Species traits and the response of open‐habitat species to forest edge in landscape mosaics

Human driven changes in land‐use have increased the need to understand how landscape structure affects species distribution. We studied how forest edges affected the distribution of birds in grasslands recently encroached by forest patches. We investigated how species’ biological traits influenced their response to vegetation change near forest edges. We censured birds along 300‐m line transects run into the open habitat perpendicularly to forest edges. We recorded habitat variables and landscape context along each transect and characterized edges and forest patches. We recorded 33 bird species in 153 transects for a total of 654 individuals. We analyzed species response to edges with generalized linear mixed models. Habitat preference was prevalent to explain species response to forest edges. The abundance of open‐habitat birds such as skylark Alauda arvensis decreased significantly in the vicinity of edges. This negative response extended within 150 m from the edge. The effect was disproportionately higher in open‐habitat species with high conservation concern. The abundance of species feeding or/and breeding in both forest and open habitat, such as woodlarks Lullula arborea, sharply increased near edges (positive edge response). Abundance of shrub and non‐shrub dependent species increased with distance to edge. The two species groups did no differ in abundance/distance to edge relationship. Intensity of species response to forest edges varied among transects in relation to transect vegetation characteristics. Edge length or aspect, diet and nest height had no direct effect. We discuss the possible role of variation in resources and nest predation risk to explain observed patterns.     

Spatial variation in insect community and species responses to habitat loss and plant community composition

Several experimental studies have examined species responses to manipulations of habitat area and spatial arrangement, but plant composition and spatial variation in species distributions also affect animal responses to habitat alteration. We used an experimental approach to study the combined effects of habitat area, edge, and plant community composition on the spatial structure of insect species richness and composition. The abundance of three guilds (herbivores, predators and parasitoids) and individual species were also analyzed. Habitat patches were created that differed in area and edge by selectively mowing portions of 15 m×15 m plots in a 1.7-ha old field. Spatial and environmental variables were used to predict insect responses in separate multiple regression and ordination models. The variation in species responses due to spatial and environmental variables was then partitioned by combining these variables into an overall regression or ordination. Spatial and environmental variables contributed similar percentages to the total variance in insect species richness, abundance or composition. No significant effects of habitat area were observed in any response variable. Herbivore abundance showed positive responses to legume or grass cover, as well as spatial variation that was unrelated to environmental variables. Predators and parasitoids had greater effects of plant species richness and habitat edge, and less unexplained spatial variation. Individual species differed in their responses to plant variables, depending on host specialization or intraspecific aggregation. Our study highlights the importance of plant community composition and spatial variation apart from environmental variables. Spatial variation stems both from species responses to environmental features as well as species differences in habitat specialization and intraspecific aggregation.     

Of mice and mallards: positive indirect effects of coexisting prey on waterfowl nest success

Coexisting prey species interact indirectly via their shared predators when one prey type influences predation rates of the second prey type. In a temperate system where the predominant shared predator is a generalist, I studied the indirect effects of rodent populations on waterfowl nest success, both within the nesting season among sites and among years. Among six to ten upland fields (14 to 27 ha), mallard ( Anas platyrhynchos ) nest success was positively correlated with rodent abundance in all three years of the study. After removing year effects, mallard nest success remained positively correlated with the relative abundance of rodents. Of the rodent species present, California voles ( Microtus californicus ) were the most important coexisting prey type influencing nest success. Among years, mallard nest success was positively correlated with vole abundance; the asymptotic relationship suggests a threshold response to vole abundance, beyond which predators become satiated and additional voles do little to affect nest success. I tested and rejected three alternative explanations for the observed positive correlation between mallard nest success and rodent abundance that do not involve an indirect effect of coexisting prey populations. The influences of dense nesting cover, nesting density, and predator activity did not explain the observed patterns of nest success. These results suggest that rodent populations buffer predation on waterfowl nests, both within and among years, via the behavioral responses of shared predators to coexisting prey.     

