Beyond the Fragmentation Threshold Hypothesis: Regime Shifts in Biodiversity Across Fragmented Landscapes

Ecological systems are vulnerable to irreversible change when key system properties are pushed over thresholds, resulting in the loss of resilience and the precipitation of a regime shift. Perhaps the most important of such properties in human-modified landscapes is the total amount of remnant native vegetation. In a seminal study Andren proposed the existence of a fragmentation threshold in the total amount of remnant vegetation, below which landscape-scale connectivity is eroded and local species richness and abundance become dependent on patch size. Despite the fact that species patch-area effects have been a mainstay of conservation science there has yet to be a robust empirical evaluation of this hypothesis. Here we present and test a new conceptual model describing the mechanisms and consequences of biodiversity change in fragmented landscapes, identifying the fragmentation threshold as a first step in a positive feedback mechanism that has the capacity to impair ecological resilience, and drive a regime shift in biodiversity. The model considers that local extinction risk is defined by patch size, and immigration rates by landscape vegetation cover, and that the recovery from local species losses depends upon the landscape species pool. Using a unique dataset on the distribution of non-volant small mammals across replicate landscapes in the Atlantic forest of Brazil, we found strong evidence for our model predictions - that patch-area effects are evident only at intermediate levels of total forest cover, where landscape diversity is still high and opportunities for enhancing biodiversity through local management are greatest. Furthermore, high levels of forest loss can push native biota through an extinction filter, and result in the abrupt, landscape-wide loss of forest-specialist taxa, ecological resilience and management effectiveness. The proposed model links hitherto distinct theoretical approaches within a single framework, providing a powerful tool for analysing the potential effectiveness of management interventions.

Habitat heterogeneity and pollinators: Bees in fragmented landscapes (Hymenoptera : Apidae)

This text presents the main studies that report the effect of habitat degradation upon plant-pollinator interactions and the importance of the presence of native vegetation remnants to pollinators, which leads to the need for conservation measures.

Patch size, functional isolation, visibility and matrix permeability influences neotropical primate occurrence within highly fragmented landscapes

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Unraveling the drivers of community dissimilarity and species extinction in fragmented landscapes

Communities in fragmented landscapes are often assumed to be structured by species extinction due to habitat loss, which has led to extensive use of the species-area relationship (SAR) in fragmentation studies. However, the use of the SAR presupposes that habitat loss leads species to extinction but does not allow for extinction to be offset by colonization of disturbed-habitat specialists. Moreover, the use of SAR assumes that species richness is a good proxy of community changes in fragmented landscapes. Here, we assessed how communities dwelling in fragmented landscapes are influenced by habitat loss at multiple scales; then we estimated the ability of models ruled by SAR and by species turnover in successfully predicting changes in community composition, and asked whether species richness is indeed an informative community metric. To address these issues, we used a data set consisting of 140 bird species sampled in 65 patches, from six landscapes with different proportions of forest cover in the Atlantic Forest of Brazil. We compared empirical patterns against simulations of over 8 million communities structured by different magnitudes of the power-law SAR and with species-specific rules to assign species to sites. Empirical results showed that, while bird community composition was strongly influenced by habitat loss at the patch and landscape scale, species richness remained largely unaffected. Modeling results revealed that the compositional changes observed in the Atlantic Forest bird metacommunity were only matched by models with either unrealistic magnitudes of the SAR or by models ruled by species turnover, akin to what would be observed along natural gradients. We show that, in the presence of such compositional turnover, species richness is poorly correlated with species extinction, and z values of the SAR strongly underestimate the effects of habitat loss. We suggest that the observed compositional changes are driven by each species reaching its individual extinction threshold: either a threshold of forest cover for species that disappear with habitat loss, or of matrix cover for species that benefit from habitat loss.

Higher temporal variability of forest breeding bird communities in fragmented landscapes

Understanding the relationship between animal community dynamics and landscape structure has become a priority for biodiversity conservation. In particular, predicting the effects of habitat destruction that confine species to networks of small patches is an important prerequisite to conservation plan development. Theoretical models that predict the occurrence of species in fragmented landscapes, and relationships between stability and diversity do exist. However, reliable empirical investigations of the dynamics of biodiversity have been prevented by differences in species detection probabilities among landscapes. Using long-term data sampled at a large spatial scale in conjunction with a capture-recapture approach, we developed estimates of parameters of community changes over a 22-year period for forest breeding birds in selected areas of the eastern United States. We show that forest fragmentation was associated not only with a reduced number of forest bird species, but also with increased temporal variability in the number of species. This higher temporal variability was associated with higher local extinction and turnover rates. These results have major conservation implications. Moreover, the approach used provides a practical tool for the study of the dynamics of biodiversity.

