐to‐area ratio, and within‐patch distance from an edge on plant reproductive output. Using five experimentally planted species of restoration interest—three wind‐pollinated grass species and two insect‐pollinated forb species—we measured plant flowering, pollination rate, and seed production.All plant species were more likely to flower and produce more flowering structures farther from the forest edge. Connectivity and distance from an edge, however, had no effect on the pollination rate (regardless of pollination mode). Despite no influence of fragmentation on pollination, plant seed production increased farther from the edge for four of five species, driven by the increase in flower production.Synthesis. Altogether, we demonstrate that plant reproductive output (seed production) is decreased by habitat fragmentation through edge effects on flowering. Our work provides evidence that an important contributor to plant demography, reproductive output, is altered by edge effects in fragmented patches. These species‐level impacts of fragmentation may provide insight into the mechanisms of fragmentation effects on community‐level changes in biodiversity. 

Abstract Fragmentation and scaleAlthough habitat loss has well‐known impacts on biodiversity, the effects of habitat fragmentation remain intensely debated. It is often argued that the effects of habitat fragmentation, or the breaking apart of habitat for a given habitat amount, can be understood only at the scale of entire landscapes composed of multiple habitat patches. Yet, fragmentation also impacts the size, isolation and habitat edge for individual patches within landscapes. Addressing the problem of scale on fragmentation effects is crucial for resolving how fragmentation impacts biodiversity. Scaling frameworkWe build upon scaling concepts in ecology to describe a framework that emphasizes three “dimensions” of scale in habitat fragmentation research: the scales of phenomena (or mechanisms), sampling and analysis. Using this framework, we identify ongoing challenges and provide guidance for advancing the science of fragmentation. ImplicationsWe show that patch‐ and landscape‐scale patterns arising from habitat fragmentation for a given amount of habitat are fundamentally related, leading to interdependencies among expected patterns arising from different scales of phenomena. Aggregation of information when increasing the grain of sampling (e.g., from patch to landscape) creates challenges owing to biases created from the modifiable areal unit problem. Consequently, we recommend that sampling strategies use the finest grain that captures potential underlying mechanisms (e.g., plot or patch). Study designs that can capture phenomena operating at multiple spatial extents offer the most promise for understanding the effects of fragmentation and its underlying mechanisms. By embracing the interrelationships among scales, we expect more rapid advances in our understanding of habitat fragmentation. 

Abstract ContextHabitat fragmentation is a leading threat to biodiversity, yet the impacts of fragmentation on most taxa, let alone interactions among those taxa, remain largely unknown. ObjectivesWe studied how three consequences of fragmentation—reduced patch connectivity, altered patch shape, and edge proximity—impact plant-dwelling mite communities and mite-plant-fungus interactions within a large-scale habitat fragmentation experiment. MethodsWe sampled mite communities from the leaves ofQuercus nigra(a plant species that has foliar domatia which harbor fungivorous and predacious mites) near and far from edge within fragments of varying edge-to-area ratio (shape) and connectivity via corridors. We also performed a mite-exclusion experiment across these fragmentation treatments to test the effects of mite presence and fungal hyphal abundance on leaf surfaces. ResultsHabitat edges influenced the abundance and richness of leaf-dwelling mites; plants closer to the edge had higher mite abundance and species richness. Likewise, hyphal counts were higher on leaves near patch edges. Despite both mite and fungal abundance being higher at patch edges, leaf hyphal counts were not impacted by mite abundance on those leaves. Neither patch shape nor connectivity influenced mite abundance, mite species richness, or the influence of mites on leaf surface fungal abundance. ConclusionOur results suggest that mites and foliar fungi may be independently affected by edge-structured environmental gradients, like temperature, rather than trophic effects. We demonstrate that large-scale habitat fragmentation and particularly edge effects can have impacts on multiple levels of microscopic communities, even in the absence of cascading trophic effects. 