The nest predator community of grassland birds responds to agroecosystem habitat at multiple scales

Nest predation is the leading cause of reproductive failure for grassland birds of conservation concern. Understanding variation in nest predation rates is complicated by the diverse assemblage of species known to prey on nests. As part of a long‐term study of grassland bird ecology, we monitored populations of predators known to prey on grassland bird nests. We used information theoretic approach to examine the predator community's association with habitat at multiple scales, including local vegetation structure of grassland patches, spatial attributes of grassland patches (size and shape), and landscape composition surrounding grassland patches (land cover within 400 and 1600 m). Our results confirmed that nest predators respond to habitat at multiple scales and different predator species respond to habitat in different ways. The most informative habitat models we selected included variability in local vegetation (CV in the density of forbs), local patch (area and edge‐to‐interior ratio), and landscape within a 1600 m buffer around grasslands (percent of land covered by human structures and development). As a separate question, we asked if models that incorporated information from multiple scales simultaneously might improve the ability to explain variation in the predator community. Multi‐ scale models were not consistently superior to models derived from variables focused at a single spatial scale. Our results suggest that minimizing human development on and surrounding conservation land and the management of the vegetation structure on grassland fragments both may benefit grassland birds by decreasing the risk of nest predation.     

Predator functional response and prey survival: direct and indirect interactions affecting a marked prey population

1. Predation plays an integral role in many community interactions, with the number of predators and the rate at which they consume prey (i.e. their functional response) determining interaction strengths. Owing to the difficulty of directly observing predation events, attempts to determine the functional response of predators in natural systems are limited. Determining the forms that predator functional responses take in complex systems is important in advancing understanding of community interactions. 2. Prey survival has a direct relationship to the functional response of their predators. We employed this relationship to estimate the functional response for bald eagle Haliaeetus leucocepalus predation of Canada goose Branta canadensis nests. We compared models that incorporated eagle abundance, nest abundance and alternative prey presence to determine the form of the functional response that best predicted intra-annual variation in survival of goose nests. 3. Eagle abundance, nest abundance and the availability of alternative prey were all related to predation rates of goose nests by eagles. There was a sigmoidal relationship between predation rate and prey abundance and prey switching occurred when alternative prey was present. In addition, predation by individual eagles increased as eagle abundance increased. 4. A complex set of interactions among the three species examined in this study determined survival rates of goose nests. Results show that eagle predation had both prey- and predator-dependent components with no support for ratio dependence. In addition, indirect interactions resulting from the availability of alternative prey had an important role in mediating the rate at which eagles depredated nests. As a result, much of the within-season variation in nest survival was due to changing availability of alternative prey consumed by eagles. 5. Empirical relationships drawn from ecological theory can be directly integrated into the estimation process to determine the mechanisms responsible for variation in observed survival rates. The relationship between predator functional response and prey survival offers a flexible and robust method to advance our understanding of predator-prey interactions in many complex natural systems where prey populations are marked and regularly visited.     

Review of the effects of within-patch scale structural complexity on seagrass fishes

Studies on the effects of within-patch scale structure of seagrass habitats on predator–prey fish interactions and abundance/habitat use patterns were reviewed. Most laboratory experiments have employed chase-and-attack predators, usually resulting in lower foraging efficiency in (denser) seagrass. However, a few laboratory procedures employed alternative foraging tactics, resulting in no differences in prey mortality rates. Field studies did not always result in lower prey mortality rates in seagrass habitats. Accordingly, it is premature to conclude that seagrass presence is almost always negatively related to predator foraging efficiency or that increasing seagrass abundance is usually associated with a decrease in predator efficiency. Because several categories of predator and prey fishes occur in seagrass habitats, further studies are needed with all of these predator–prey combinations, in order to fully clarify predator–prey fish interactions in association with seagrass structure. Seagrass fishes have been shown to respond to alterations in seagrass structure in various ways: seagrass height and/or density reduction or clearance resulted in decreased abundance of some species but increases or no change in others. Some explanations have been proposed, not all mutually exclusive, for these phenomena. Although within-patch scale processes have been well studied, room exists for improvement. For example, predator–prey fish interactions in relation to varying within-patch scale complexity is not yet fully understand. The relationships of patch size, edge effects and within-patch scale complexity also still remain unclear. Further studies, which add to the clarification of within-patch scale process, will in turn improve our understanding of larger spatial scale processes.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

