Biodiversity extinction thresholds are modulated by matrix type

Biodiversity extinction thresholds are abrupt declines in biological diversity that occur with habitat loss, associated with a decline in habitat connectivity. Matrix quality should influence the location of thresholds along habitat loss gradients through its effects on connectivity; however these relationships have seldom been explored empirically. Using field data from 23 independent 1254 ha landscapes in the Brazilian Atlantic Forest, we evaluated how tropical avian biodiversity responds to native forest loss within habitat patches embedded either in homogeneous pasture matrix context (with a high proportion of cattle pastures), and heterogeneous coffee matrix context (with high abundance of sun coffee plantations). We considered taxonomic, functional, and phylogenetic diversity, and tested if matrix type and choice of diversity metric influenced the location of biodiversity thresholds along the forest cover gradient. We found that matrix type postponed the abrupt loss of taxonomic diversity, from a threshold of 35% of forest cover in homogeneous pasture matrix to 19% in heterogeneous coffee matrix. Phylogenetic diversity responded similarly, with thresholds at 30 and 24% in homogeneous‐pasture and heterogeneous‐coffee matrices, respectively, but no relationship with forest cover was detected when corrected for richness correlation. Despite the absence of a threshold for functional diversity in either matrix types, a strong decline below 20% of habitat amount was detected. Finally, below 20% native habitat loss, all diversity indices demonstrated abrupt declines, indicating that even higher‐quality matrices cannot postpone diversity loss below this critical threshold. These results highlight that taxonomic diversity is a more sensitive index of biodiversity loss in fragmented landscapes, which may be used as a benchmark to prevent subsequent functional and phylogenetic losses. Furthermore, increasing matrix quality appears an efficient conservation strategy to maintain higher biodiversity levels in fragmented landscapes over a larger range of habitat loss.     

Matrix heterogeneity affects population size of the harvest mice: Bayesian estimation of matrix resistance and model validation

It has been increasingly recognized that landscape matrices are an important factor determining patch connectivity and hence the population size of organisms living in highly fragmented landscapes. However, most previous studies estimated the effect of matrix heterogeneity using prior information regarding dispersal or habitat preferences of a focal organism. Here we estimated matrix resistance of harvest mice in agricultural landscapes using a novel pattern‐oriented modeling with Bayesian estimation and no prior information, and then conducted model validation using data sets independent from those used for model construction. First, we investigated the distribution patterns of harvest mice for approximately 400 habitat patches, and estimated matrix resistance for different matrix types using statistical models incorporating patch size, patch environment, and patch connectivity. We used Bayesian estimation with a Markov chain Monte Carlo algorithm, and searched for appropriate matrix resistance that best explained the distribution pattern. Patch connectivity as well as patch quality was an important determinant of local population size for the harvest mice. Moreover, matrix resistance was far from uniform, with rice and crop fields exhibiting low resistance and forests, creeks, roads and residential areas showing much higher resistance. The deviance explained by this model (heterogeneous matrix model) was much larger than that obtained by the model with no consideration of matrix heterogeneity (homogeneous matrix model). Second, we obtained distribution data from five additional landscapes that were more fragmented than that used for model construction, and used them for model validation. The heterogeneous matrix model well predicted the population size for four out of five landscapes. In contrast, the homogeneous model considerably overestimated population sizes in all cases. Our approach is widely applicable to species living in fragmented landscapes, especially those for which prior information regarding movement or dispersal is difficult to obtain.     

