Effects of structural and functional connectivity and patch size on the abundance of seven Atlantic Forest bird species

We studied the importance of fragment size and structural and functional connectivity on the occurrence and abundance of seven Atlantic Forest bird species in 13 patches (13-275 ha) and three sites within a continuous forest (10,000 ha). We sampled birds with point counts and evaluated structural connectivity considering the presence of corridors and the degree of isolation. We defined functional connectivity by analyzing species movements using playbacks in forest corridors between fragments and in the surrounding matrix. Species differed in their responses to fragmentation. For the frugivorous species, Trogon surrucura, Carpornis cucullatus and Triclaria malachitacea, patch size was the main factor determining abundance. Two understory insectivorous species, Basileuterus leucoblepharus and Pyriglena leucoptera, were more affected by the degree of patch connectivity, the former by the presence of corridors and the latter by the distance between patches. The capacity of P. leucoptera to use corridors and open areas (i.e. functional connectivity) shaped its abundance pattern. Fragmentation had no effect on the abundance of Chiroxiphia caudata and had a positive effect on Batara cinerea. This study emphasizes the importance of considering species’ perceptions of landscape, especially functional connectivity, in understanding the effects of habitat fragmentation.     

The role of forest structure, fragment size and corridors in maintaining small mammal abundance and diversity in an Atlantic forest landscape

Using the abundance and distribution of small mammals at 26 sites in an Atlantic forest landscape, we investigated how species abundance and alpha and beta diversity are affected by fragment size and the presence of corridors. To account for the variability in forest structure among fragments, we described and minimized the influence of foliage density and stratification on small mammal data. Sites were distributed among three categories of fragment size and in continuous forest. For small and medium-sized categories, we considered isolated fragments and fragments connected by corridors to larger remnants. Small mammal abundance and alpha and beta diversity were regressed against site scores from the first axis of a Principal Component Analysis on forest structure variables. Residuals were used in analyses of variance to compare fragment size and connectivity categories. Forest structure influenced total abundance and abundance of some species individually, but not the diversity of small mammal communities. Total abundance and alpha diversity were lower in small and medium-sized fragments than in large fragments and continuous forest, and in isolated compared to connected fragments. Three species were less common, but none was more abundant in smaller fragments. At least one species was more abundant in connected compared to isolated fragments. Beta diversity showed an opposite relationship to fragment size and corridors, increasing in small and isolated fragments. Results highlight the importance of secondary forest for the conservation of tropical fauna, the hyper-dynamism of small isolated fragments and the potential of corridors to buffer habitat fragmentation effects in tropical landscapes.     

Mixed-species bird flocks from Brazilian Atlantic forest: the effects of forest fragmentation and seasonality on their size, richness and stability

The effects of forest fragmentation on species richness, size and stability of bird mixed-species flocks were studied during rainy and dry seasons at nine Atlantic Forest fragments in Minas Gerais state, southeastern Brazil. Two distinct kinds of mixed-species flocks were identified at the study area: heterogeneous flock and understory flocks. The first type of association was observed in all forest fragments and was composed mainly of canopy and midstory species, but may include a few understory species. Understory flocks were composed mainly of understory species and were detected in only three forest fragments. Forest fragment area and season significantly affected species richness, size and stability of heterogeneous flocks. Moreover, species richness in heterogeneous flocks was correlated with total species pool in forest fragments. Species richness and size of understory flocks were different among the three forest fragments, but season influenced only flock size. Understory flock stability differed neither among forest fragments nor between seasons. The red-crowned ant-tanager Habia rubica was the nuclear and leader species of understory flocks, and determined the occurrence of this association in forest fragments. Our results show that forest fragmentation can affect species’ interactions in foraging associations, and that conservation plans for the region should preserve nuclear species of mixed-species flocks and their preferred habitats.     