陕西红碱淖同域分布遗鸥与棕头鸥巢址选择比较

2009—2011年4—7月在陕西红碱淖对同域分布的遗鸥和棕头鸥巢参数特征、巢群空间分布和巢址生境选择进行了对比分析。结果表明:巢外径、巢深、巢高等参数均存在显著差异,这主要是与营巢生境和巢址空间分布上有关;从巢群空间分布上来看,两者均选择于湖心岛上营巢繁殖,巢群呈斑块状分布,但彼此没有混巢现象,有时甚至通过压缩种间巢间距来达到共存,由于遗鸥繁殖群体数量占优势,遗鸥总是占据最佳的巢域,棕头鸥巢群通常被排挤至湖心岛边缘部分;在巢址生境选择上,植被类型、植被高度、植被盖度、巢离水面高度和巢距最近水边距离均存在显著差异。这表明两者巢址生境选择上具有特异性,这种特异性使得巢址生境选择上存在生态位差异,促成了遗鸥和棕头鸥能在同一湖心岛上营巢繁殖因素之一。     

Contrast in edge vegetation structure modifies the predation risk of natural ground nests in an agricultural landscape

Nest predation risk generally increases nearer forest-field edges in agricultural landscapes. However, few studies test whether differences in edge contrast (i.e. hard versus soft edges based on vegetation structure and height) affect edge-related predation patterns and if such patterns are related to changes in nest conspicuousness between incubation and nestling feeding. Using data on 923 nesting attempts we analyse factors influencing nest predation risk at different edge types in an agricultural landscape of a ground-cavity breeding bird species, the Northern Wheatear (Oenanthe oenanthe). As for many other bird species, nest predation is a major determinant of reproductive success in this migratory passerine. Nest predation risk was higher closer to woodland and crop field edges, but only when these were hard edges in terms of ground vegetation structure (clear contrast between tall vs short ground vegetation). No such edge effect was observed at soft edges where adjacent habitats had tall ground vegetation (crop, ungrazed grassland). This edge effect on nest predation risk was evident during the incubation stage but not the nestling feeding stage. Since wheatear nests are depredated by ground-living animals our results demonstrate: (i) that edge effects depend on edge contrast, (ii) that edge-related nest predation patterns vary across the breeding period probably resulting from changes in parental activity at the nest between the incubation and nestling feeding stage. Edge effects should be put in the context of the nest predator community as illustrated by the elevated nest predation risk at hard but not soft habitat edges when an edge is defined in terms of ground vegetation. These results thus can potentially explain previously observed variations in edge-related nest predation risk.     

Alternative causes of edge-abundance relationships in birds and small mammals of California coastal sage scrub

Changes in the distribution and abundance of bird and small mammal species at urban-wildland edges can be caused by different factors. Edges can affect populations directly if animals respond behaviorally to the edge itself or if proximity to edge directly affects demographic vital rates (an "ecotonal" effect). Alternatively, urban edges can indirectly affect populations if edges alter the characteristics of the adjacent wildland vegetation, which in turn prompts a response to the altered habitat (a "matrix" or "habitat" effect). We studied edge effects of birds and small mammals in southern Californian coastal sage scrub, and assessed whether edge effects were attributable to direct behavioral responses to edges or to animal responses to changes in habitat at edges. Vegetation species composition and structure varied with distance from edge, but the differences varied among study sites. Because vegetation characteristics were correlated with distance from edge, responses to habitat were explored by using independently-derived models of habitat associations to calibrate vegetation measurements to the habitat affinities of each animal species. Of sixteen species examined, five bird and one small mammal species responded to edge independently of habitat features, and thus habitat restoration at edges is expected to be an ineffective conservation measure for these species. Two additional species of birds and one small mammal responded to habitat gradients that coincided with distance from edge, such that the effect of edge on these species was expressed via potentially reversible habitat degradation.     