Edge effects as the principal cause of area effects on birds in fragmented secondary forest

Bird communities in tropical forests are strongly affected by both patch area and habitat edges. The fact that both effects are intrinsically confounded in space raises questions about how these two widely reported ecological patterns interact, and whether they are independent or simply different spatial manifestations of the same phenomenon. Moreover, do small patches of secondary forest, in landscapes where the most sensitive species have gone locally extinct, exhibit similar patterns to those previously observed in fragmented and continuous primary forests? We addressed these questions by testing edge‐related differences in vegetation structure and bird community composition at 31 sites in fragmented and continuous landscapes in the imperilled Atlantic forest of Brazil. Over a two‐year period, birds were captured with mist nets to a standardized effort of 680 net‐hours at each site (～22 000 net‐hours resulting in 3381 captures from 114 species). We found that the bird community in patches of secondary forest was degraded in species composition compared to primary continuous forest, but still exhibited a strong response to edge effects. In fragmented secondary forests, edge and area effects also interacted, such that the magnitude of edge to interior differences on bird community composition declined markedly with patch size. The change in bird species composition between forest interiors and edges was similar to the change in community composition between large and small patches (because species had congruent responses to edge and area), but after controlling for edge effects community composition was no longer affected by patch area. Our results show that although secondary forests hold an impoverished bird community, ecological patterns such as area and edge effects are similar to those reported for primary forests. Our data provide further evidence that edge effects are the main drivers of area effects in fragmented landscapes.     

The role of habitat connectivity and landscape geometry in experimental zooplankton metacommunities

Theory predicts that inter-patch dispersal rates and patterns of patch heterogeneity both have the potential to alter patterns of local and regional species diversity. To test this, we manipulated both rates of habitat connectivity and the geometric arrangement of habitat heterogeneity within regions of experimental zooplankton communities. We found no effects of habitat geometry on any metric of species diversity or composition. Additionally, we found no effect of habitat connectivity rate on local species diversity. We did, however, find that increasing connectivity led to a decrease in regional diversity, as well as an increase in the percent similarity of local communities within regions. Of all of the species in these communities, the relatively large cladoceran Ceriodaphnia reticulata significantly responded to the treatments, and had a higher probability of achieving high densities when connectance was high. As such, we suggest that this species played a large role in driving the increased local community similarity and decreased regional species richness as connectivity increased. These findings are in opposition to previous experimental studies of metacommunities, but support the notion that increased connectance among local patches may decrease regional diversity when patches are heterogeneous.     

The social elite: Habitat heterogeneity,complexity and quality in granite inselbergs influence patterns of aggregation in Egernia striolata (Lygosominae: Scincidae)

Habitat heterogeneity, structural complexity and habitat quality are key features of the environment that drive species' distribution and patterns of biological organization. Traditionally, pattern‐based studies have focused on faunal responses to biological systems. However, the influence of non‐biological environments such as insular rock outcrops on patterns of vertebrate distribution is conceivably as important, but has received less attention. Granite inselbergs are a naturally heterogeneous and spatially‐limited habitat. As such, they provide an opportunity for investigating whether environmental attributes influence social behaviour in animals that use these kinds of habitat, particularly lizards that are well adapted to saxicoline environments. We applied ecological theory to investigate the influence of habitat heterogeneity, structural complexity and habitat quality on patterns of home‐site occupancy in the crevice skink Egernia striolata (Lygosominea: Scincidae) from insular granite outcrops located within fragmented agricultural landscapes. We compared home‐site occupancy among solitary juveniles, solitary adults and lizard aggregations. We found significant differences in home‐site occupancy between aggregations and solitary lizard outcrop attributes measured at multiple spatial scales. The probability of a home‐site being occupied by an aggregation increased where large rock masses were present, on northern aspects near the core of the outcrop and in structurally variegated landscapes. Significantly more aggregations occupied home‐sites surrounded by high boulder cover and crevice microhabitat. We provide evidence that geophysical attributes of granite inselbergs and landscape context can influence patterns of lizard aggregation. Thus, we clearly document the environmental correlations of variability in sociality among subpopulations of Egernia striolata.     

