Detecting Biodiversity Hotspots by Species-Area Relationships: a Case Study of Mediterranean Beetles

Abstract:  Any method of identifying hotspots should take into account the effect of area on species richness. I examined the importance of the species-area relationship in determining tenebrionid (Coleoptera: Tenebrionidae) hotspots on the Aegean Islands (Greece). Thirty-two islands and 170 taxa (species and subspecies) were included in this study. I tested several species-area relationship models with linear and nonlinear regressions, including power, exponential, negative exponential, logistic, Gompertz, Weibull, Lomolino, and He-Legendre functions. Islands with positive residuals were identified as hotspots. I also analyzed the values of the C parameter of the power function and the simple species-area ratios. Species richness was significantly correlated with island area for all models. The power function model was the most convenient one. Most functions, however, identified certain islands as hotspots. The importance of endemics in insular biotas should be evaluated carefully because they are of high conservation concern. The simple use of the species-area relationship can be problematic when areas with no endemics are included. Therefore the importance of endemics should be evaluated according to different methods, such as percentages, to take into account different levels of endemism and different kinds of "endemics" (e.g., endemic to single islands vs. endemic to the archipelago). Because the species-area relationship is a key pattern in ecology, my findings can be applied at broader scales.     

Effects of Habitat Fragmentation on Bird Species in a Relict Temperate Forest in Semiarid Chile

Abstract: We analyzed the structure and composition of a bird assemblage in a fragmented relict temperate forest located in northcentral Chile ( Fray Jorge National Park). In terms of species composition, the bird assemblage we found in Fray Jorge was more similar to southern temperate forest sites, located more than 1200 km south of Fray Jorge, than to localities found in nearby scrub habitats. The relict character and long-term isolation of the Fray Jorge forest provides a natural experiment with which to establish the potential long-term effects of fragmentation and isolation on southern Chilean temperate forests. Between May 1996 and March 1997, we conducted seasonal surveys of birds in six forest fragments, ranging in size from 0.5 to 22.5 ha, at Fray Jorge. The number of bird species at each forest fragment was positively correlated with fragment area during all seasons. The relict forest system had a steeper species-area slope than that reported for similar temperate-forest bird assemblages in forest fragments within Chiloé Island and for islands across the Chiloé Archipelago in southern Chile. In this regard, this bird fauna resembled a depauperate oceanic archipelago. This difference in area effects is likely a consequence of the minimization of rescue effects because of the absence of large source forest areas nearby and the long-term isolation of the system. In addition, the distribution of species among forest fragments in Fray Jorge was not random, showing a nested subset pattern. Thus, some species occur across all fragments, regardless of their area, and therefore are less affected by habitat fragmentation and less prone to local extinction. These results suggest that, for south-temperate forest birds, large fragments (or reserves) should afford better protection against extinction than small forest patches.     

Habitat fragmentation and effects of herbivore (howler monkey) abundances on bird species richness

Habitat fragmentation can alter herbivore abundances, potentially causing changes in the plant community that can propagate through the food web and eventually influence other important taxonomic groups such as birds. Here we test the relationship between the density of red howler monkeys (Alouatta seniculus) and bird species richness on a large set of recently isolated land-bridge islands in Lago Guri, Venezuela (n = 29 islands). Several of these islands host relict populations of howler monkeys at densities up to more than 30 times greater than those on the mainland. These "hyperabundant" herbivores previously have been shown to have a strong positive influence on aboveground plant productivity. We predicted that this should lead to a positive, indirect effect of howler monkey density on bird species richness. After accounting for passive sampling (the tendency for species richness to be positively associated with island area, regardless of differences in habitat quality) we found a significant positive correlation between howler monkey density and bird species richness. A path analysis incorporating data on tree growth rates from a subset of islands (n = 9) supported the hypothesis that the effect of howler monkeys on the resident bird communities is indirect and is mediated through changes in plant productivity and habitat quality. These results highlight the potential for disparate taxonomic groups to be related through indirect interactions and trophic cascades.     

