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 Andrén 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.     

Forest Loss and the Biodiversity Threshold: An Evaluation Considering Species Habitat Requirements and the Use of Matrix Habitats

Habitat loss is the main driver of the current biodiversity crisis, a landscape-scale process that affects the survival of spatially-structured populations. Although it is well-established that species responses to habitat loss can be abrupt, the existence of a biodiversity threshold is still the cause of much controversy in the literature and would require that most species respond similarly to the loss of native vegetation. Here we test the existence of a biodiversity threshold, i.e. an abrupt decline in species richness, with habitat loss. We draw on a spatially-replicated dataset on Atlantic forest small mammals, consisting of 16 sampling sites divided between forests and matrix habitats in each of five 3600-ha landscapes (varying from 5% to 45% forest cover), and on an a priori classification of species into habitat requirement categories (forest specialists, habitat generalists and open-area specialists). Forest specialists declined abruptly below 30% of forest cover, and spillover to the matrix occurred only in more forested landscapes. Generalists responded positively to landscape heterogeneity, peaking at intermediary levels of forest cover. Open area specialists dominated the matrix and did not spillover to forests. As a result of these distinct responses, we observed a biodiversity threshold for the small mammal community below 30% forest cover, and a peak in species richness just above this threshold. Our results highlight that cross habitat spillover may be asymmetrical and contingent on landscape context, occurring mainly from forests to the matrix and only in more forested landscapes. Moreover, they indicate the potential for biodiversity thresholds in human-modified landscapes, and the importance of landscape heterogeneity to biodiversity. Since forest loss affected not only the conservation value of forest patches, but also the potential for biodiversity-mediated services in anthropogenic habitats, our work indicates the importance of proactive measures to avoid human-modified landscapes to cross this threshold.     

Long-term persistence of midsized to large-bodied mammals in Amazonian landscapes under varying contexts of forest cover

Both forest fragmentation and overhunting have profound effects on the structure of large-vertebrate assemblages in neotropical forests. However, the long-term value of habitat fragments for forest mammals remains poorly understood and few regional scale studies have replicated sampling across spatially independent landscapes. Here, we assess the species occupancy and abundance of midsized to large-bodied mammals within three neighbouring Amazonian forest landscapes varying widely in extent of forest cover. One of these consisted of forest fragments surrounded by semi-natural scrub savannahs that had been occupied by paleoindian populations for at least 7,000 years, whereas forest cover in the other two landscapes was either variegated or continuous. Data on species occurrence and abundance from diurnal and nocturnal line-transect surveys and local interviews in each landscape were used to examine the effects of forest cover and hunting pressure on mammal persistence within forest patches. The extent of forest cover was a key determinant of species persistence across the three landscapes, but populations of large-bodied species were either reduced or driven to local extinction by hunting even in the most forested and least fragmented landscape. Many game and non-game species persisted in forest isolates, even though, individually, these were likely too small to support viable populations. This study indicates that even small, long-term forest fragments may retain significant conservation value if they can be managed within the context of enhanced connectivity across wider fragmented landscapes.     

Ferns,frogs, lizards,birds and bats in forest fragments and shade cacao plantations in two contrasting landscapes in the Atlantic forest,Brazil