Anthropogenic landscape change promotes asymmetric dispersal and limits regional patch occupancy in a spatially structured bird population

1. Local extinctions in habitat patches and asymmetric dispersal between patches are key processes structuring animal populations in heterogeneous environments. Effective landscape conservation requires an understanding of how habitat loss and fragmentation influence demographic processes within populations and movement between populations. 2. We used patch occupancy surveys and molecular data for a rainforest bird, the logrunner (Orthonyx temminckii), to determine (i) the effects of landscape change and patch structure on local extinction; (ii) the asymmetry of emigration and immigration rates; (iii) the relative influence of local and between-population landscapes on asymmetric emigration and immigration; and (iv) the relative contributions of habitat loss and habitat fragmentation to asymmetric emigration and immigration. 3. Whether or not a patch was occupied by logrunners was primarily determined by the isolation of that patch. After controlling for patch isolation, patch occupancy declined in landscapes experiencing high levels of rainforest loss over the last 100 years. Habitat loss and fragmentation over the last century was more important than the current pattern of patch isolation alone, which suggested that immigration from neighbouring patches was unable to prevent local extinction in highly modified landscapes. 4. We discovered that dispersal between logrunner populations is highly asymmetric. Emigration rates were 39% lower when local landscapes were fragmented, but emigration was not limited by the structure of the between-population landscapes. In contrast, immigration was 37% greater when local landscapes were fragmented and was lower when the between-population landscapes were fragmented. Rainforest fragmentation influenced asymmetric dispersal to a greater extent than did rainforest loss, and a 60% reduction in mean patch area was capable of switching a population from being a net exporter to a net importer of dispersing logrunners. 5. The synergistic effects of landscape change on species occurrence and asymmetric dispersal have important implications for conservation. Conservation measures that maintain large patch sizes in the landscape may promote asymmetric dispersal from intact to fragmented landscapes and allow rainforest bird populations to persist in fragmented and degraded landscapes. These sink populations could form the kernel of source populations given sufficient habitat restoration. However, the success of this rescue effect will depend on the quality of the between-population landscapes.     

Landscape-level comparison of genetic diversity and differentiation in a small mammal inhabiting different fragmented landscapes of the Brazilian Atlantic Forest

Habitat loss and fragmentation can have detrimental effects on all levels of biodiversity, including genetic variation. Most studies that investigate genetic effects of habitat loss and fragmentation focus on analysing genetic data from a single landscape. However, our understanding of habitat loss effects on landscape-wide patterns of biodiversity would benefit from studies that are based on quantitative comparisons among multiple study landscapes. Here, we use such a landscape-level study design to compare genetic variation in the forest-specialist marsupial Marmosops incanus from four 10,000-hectare Atlantic forest landscapes which differ in the amount of their remaining native forest cover (86, 49, 31, 11 %). Additionally, we used a model selection framework to evaluate the influence of patch characteristics on genetic variation within each landscape. We genotyped 529 individuals with 12 microsatellites to statistically compare estimates of genetic diversity and genetic differentiation in populations inhabiting different forest patches within the landscapes. Our study indicates that before the extinction of the specialist species (here in the 11 % landscape) genetic diversity is significantly reduced in the 31 % landscape, while genetic differentiation is significantly higher in the 49 and 31 % landscapes compared to the 86 % landscape. Results further provide evidence for non-proportional responses of genetic diversity and differentiation to increasing habitat loss, and suggest that local patch isolation impacts gene flow and genetic connectivity only in the 31 % landscape. These results have high relevance for analysing landscape genetic relationships and emphasize the importance of landscape-level study designs for understanding habitat loss effects on all levels of biodiversity.     

Relative importance of quantity, quality and isolation of patches for butterfly diversity in fragmented urban forests

The majority of forests in urban areas are small and isolated. Improving habitat quality of small forests instead of increasing habitat size and connectivity could be an effective means of conserving the biodiversity of such highly fragmented landscapes. In this study, we investigated the relative importance of habitat quantity, quality and isolation on butterfly assemblages in urban fragmented forests in Tokyo, Japan. We used four habitat geographic parameters: (1) fragment size, (2) shape index, (3) isolation (distance to the mainland), and (4) connectivity; and three habitat quality parameters: (1) herbaceous nectar plant abundance, (2) herbaceous nectar plant diversity, and (3) larval host plant diversity. We surveyed butterfly assemblages along transects in 20 forest fragments that ranged in size from 1 to 122 ha. We used generalized linear models to relate the number of species in a fragment to four habitat geographic parameters and three habitat quality parameters. The averaged models based on AIC_c showed that fragment size had a strong positive effect on butterfly species richness. There was also a positive effect of herbaceous nectar plant abundance on species diversity. These findings suggest that improving the habitat quality of small and isolated forests in highly fragmented landscapes may be capable of maintaining levels of butterfly diversity comparable to those of large fragments.     