Is the effect of forest structure on bird diversity modified by forest productivity?

Currently, the most common strategy when managing forests for biodiversity at the landscape scale is to maintain structural complexity within stands and provide a variety of seral stages across landscapes. Advances in ecological theory reveal that biodiversity at continental scales is strongly influenced by available energy (i.e., climate factors relating to heat and light and primary productivity). This paper explores how available energy and forest structural complexity may interact to drive biodiversity at a regional scale. We hypothesized that bird species richness exhibits a hump-shaped relationship with energy at the regional scale of the northwestern United States. As a result, we hypothesized that the relationship between energy and richness within a landscape is positive in energy-limited landscapes and flat or decreasing in energy-rich landscapes. Additionally, we hypothesized that structural complexity explains less of the variation in species richness in energy-limited environments and more in energy-rich environments and that the slope of the relationship between structural complexity and richness is greatest in energy-rich environments. We sampled bird communities and vegetation across seral stages and biophysical settings at each of five landscapes arrayed across a productivity gradient from the Pacific Coast to the Rocky Mountains within the five northwestern states of the contiguous United States. We analyzed the response of richness to structural complexity and energy covariates at each landscape. We found that (1) richness had a hump-shaped relationship with available energy across the northwestern United States, (2) the landscape-scale relationships between energy and richness were positive or hump shaped in energy-limited locations and were flat or negative in energy-rich locations, (3) forest structural complexity explained more of the variation in bird species richness in energy-rich landscapes, and (4) the slope of the relationship between forest structural complexity and richness was steepest in energy-limited locations. In energy-rich locations, forest managers will likely increase landscape-scale bird diversity by providing a range of forest structural complexity across all seral stages. In low-energy environments, bird diversity will likely be maximized by managing local high-energy hotspots judiciously and adjusting harvest intensities in other locations to compensate for slower regeneration rates.     

A hidden species-area curve

The species-area curve, which describes the relationship between the number of observed species in a geographical region and the area of the region, plays a central role in biogeography. Beyond its scientific importance, the species-area curve is commonly used to assess the loss of species due to habitat loss. When the species-area curve is estimated from spatial samples, the existence of species with low or highly spatially variable abundance exaggerates the true rate at which species accumulate with area. Here, a hidden species-area curve is defined that accounts for this sampling effect and its estimation by maximum likelihood is outlined. Both the species-area curve and the hidden species-area curve are conditioned on the observed species list; thus the analysis does not depend on the total number of observed and unobserved species, that is, species richness. The method is tested by sub-sampling some tropical forest data and found to work well. It is also applied to a classic data set from the deep sea. For these data, accounting for this sampling effect has a large impact.     

Deforestation Predicts the Number of Threatened Birds in Insular Southeast Asia

The world's tropical forests are being cleared rapidly, and ecologists claim this is causing a massive loss of species. This claim has its critics. Can we predict extinctions from the extent of deforestation? We mapped the percentage of deforestation on the islands of the Philippines and Indonesia and counted the number of bird species found only on these islands. We then used the species-area relationship to calculate the number of species predicted to become globally extinct following deforestation on these islands. Next, we counted the numbers of insular southeast Asian endemic bird species considered threatened—i.e., those having "a high probability of extinction in the wild in the medium-term future"—in the latest summary Red Data Book. The numbers of extinctions predicted from deforestation and the numbers of species actually threatened are strikingly similar. This suggests we can estimate the size of the extinction crisis in once-forested regions from the extent of deforestation. The numbers of extinctions will be large. Without rapid and effective conservation, many of the species endemic to insular southeast Asia will soon be lost.     

