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.     

Bird conservation in fragmented Mediterranean forests of Spain: effects of geographical location, habitat and landscape degradation

The effects of habitat fragmentation on forest bird assemblages were analysed in 214 holm oak (Quercus ilex) remnants spread across the northern and southern plateaux of central Spain. Bird richness was highly dependent on fragment area for all species regardless of isolation, and barely affected by habitat traits. Geographical location was associated with high differences in richness of bird assemblages, which included 17 species exclusive to northern remnants and one exclusive to southern remnants. This supports the hypothesis that habitat suitability deteriorates sharply from north to south for forest birds in Spain. The species-area relationships of bird assemblages sampled in fragmented forests along a broad continental gradient (from Norway to southern Spain) showed that true forest birds only nest in woodlands >100 ha in southern Spain, whereas the full complement of forest species occurs in much smaller fragments in central-western Europe. Loss of species that are particularly sensitive to habitat fragmentation accounts for these differences between dry Spanish and mesic European woodlands. These results are explained by the low habitat suitability of Spanish woodlands, associated with the restrictive conditions for plant regeneration in the Mediterranean climate and long-standing human usage. There is, therefore, a particular need to develop management strategies that conserve birds, and probably other forest organisms, in Mediterranean regions by preventing habitat deterioration and decreases in fragment size, and by conserving all woods >100 ha.     

Effects of age and intensity of urbanization on farmland bird communities

Urban sprawl is now occurring worldwide and considered as a major large-scale perturbation on ecosystems. Consequently, urban territory is replacing other habitats such as agricultural areas. As farmland biotic communities are already reported to be declining, it seems necessary to assess the urbanization impact on them. We conducted a bird survey on 92 plots of 1 × 1 km chosen after stratification on the proportion of urban area and farmland habitat (either 0%, 25%, 50%, 75%), focusing on farmland habitat. Two aspects of urbanization were studied: its intensity and its age. We found that farmland bird species richness did not vary with increasing proportion of urbanized habitat. Non-farmland bird species richness increased from 0% to 25% classes and was constant for other classes. No effect of the urbanization age on farmland bird species richness was found, whereas a positive one was found on the non-farmland birds’ species richness. Abundance of the most specialized farmland birds decreases with urbanization intensity and age. We also found that, the more urbanized and the more recently urbanized the plots, the more similar bird communities. A strong difference in farmland bird’s communities’ compositions was found between 0% and 25% of urbanization, whereas no distinction was found between 50% and 75%. Altogether, our results suggest that to maintain for farmland birds, it is better to add new urban habitat in place where it already exist, rather than to spread it in small lots throughout the landscape.     

Scale-dependent homogenization: Changes in breeding bird diversity in the Netherlands over a 25-year period

Changes in breeding bird diversity in the Netherlands between 1973-1977 and 1998-2000 were evaluated by testing three hypotheses related to the loss of biodiversity: (1) species diversity is declining, (2) biotic homogenization is increasing and (3) rare species are declining more severely than abundant species. Using data collected for two successive national breeding bird atlases, changes in diversity were assessed at different spatial scales (local, regional and national) and among species characteristic for different landscapes (farmland, woodland, heathland, wetland, coastal habitats and urban habitats). National species richness, diversity and equitability had increased between the two atlas periods, with more species increasing than decreasing in range and abundance. Most species in the large groups of woodland and wetland birds showed positive trends, whereas most in the smaller groups of heathland, reed-breeding and meadow birds showed negative trends. However, findings varied between regions and localities. Increases in species richness occurred mainly in regions in the low-lying, western part of the country which were previously relatively poor in species. By contrast, species richness decreased in some previously species-rich regions in the eastern part of the country. This has resulted in a homogenization of breeding bird communities between regions. We advocate the conservation and restoration of regional identity as a priority for landscape planning in the Netherlands. We did not find a clear relation between species abundance and trends, although both rare and very abundant species tended to decrease on average.     

Does variation in garden characteristics influence the conservation of birds in suburbia?