Metacommunity,mainland‐island system or island communities? Assessing the regional dynamics of plant communities in a fragmented landscape

Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among‐patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape‐scale. In this study, we used extensive field data from a fragmented, semi‐arid landscape in Israel to parameterize a multi‐species incidence‐function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics – the metacommunity, the mainland‐island, or the island communities type – best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch‐matrix study landscape is best represented as a system of highly isolated ‘island’ communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33–60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.     

Stream communities across a rural–urban landscape gradient

Rapid urbanization throughout the world is expected to cause extensive loss of biodiversity in the upcoming decades. Disturbances associated with urbanization frequently operate over multiple spatial scales such that local species extirpations have been attributed both to localized habitat degradation and to regional changes in land use. Urbanization also may shape stream communities by restricting species dispersal within and among stream reaches. In this patch-dynamics view, anthropogenic disturbances and isolation jointly reduce stream biodiversity in urbanizing landscapes. We evaluated predictions of stream invertebrate community composition and abundance based on variation in environmental conditions at five distinct spatial scales: stream habitats, reaches, riparian corridors and watersheds and their spatial location within the larger three-river basin. Despite strong associations between biodiversity loss and human density in this study, local stream habitat and stream reach conditions were poor predictors of community patterns. Instead, local community diversity and abundance were more accurately predicted by riparian vegetation and watershed landscape structure. Spatial coordinates associated with instream distances provided better predictions of stream communities than any of the environmental data sets. Together, results suggest that urbanization in the study region was associated with reduced stream invertebrate diversity through the alteration of landscape vegetation structure and patch connectivity. These findings suggest that maintaining and restoring watershed vegetation corridors in urban landscapes will aid efforts to conserve freshwater biodiversity.     

Patch occupancy and abundance of local populations in landscapes differing in degree of habitat fragmentation: a case study of the colonial black‐headed gull,Chroicocephalus ridibundus

Aim This study investigated whether habitat fragmentation at the landscape level influences patch occupancy and abundance of the black‐headed gull, Chroicocephalus ridibundus, and whether the response of the species to environmental factors is consistent across replicated landscape plots. Location Water bodies (habitat patches) in southern Poland. Methods Surveys were conducted in two landscape types (four plots in each): (1) more‐fragmented landscape, in which habitat patches were small (mean size 2.2–6.2 ha) and far apart (mean distance 2.5–3.1 km); and (2) less‐fragmented landscape, in which habitat patches were large (mean size 9.2–16.5 ha) and separated by short distances (mean 0.9–1.4 km). Observations were performed twice in 284 potential habitat patches during the 2007 breeding season. Results Colonies were significantly more frequent and larger in the less‐fragmented landscapes than in the more‐fragmented ones. Probability of patch occupancy and number of breeding birds were positively related with patch size and these relationships were especially strong in the more‐fragmented landscapes. In the less‐fragmented landscapes, the occurrence of black‐headed gulls was negatively related to the distance to the nearest local population, but in the more‐fragmented landscapes such a relationship was not detected. As distance to the nearest habitat patch increased, the probability of the patch occupancy decreased in the more‐fragmented landscapes. Moreover, abundance was negatively influenced by distance to the nearest habitat patch, especially strongly in more‐fragmented landscapes. Proximity of corridors (rivers) positively influenced the occupation of patches regardless of landscape type. The number of islets positively influenced occupancy and abundance of local populations, and this relationship was stronger in the more‐fragmented landscapes. Main conclusions Our results are in agreement with predictions from metapopulation theory and are the first evidence that populations of black‐headed gulls may have a metapopulation structure. However, patch occupancy and abundance were differentially affected by explanatory variables in the more‐fragmented landscapes than in the less‐fragmented ones. This implies that it is impossible to derive, a priori, predictions about presence/abundance patterns based on only a single landscape.     