The traditional shade cacao plantations (cabrucas) of southern Bahia, Brazil, are biologically rich habitats, encompassing many forest-dwelling species. However, a critical question for the conservation management of this specific region, and the highly fragmented Atlantic forest in general, is to what extent the conservation value of cabrucas relies on the presence of primary forest habitat in the landscape. We investigated the relative importance of cabrucas and forests for the conservation of five diverse biological groups (ferns, frogs, lizards, birds and bats) in two contrasting landscapes in southern Bahia, one dominated by forest with some interspersed cabrucas, and one dominated by cabrucas with interspersed forest fragments. The community structure (richness, abundance and diversity) of all biological groups differed between cabrucas and forests, although these differences varied among groups. A high number of forest species was found in the cabrucas. However, there were pronounced differences between the two landscapes with regard to the ability of cabrucas to maintain species richness. Irrespective of the biological group considered, cabrucas located in the landscape with few and small forest fragments supported impoverished assemblages compared to cabrucas located in the landscape with high forest cover. This suggests that a greater extent of native forest in the landscape positively influences the species richness of cabrucas. In the landscape with few small forest fragments interspersed into extensive areas of shade cacao plantations, the beta diversity of birds was higher than in the more forested landscape, suggesting that forest specialist species that rarely ventured into cabrucas were randomly lost from the fragments. These results stress both the importance and the vulnerability of the small forest patches remaining in landscapes dominated by shade plantations. They also point to the need to preserve sufficient areas of primary habitat even in landscapes where land use practices are generally favorable to the conservation of biodiversity.     

Fine-scale movement behaviors of calopterygid damselflies are influenced by landscape structure: an experimental manipulation

We explore the effect of differences in landscape structure, arising from habitat loss, on the fine-scale movement behaviors of two congeneric damselflies – Calopteryx aequabilis and C. maculata . Both species require streams for breeding and naiad development and both often use forest for foraging. We compare movement behaviors across three types of landscape: forested landscapes, where stream and forest habitat are adjacent; partially forested landscapes, where streams and forest habitat are disjunct, and non-forested landscapes, where little to no forest habitat is available. We employ a reciprocal transplant experiment to determine the extent to which movement along and away from streams is influenced by landscape structure and historical behavior or morphological adaptations. For both species, we show that both the propensity to move away from streams and rates of net displacement differ among landscape types. Both species move away from streams on landscapes with high or moderate levels of forest cover but neither moves away from streams on landscapes with little or no forest. Furthermore, C. maculata native to predominantly forested landscapes are more likely to move away from streams, regardless of the landscape structure they encounter, than are individuals native to moderately forested or non-forested landscapes. There was no effect of natal landscape on C. aequabilis . Comparisons with microlandscape studies suggest that there may be some general similarities among the different systems but these are clouded by uncertainty regarding the similarity of the underlying processes responsible for observed behavioral responses to landscape structure. Despite this uncertainty, animal movement behaviors are contingent upon the structure of the broader landscape, regardless of the absolute scale of the landscape.     

Isolation determines patterns of species presence in highly fragmented landscapes

In fragmented landscapes, changes in habitat availability, patch size, shape and isolation may affect survival of local populations. Proposing efficient conservation strategies for such species relies initially on distinguishing the particular effects of those factors. To address these issues, we investigated the occurrence of 3 bird species in fragmented Brazilian Atlantic Forest landscapes. Playback techniques were used to collect presence/absence data of these species inside 80 forest patches, and incidence models were used to infer their occupancy pattern from landscape spatial structure. The relative importance of patch size, shape and surrounding forest cover and isolation was assessed using a model selection approach based on maximum likelihood estimation. The presence of all species was in general positively affected by the amount of surrounding habitat and negatively affected by inter‐patch distances. The joint effects of patch size and the surrounding landscape characteristics were important determinants of occupancy for two species. The third species was affected only by forest cover and mean patch isolation. Our results suggest that local species presence is in general more influenced by the isolation from surrounding forests than by patch size alone. We found evidence that, in highly fragmented landscapes, birds that can not find patches large enough to settle may be able to overcome short distances through the matrix and include several nearby patches within their home‐ranges to complement their resource needs. In these cases, patches must be defined as functionally connected habitat networks rather than mere continuous forest segments. Bird conservation strategies in the Atlantic forest should focus on increasing patch density and connectivity, in order to implement forest networks that reduce the functional isolation between large remnants with remaining core habitat.     

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.     