Can enhancement of garden habitat for native birds have conservation benefits, or are garden bird assemblages determined by landscape and environmental characteristics? The relative roles of vegetation structure, floristics and other garden attributes, and environmental and landscape controls, on the abundance and richness of bird species in 214 back or front gardens in 10 suburbs of Hobart, Tasmania, Australia, are addressed to answer this question. Birds were counted in each garden and the resources they utilized noted. Vascular plant species and other attributes of the garden were noted, along with rainfall, altitude, distance from natural vegetation, distance from the city and garden size. Garden floristics and bird assemblages were ordinated, and garden groups characterized by particular assemblages of birds identified. General linear modelling was used to determine the combinations of independent variables that best predicted the richness of birds and the abundance of individual bird species and groups of species. The models for bird richness, bird species and groups of bird species were highly individualistic. Although native birds showed a preference for native plants, they also utilized many exotic plants. Exotic birds largely utilized exotic plants. Variation in garden characteristics does substantially affect the nature of garden bird assemblages in Hobart, with weaker environmental and landscape influences. The fact that gardens can be designed and managed to favour particular species and species assemblages gives gardeners a potentially substantial role in the conservation of urban native avifauna.     

Environmental monitoring using populations of birds and small mammals: Analyses of sampling effort

Characteristics of bird and small mammal communities can be valuable in monitoring for impacts on specific plots of land. In Utah (USA), significant differences were found among years, habitats and seasons in bird and small mammal abundance, species diversity, and species richness; thus, estimates of these characteristics are consistent and repeatable and can be used for detecting change. Also, bird and small mammal communities are dependent on a wide variety of food resources and habitat characteristics; thus they should reflect a variety of impacts.A comparison is made between two methods of handling bird data from transects: enumeration (individual birds per km) and density (birds per unit area). Correlation between the two was high (r = 0·93). However, density estimates may be necessary to trace an impact to a particular group, guild or species and possibly to its cause.An analysis is made of how much sampling is necessary to make an adequate estimate of the characteristics of bird and small mammal communities. Estimates of bird abundance, species richness and species diversity (from enumeration) may be obtained with three repetitions of 2 km of transect. Three repetitions are also sufficient to estimate abundance of small mammals on 12 by 12 trapping grids. To estimate species richness and species diversity, however, four or more repetitions are needed. Small mammal community characteristics are not well predicted by trapping on transects or small grids except in grids 9 by 9 or larger. These results of sampling effort may only apply to certain habitats but data from bird and small mammal communities should be useful in environmental monitoring at any site.     

Small isolated aspen stands enrich bird communities in southwestern ponderosa pine forests

Small aspen stands are disappearing from the landscape in the Southwest, so it is important to understand their contribution to the avian community. We sampled birds in 53 small, isolated aspen stands and 53 paired plots within the ponderosa pine forest in northern Arizona, during the 1996 and 1997 breeding seasons. Bird species richness and abundance were higher in aspen than in pine. However, bird species richness and abundance did not vary with size of the aspen patch or isolation index. In addition, direct ordination of species distributions with habitat factors suggested no distinct avian communities. This suggests that aspen stands do not harbor separate populations, but rather are locations where the regional avifauna reaches high local density and richness and may be crucial to birds in years of resource scarcity. Thus it is important for avian conservation to maintain many aspen stands across the landscape, encompassing a diversity of vegetation structure and composition.     

The search for fragmentation thresholds in a Southern Sydney Suburb

Fragmentation of habitat is recognised as the number one threat to biodiversity and as such has attracted considerable research. However, much of this research has been conducted in forestry and agricultural environments, with little research in urban areas. In this study, field surveys were conducted measuring the impact of fragmentation on bird, frog, plant and fungi species richness, within the fragmented urban landscape of southern Sydney. Of all fragmentation parameters examined, remnant area was the best and most significant predictor of species richness for all taxa studied. Remnant size thresholds, below which biodiversity declined rapidly, were observed at approximately 4 ha for bird and frog species richness and approximately 2 ha for plant and fungal species richness. A further threshold appears to exist at approximately 50 ha for the dominance of forest interior species. Further relationships were also observed for perimeter:area ratio, indicating the influence of various edge effects on all taxa. Isolation effects were observed in the form of an inverse linear relationship between distance to other large reserves and species richness for fungi, birds and frogs. Corridor connectivity also produced an overall positive relationship for birds, frogs and plants. It is concluded that the identification of fragmentation thresholds and relationships provides an important management tool for the design of networks aimed at conserving biodiversity in fragmented urban environments.     