Multifaceted Analysis of Patch-Level Plant Diversity in Response to Landscape Spatial Pattern and History on Mediterranean Dunes

Landscape structure is known to critically affect biodiversity. However, although the multi-facetted character of biodiversity is widely recognized, few studies have linked landscape spatial pattern and history simultaneously to multiple facets (taxonomic, functional, and phylogenetic) and spatial components (α, β, and γ) of plant diversity. We set out to reveal whether landscape parameters have specific effects on the separate diversity facets and components of plant diversity at a patch scale on coastal dune landscapes of Central Italy. For each landscape patch, we computed a set of patch-based metrics relying on multi-temporal land-cover maps. Based on a database of plant community plots, on functional traits from field measurements and on a dated phylogenetic tree, we calculated taxonomic (TD), functional (FD), and phylogenetic diversity (PD) within each patch at α, β, and γ level. Diversity measures were then related to the landscape metrics via linear mixed-effect models. Landscape pattern and transformations affected TD only moderately in coastal dune ecosystems. We found much stronger and contrasted effects on FD and PD. FD increased in patches surrounded by human-dominated habitats; PD was higher in fragmented patches, particularly in the Mediterranean macchia. Moreover, landscape pattern affected differently the single communities, the turnover among communities and the pool of species within the patch (α, β, and γ components). Our results call for the combined inclusion of FD and PD and their partitions into ecological analyses, being TD too crude to capture the comprehensive and contrasted response of plant diversity to landscape spatial pattern.     

Thresholds of riparian forest use by terrestrial mammals in a fragmented Amazonian deforestation frontier

Species persistence in fragmented landscapes is intimately related to the quality, structure, and context of remaining habitat remnants. Riparian vegetation is legally protected within private landholdings in Brazil, so we quantitatively assessed occupancy patterns of terrestrial mammals in these remnants, examining under which circumstances different species effectively use them. We selected 38 riparian forest patches and five comparable riparian sites within continuous forest, at which we installed four to five camera-traps per site (199 camera-trap stations). Terrestrial mammal assemblages were sampled for 60 days per station during the dry seasons of 2013 and 2014. We modelled species occupancy and detection probabilities within riparian forest remnants, and examined the effects of patch size, habitat quality, and landscape structure on occupancy probabilities. We then scaled-up modelled occupancies to all 1915 riparian patches throughout the study region to identify which remnants retain the greatest potential to work as habitat for terrestrial vertebrates. Of the ten species for which occupancy was modelled, six responded to forest quality (remnant degradation, cattle intrusion, palm aggregations, and understorey density) or structure (remnant width, isolation, length, and area of the patch from which it originates). Patch suitability was lower considering habitat quality than landscape structure, and virtually all riparian remnants were unsuitable to maintain a high occupancy probability for all species that responded to forest patch quality or structure. Beyond safeguarding legal compliance concerning riparian remnant amount, ensuring terrestrial vertebrate persistence in fragmented landscapes will require curbing the drivers of forest degradation within private landholdings.     

Salinisation reduces biodiversity in neighbouring freshwater patches in experimental metacommunities