Time-lag in biological responses to landscape changes in a highly dynamic Atlantic forest region

Time-lagged responses of biological variables to landscape modifications are widely recognized, but rarely considered in ecological studies. In order to test for the existence of time-lags in the response of trees, small mammals, birds and frogs to changes in fragment area and connectivity, we studied a fragmented and highly dynamic landscape in the Atlantic forest region. We also investigated the biological correlates associated with differential responses among taxonomic groups. Species richness and abundance for four taxonomic groups were measured in 21 secondary forest fragments during the same period (2000-2002), following a standardized protocol. Data analyses were based on power regressions and model selection procedures. The model inputs included present (2000) and past (1962, 1981) fragment areas and connectivity, as well as observed changes in these parameters. Although past landscape structure was particularly relevant for trees, all taxonomic groups (except small mammals) were affected by landscape dynamics, exhibiting a time-lagged response. Furthermore, fragment area was more important for species groups with lower dispersal capacity, while species with higher dispersal ability had stronger responses to connectivity measures. Although these secondary forest fragments still maintain a large fraction of their original biodiversity, the delay in biological response combined with high rates of deforestation and fast forest regeneration imply in a reduction in the average age of the forest. This also indicates that future species losses are likely, especially those that are more strictly-forest dwellers. Conservation actions should be implemented to reduce species extinction, to maintain old-growth forests and to favour the regeneration process. Our results demonstrate that landscape history can strongly affect the present distribution pattern of species in fragmented landscapes, and should be considered in conservation planning.     

Plantation rows as dispersal routes: A test with didelphid marsupials in the Atlantic Forest, Brazil

Anecdotal observations suggest that some vertebrates follow plantation rows when crossing between habitat patches in fragmented landscapes, but the frequency of such behavior was never formally tested despite its potential implications for landscape management. We experimentally tested if the didelphid marsupials Didelphis aurita and Philander frenatus use plantation rows as guidelines when searching for habitat patches in a landscape of fragmented Atlantic Forest in Brazil. Thirty-seven individuals of P. frenatus and 24 of D. aurita were captured in forest fragments and released carrying spool-and-line tracking devices at four distances (30-200 m) from a forest fragment, in a matrix of manioc plantations. Animals of both species used plantation rows to orient their initial movements and used rows as guidelines along their paths for navigating in the matrix. By moving predominantly parallel to the plantation rows animals reduced the tortuosity of their paths and obtained an unobstructed view of the landscape ahead. These results demonstrate that the orientation of linear plantations can strongly influence functional connectivity between habitat fragments. Plantation rows should be oriented to maximize connection between nearby fragments, enhancing dispersal success of these species and probably many other terrestrial vertebrates in agricultural landscapes.     

Structure of the advanced regeneration community in tropical rain forest fragments of Los Tuxtlas, Mexico

We characterized the tree seedling community within seven forest fragments of different size (3-640 ha) at Los Tuxtlas, Mexico. We considered individual density, species richness, diversity, composition and functional group (shade tolerant, non-pioneer-light demander and pioneer species). We recorded 8038 individuals, belonging to 128 species and 40 morphospecies. Overall, shade-tolerant species had the highest density and diversity followed by non-pioneer-light demanders and pioneer species. Mean individual density per plot was not related to fragment size, but mean species richness per plot, total observed species richness and diversity were inversely related to fragment size, with the smallest fragments having the highest values. Species composition comparisons highlighted low similarity among fragments. Furthermore, larger fragments shared more species with the smallest fragments than between each other. Our results show that the largest fragments do not always hold the highest species richness; that small fragments comprise of high seedling species diversity, and represent a high regeneration potential for tree native species in the study landscape.     

Tree species composition and similarity in semideciduous forest fragments of southeastern Brazil

This study investigates the influence of fragment size on tree species composition, species richness and on individual groups of species within 11 semideciduous forest fragments in southeastern Brazil. We compared same-sized samples of 500 trees from 10 fragments <100 ha, allocated to three size categories, with three 500-tree samples collected in different areas within a large forest, used as a reference forest. The variation in species richness, in the proportion of species/individuals in dispersal modes, and in the proportion of rare species was not related to fragment size. Nonetheless, comparisons between the mean values of these variables of each fragment size categories, using 95% confidence intervals, showed a wider range of variation within the category of small-sized fragments than in the other size categories. NMS ordination plot also suggested the absence of a relationship between fragment area and tree species composition. However, multiple regressions using the scores from the NMS ordination as response variables, and area and disturbance intensities as predictor variables, suggested that the interaction of area and disturbance might be a good predictor of species composition. Pre-existing environmental heterogeneity and geographical proximity also appeared to play a role in the variations of forest composition among the fragments studied. Our results demonstrated the high conservation value of small forest fragments.     