Land use and habitat gradients determine bird community diversity and abundance in suburban, rural and reserve landscapes of Minnesota, USA

Bird species’ community responses to land use in the suburbanizing Twin Cities, Minnesota, USA, were contrasted among reserves, rural lands, and suburbs. For each land use type, bird composition, diversity, and abundance were recorded for 2 years in ≈99 plots in three sampling units (each ≈4500 ha). A habitat gradient defined by canopy structure (grasslands to savannas to forests) was influenced by land use, so ≈300 plots were used to characterize simultaneous variation in bird communities along land use and habitat gradients. At broad scales (aggregate of 33 plots covering ≈4500 ha) suburbs supported the lowest bird richness and diversity and rural landscapes the most, with reserves slightly below rural. Although reserves were like rural lands in diversity of bird communities, they supported more species of conservation concern, particularly of grasslands and savannas. Differences among land use types varied with habitat structure. Suburbs, rural lands, and reserves had similar forest bird communities, but differed in grassland and savanna bird communities. The extensive rural forests are important for the region’s forest birds. Suburban grasslands and savannas had low shrub abundance, low native bird richness and high non-native bird richness and abundance. However, total bird richness and diversity were as high in suburban as in rural and reserve plots because high native richness in suburban forests and high non-native species richness in suburban grasslands and savannas compensated for lower native richness in suburban grasslands and savannas. Bird conservation here and in the Midwest USA should protect rural forests, expand grasslands and savannas in reserves, and improve habitat quality overall.     

Spatial and temporal variation in the Argentine ant edge effect: Implications for the mechanism of edge limitation

One consequence of human land-use is the exposure of native communities to invasive human commensal species along edges. Argentine ants (Linepithema humile) invade a variety of habitats in California with consequent dramatic declines in native ants. In coastal southern California, USA Argentine ants appear to be an edge effect in small habitat fragments [Suarez, A.V., Bolger, D.T., Case, T.J., 1998. Effects of fragmentation and invasion on native ant communities in coastal southern California. Ecology 79, 2041-2056]. They invade fragments from the urban edge, but only penetrate coastal sage scrub (CSS) habitat to a distance of approximately 200 m. Using pitfall sampling in edge (<250 m from the urban edge) and interior (>600 m from the edge) habitat I tested whether there is also an Argentine ant edge effect in the largest blocks of habitat in the landscape and investigated patterns of spatial and temporal variation in native and Argentine ants. Argentine ants were common in coastal sage scrub habitat within 250 m of urban edges, but rare in interior areas. Correspondingly, native ants were significantly less abundant and diverse in edge areas as compared to interior. Over the period 1997-2000 Argentine ants did not become more abundant in interior habitat suggesting it will remain a refuge for native ants. Argentine ant abundance in edge plots varied greatly among years and sites. Annual variation in abundance was positively related to annual rainfall. Increased soil moisture near edges due to urban runoff has often been suggested as the mechanism that allows the invasion of edge but not interior habitat. This hypothesis predicts that edge habitat downslope of the urban edge should support invasions of higher abundance and greater spatial penetration than habitat upslope. However, I found that edge slope did not predict the extent of invasion, whereas, soil type did. Coarse, well-drained soils supported an Argentine ant invasion of lower abundance and lesser spatial penetration than soils that should retain more moisture. These patterns of spatial and annual variation are more consistent with a biotic flow mechanism where ants move from urban habitat into CSS sites that are temporarily favorable, rather than an abiotic flow where urban runoff causes a physical change to CSS habitat near edges. This invasion affects a substantial area of habitat, however, the inability of Argentine ants to invade interior habitat suggests that refugia for native ants will persist if large unfragmented blocks of habitat are maintained.     

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.     

Spatial and temporal variations in the use of rice-paddy dominated landscapes by birds in Japan

Agricultural landscapes with spatial and temporal variations interact with each other to affect the existing biodiversity. Though rice fields provide important habitats for birds all over the world, studies so far have rarely explored the effects of landscape heterogeneity on bird species in rice paddy areas. This study investigated the effects of habitat cover and landscape variables on the species richness and the abundance of birds in rice paddy areas in Japan. Data on bird occurrence and the environment were collected at 32 grid squares (1 × 1 km) in the Tone River basin. The richness and the abundance of agricultural wetland species were particularly high in landscapes with large areas of rice fields in summer, when rice fields were irrigated, but in those with large areas of open water in winter, when rice fields were drained. It is important to maintain a combination of rice fields and open water to satisfy multiple habitat requirements by agricultural wetland species throughout the year. Grassland species were positively associated with a rich diversity of land cover including fallow fields and open water, indicating the importance of a simultaneous existence of multiple landscape elements. Forest cover in landscapes positively affected edge species and woodland species. Since forest cover had a relatively strong correlation with edge density, the responses of bird species to changes in forest cover and edge density need to be explored further. This study illustrates the importance of spatial and temporal landscape complementation for bird species in rice paddy areas.     