Modeling biodiversity dynamics in countryside landscapes

The future of biodiversity hinges to a great extent on the conservation value of countryside, the growing fraction of Earth's surface heavily influenced by human activities. How many species, and which species, can persist in such landscapes (and analogous seascapes) are open questions. Here we explore two complementary theoretical frameworks to address these questions: species-area relationships and demographic models. We use the terrestrial mammal fauna of Central America to illustrate the application of both frameworks. We begin by proposing a multi-habitat species-area relationship, the countryside species-area relationship, to forecast species extinction rates. To apply it, we classify the mammal fauna by affinity to native and human-dominated habitats. We show how considering the conservation value of countryside habitats changes estimates derived from the classic species-area approach We also examine how the z value of the species-area relationship affects extinction estimates. Next, we present a framework for assessing the relative vulnerability of species to extinction in the countryside, based on the Skellam model of population dynamics. This model predicts the minimum area of contiguous native habitat required for persistence of a species, which we use as an indicator of vulnerability to habitat change. To apply the model, we use our habitat affinity classification of mammals and we estimate life-history parameters by species and habitat type. The resulting ranking of vulnerabilities is significantly correlated with the World Conservation Union (IUCN) Red List assessment.     

Metacommunity models predict the local-regional species richness relationship in a natural system

Many natural communities exhibit positive relationships between local and regional species richness (LSR-RSR relationships), which can be either linear or curvilinear. Previous models have shown that the form of this relationship depends on the relative rates of colonization and extinction and the sensitivity of these rates to competition. We use simple models to show that the LSR-RSR relationship also depends on the type of metacommunity structure (Levins-like or mainland-island), and our models generate a wider range of realistic forms than do most previous models. We parameterize and test our models with two independent data sets for Daphnia in rock pools on islands in Finland and Sweden. We find that the Levins-like model with competition correctly predicts the observed LSR-RSR relationship and provides the best fit to the average local species richness per island. Simulations show that our models are robust to relaxing our assumption of identical species properties. Our study is one of the first to make and successfully test quantitative predictions for how a widely studied community pattern, the LSR-RSR relationship, arises from metacommunity dynamics.     

A comparison of land-sharing and land-sparing strategies for plant richness conservation in agricultural landscapes

Strategies for conserving plant diversity in agroecosystems generally focus on either expanding land area in non-crop habitat or enhancing diversity within crop fields through changes in within-field management practices. In this study, we compare effects on landscape-scale species richness from such land-sharing or land-sparing strategies. We collected data in arable field, grassland, pasture, and forest habitat types (1.6 ha sampled per habitat type) across a 100-km2 region of farmland in Lancaster County, Pennsylvania, USA. We fitted species-area relationships (SARs) for each habitat type and then combined extrapolations from the curves with estimates of community overlap to estimate richness in a 314.5-ha landscape. We then modified these baseline estimates by adjusting parameters in the SAR models to compare potential effects of land-sharing and land-sparing conservation practices on landscape richness. We found that species richness of the habitat types showed a strong inverse relationship to the relative land area of each type in the region, with 89 species in arable fields (66.5% of total land area), 153 in pastures (6.7%), 196 in forests (5.2%), and 213 in grasslands (2.9%). Relative to the baseline scenario, major changes in the richness of arable fields produced gains in landscape-scale richness comparable to a conversion of 3.1% of arable field area into grassland habitat. Sensitivity analysis of our model indicated that relative gains from land sparing would be greatest in landscapes with a low amount of non-crop habitat in the baseline scenario, but that in more complex landscapes land sharing would provide greater gains. These results indicate that the majority of plant species in agroecosystems are found in small fragments of non-crop habitat and suggest that, especially in landscapes with little non-crop habitat, richness can be more readily conserved through land-sparing approaches.     