Patch size and isolation effects on epiphytic and epiphyllous bryophytes in the fragmented Brazilian Atlantic forest

An inventory of the understory forest epiphyte and epiphyllous bryophyte floras was carried out in eight Atlantic rain forest fragments. The fragments were between 7 and 500 ha in size and belonged to two areas (lowland and submontane) of the state of Pernambuco, northeastern Brazil. The aim of this study was to investigate the effect of habitat fragmentation and changes in the natural landscape on community structure (composition, richness, diversity, and abundance). Although the influence of altitude was noted in this study by increasing richness, diversity, and abundance, it was clear that in some fragments the influence of fragment size and isolation can be more important than this positive environmental factor. Fragment size and isolation affect both communities but the last one seems to be a stronger threatening factor for the epiphylls. The least isolated and the largest fragments housed the richest floras - especially in relation to the epiphylls - and had the greatest proportion of shade species. Habitat fragmentation negatively affected epiphytic and epiphyllous bryophytes and increased the representation of species with larger niches (generalists) while decreasing the representation of species with smaller niches (typically found in shady or in sunny areas). The results suggests that the critical fragment size necessary for bryophyte preservation must be correlated with insularity levels; for epiphytes, however, it is likely that low levels of isolation cannot compensate borderline effects as with the epiphyllous bryophytes.     

The influence of patch area and connectivity on avian communities in urban revegetation

Landscape restoration through revegetation is being increasingly used in the conservation management of degraded landscapes. To effectively plan restoration programs information is required on how the landscape context of revegetation influences biodiversity gains. Here, we investigate the relative influence of patch area and connectivity on bird species richness and abundance within urban revegetation patches in Brisbane, Australia. We carried out bird surveys at 20 revegetation sites, and used hierarchical partitioning and model selection to test the relative importance of patch area (the area of revegetation including all directly connected remnant vegetation) and landscape connectivity (the vegetated area connected by less than 10 m, 20 m, 30 m, 40 m and 50 m cleared gaps). We controlled for a number of possible confounding variables within the hierarchical partitioning procedure. Both the hierarchical partitioning and model selection procedures indicated that connectivity had an important influence on bird species richness. Patch area in combination with connectivity were important influencing factors on overall bird abundance. We also carried out the hierarchical partitioning procedure for bird abundance data within a range of feeding guilds, yielding results specific to species groups. Overall our data suggest that greater connectivity enhances the habitat area that colonists can arrive from (resulting in greater species richness), whereas increased patch area allows for increased abundance by expanding the habitat available to species already present in a patch. A combined approach where connectivity and overall habitat area is enhanced across the landscape is likely to be necessary to meet long-term conservation objectives.     

Low impact of present and historical landscape configuration on the genetics of fragmented Anthyllis vulneraria populations

Decreasing habitat fragment area and increasing isolation may cause loss of plant population genetic diversity and increased genetic differentiation between populations. We studied the relation between the historical and the present landscape configuration (i.e., patch area and patch connectivity), and the present management of calcareous grassland fragments on the one hand, and the within and between population genetic structure of 18 Anthyllis vulneraria populations on the other hand. Despite the long-time fragmentation history and the mainly selfing breeding system of the species, we detected very low genetic differentiation (Φ_st = 0.056) among habitat fragments and no significant isolation-by-distance relation. Average within fragment genetic diversity measured as molecular variance and expected heterozygosity, were relatively high (16.46 and 0.28, respectively), and weakly positively correlated with the current fragment area, most likely because larger fragments contained larger populations. We found no effects of the historical landscape configuration on the genetic diversity of the populations. Our data suggest that the consequences of habitat fragmentation for genetic differentiation and genetic diversity of A. vulneraria are relatively minor which is very likely due to the historical high levels of seed exchange among fragments through grazing and roaming livestock. This study provides indirect evidence that nature management by grazing not only positively affects habitat quality but that it might also mitigate the genetic consequences of habitat fragmentation. From the conservation point of view, this study illustrates the importance of grazing and of the regular transport of livestock between fragments to prevent the long-term effects of fragmentation on the genetic diversity of the populations studied.     