How population loss through habitat boundaries determines the dynamics of a predator–prey system

The increased persistence of predator–prey systems when interactions are distributed through the space has been acknowledged by both empirical and theoretical studies. One salient feature of predator–prey interactions in heterogeneous space, for example, is the existence of cycles with reduced amplitude when compared with a homogeneous landscape. Although the role of spatial interactions in shaping the dynamics of predator–prey systems has been extensively studied, still very few works have focused on the effects of habitat loss and fragmentation on these systems. In this work, we study the population dynamics of a predator–prey system in a single finite habitat with flux at the boundaries. Species movement and growth are described through a reaction–diffusion model with Rosenzweig–MacArthur type local interactions. Conforming with the existing literature, we find that the reduction of habitat size, or increasing of species movement rates equivalently, has the potential to decrease the amplitude of oscillations and even bring the system to a steady coexistence equilibrium above a threshold. We observe, however, situations in which this trend is reversed. This occurs when species movement rates and response at patch boundaries interact to induce non-trivial patterns of species distributions. These distributions are characterized by anti-correlation between predator and prey, creating then spatial refugia for prey. Our results highlight the role of population loss through habitat boundaries in determining the dynamics of predator–prey interactions.     

Urbanization Impacts on Mammals across Urban-Forest Edges and a Predictive Model of Edge Effects

With accelerating rates of urbanization worldwide, a better understanding of ecological processes at the wildland-urban interface is critical to conserve biodiversity. We explored the effects of high and low-density housing developments on forest-dwelling mammals. Based on habitat characteristics, we expected a gradual decline in species abundance across forest-urban edges and an increased decline rate in higher contrast edges. We surveyed arboreal mammals in sites of high and low housing density along 600 m transects that spanned urban areas and areas turn on adjacent native forest. We also surveyed forest controls to test whether edge effects extended beyond our edge transects. We fitted models describing richness, total abundance and individual species abundance. Low-density housing developments provided suitable habitat for most arboreal mammals. In contrast, high-density housing developments had lower species richness, total abundance and individual species abundance, but supported the highest abundances of an urban adapter (Trichosurus vulpecula). We did not find the predicted gradual decline in species abundance. Of four species analysed, three exhibited no response to the proximity of urban boundaries, but spilled over into adjacent urban habitat to differing extents. One species (Petaurus australis) had an extended negative response to urban boundaries, suggesting that urban development has impacts beyond 300 m into adjacent forest. Our empirical work demonstrates that high-density housing developments have negative effects on both community and species level responses, except for one urban adapter. We developed a new predictive model of edge effects based on our results and the literature. To predict animal responses across edges, our framework integrates for first time: (1) habitat quality/preference, (2) species response with the proximity to the adjacent habitat, and (3) spillover extent/sensitivity to adjacent habitat boundaries. This framework will allow scientists, managers and planners better understand and predict both species responses across edges and impacts of development in mosaic landscapes.     

Edge effects influence the composition and density of reef residents on subtidal restored oyster reefs

Within estuarine and coastal ecosystems globally, extensive habitat degradation and loss threaten critical ecosystem functions and necessitate widescale restoration efforts. There is abundant evidence that ecological processes and species interactions can vary with habitat characteristics, which has important implications for the design and implementation of restoration efforts aimed at enhancing specific ecosystem functions and services. We conducted an experiment examining how habitat characteristics (presence; edge vs. interior) influence the communities of resident fish and mobile invertebrates on restored oyster (Crassostrea virginica) reefs. Similar to previous studies, we found that restored reefs altered community composition and augmented total abundance and biomass relative to unstructured sand habitat. Community composition and biomass also differed between the edge and interior of individual reefs as a result of species-specific patterns over small spatial scales. These patterns were only weakly linked to oyster density, suggesting that other factors that vary between edge and interior (e.g. predator access or species interactions) are likely more important for community structure on oyster reefs. Fine-scale information on resident species' use of oyster reefs will help facilitate restoration by allowing decision makers to optimize the amount of edge versus interior habitat. To improve the prediction of faunal use and benefits from habitat restoration, we recommend investigations into the mechanisms shaping edge and interior preferences on oyster reefs.     