Benefits to poorly studied taxa of conservation of bird and mammal diversity on islands

Protected area delineation and conservation action are urgently needed on marine islands, but the potential biodiversity benefits of these activities can be difficult to assess due to lack of species diversity information for lesser known taxa. We used linear mixed effects modeling and simple spatial analyses to investigate whether conservation activities based on the diversity of well‐known insular taxa (birds and mammals) are likely to also capture the diversity of lesser known taxa (reptiles, amphibians, vascular land plants, ants, land snails, butterflies, and tenebrionid beetles). We assembled total, threatened, and endemic diversity data for both well‐known and lesser known taxa and combined these with physical island biogeography characteristics for 1190 islands from 109 archipelagos. Among physical island biogeography factors, island area was the best indicator of diversity of both well‐known and little‐known taxa. Among taxonomic factors, total mammal species richness was the best indicator of total diversity of lesser known taxa, and the combination of threatened mammal and threatened bird diversity was the best indicator of lesser known endemic richness. The results of other intertaxon diversity comparisons were highly variable, however. Based on our results, we suggest that protecting islands above a certain minimum threshold area may be the most efficient use of conservation resources. For example, using our island database, if the threshold were set at 10 km² and the smallest 10% of islands greater than this threshold were protected, 119 islands would be protected. The islands would range in size from 10 to 29 km² and would include 268 lesser known species endemic to a single island, along with 11 bird and mammal species endemic to a single island. Our results suggest that for islands of equivalent size, prioritization based on total or threatened bird and mammal diversity may also capture opportunities to protect lesser known species endemic to islands. Beneficios de los Taxa Poco Estudiados para la Conservación de la Diversidad de Aves y Mamíferos en Islas     

Threats to Avifauna on Oceanic Islands

Abstract:  Results of the study by Blackburn et al. (2004 a ) of avifauna on oceanic islands suggest that distance from the mainland and time since European colonization have major influences on species extinctions and that island area is a significant but secondary contributing factor. After augmenting the data of the study on geographical properties for some of the islands they examined, we used a causal analysis approach with structural equation modeling to reexamine their conclusions. In our model geographical properties of islands, such as island area and isolation, were considered constraints on biological factors, such as the number of introduced mammalian predators and existing number of avifauna, that can directly or indirectly influence extinction. Of the variables we tested, island area had the greatest total influence on the threat of extinction due to its direct and indirect effects on the size of island avifauna. Larger islands had both a greater number of threatened bird species and more avifauna, increasing the number of species that could become threatened with extinction. Island isolation also had a significant, positive, and direct effect on threats to island avifauna because islands farther from the mainland had fewer current extant avifauna. Time since European colonization had a significant negative, but relatively weaker, influence on threats compared with the traditional biogeographic factors of island area and distance to the mainland. We also tested the hypothesis that the amount of threat is proportionally lower on islands that have had more extinctions (i.e., there is a "filter effect"). Because the proportion of bird extinctions potentially explained only 2.3% of the variation in the proportion of threatened species on islands, our results did not support this hypothesis. Causal modeling provided a powerful tool for examining threat of extinction patterns of known and hypothesized pathways of influence.     

Patterns of sponge biodiversity and abundance across different biogeographic regions

Few studies have compared the underlying nature or structure of marine communities and assemblages across broad spatial scales, despite the importance of such comparisons in understanding global scale responses to environmental change and biodiversity conservation. The aim of this study was to examine the consistency of relationships between sponge abundance and richness on the undersides of boulders (to control for multiple confounding factors) in relation to space availability (boulder size) in three widely separated biogeographic regions. Sponges followed typical species-area (SA) relationships irrespective of site or ocean and we consistently found no decrease in sponge richness at larger boulder sizes. We also found reliable underlying density–area (DA) relationships, with larger boulders supporting more sponge patches at all sites. Although the general SA and DA relationships were similar between oceans and sites, the exact nature of these relationships in terms of the actual species present per unit area, total number of species present or density differed between sites. There were no consistent differences between SA and DA relationships at each locality, suggesting that although disturbance (particularly wave action) regimes may play an important role in controlling sponge biodiversity and abundance, its effects are manifested differently at local-scales. Even though boulders have received a considerable amount of research effort over the past three decades, this is the first study to examine the consistency of DA and SA relationships between ocean basins, finding similar relationships.     