Comparing bird community responses to forest fragmentation in two lowland Central American reserves

Habitat loss and fragmentation reduce diversity of tropical bird communities, but the predictability of how communities in fragments disintegrate over time remains unclear. We compared bird community changes of two lowland forest reserves, La Selva Biological Station (LSBS), Costa Rica and Barro Colorado Island (BCI), Panama, both approximately the same size (1500 ha) and at similar latitude (9-10 N) in Central America. Both reserves are losing bird species despite LSBS’s connection to an existing large park (incomplete isolation) and BCI’s favorable location within a largely forested landscape. We examined characteristics of guilds and species lost from the sites to determine whether patterns were similar, and thus predictable. Some of the same guilds declined at both reserves, particularly insectivores and ground/understory nesters. At LSBS mixed-species flock participants, forest species, and burrow-nesters also declined or became extirpated disproportionately. At BCI edge species became extirpated. Body mass was a poor predictor of species and guild loss at both sites, except for carnivores at La Selva. Thus, fragmentation appears to influence some guilds more than others, but which species decline or disappear in tropical forest fragments is also influenced by site-specific factors, mostly yet to be determined. We need to understand such idiosyncratic effects of fragmentation better, rather than rely on one-size-fits-all management plans to conserve bird communities in tropical forest fragments.     

Landscape patterns influencing bird assemblages in a fragmented neotropical cloud forest

The cloud forest is one of the rarest and most threatened ecosystems in Mexico, although it contributes highly to the country's biological diversity and provides important ecological services. It is a naturally fragmented ecosystem, but anthropogenic deforestation and fragmentation has been severe. Consequently, it is essential to identify landscape patterns critical for the conservation of cloud forest. In order to understand how landscape patterns affect diversity in this ecosystem, this study explores the consequences of cloud forest fragmentation on bird diversity in eastern Mexico. I analysed the response of bird species richness and abundance as a function of forest fragment size, shape, topographical complexity, altitudinal range, connectivity, and proportion of landscape forested area in a system of 13 cloud forest fragments. Fragment shape was the main characteristic positively related to species richness in the bird community, but a differential response to landscape patterns was also detected. Fragment size was the main characteristic influencing the segment of the bird community depending mostly on forest, that is to say, forest interior and generalist species. In contrast, the extent of forest edge, expressed as fragment shape, produced a positive response of forest border species. Both, forest dependent and border dependent species positively responded to the extent of their suitable habitat. The immediate and most effective ecologically oriented conservation strategy for the region is the conservation of the largest cloud forest fragments.     

No evidence of a plant extinction debt in highly fragmented calcareous grasslands in Belgium

Long transient times in response to decreasing habitat area and increasing isolation may cause the present plant species distribution to reflect the historical rather than the present landscape configuration, resulting in a so-called extinction debt. To investigate how plant species richness is shaped by both present and historical landscape configuration and local environmental conditions in fragmented calcareous grasslands, 64 sites in Southern Belgium were surveyed for their total species richness and environmental characteristics. Species were subdivided in specialist and generalist groups and the former were clustered into emergent groups (EGs) according to 16 relevant life-history traits. Four specialist emergent groups were derived: orchids, rosette species, annuals and half-rosette species. Both specialist and generalist species richness increased with present fragment area, while present fragment connectivity only affected the number of specialist species. This trend was maintained for most EGs, although habitat area was mostly more important than connectivity. All species groups responded to an insolation gradient, except EGs of rosette and annual species. Differential response to fragmentation among emergent groups could be interpreted in terms of dispersal and persistence traits. No relation between species richness and historical area or connectivity was found, making an extinction debt unlikely. Hence, present-day variation in plant species diversity seems to be no legacy of past landscape configuration. Our results clearly indicate that plant species may be quickly lost in response to new fragmentation events. To restore plant diversity, management should focus both on mitigating landscape fragmentation and restoring habitat quality.     

Drastic erosion in functional attributes of tree assemblages in Atlantic forest fragments of northeastern Brazil

The long-term dynamics of plant communities remain poorly understood in isolated tropical forest fragments. Here we test the hypothesis that tropical tree assemblages in both small forest fragments and along forest edges of very large fragments are functionally much more similar to stands of secondary growth (5-65-yr old) than to core primary forest patches. The study was carried out in a severely fragmented landscape of the Brazilian Atlantic forest. Nine functional attributes of tree assemblages were quantified by sampling all trees (DBH ? 10 cm) within 75 plots of 0.1 ha distributed in four forest habitats: small forest fragments (3.4-79.6 ha), forest edges, second-growth patches, and primary forest interior areas within a large forest fragment (3500 ha). These habitats were markedly different in terms of tree species richness, and in the proportion of pioneer, large-seeded, and emergent species. Age of second-growth stands explained between 31.4% and 88.2% of the variation in the functional attributes of tree assemblages in this habitat. As expected, most traits associated with forest edges and small forest fragments fell within the range shown by early (<25-yr old) and intermediate-aged secondary forest stands (25-45-yr old). In contrast to habitat type, tree assemblage attributes were not affected by vegetation type, soil type and the spatial location of plots. An ordination analysis documented a striking floristic drift in edge-affected habitats. Our results suggest that conservation policy guidelines will fail to protect aging, hyper-fragmented landscapes from drastic impoverishment if the remaining forest patches are heavily dominated by edge habitat.     