A spatial theory for characterizing predator–multiprey interactions in heterogeneous landscapes

Trophic interactions in multiprey systems can be largely determined by prey distributions. Yet, classic predator–prey models assume spatially homogeneous interactions between predators and prey. We developed a spatially informed theory that predicts how habitat heterogeneity alters the landscape-scale distribution of mortality risk of prey from predation, and hence the nature of predator interactions in multiprey systems. The theoretical model is a spatially explicit, multiprey functional response in which species-specific advection–diffusion models account for the response of individual prey to habitat edges. The model demonstrates that distinct responses of alternative prey species can alter the consequences of conspecific aggregation, from increasing safety to increasing predation risk. Observations of threatened boreal caribou, moose and grey wolf interacting over 378 181 km² of human-managed boreal forest support this principle. This empirically supported theory demonstrates how distinct responses of apparent competitors to landscape heterogeneity, including to human disturbances, can reverse density dependence in fitness correlates.     

Influence of Vegetation on Bat Use of Riparian Areas at Multiple Spatial Scales

ABSTRACT Research on habitat use by bats typically occurs at a single fine spatial scale, despite recent work demonstrating the importance of considering multiple spatial scales when investigating vertebrate habitat selection. We assessed bat use of 118 stream reaches located throughout the Oregon Coast Range, USA, and measured vegetation characteristics at 3 spatial scales surrounding each of these locations. We used an information-theoretic approach to determine vegetation characteristics most closely related to bat activity and a multilevel modeling approach to evaluate variation in bat activity at different spatial scales. Characteristics of vegetation at the finest spatial scale explained more variation in bat activity than did characteristics of vegetation at broader spatial scales, suggesting that fine-scale anthropogenic or natural disturbance events that alter cover of shrubs or trees in riparian areas are likely to influence bat habitat use. The influence of vegetation on activity varied by species of bat and appeared to operate more strongly through constraints imposed by vegetation architecture than through influences on abundance of insect prey. This diversity of responses to vegetation characteristics among bat species suggests that the best strategy for biodiversity conservation over broad spatial scales is maintenance or creation of a diversity of riparian vegetation conditions. We recommend that land managers planning to manipulate riparian vegetation strive to create diversity in shrub coverage, canopy coverage, and open space above the stream channel to promote foraging habitat for all species.     

Testing mechanistic explanations for mammalian predator responses to habitat edges

Increased predator activity along habitat edges (the edge effect) is often documented in the temperate zone, but earlier studies have rarely been able to explicitly test the suggested mechanisms underlining this phenomenon. In this study, we measured the distribution of mammalian predators by scent stations and their main prey (rodents) in four types of landscape elements corresponding to an edge gradient between two habitat types; grassland and forest. We found a contrasting pattern in carnivore activities between years. Whereas carnivores did not exhibit a significant occurrence along the forest–grassland edge in the first year with low prey abundance, they were more likely to be detected along habitat edges in the subsequent year with high prey abundance. Our results do not suggest that the increased activity of carnivores at habitat edges arises as a consequence of predator overflow from higher quality habitat through the edge into lower quality habitat, but showed that most predator species focus their activity specifically to the edge structure. On the other hand, our data do not provide straightforward support for the hypothesis that predator edge preferences are caused by a spatial gradient in carnivores’ main prey—rodents.     

Intraguild dynamics of understudied carnivores in a human‐altered landscape

Mesocarnivores constitute a diverse and often abundant group of species, which are increasingly occupying hweigher trophic levels within multi‐use landscapes. Yet, we know relatively little about their interactions with each other, especially in human‐altered areas. Using camera trap data collected in a forestry concession in the Greater Gorongosa ecosystem of central Mozambique, we examined the spatiotemporal relationships and potential for intraguild competition among three understudied African carnivores: African civets (Civettictis civetta), bushy‐tailed mongooses (Bdeogale crassicauda), and large‐spotted genets (Genetta maculata). After accounting for habitat preferences and tolerance to anthropogenic factors, we found that African civets and bushy‐tailed mongooses avoid each other spatially and temporally. Additionally, civets and mongooses were also both more likely to use sites farther away from human settlements, possibly decreasing the total available habitat for each species if competition is driving this spatial partitioning. In contrast, we did not find evidence for spatial or temporal partitioning between large‐spotted genets and African civets, but bushy‐tailed mongooses altered their activity patterns where they co‐occurred with genets. Our study contributes to scant ecological knowledge of these mesocarnivores and adds to our understanding of community dynamics in human‐altered ecosystems.     