Bird metacommunities in temperate South American forest: vegetation structure, area, and climate effects

Spatial structure in metacommunities and their relationships to environmental gradients have been linked to opposing theories of community assembly. In particular, while the species sorting hypothesis predicts strong environmental influences, the neutral theory, the mass effect, and the patch dynamics frameworks all predict differing degrees of spatial structure resulting from dispersal and competition limitations. Here we study the relative influence of environmental gradients and spatial structure in bird assemblages of the Chilean temperate forest. We carried out bird and vegetation surveys in South American temperate forests at 147 points located in nine different protected areas in central Chile, and collected meteorological and productivity data for these localities. Species composition dissimilarities between sites were calculated, as well as three indices of bird local diversity: observed species richness, Chao estimate of richness, and Shannon diversity. A stepwise multiple regression and partial regression analyses were used to select a small number of environmental factors that predicted bird species diversity. Although diversity indices were spatially autocorrelated, environmental factors were sufficient to account for this autocorrelation. Moreover, community dissimilarities were not significantly related to distance between sites. We then tested a multivariate hypothesis about climate, vegetation, and avian diversity interactions using a structural equation modeling (SEM) approach. The SEM showed that climate and area of fragments have important indirect effects on avian diversity, mediated through changes in vegetation structure. Given the scale of this study, the metacommunity framework provides useful insights into the mechanisms driving bird assemblages in this region. Taken together, the weak spatial structure of community composition and diversity, as well as the strong environmental effects on bird diversity, support the interpretation that species sorting has a predominant role in structuring avian assemblages in the region.     

Systematic Temporal Patterns in the Relationship Between Housing Development and Forest Bird Biodiversity

As people encroach increasingly on natural areas, one question is how this affects avian biodiversity. The answer to this is partly scale‐dependent. At broad scales, human populations and biodiversity concentrate in the same areas and are positively associated, but at local scales people and biodiversity are negatively associated with biodiversity. We investigated whether there is also a systematic temporal trend in the relationship between bird biodiversity and housing development. We used linear regression to examine associations between forest bird species richness and housing growth in the conterminous United States over 30 years. Our data sources were the North American Breeding Bird Survey and the 2000 decennial U.S. Census. In the 9 largest forested ecoregions, housing density increased continually over time. Across the conterminous United States, the association between bird species richness and housing density was positive for virtually all guilds except ground nesting birds. We found a systematic trajectory of declining bird species richness as housing increased through time. In more recently developed ecoregions, where housing density was still low, the association with bird species richness was neutral or positive. In ecoregions that were developed earlier and where housing density was highest, the association of housing density with bird species richness for most guilds was negative and grew stronger with advancing decades. We propose that in general the relationship between human settlement and biodiversity over time unfolds as a 2‐phase process. The first phase is apparently innocuous; associations are positive due to coincidence of low‐density housing with high biodiversity. The second phase is highly detrimental to biodiversity, and increases in housing density are associated with biodiversity losses. The long‐term effect on biodiversity depends on the final housing density. This general pattern can help unify our understanding of the relationship of human encroachment and biodiversity response. Patrones Sistemáticos Temporales en la Relación entre Desarrollos Urbanos y la Biodiversidad de Aves de Bosque     

Evidence of dispersal limitation in soil microorganisms: isolation reduces species richness on mycorrhizal tree islands