Long-term effects of fragmentation and fragment properties on bird species richness in Hawaiian forests

Forest fragmentation is a common disturbance affecting biological diversity, yet the impacts of fragmentation on many forest processes remain poorly understood. Forest restoration is likely to be more successful when it proceeds with an understanding of how native and exotic vertebrates utilize forest patches of different size. We used a system of forest fragments isolated by volcanic activity 153 years ago in Hawaii to examine how long-term fragmentation, as well as fragment size and structural features affect the richness of native and exotic bird species. The total number of bird species increased rapidly with forest fragment size, with most of the native species pool found in patches <3 ha. Smaller fragments were dominated by native bird species with several exotic bird species found only in the largest fragments, suggesting that exotic bird species in this landscape show greater area-sensitivity than native species. We used airborne scanning light detection and ranging (LiDAR) to assess whether fragment area was correlated with estimates of fragment vegetation volume as well as measures of tree height. Fragment area was highly correlated with vegetation volume, maximum tree height, and canopy height heterogeneity, and these variables were strong predictors of bird richness, demonstrating that remote sensing can provide key insights into the relationship between fragment structural attributes and biodiversity indicators. Overall, this work demonstrates the value of conserving small remnant mid-elevation forest patches for native birds in Hawaii. This work also provides insight into how newly created forest patches might be used by native and exotic bird species in Hawaii.     

Movements of neotropical understory passerines affected by anthropogenic forest edges in the Brazilian Atlantic rainforest

Edge effects are suggested to have great impact on the persistence of species in fragmented landscapes. We tested edge avoidance by forest understory passerines in the Brazilian Atlantic Rainforest and also compared their mobility and movement patterns in contiguous and fragmented landscapes to assess whether movements would increase in the fragmented landscape. Between 2003 and 2005, 96 Chiroxiphia caudata, 38 Pyriglena leucoptera and 27 Sclerurus scansor were radio-tracked. The most strictly forest species C. caudata and S. scansor avoided forest edges, while P. leucoptera showed affinities for the edge. Both sensitive species showed larger mean step length and maximal observed daily distance in the fragmented forest versus the unfragmented forest. P. leucoptera did not show any significant difference. There were no significant differences in proportional daily home range use for any of the three species. Our results suggested that fragmentation and the consequent increase in edge areas do influence movement behavior of sensitive forest understory birds that avoided the use of edges and increased the speed and distance they covered daily. For the most restricted forest species, it would be advisable to protect larger patches of forest instead of many small or medium fragments connected by narrow corridors. However, by comparing our data with that obtained earlier, we concluded that movement behavior of resident birds differs from that of dispersing birds and might not allow to infer functional connectivity or landscape-scale sensitivity to fragmentation; a fact that should be taken into consideration when suggesting conservation strategies.     

Habitat fragmentation reduces genetic diversity and connectivity among toad populations in the Brazilian Atlantic Coastal Forest

Tropical rainforests are becoming increasingly fragmented and understanding the genetic consequences of fragmentation is crucial for conservation of their flora and fauna. We examined populations of the toad Rhinella ornata, a species endemic to Atlantic Coastal Forest in Brazil, and compared genetic diversity among small and medium forest fragments that were either isolated or connected to large forest areas by corridors. Genetic differentiation, as measured by F_ST, was not related to geographic distance among study sites and the size of the fragments did not significantly alter patterns of genetic connectivity. However, population genetic diversity was positively related to fragment size, thus haplotype diversity was lowest in the smallest fragments, likely due to decreases in population sizes. Spatial analyses of genetic discontinuities among groups of populations showed a higher proportion of barriers to gene flow among small and medium fragments than between populations in continuous forest. Our results underscore that even species with relatively high dispersal capacities may, over time, suffer the negative genetic effects of fragmentation, possibly leading to reduced fitness of population and cases of localized extinction.     