Factors affecting species number and density of dabbling duck guilds in North Europe

We addressed how species number and pair density in guilds of co-existing species is related to habitat structure, and to the abundance and diversity of food resources using the assemblage of seven species of dabbling ducks (genus Anas ) breeding in 60 lakes distributed over six regions in temperate north Europe
Partial correlation and multiple regression revealed that species richness was best predicted by habitat structural diversity as indexed by a principal component analysis based on 18 vegetation and lake characteristics, and by the abundance of aquatic and emergent prey We found no effect of lake size or prey size diversity on species richness Pair density was correlated with the percentage of shoreline with horsetails ( Equisetum ), by habitat structural diversity and by the abundance of emergent invertebrate prey Neither prey size diversity nor abundance of aquatic prey correlated with pair density Species richness and pair density in North European duck guilds vary both with habitat structure and prey availability     

Aggregative response in bats: prey abundance versus habitat

In habitats where prey is either rare or difficult to predict spatiotemporally, such as open habitats, predators must be adapted to react effectively to variations in prey abundance. Open-habitat foraging bats have a wing morphology adapted for covering long distances, possibly use information transfer to locate patches of high prey abundance, and would therefore be expected to show an aggregative response at these patches. Here, we examined the effects of prey abundance on foraging activities of open-habitat foragers in comparison to that of edge-habitat foragers and closed-habitat foragers. Bat activity was estimated by counting foraging calls recorded with bat call recorders (38,371 calls). Prey abundance was estimated concurrently at each site using light and pitfall traps. The habitat was characterized by terrestrial laser scanning. Prey abundance increased with vegetation density. As expected, recordings of open-habitat foragers clearly decreased with increasing vegetation density. The foraging activity of edge- and closed-habitat foragers was not significantly affected by the vegetation density, i.e., these guilds were able to forage from open habitats to habitats with dense vegetation. Only open-habitat foragers displayed a significant and proportional aggregative response to increasing prey abundance. Our results suggest that adaptations for effective and low-cost foraging constrains habitat use and excludes the guild of open-habitat foragers from foraging in habitats with high prey abundance, such as dense forest stands.     

The Seasonal Dynamics of Artificial Nest Predation Rates along Edges in a Mosaic Managed Reedbed

Boundaries between different habitats can be responsible for changes in species interactions, including modified rates of encounter between predators and prey. Such ‘edge effects’ have been reported in nesting birds, where nest predation rates can be increased at habitat edges. The literature concerning edge effects on nest predation rates reveals a wide variation in results, even within single habitats, suggesting edge effects are not fixed, but dynamic throughout space and time. This study demonstrates the importance of considering dynamic mechanisms underlying edge effects and their relevance when undertaking habitat management. In reedbed habitats, management in the form of mosaic winter reed cutting can create extensive edges which change rapidly with reed regrowth during spring. We investigate the seasonal dynamics of reedbed edges using an artificial nest experiment based on the breeding biology of a reedbed specialist. We first demonstrate that nest predation decreases with increasing distance from the edge of cut reed blocks, suggesting edge effects have a pivotal role in this system. Using repeats throughout the breeding season we then confirm that nest predation rates are temporally dynamic and decline with the regrowth of reed. However, effects of edges on nest predation were consistent throughout the season. These results are of practical importance when considering appropriate habitat management, suggesting that reed cutting may heighten nest predation, especially before new growth matures. They also contribute directly to an overall understanding of the dynamic processes underlying edge effects and their potential role as drivers of time-dependent habitat use.