Dispersal limitation plays an important role in a number of equilibrium and nonequilibrium theories about community ecology. In this study we use the framework of island biogeography to look for evidence of dispersal limitation in ectomycorrhizal fungal assemblages on "tree islands," patches of host trees located in a non-host vegetation matrix. Because of the potentially strong effects of island area on species richness and immigration, we chose to control island size by sampling tree islands consisting of a single host individual. Richness on tree islands was high, with estimates ranging up to 42 species of ectomycorrhizal fungi associating with a single host individual. Species richness decreased significantly with increasing isolation of tree islands, with our regression predicting a 50% decrease in species richness when tree islands are located distances of approximately 1 km from large patches of contiguous forests. Despite the fact that fungal fruit bodies produce large numbers of spores with high potential for long-distance travel, these results suggest that dispersal limitation is significant in ectomycorrhizal assemblages. There were no discernible effects of isolation or environment on the species identity of tree island fungal colonists. In contrast to the highly predictable patterns of tree island colonization we observed in a previous study on early successional forests, we suggest that over longer time periods the community assembly process becomes more dominated by stochastic immigration and local extinction events.     

Species richness patterns along environmental gradients in island land molluscan fauna

The relationship between species richness and environmental variables may change depending on habitat structure, dispersal ability, species mixing, and community adaptation to the environment. It is crucial to know how these factors regulate the environment-diversity relationship. The land molluscan fauna of the Ogasawara Islands in the West Pacific is an excellent model system to address this question because of the high species endemicity (> 90%), small area, and simple habitat structure of the islands. I examined relationships among indigenous species composition, richness, and habitat condition, and especially productivity and forest moisture on the island of Anijima. Two major communities of snails could be distinguished by detrended correspondence analysis (DCA): one group dominated in a moist habitat with high productivity, and the other group dominated in a dry habitat with low productivity. However, species richness became highest at the intermediate condition between the habitats in which the two snail communities were dominant, so that species richness showed a hump-shaped relationship with moisture and productivity. In contrast, the species richness of the snail community in the moist habitat showed a monotonically positive correlation, and that in the dry habitat showed a monotonically negative correlation with moisture and productivity. Thus, the greater species richness in intermediate moisture and productivity resulted from the ecotone effect or community overlap at the transitional areas, where faunas with different ecologies can meet in a single site. These findings suggest that hump-shaped productivity-diversity relationships in land Mollusca would reflect the ecotone effect as a result of the mixing of species adapted to either fertile habitats or sterile habitats.     

Variation in species losses from islands: artifacts, extirpation rates, or pre-fragmentation diversity?

Species are being lost from isolated reserves as predicted by ecological theory, prompting calls for larger reserves with higher species immigration rates. However, some large islands have lost a large proportion of their species, whereas some small islands have not lost any. Conservation efforts would be more efficient if the cause of such variation in the relationships among number of species lost, island size, and immigration rate were known. Observed species losses could be affected by the time since islands were isolated, species immigration rates, species extirpation rates, the pre-fragmentation diversity of the region relative to steady state, or overestimation of the pre-fragmentation diversity of islands. To test the last three hypotheses, I compared the intersection points of the island, intraprovincial, and interprovincial species-area relationships of terrestrial mammals from nine archipelagos of land-bridge islands and terrestrial habitat isolates. Species losses from three archipelagos were greater than expected due to reduced immigration rates alone, although I could not resolve if this was due to increased extirpation rates or overestimation of the pre-fragmentation diversity of the islands. Analysis of six archipelagos indicates that the diversity of mammals in two regions of North America is currently below steady state, probably due to the extinction of mammals and glacial retreat during the late Pleistocene. These results have direct implications for reserve planning. When provincial diversity is below steady state, some combinations of reserve size and species immigration rate will allow reserves to maintain their pre-isolation diversity. However, the diversity of provinces relative to steady state is likely to vary, so conservation of a given proportion of a province may not always conserve the same proportion of its species. I present a new species-area relationship for islands formed by fragmentation that replaces the parameter c (fitted constant) with a rotation point. Estimation of this rotation point will allow reserve planners to separate the effects of extirpation and immigration rates on species losses from islands, identify provinces that are below steady-state diversity, and estimate the combinations of reserve size and immigration rate that will prevent loss of species from reserves.     