Designing optimal human‐modified landscapes for forest biodiversity conservation

Agriculture and development transform forest ecosystems to human‐modified landscapes. Decades of research in ecology have generated myriad concepts for the appropriate management of these landscapes. Yet, these concepts are often contradictory and apply at different spatial scales, making the design of biodiversity‐friendly landscapes challenging. Here, we combine concepts with empirical support to design optimal landscape scenarios for forest‐dwelling species. The supported concepts indicate that appropriately sized landscapes should contain ≥ 40% forest cover, although higher percentages are likely needed in the tropics. Forest cover should be configured with c. 10% in a very large forest patch, and the remaining 30% in many evenly dispersed smaller patches and semi‐natural treed elements (e.g. vegetation corridors). Importantly, the patches should be embedded in a high‐quality matrix. The proposed landscape scenarios represent an optimal compromise between delivery of goods and services to humans and preserving most forest wildlife, and can therefore guide forest preservation and restoration strategies.     

Avian species richness in relation to intensive forest management practices in early seral tree plantations

Background

Managers of landscapes dedicated to forest commodity production require information about how practices influence biological diversity. Individual species and communities may be threatened if management practices truncate or simplify forest age classes that are essential for reproduction and survival. For instance, the degradation and loss of complex diverse forest in young age classes have been associated with declines in forest-associated Neotropical migrant bird populations in the Pacific Northwest, USA. These declines may be exacerbated by intensive forest management practices that reduce hardwood and broadleaf shrub cover in order to promote growth of economically valuable tree species in plantations.

Methodology and Principal Findings

We used a Bayesian hierarchical model to evaluate relationships between avian species richness and vegetation variables that reflect stand management intensity (primarily via herbicide application) on 212 tree plantations in the Coast Range, Oregon, USA. Specifically, we estimated the influence of broadleaf hardwood vegetation cover, which is reduced through herbicide applications, on bird species richness and individual species occupancy. Our model accounted for imperfect detection. We used average predictive comparisons to quantify the degree of association between vegetation variables and species richness. Both conifer and hardwood cover were positively associated with total species richness, suggesting that these components of forest stand composition may be important predictors of alpha diversity. Estimates of species richness were 35–80% lower when imperfect detection was ignored (depending on covariate values), a result that has critical implications for previous efforts that have examined relationships between forest composition and species richness.

Conclusion and Significance

Our results revealed that individual and community responses were positively associated with both conifer and hardwood cover. In our system, patterns of bird community assembly appear to be associated with stand management strategies that retain or increase hardwood vegetation while simultaneously regenerating the conifer cover in commercial tree plantations.     

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.     

Influence of within‐patch habitat quality on high‐Andean Polylepis bird abundance

Habitat restoration strategies for fragmented high Andean forest landscapes must consider the influence of within‐patch habitat quality on bird abundance. I examined vegetation and bird abundance at three locations within a highly fragmented Polylepis forest landscape in the Cordillera Vilcanota, southern Peru. Across the landscape, there was significant variation in the vegetation structure of Polylepis forest patches of different size categories, especially in terms of tree girth, tree height, tree density, and canopy vegetation structure. Principal Component Analysis extracted five factors of habitat quality, which together accounted for 74.2% of the variability within 15 habitat variables. Polylepis bird species differed in their responses to habitat quality but, overall, variation in Polylepis bird abundance was not fully captured by the range of habitat quality variables. Tall, dense vegetation cover was clearly important for 11 conservation‐important species, a high density of large trees was important for 10 species and primary forest ground cover was important for eight species. Habitat quality exhibited no significant influence on the abundance of only one species –Asthenes urubambensis. The abundance of seven species was associated with lower elevation forest, but only one species was associated with higher elevation forest. Management of habitat quality in large and medium remnant forest patches throughout the Cordillera Vilcanota, particularly in the 3800–4200 m elevation range, will be a cornerstone in ensuring the persistence of the majority of conservation‐important bird species populations.     