Long-term erosion of tree reproductive trait diversity in edge-dominated Atlantic forest fragments

Habitat loss and fragmentation promote relatively predicable shifts in the functional signature of tropical forest tree assemblages, but the full extent of cascading effects to biodiversity persistence remains poorly understood. Here we test the hypotheses that habitat fragmentation (a) alters the relative contribution of tree species exhibiting different reproductive traits; (b) reduces the diversity of pollination systems; and (c) facilitates the functional convergence of reproductive traits between edge-affected and early-secondary forest habitats (5-32 years old). This study was carried out in a severely fragmented 670-km² forest landscape of the Atlantic forest of northeastern Brazil. We assigned 35 categories of reproductive traits to 3552 trees (DBH ? 10 cm) belonging to 179 species, which described their pollination system, floral biology, and sexual system. Trait abundance was calculated for 55 plots of 0.1 ha across four habitats: forest edges, small forest fragments (3.4-83.6 ha), second-growth patches, and core tracts of forest interior within the largest available primary forest fragment (3500 ha) in the region. Edge-affected and secondary habitats showed a species-poor assemblage of trees exhibiting particular pollination systems, a reduced diversity of pollination systems, a higher abundance of reproductive traits associated with pollination by generalist diurnal vectors, and an elevated abundance of hermaphroditic trees. As expected, the reproductive signature of tree assemblages in forest edges and small fragments (edge-affected habitats), which was very similar to that of early second-growth patches, was greatly affected by both habitat type and plot distance to the nearest forest edge. In hyper-fragmented Atlantic forest landscapes, we predict that narrow forest corridors and small fragments will become increasingly dominated by edge-affected habitats that can no longer retain the full complement of tree life-history diversity and its attendant mutualists.     

The effects of habitat connectivity on cryptogam richness in boulder metacommunity

We investigated the effects of habitat patch size and connectivity on cryptogam richness on 288 calcareous boulders. The studied boulders were found to be a particularly suitable ‘metacommunity model system’ and allowed comparison of species richness among cryptogams with differing substrate specificity, life history and red list status. The dependence of species richness on boulder surface area was modelled using Arrhenius power model, and contrasts were analysed between connected and isolated boulders. Species richness was clearly related to boulder size, but the rates of increase of species richness with increasing boulder area (z-values) were highly variable among species groups. Highest value of z (1.17) was found for species with large propagules, and values around 0.5 were obtained for calciphilous and regionally red-listed species and z = 0.42 for perennials. The lowest value of z (0.27) occurred in calcicolous species while for all species, colonists, and species with small propagules the values were 0.3-0.4. The richness of all species was 11% higher on connected than isolated boulders, while connected boulders had 31% more of both calcicolous and regionally red-listed species and 19% more of colonists. The result suggests that in the investigated system that boulder size is a crucial determinant of cryptogam richness, and their spatial configuration in terms of connectivity may further contribute to cryptogam richness. The latter result lends support to the importance of metacommunity processes for the maintenance of cryptogam richness.     

Species diversity and habitat fragmentation: frogs in a tropical montane landscape in Mexico

We evaluate the alpha (within patch species richness), beta (spatial turnover among patches) and gamma (landscape) diversity of frogs in a tropical montane cloud forest (TMCF) in central Veracruz, Mexico in order to assess (1) the influence of forest fragmentation on frog assemblages, (2) the importance to diversity of the various elements of the landscape matrix, including the shaded coffee plantations and cattle pastures that surround TMCF and (3) to identify the frog guilds most affected by habitat transformation. We sampled ten sites between May 1998 and November 2000: five TMCF fragments and five anthropogenic habitats. For the entire landscape, we registered 21 species belonging to six families. 100% of these were found in the TMCF fragments and 62% in the surrounding mosaic of anthropogenic habitats. Gamma diversity (γ) is determined to a greater extent by species exchange (β) than by local species richness (α). Elevational variation, the degree of conservation of the vegetation canopy and fragment size appear to determine the species diversity of this landscape. Large species, terrestrial species, those whose eggs develop outside water, and those whose larvae develop in the water seemed to be most affected by habitat transformation. On its own, even the largest and most species-rich cloud forest fragment is not capable of preserving the current anuran diversity. Neither are the shaded coffee plantations that are interspersed among and link the patches of TMCF. However together they form a diverse system of habitats crucial to species conservation in this landscape.