Six years of habitat modification in a tropical rainforest margin of Indonesia do not affect bird diversity but endemic forest species

Studies on temporal changes of tropical bird communities in response to habitat modification are rare. We quantified changes in bird assemblages at the rainforest margin of Lore Lindu National Park, Central Sulawesi, over an interval of 6 years. Standardized bird counts were conducted in the years 2001/2002 and 2008 at 15 census points representing natural forest, secondary forest, agroforest and openland sites. Although overall species richness remained nearly identical, different species groups were affected unequally by habitat modification within the forest margin landscape. The mostly endemic forest species declined in abundance (72.0% of forest species) and were detected at fewer census points in 2008 (56.0%). In contrast, 81.8% of the solely widespread openland birds became more abundant and 63.6% of the species were recorded at a larger number of census points. Hence, recent human activities in the forest margin ecotone negatively affected species of high conservation value. Species richness turned out to be a poor indicator of habitat change, and our results underline the importance of considering species identities. Biotic homogenization as result of habitat conversion is a global phenomenon. In our study, the winners were widespread openland species, while the losers were endemic forest birds. In conclusion, our study shows that 6 years of land-use change had negative impacts on bird community structure and endangered species, but not on overall bird species richness.     

Avifauna homogenisation by urbanisation: Analysis at different European latitudes

We studied the homogenisation effects of urbanisation on avifauna in towns of three countries selected along a latitudinal gradient: Italy, France and Finland. In each town (n = 19), numbers of species were analysed along an urbanisation gradient using two urban sectors (centre and suburban) and one non-urban sector (periurban), representing the regional species pool. Firstly, we compared the avifauna by using species richness and similarity indexes along the urban gradient (S) and latitudinal gradient (L). In Europe, the number of exotic bird species in towns was low. The number of species decreased from the periurban and suburban sectors to the centre sector. Thus, the generally low number of species and few dominant birds indicate that urban bird communities are structurally simple. In addition, many habitat specialists were lacking from urban centres. The centre species represented about 43% of periurban species (similarity S). There was no correlation between town size and species trend in sectors. However, bird community similarity L was lower between town centres than between periurban areas. Latitude explained 89% of the species difference in periurban sector but only 52% in the centre, supporting the homogenisation effect of urbanisation. Secondly, we examined the homogenisation effect through the variability of some specific life-history traits (diets, nest heights, feeding habitats) by using data on Passeriformes. Our results suggested that urbanisation might cause homogenisation by decreasing the abundance of ground nesting bird species and bird species preferring bush-shrub habitats. Urbanisation appeared a cause of taxonomic homogenisation of the avifauna but the effects of latitude and urban habitat diversity may make generalisation difficult.     

Relationships between avian diversity, neighborhood age, income, and environmental characteristics of an urban landscape

Maintaining biodiversity in urbanizing landscapes has become a top conservation priority. We examined variation in bird communities across a diverse array of urban and suburban neighborhoods in the Chicago, Illinois, metropolitan region. Rather than taking the usual approach of focusing solely on natural features of the urban landscape, we investigated how urban bird communities were related to neighborhood age and income, as well as environmental characteristics. We found that median housing age was strongly related to avian species richness, with newer neighborhoods supporting more species. Housing age was an important correlate of abundance for several species as well as abundance of exotic, migratory, and non-migratory species groups. Per capita income was inversely related to richness of native bird species and positively related to exotic richness. Total richness was higher in urban sites with undeveloped patches and heterogeneous land cover types; moreover, richness decreased with increasing distance from natural areas greater than 1 km². Our findings suggest that bird richness is enhanced both by small patches of natural land within the urban matrix and by close proximity to large natural preserves. Furthermore, these results suggest that investigating a combination of abiotic and environmental features of the built landscape, rather than focusing solely on environmental features, may provide a more complete understanding of the factors influencing avian diversity in human-dominated landscapes.     