Plant recolonization of reclamation areas from patches of salvaged forest floor material

Question

Understorey development is a great challenge in the restoration of many forest sites, particularly when sources of vegetation propagules are scarce. Can placement of propagule‐rich soil patches within reclaimed landscapes otherwise covered with propagule‐poor material promote the dispersal of vegetation from the patches into the surrounding areas?

Location

Large reclamation site in the Canadian (Alberta) boreal forest.

Method

Patches of propagule‐rich forest floor material were placed within a matrix of propagule‐poor peat material. Vegetation assessments (cover estimates, seed rain) were done surrounding these patches in the third and fourth growing seasons.

Results

There was significant egress of species from the patches into the peat after four growing seasons, and overall species associated with the patches had higher cover in the peat than species that were associated with the peat itself. While wind‐dispersed herbaceous species from the patches were found at the leading edge of the egressing community, most species used vegetative propagation, resulting in short egress distances. Several patch‐associated species were found in seed rain collected on the peat areas but were not observed in this material, suggesting seedbed limitations.

Conclusion

Despite the relatively short distance of egress, this experiment suggests that placement of propagule‐rich soil material within reclaimed landscapes will promote egress into adjacent propagule‐poor soil material.     

Can agroforest woodlots work as stepping stones for birds in the Atlantic forest region?

In fragmented landscapes, agroforest woodlots can potentially act as stepping stones facilitating movement between forest fragments. We assessed the influence of agroforest woodlots on bird distribution and diversity in the Atlantic forest region (SE Brazil), and also tested which categories of species can use different types of connection elements, and whether this use is influenced by the distance to large forest patches. We studied two fragmented landscapes, with and without stepping stones linking large fragments, and one forested landscape. Using a point count, a bird survey was undertaken in the fragmented landscapes in five different elements: large remnants (>400 ha), agroforest woodlots (0.4–1.1 ha), small patches (0.5–7 ha), riparian corridor, and pasture areas (the main matrix). Generalist and open-area species were commonly observed in the agroforest system or other connection elements, whereas only a few forest species were present in these connections. For the latter species, the distance of woodlots to large patches was essential to determine their richness and abundance. Based on our results and data from literature, we suggest that there is an optimal relationship between the permeability of the matrix and the efficiency of stepping stones, which occurs at intermediate degrees of matrix resistance, and is species-dependent. Because the presence of agroforest system favors a higher richness of generalist species, they appeared to be more advantageous for conservation than the monoculture system; for this reason, they should be considered as a management alternative, particularly when the matrix permeability requirement is met.     

Multi-Scale Associations between Vegetation Cover and Woodland Bird Communities across a Large Agricultural Region

Improving biodiversity conservation in fragmented agricultural landscapes has become an important global issue. Vegetation at the patch and landscape-scale is important for species occupancy and diversity, yet few previous studies have explored multi-scale associations between vegetation and community assemblages. Here, we investigated how patch and landscape-scale vegetation cover structure woodland bird communities. We asked: (1) How is the bird community associated with the vegetation structure of woodland patches and the amount of vegetation cover in the surrounding landscape? (2) Do species of conservation concern respond to woodland vegetation structure and surrounding vegetation cover differently to other species in the community? And (3) Can the relationships between the bird community and the woodland vegetation structure and surrounding vegetation cover be explained by the ecological traits of the species comprising the bird community? We studied 103 woodland patches (0.5 - 53.8 ha) over two time periods across a large (6,800 km²) agricultural region in southeastern Australia. We found that both patch vegetation and surrounding woody vegetation cover were important for structuring the bird community, and that these relationships were consistent over time. In particular, the occurrence of mistletoe within the patches and high values of woody vegetation cover within 1,000 ha and 10,000 ha were important, especially for bird species of conservation concern. We found that the majority of these species displayed similar, positive responses to patch and landscape vegetation attributes. We also found that these relationships were related to the foraging and nesting traits of the bird community. Our findings suggest that management strategies to increase both remnant vegetation quality and the cover of surrounding woody vegetation in fragmented agricultural landscapes may lead to improved conservation of bird communities.     