Deforestation and Avian Extinction on Tropical Landbridge Islands

Abstract: There are few empirical data, particularly collected simultaneously from multiple sites, on extinctions resulting from human‐driven land‐use change. Southeast Asia has the highest deforestation rate in the world, but the resulting losses of biological diversity remain poorly documented. Between November 2006 and March 2008, we conducted bird surveys on six landbridge islands in Malaysia and Indonesia. These islands were surveyed previously for birds in the early 1900s, when they were extensively forested. Our bird inventories of the islands were nearly complete, as indicated by sampling saturation curves and nonparametric true richness estimators. From zero (Pulau Malawali and Pulau Mantanani) to 15 (Pulau Bintan) diurnal resident landbird species were apparently extirpated since the early 1900s. Adding comparable but published extinction data from Singapore to our regression analyses, we found there were proportionally fewer forest bird extinctions in areas with greater remaining forest cover. Nevertheless, the statistical evidence to support this relationship was weak, owing to our unavoidably small sample size. Bird species that are restricted to the Indomalayan region, lay few eggs, are heavier, and occupy a narrower habitat breadth, were most vulnerable to extinction on Pulau Bintan. This was the only island where sufficient data existed to analyze the correlates of extinction. Forest preservation and restoration are needed on these islands to conserve the remaining forest avifauna. Our study of landbridge islands indicates that deforestation may increasingly threaten Southeast Asian biodiversity.     

Choice of Species-Area Function Affects Identification of Hotspots

Abstract: I tested the reliability of species-area curves for use in identifying hotspots, political or geographical regions of high species richness. On a species-area plot, hotspots are points (regions) that appear above the curve to a greater extent than other points. Because several different curves can be fit to species-area data, identification of hotspots may differ depending on the curve-fitting function used. I tested this hypothesis by comparing hotspots identified by the power function, the extreme value function, a linear function, and the exponential function. I examined several species-area data sets varying in size and in the presence of endemics. I defined hotspots as the highest 25% for small data sets and highest 10% and 25% for large data sets of standardized residuals from each function fitted to each data set. For some data sets, the functions agreed in identification of hotspots in that they identified 75% or more of the same hotspots. The extreme value function tended to identify hotspots not identified by the other three functions. For most data sets, the functions did not agree completely in identifying hotspots. Therefore, species-area curves should not be used as the sole means of identifying hotspots of species richness, although they can be used to examine the effect hotspot area has on richness for hotspots identified by other methods.     

Fragments as Islands: a Synthesis of Faunal Responses to Habitat Patchiness

Abstract:  Scientific interest in the impact of habitat fragmentation on biodiversity is increasing, but our understanding of fragmentation is clouded by a lack of appreciation for fundamental similarities and differences across studies representing a wide range of taxa and landscape types. In an effort to synthesize data describing ecological responses of animals to fragmentation across two classes of independent variables (taxonomic group and landscape), we sampled 148 studies of five major faunal groups from the primary literature and analyzed data on 13 variables extracted from those studies. We focused our analyses on three classes of dependent variables (effects of area and isolation on species richness, z values, and nestedness and species composition). Area ranged over more orders of magnitude than isolation and tended to explain more variation in species richness than isolation. There were few matrix or taxon effects on the patterns we investigated, although we did find that sky islands tended to manifest isolation effects on both species richness and nestedness more frequently than other patch types. Sky islands may offer insight into the future of habitat patches fragmented by contemporary habitat loss, and because they show a stronger effect of isolation than other patch types, we suggest that isolation will play an increasing role in the biology of habitat fragments. We use multiple lines of evidence to suggest that our understanding of the role of isolation on community assembly in fragmented landscapes is inadequate. Finally, our observation that consistent taxonomic differences in community patterns were minimal suggests that conservation actions intended to mitigate the negative effects of extinction may have far-reaching effects across taxonomic groups.