Riparian zones provide for distinct bird assemblages in forest mosaics of south-east Australia

Riparian zones are a characteristic component of many landscapes throughout the world and increasingly are recognised as key areas for biodiversity conservation. Their importance for bird communities has been well recognised in semi-arid environments and in modified landscapes where there is a marked contrast between riparian and adjacent upslope vegetation. The value of riparian zones in largely intact landscapes with continuous vegetation cover is less well understood. In this study, birds were surveyed at 30 pairs of riparian and adjacent non-riparian sites in extensive mesic forests of the Victorian Highlands, Australia. Riparian sites were floristically distinct from non-riparian sites and had a more complex vegetation structure, including a mid-storey tree layer mostly absent from non-riparian sites. Bird assemblages at riparian sites had significantly greater richness, abundance and diversity of species than was recorded at adjacent non-riparian sites. Species composition also differed significantly between these habitat types. Compositional differences in assemblages were due to a suite of distinctive species in each habitat and to significant contrasts in the densities of species that occurred in both habitat types. Many species (36%) attained a significantly greater abundance in riparian habitats. The distinctiveness and richness of the riparian avifauna contribute to the diversity of continuous forest landscapes. The spatial patterning of the avifauna, the occurrence of complementary assemblages, the presence of rare species and the potential for riparian habitats to serve as refuges, all point to the value of riparian zones and highlight the importance of landscape-level planning and management for avifaunal conservation.     

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.     

Composition and abundance of birds in Andean alder (Alnus acuminata) patches with past and present harvest in Bolivia

We compared species richness and abundance of birds between five patches under selective Alnus exploitation and five patches that have not been harvested for at least 10 years prior to our study, during the early dry season (April-July 2001), in Cotapata National Park. Using “point counts” we recorded birds and their distribution in two (<1.5 m and >1.5 m) forest layers. Simultaneously we evaluated the floristic structure (size [dbh] distribution, basal area, tree density, tree height, and vegetation cover) and composition (diversity) on three transects placed within each Alnus patch. Both bird diversity and vegetation cover were significantly higher in not presently used patches but only for the higher layer of the forest, whereas plant diversity was higher in presently used patches. Lack of differences between the two types of Alnus patches in any of the vegetation parameters measured in the lower layer was coupled with an indistinct avifauna. Small changes in habitat characteristics following a perturbation like selective logging have the potential to affect richness and abundance of birds, at least within the habitats directly affected by the perturbation.     

Long-term consequences of habitat fragmentation—highland birds in Oaxaca, Mexico

Studies of habitat fragmentation have been restricted primarily to anthropogenically-altered habitats, with most research conducted 60-90 years post-fragmentation. It is unclear whether patterns in older systems concur with results from these dynamic landscapes, and hence the long-term viability of populations inhabiting habitat fragments remains largely unexplored. I focused on resident birds in fragments of humid pine-oak forest in Oaxaca, southern Mexico, isolated over 5000 years ago by climate-change. Seventeen fragments, ranging from 2 ha to over 150,000 ha were sampled in 1997 and 1998 yielding 141 species, of which 60 residents were used for analysis. Avian assemblages exhibited a highly nested structure and, with several notable exceptions, assemblages of birds in low-richness fragments were predictable subsets of those in more diverse fragments. Patch-scale factors—area, shape, elevation, habitat diversity and fractal dimension of edge—all exerted strong univariate influence on avian richness but were so closely inter-related that none had a significant independent effect. Thus, larger fragments were more complex in shape, included higher peaks, supported more diverse forests, and contained higher diversities of resident species. In contrast, the landscape-scale index used—distance from nearest large fragment (>50,000 ha)—had little effect on richness. This was reinforced by species-level analyses—one species was significantly influenced by isolation, compared with 31 species that displayed significant minimum-area distributions, restricted to patches larger than a particular threshold value. In terms of autecology, vagility, relative abundance and elevational breadth were closely related to distribution—those species with greater mobility, higher abundances and broader elevational tolerances were consistently more widespread. I suggest that more abundant species were less prone to extinction initially, more vagile species were better dispersers and species with broader elevational tolerances more likely to be successful colonists. As with previous research from older landscapes, patch-scale factors were consistently found to be influential, with high quality fragments supporting diverse communities regardless of landscape context. This suggests that the influence of landscape-scale factors noted in younger, anthropogenically fragmented systems may be transitory, overwhelmed by patch-scale factors with time. Which patch attributes are most influential could not be resolved, however, indicating that even thousands of years after fragmentation, they affect diversity patterns in concert. Rather than differentiating effects of area from habitat heterogeneity and other patch-level factors, I advocate resource-based approaches to understand and manage diversity in habitat fragments.     

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.