Changes in transmission of Baylisascaris procyonis to intermediate hosts as a function of spatial scale

Physical changes in landscapes alter the abundance and distribution of species. Higher-order effects can occur when changes in ecological processes result in altered interspecific interactions and subsequent changes in a species' abundance or persistence. Baylisascaris procyonis , a roundworm parasite of raccoons ( Procyon lotor ), is pathogenic to numerous small vertebrates that serve as intermediate hosts, including white-footed mice ( Peromyscus leucopus ). Raccoons have responded positively to agriculturally induced changes in landscapes, with potential consequences for intermediate hosts of B . procyonis . We examined white-footed mice from a homogeneous, predominantly forested landscape in south-central Indiana and a heterogeneous, predominantly agricultural landscape in northwestern Indiana for presence of larval B . procyonis . We compared prevalence of infection, intensity of infection, and average number of larvae per mouse between the landscapes, and among forest patches within the highly fragmented agricultural landscape. Prevalence, intensity of infection, and average number of larvae were significantly higher in the highly fragmented landscape. Within the agricultural landscape, regression models predicted probability of infection, intensity of infection, and average number of larvae per mouse per patch as functions of forest patch area and isolation. We conclude that positive responses of raccoons to agriculturally induced fragmentation of forests have resulted in increased encounter rates between white-footed mice and infective stages of B . procyonis between and within landscapes, resulting in increased transmission of the parasite to intermediate hosts.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

Modelling the Species Distribution of Flat-Headed Cats (Prionailurus planiceps), an Endangered South-East Asian Small Felid

Background

The flat-headed cat (Prionailurus planiceps) is one of the world''s least known, highly threatened felids with a distribution restricted to tropical lowland rainforests in Peninsular Thailand/Malaysia, Borneo and Sumatra. Throughout its geographic range large-scale anthropogenic transformation processes, including the pollution of fresh-water river systems and landscape fragmentation, raise concerns regarding its conservation status. Despite an increasing number of camera-trapping field surveys for carnivores in South-East Asia during the past two decades, few of these studies recorded the flat-headed cat.

Methodology/Principal Findings

In this study, we designed a predictive species distribution model using the Maximum Entropy (MaxEnt) algorithm to reassess the potential current distribution and conservation status of the flat-headed cat. Eighty-eight independent species occurrence records were gathered from field surveys, literature records, and museum collections. These current and historical records were analysed in relation to bioclimatic variables (WorldClim), altitude (SRTM) and minimum distance to larger water resources (Digital Chart of the World). Distance to water was identified as the key predictor for the occurrence of flat-headed cats (>50% explanation). In addition, we used different land cover maps (GLC2000, GlobCover and SarVision LLC for Borneo), information on protected areas and regional human population density data to extract suitable habitats from the potential distribution predicted by the MaxEnt model. Between 54% and 68% of suitable habitat has already been converted to unsuitable land cover types (e.g. croplands, plantations), and only between 10% and 20% of suitable land cover is categorised as fully protected according to the IUCN criteria. The remaining habitats are highly fragmented and only a few larger forest patches remain.

Conclusion/Significance

Based on our findings, we recommend that future conservation efforts for the flat-headed cat should focus on the identified remaining key localities and be implemented through a continuous dialogue between local stakeholders, conservationists and scientists to ensure its long-term survival. The flat-headed cat can serve as a flagship species for the protection of several other endangered species associated with the threatened tropical lowland forests and surface fresh-water sources in this region.     