Landscape connectivity: a key to effective habitat restoration in lowland agricultural landscapes

Landscape scale habitat restoration has the potential to reconnect habitats in fragmented landscapes. This study investigates landscape connectivity as a key to effective habitat restoration in lowland agricultural landscapes and applies these findings to transferable management recommendations. The study area is the Stonehenge World Heritage Site, UK, where landscape scale chalk grassland restoration has been implemented. Here, the ecological benefits of landscape restoration and the species, habitat and landscape characteristics that facilitate or impede the enhancement of biodiversity and landscape connectivity were investigated. Lepidoptera were used as indictors of restoration success and results showed restoration grasslands approaching the ecological conditions of the target chalk grassland habitat and increasing in biodiversity values within a decade. Restoration success is apparent for four species with a broad range of grass larval host plants (e.g. Melanargia galathea, Maniola jurtina) or with intermediate mobility (Polyommatus icarus). However, two species with specialist larval host plants and low mobility (Lysandra bellargus), are restricted to chalk grassland fragments. Studies of restoration grassland of different ages show that recent grassland restoration (1 or 2 years old) may reduce the functional isolation of chalk grassland fragments. A management experiment showed that mowing increases boundary following behaviour in two species of grassland Lepidoptera; Maniola jurtina and Zygaena filipendulae. Analysis of the landscape scale implications of the grassland restoration illustrates an increase in grassland habitat network size and in landscape connectivity, which is likely to benefit the majority of grassland associated Lepidoptera. Landscape and habitat variables can be managed to increase the success of restoration projects including the spatial targeting of receptor sites, vegetation structure and selection of seed source and management recommendations are provided that are transferrable to other species-rich grassland landscape scale restoration projects. Overall results show restoration success for some habitats and species within a decade. However, additional management is required to assist the re-colonisation of specialist species. Despite this, habitat restoration at the landscape scale can be an effective, long term approach to enhance butterfly biodiversity and landscape connectivity.

The compositional and configurational heterogeneity of matrix habitats shape woodland carabid communities in wooded-agricultural landscapes

Landscape heterogeneity (the composition and configuration of matrix habitats) plays a major role in shaping species communities in wooded-agricultural landscapes. However, few studies consider the influence of different types of semi-natural and linear habitats in the matrix, despite their known ecological value for biodiversity. Objective To investigate the importance of the composition and configuration of matrix habitats for woodland carabid communities and identify whether specific landscape features can help to maintain long-term populations in wooded-agricultural environments. Methods Carabids were sampled from woodlands in 36 tetrads of 4 km2 across southern Britain. Landscape heterogeneity including an innovative representation of linear habitats was quantified for each tetrad. Carabid community response was analysed using ordination methods combined with variation partitioning and additional response trait analyses. Results Woodland carabid community response was trait-specific and better explained by simultaneously considering the composition and configuration of matrix habitats. Semi-natural and linear features provided significant refuge habitat and functional connectivity. Mature hedgerows were essential for slow-dispersing carabids in fragmented landscapes. Species commonly associated with heathland were correlated with inland water and woodland patches despite widespread heathland conversion to agricultural land, suggesting that species may persist for some decades when elements representative of the original habitat are retained following landscape modification. Conclusions Semi-natural and linear habitats have high biodiversity value. Landowners should identify features that can provide additional resources or functional connectivity for species relative to other habitat types in the landscape matrix. Agri-environment options should consider landscape heterogeneity to identify the most efficacious changes for biodiversity.

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 (similar to 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.

Importance of estimating matrix quality for modeling species distribution in complex tropical landscapes: a test with Atlantic forest small mammals