Abstract Ecological restoration is beneficial to ecological communities in this era of large‐scale landscape change and ecological disruption. However, restoration outcomes are notoriously variable, which makes fine‐scale decision‐making challenging. This is true for restoration efforts that follow large fires, which are increasingly common as the climate changes.Post‐fire restoration efforts, like tree planting and seeding have shown mixed success, though the causes of the variation in restoration outcomes remain unclear. Abiotic factors such as elevation and fire severity, as well as biotic factors, such as residual canopy cover and abundance of competitive understorey grasses, can vary across a burned area and may all influence the success of restoration efforts to re‐establish trees following forest fires.We examined the effect of these factors on the early seedling establishment of a tree species—māmane (Sophora chrysophylla)—in a subtropical montane woodland in Hawaiʻi. Following a human‐caused wildfire, we sowed seeds of māmane as part of a restoration effort. We co‐designed a project to examine māmane seedling establishment.We found that elevation was of overriding importance, structuring total levels of plant establishment, with fewer seedlings establishing at higher elevations. Residual canopy cover was positively correlated with seedling establishment, while cover by invasive, competitive understorey grasses very weakly positively correlated with increased seedling establishment.Our results point to specific factors structuring plant establishment following a large fire and suggest additional targeted restoration actions within this subtropical system. For example, if greater native woody recruitment is a management goal, then actions could include targeted seed placement at lower elevations where establishment is more likely, increased seeding densities at high elevation where recruitment rates are lower, and/or invasive grass removal prior to seeding. Such actions may result in faster native ecosystem recovery, which is a goal of local land managers. 

Abstract Habitat fragmentation impacts ecosystems worldwide through habitat loss, reduced connectivity, and edge effects. Yet, these landscape factors are often confounded, leaving much to be investigated about their relative effects, especially on species interactions. In a landscape experiment, we investigated the consequences of connectivity and edge effects for seed dispersal by ants. We found that ants dispersed seeds farther in habitat patches connected by corridors, but only in patch centers. We did not see an effect on the total number of seeds moved or the rate ants detected seeds. Furthermore, we did not see any differences in ant community composition across patch types, suggesting that shifts in ant behavior or other factors increased ant seed dispersal in patches connected by corridors. Long‐distance seed dispersal by ants that requires an accumulation of short‐distance dispersal events over generations may be an underappreciated mechanism through which corridors increase plant diversity. 

Abstract Although corridors are frequently regarded as a way to mitigate the negative effects of habitat fragmentation, concerns persist that corridors may facilitate the spread of invasive species to the detriment of native species.The invasive fire ant,Solenopsis invicta,has two social forms. The polygyne form has limited dispersal abilities relative to the monogyne form. Our previous work in a large‐scale corridor experiment showed that in landscapes dominated by the polygyne form, fire ant density was higher and native ant species richness was lower in habitat patches connected by corridors than in unconnected patches.We expected that these observed corridor effects would be transient, that is, that fire ant density and native ant species richness differences between connected and unconnected patches would diminish over time as fire ants eventually fully established within patches. We tested this prediction by resampling the three landscapes dominated by polygyne fire ants 6 to 11 years after our original study.Differences in fire ant density between connected and unconnected habitat patches in these landscapes decreased, as expected. Differences in native ant species richness were variable but lowest in the last 2 years of sampling.These findings support our prediction of transient corridor effects on this invasive ant and stress the importance of temporal dynamics in assessing population and community impacts of habitat connectivity. 

Abstract Habitat loss and fragmentation are leading causes of species declines, driven in part by reduced dispersal. Isolating the effects of fragmentation on dispersal, however, is daunting because the consequences of fragmentation are typically intertwined, such as reduced connectivity and increased prevalence of edge effects. We used a large‐scale landscape experiment to separate consequences of fragmentation on seed dispersal, considering both distance and direction of local dispersal. We evaluated seed dispersal for five wind‐ or gravity‐dispersed, herbaceous plant species that were planted at different distances from habitat edges, within fragments that varied in their connectivity and shape (edge‐to‐area ratio). Dispersal distance was affected by proximity and direction relative to the nearest edge. For four of five species, dispersal distances were greater further from habitat edges and when seeds dispersed in the direction of the nearest edge. Connectivity and patch edge‐to‐area ratio had minimal effects on local dispersal. Our findings illustrate how some, but not all, landscape changes associated with fragmentation can affect the key population process of seed dispersal.