1. Aquatic ecosystems are biodiversity hot spots across many landscapes; therefore, the degradation of these habitats can lead to decreases in biodiversity across multiple scales. Salinisation is a global issue that threatens freshwater ecosystems by reducing water quality and local biodiversity. The effects of salinity on local processes have been studied extensively; however, the effects of salinisation or similar environmental stressors within a metacommunity (a dispersal network of several distinct communities) have not been explored.
2. We tested how the spatial heterogeneity and the environmental contrast between freshwater and saline habitat patches influenced cladoceran biodiversity and species composition at local and regional scales in a metacommunity mesocosm experiment. We defined spatial heterogeneity as the proportion of freshwater to saltwater patches within the metacommunity, ranging from a freshwater-dominated metacommunity to a saltwater-dominated metacommunity. Environmental contrast was defined as the environmental distance between habitat patches along the salinity gradient in which low-contrast metacommunities consisted of freshwater and low-salinity patches and high-contrast metacommunities consisted of freshwater and high-salinity patches.
3. We hypothesised that the α-richness of freshwater patches and metacommunity γ-richness would decrease as freshwater patches became less abundant along the spatial heterogeneity gradient in both low- and high-contrast metacommunities, because there would be fewer freshwater patches that could serve as source populations for declining populations. We hypothesised that low-contrast metacommunities would support more species across the spatial heterogeneity gradient than high-contrast metacommunities, because, via dispersal, low-salinity patches can support halotolerant freshwater species that can mitigate population declines in neighbouring freshwater patches, whereas` high-salinity patches will mostly support halophilic species, providing fewer potential colonisers to freshwater patches.
4. We found that α-richness of freshwater mesocosms and metacommunity γ-richness declined in saline-dominated metacommunities regardless of the environmental contrast between the freshwater and saline mesocosms. We found that environmental contrast influenced freshwater and saline community composition in low-contrast metacommunities by increasing the abundances of species that could tolerate low-salinity environments through dispersal, whereas freshwater and high-salinity communities showed limited interactions through dispersal.
5. Freshwater mesocosms had a disproportionate effect on the local and regional biodiversity in these experimental metacommunities, indicating that habitat identity may be more important than habitat diversity for maintaining biodiversity in some metacommunities. This study further emphasises the importance in maintaining multiple species-rich habitat patches across landscapes, particularly those experiencing landscape-wide habitat degradation.

     

Using historical ecology to understand patterns of biodiversity in fragmented agricultural landscapes

Aim To enhance current attempts to understand biodiversity patterns by using an historical ecology approach to highlight the over‐riding influence of land‐use history in creating past, current and future patterns of biodiversity in fragmented agricultural landscapes. Methods We develop an integrative conceptual framework for understanding spatial and temporal variations in landscape patterns in fragmented agricultural landscapes by presenting five postulates (hypotheses) which highlight the important role of historical, anthropogenic disturbance regimes. We then illustrate each of these postulates with examples drawn from fragmented woodlands in agricultural areas of south‐eastern Australia, and discuss these findings in an international context. Location examples are drawn from agricultural areas in south‐eastern Australia. Results We conclude that there is limited potential to refine our understanding of patterns of biodiversity in human‐modified landscapes based on traditional concepts of island biogeography, or simple assumptions of ongoing destruction and degradation. Instead, we propose that in agricultural landscapes that were largely cleared over a century ago: (1) present‐day remnant vegetation patterns are not accidental, but are logically arrayed due to historic land‐use decisions, (2) historic anthropogenic disturbances have a major influence on current ecosystem conditions and diversity patterns, and (3) the condition of remnant ecosystems is not necessarily deteriorating rapidly. Main conclusions An historical ecology approach can enhance our understanding of why different species and ecosystem states occur where they do, and can explain internal variations in ecological conditions within remnant ecosystems, too often casually attributed to the ‘mess of history’. This framework emphasizes temporal changes (both past and future) in biotic patterns and processes in fragmented agricultural landscapes. Integration of spatially and temporally explicit historical land‐use information into ecological studies can prove extremely useful to test hypotheses of the effects of changes in landscape processes, and to enhance future research, restoration and conservation management activities.     

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km²) that differ in the amount and heterogeneity of semi‐natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi‐natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi‐natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi‐natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non‐crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.     