Information to guide decision making is especially urgent in human dominated landscapes in the tropics, where urban and agricultural frontiers are still expanding in an unplanned manner. Nevertheless, most studies that have investigated the influence of landscape structure on species distribution have not considered the heterogeneity of altered habitats of the matrix, which is usually high in human dominated landscapes. Using the distribution of small mammals in forest remnants and in the four main altered habitats in an Atlantic forest landscape, we investigated 1) how explanatory power of models describing species distribution in forest remnants varies between landscape structure variables that do or do not incorporate matrix quality and 2) the importance of spatial scale for analyzing the influence of landscape structure. We used standardized sampling in remnants and altered habitats to generate two indices of habitat quality, corresponding to the abundance and to the occurrence of small mammals. For each remnant, we calculated habitat quantity and connectivity in different spatial scales, considering or not the quality of surrounding habitats. The incorporation of matrix quality increased model explanatory power across all spatial scales for half the species that occurred in the matrix, but only when taking into account the distance between habitat patches (connectivity). These connectivity models were also less affected by spatial scale than habitat quantity models. The few consistent responses to the variation in spatial scales indicate that despite their small size, small mammals perceive landscape features at large spatial scales. Matrix quality index corresponding to species occurrence presented a better or similar performance compared to that of species abundance. Results indicate the importance of the matrix for the dynamics of fragmented landscapes and suggest that relatively simple indices can improve our understanding of species distribution, and could be applied in modeling, monitoring and managing complex tropical landscapes.

Frugivory, Post-feeding Flights of Frugivorous Birds and the Movement of Seeds in a Brazilian Fragmented Landscape

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Diversity of leaf-litter anurans in a fragmented landscape of the Atlantic Plateau of Sao Paulo State, southeastern Brazil

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Land-use and land-cover change in Atlantic Forest landscapes

The Atlantic Forest is one of the most threatened tropical biomes, with much of the standing forest in small (less than 50 ha), disturbed and isolated patches. The pattern of land-use and land-cover change (LULCC) which has resulted in this critical scenario has not yet been fully investigated. Here, we describe the LULCC in three Atlantic Forest fragmented landscapes (Sao Paulo, Brazil) between 1960-1980s and 1980-2000s. The three studied landscapes differ in the current proportion of forest cover, having 10%, 30% and 50% respectively. Between the 1960s and 1980s. forest cover of two landscapes was reduced while the forest cover in the third landscape increased slightly. The opposite trend was observed between the 1980s and 2000s: forest regeneration was greater than deforestation at the landscapes with 10% and 50% of forest cover and, as a consequence, forest cover increased. By contrast, the percentage of forest cover at the landscape with 30% of forest cover was drastically reduced between the 1980s and 2000s. LULCC deviated from a random trajectory, were not constant through time in two study landscapes and were not constant across space in a given time period. This landscape dynamism in single locations over small temporal scales is a key factor to be considered in models of LULCC to accurately simulate future changes for the Atlantic Forest. In general, forest patches became more isolated when deforestation was greater than forest regeneration and became more connected when forest regeneration was greater than deforestation. As a result of the dynamic experienced by the study landscapes, individual forest patches currently consist of a mosaic of different forest age classes which is likely to impact bio-diversity. Furthermore, landscape dynamics suggests the beginning of a forest transition in some Atlantic Forest regions, what could be of great importance for biodiversity conservation due to the potential effects of young secondary forests in reducing forest isolation and maintaining a significant amount of the original biodiversity. (C) 2012 Elsevier B.V. All rights reserved.

A landscape-scale test of the predictive ability of a spatially explicit model for population viability analysis

1. Although population viability analysis (PVA) is widely employed, forecasts from PVA models are rarely tested. This study in a fragmented forest in southern Australia contrasted field data on patch occupancy and abundance for the arboreal marsupial greater glider Petauroides volans with predictions from a generic spatially explicit PVA model. This work represents one of the first landscape-scale tests of its type. 2. Initially we contrasted field data from a set of eucalypt forest patches totalling 437 ha with a naive null model in which forecasts of patch occupancy were made, assuming no fragmentation effects and based simply on remnant area and measured densities derived from nearby unfragmented forest. The naive null model predicted an average total of approximately 170 greater gliders, considerably greater than the true count (n = 81). 3. Congruence was examined between field data and predictions from PVA under several metapopulation modelling scenarios. The metapopulation models performed better than the naive null model. Logistic regression showed highly significant positive relationships between predicted and actual patch occupancy for the four scenarios (P = 0.001-0.006). When the model-derived probability of patch occupancy was high (0.50-0.75, 0.75-1.00), there was greater congruence between actual patch occupancy and the predicted probability of occupancy. 4. For many patches, probability distribution functions indicated that model predictions for animal abundance in a given patch were not outside those expected by chance. However, for some patches the model either substantially over-predicted or under-predicted actual abundance. Some important processes, such as inter-patch dispersal, that influence the distribution and abundance of the greater glider may not have been adequately modelled. 5. Additional landscape-scale tests of PVA models, on a wider range of species, are required to assess further predictions made using these tools. This will help determine those taxa for which predictions are and are not accurate and give insights for improving models for applied conservation management.