Soil millipede diversity in tropical forest patches and its relation to landscape structure in northeastern Puerto Rico

Uncontrolled urban development is threatening the survival of many terrestrial species worldwide, especially those with limited dispersal capabilities. Soil invertebrates account for a high proportion of the global biodiversity but there are few studies on how soil biodiversity varies with different parameters of landscape structure, especially in fragmented tropical habitats. Millipedes are among the most abundant detritivore invertebrates in the soil-litter macrofauna. We examined the relationships between soil millipede diversity and landscape structure in 12 forest patches of approximately 30 years of age, in northeastern Puerto Rico. Spatial characteristics of the landscape were determined from aerial color photographs and were digitized into a GIS package for analysis. Millipede species diversity and composition in these forest patches showed correlations with their surroundings (e.g. amount of forest in the matrix) and with the presence of vegetation corridors that connected to other forest patches, rather than forest patch attributes such as patch area and shape. Millipede species richness correlated negatively with the degree of isolation of forest patch (within 600 m radius), while species evenness correlated positively to the amount of forest within a 50 m buffer. Millipede species composition was related with the presence of vegetation corridors and the distance to the Luquillo Experimental Forest reserve. These findings show that a low degree of patch isolation, forested buffers, and presence of vegetation corridors need to be considered for the conservation and management of forest patches surrounded by urban developments, especially to protect terrestrial invertebrate species that require forested habitats for their dispersal.     

Is the matrix a sea? Habitat specificity in a naturally fragmented landscape

Abstract.  1. Metapopulation and island biogeography theory assume that landscapes consist of habitat patches set in a matrix of non-habitat. If only a small proportion of species conform to the patch–matrix assumptions then metapopulation theory may only describe special cases rather than being of more general ecological importance.
2. As an initial step towards understanding the prevalence of metapopulation dynamics in a naturally fragmented landscape, the distribution of beetle species in three replicates of three habitat types was examined, including rainforest and eucalypt forest (the habitat patches), and buttongrass sedgeland (the matrix), in south-west Tasmania, Australia.
3. Ordination methods indicated that the buttongrass fauna was extremely divergent from the fauna of forested habitats. Permutation tests showed that the abundance of 13 of 17 commonly captured species varied significantly among habitats, with eight species confined to eucalypts or rainforest, and three species found only in buttongrass. Approximately 60% of species were confined to forested habitat implying that metapopulation theory has the potential to be very important in the forest–buttongrass landscape.
4. Although floristically the rainforest and eucalypts were extremely distinct, the beetle faunas from eucalypts and rainforests overlapped substantially. Therefore rainforest patches connected by eucalypt forest represent continuous habitat for most species.
5. Other studies report a wide range of values for the proportion of patch-specific species in fragmented landscapes. Understanding the environmental or historical conditions under which a high proportion of species become patch specialists would help to identify where spatial dynamic theory may be especially applicable, and where habitat loss and fragmentation poses the greatest threat to biodiversity.     

Impact of landscape spatial pattern on liana communities in tropical rainforests at Los Tuxtlas,Mexico

Questions: What are the species composition and species and stem densities of liana communities in tropical landscapes of different deforestation levels? Which spatial attributes (forest cover, patch area, shape and isolation) have the strongest influence on liana communities in these landscapes? Location: Forty‐five rainforest patches in Los Tuxtlas Biosphere Reserve, Mexico. Methods: In three landscapes with different deforestation levels (HDL=4%; IDL=11%; and LDL=24% of remaining forest cover) liana communities (DBH ≥2.5 cm) were characterized in 15 randomly selected patches per landscape (10 50 m × 2 m transects per patch=0.1 ha), and evaluated the effects of patch area, shape and isolation on liana species and stem density (number of species and stems per 0.1 ha). Results: A total of 64 taxa and 24 families were sampled. Species composition differed highly among landscapes, with HDL being the most dissimilar landscape. The response of lianas to landscape spatial pattern differed significantly among landscapes. Proximity to villages had a strong positive effect on species and stem densities in LDL and IDL. There was a sharp decrease in liana stem density in HDL, with four patches (27%) found to be unoccupied by lianas. Conclusions: Fragmentation may have a positive effect on lianas, partly because of edge effects. This positive effect seems to be limited by the proportion of remaining forest cover in the landscape, as the liana communities had collapsed in the most deforested landscape.