Does the type of matrix matter? A quantitative review of the evidence

It has been increasingly recognized that the type of matrix surrounding habitat patches can affect biodiversity in landscapes, but there were only qualitative reviews of the subject focused on particular taxonomic groups. We present a quantitative review of studies from 1985 to 2008 that compared effects of different matrix types on individuals, populations and communities. We compiled 104 studies, most on animals, covering a broad range of landscape types and spatial scales. Most studies were empirical, focused on individuals and communities, and evaluated abundance/richness in the patch as the dependent variable. The type of matrix surrounding habitat patches influenced the studied parameters in 95% of the studies, but such effects were overall smaller compared to patch size or isolation effects. Matrix type effects were strongly species-specific, with different species responding differently to matrix type in 96% of studies comparing species or group of species. In 88% of studies, matrix types more similar in structure to the patch had higher quality for the studied organisms from the point of view of functional connectivity. Overall, the type of matrix is important, but patch size and isolation are the main determinants of ecological parameters in landscapes. Matrix quality generally increases with increasing structural similarity with habitat patches, a pattern that could be used as a general guideline for management of the matrix in fragmented landscapes.     

Meta‐analysis reveals the importance of matrix composition for animals in fragmented habitat

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

The role of habitat quality in fragmented landscapes: a conceptual overview and prospectus for future research

There is increasing empirical evidence that the quality of habitat patches (determined by either habitat degradation or natural heterogeneity in the quality of habitat) plays an important role in determining species distribution patterns and in regulating spatial dynamics in fragmented landscapes. However, to date, most of the debate has focused on whether or not to include habitat variables in fragmentation studies, and we still lack general conclusions as well as standard and robust research approaches. In this paper we show how a weak conceptualization of “patch quality” and the inappropriate choice of target surrogate variables (e.g., density is often used as an indicator of patch quality) have mainly produced case-specific results, rather than general conclusions. We then identify weaknesses in the inclusion of habitat quality measurements within fragmentation studies. In particular, we focus on: (1) the lack of appropriate experimental design, outlining how few studies have actually included a gradient of habitat quality in their sample; (2) the lack of fundamental information provided (e.g., lack of standard outputs), which in turn hampers the possibility of carrying out meta-analyses. We finally synthesize available knowledge from empirical studies and highlight the different conceptual frameworks needed for patch occupancy versus patch use studies.     

Scale dependent drivers of wild bee diversity in tropical heterogeneous agricultural landscapes

Factors associated with agricultural intensification, for example, loss of seminatural vegetation and pesticide use has been shown to adversely affect the bee community. These factors may impact the bee community differently at different landscape scales. The scale dependency is expected to be more pronounced in heterogeneous landscapes. However, the scale‐dependent response of the bee community to drivers of its decline is relatively understudied, especially in the tropics where the agricultural landscape is often heterogeneous. This study looked at effects of agricultural intensification on bee diversity at patch and landscape scales in a tropical agricultural landscape. Wild bees were sampled using 12 permanent pan trap stations. Patch and landscape characteristics were measured within a 100 m (patch scale) and a 500 m (landscape scale) radius of pan trap stations. Information on pesticide input was obtained from farmer surveys. Data on vegetation cover, productivity, and percentage of agricultural and fallow land (FL) were collected using satellite imagery. Intensive areas in a bee‐site network were less specialized in terms of resources to attract rare bee species while the less intensive areas, which supported more rare species, were more vulnerable to disturbance. A combination of patch quality and diversity as well as pesticide use regulates species diversity at the landscape scale (500 m), whereas pesticide quantity drove diversity at the patch scale (100 m). At the landscape scale, specialization of each site in terms of resources for bees increased with increasing patch diversity and FL while at the patch scale specialization declined with increased pesticide use. Bee functional groups responded differentially to landscape characteristics as well as pesticide use. Wood nesting bees were negatively affected by the number of pesticides used but other bee functional groups were not sensitive to pesticides. Synthesis and Applications: Different factors affect wild bee diversity at the scale of landscape and patch in heterogeneous tropical agricultural systems. The differential response of bee functional groups to agricultural intensification underpins the need for guild‐specific management strategies for wild bee conservation. Less intensively farmed areas support more rare species and are vulnerable to disturbance; consequently, these areas should be prioritized for conservation to maintain heterogeneity in the landscape. It is important to conserve and restore seminatural habitats to maintain complexity in the landscapes through participatory processes and to regulate synthetic chemical pesticides in farm operations to conserve the species and functional diversity of wild bees.