Responses of insect herbivores and herbivory to habitat fragmentation: a hierarchical meta‐analysis

Loss and fragmentation of natural habitats can lead to alterations of plant–animal interactions and ecosystems functioning. Insect herbivory, an important antagonistic interaction is expected to be influenced by habitat fragmentation through direct negative effects on herbivore community richness and indirect positive effects due to losses of natural enemies. Plant community changes with habitat fragmentation added to the indirect effects but with little predictable impact. Here, we evaluated habitat fragmentation effects on both herbivory and herbivore diversity, using novel hierarchical meta‐analyses. Across 89 studies, we found a negative effect of habitat fragmentation on abundance and species richness of herbivores, but only a non‐significant trend on herbivory. Reduced area and increased isolation of remaining fragments yielded the strongest effect on abundance and species richness, while specialist herbivores were the most vulnerable to habitat fragmentation. These fragmentation effects were more pronounced in studies with large spatial extent. The strong reduction in herbivore diversity, but not herbivory, indicates how important common generalist species can be in maintaining herbivory as a major ecosystem process.     

Experimental effects of habitat fragmentation on rove beetles and ants: patch area or edge?

The effects of habitat fragmentation may include the loss of species from isolated fragments or changes in species abundances among habitats that differ in area, structure, or edge characteristics. We measured the species richness and abundance of ground‐dwelling insects in a 1.14‐ha old field that was mowed to produce patches of unmowed vegetation which differed in size, degree of isolation, and the amount of habitat edge. Four treatments – ranging from unfragmented (169‐m²) to highly fragmented (1‐m²) patches – were replicated four times in a Latin square design, and insects were sampled twice during 1995 using 177 pitfall traps. Species richness showed a non‐monotonic response to fragmentation, with the fewest species occurring in the slightly fragmented treatment. Responses of rove beetles and ants, the most species‐rich and abundant taxa, respectively, were similar to the overall insect community but ants had a stronger and more consistent treatment effect in both sample months. Ordinations of ant and rove‐beetle assemblages using nonmetric multidimensional scaling showed that the slightly fragmented treatment differed from other treatments in species occurrence and abundance. The lower species richness in the slightly fragmented treatment was primarily due to a subset of ant and rove beetle species that showed a lower abundance than in other treatments, possibly because this treatment had the greatest amount of habitat edge. We hypothesize that the non‐monotonic species response to fragmentation was due to the differential effects of habitat edge on species movements across the habitat boundary between unmowed patches and mowed areas. A greater effect due to the amount of habitat edge rather than total patch area, at least among the range of patch sizes studied, suggests that the length of habitat edge may be quite important to the distribution and abundance of ground‐dwelling animals in fragmented habitats.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

Increased abundance of native and non-native spiders with habitat fragmentation

Habitat fragmentation and invasive species often contribute to the decline of native taxa. Since the penetration of non‐native species into natural habitat may be facilitated by habitat fragmentation, it is important to examine how these two factors interact. Previous research documented that, in contrast to most other arthropod taxa, spiders increased in density and morphospecies richness with decreasing fragment area and increasing fragment age (time since insularization) in urban habitat fragments in San Diego County, California, USA. We tested whether a specific mechanism, an increase in non‐native species with fragmentation, is responsible for this pattern. We found that both native and non‐native taxa contributed to the pattern. Abundance of native spiders per pitfall trap sample increased significantly with decreasing fragment size (i.e. a negative density–area relationship) and abundance of non‐natives increased significantly with increasing fragment age. The proportion of non‐native individuals also increased significantly with age. One non‐native species, Oecobius navus, comprised the majority of non‐native individuals (82.2%) and a significant proportion of total individuals (25.1%). Richness of spider families per sample (family density) increased with fragment age due to an increase in the occurrence of non‐natives in older fragments, however, native family richness did not vary with age or area. Due to increasing dominance by non‐native and some native families, family evenness declined with decreasing fragment size and increasing fragment age. Native and non‐native abundance covaried positively arguing against strong negative interactions between the two groups. O. navus had a strong positive association with another common non‐native arthropod, the Argentine ant (Linepitheme humile), suggesting a possible direct interaction. In contrast, abundance of native spiders was negatively correlated with Argentine ant abundance. We hypothesize that fragmentation in this semiarid habitat increases productivity in smaller and older fragments enhancing the density of both native and non‐native taxa.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

The effect of fragment area on site‐level biodiversity

Habitat fragmentation accompanies habitat loss, and drives additional biodiversity change; but few global biodiversity models explicitly analyse the effects of both fragmentation and loss. Here we propose and test the hypothesis that, as fragment area increases, species density (the number of species in a standardised plot) will scale with an exponent given by the difference between the exponents of the species–area relationships for islands (z ~ 0.25) and in contiguous habitat (z ~ 0.15), and test whether scaling varies between land uses. We also investigate the scaling of overall abundance and rarefaction‐based richness, as some mechanisms make different predictions about how fragment area should affect them. The relevant data from the taxonomically and geographically broad PREDICTS database were used to model the three diversity measures, testing their scaling with fragment area and whether the scaling exponent varied among land uses (primary forest, secondary forest, plantation forest, cropland and pasture). In addition, the consistency of the response of species density to fragment area was tested across three well represented taxa (Magnoliopsida, Hymenoptera and ‘herptiles’). Species density and total abundance showed area‐scaling exponents of 0.07 and 0.16, respectively, and these exponents did not vary significantly among land uses; rarefaction‐based richness by contrast did not increase consistently with area. These results suggest that the area‐scaling of species density is driven by the area‐scaling of total abundance, with additive edge effects (species moving into the small fragments from the surroundings) opposing – but not fully overcoming – the effect of fragment area on overall density of individuals. The interaction between fragment area and higher taxon (plants, vertebrates and invertebrates), which remained in the rarefied richness model, indicates that mechanisms may vary among groups.     

Local native plant diversity responds to habitat loss and fragmentation over different time spans and spatial scales

Separating the threats that habitat loss and habitat fragmentation pose to biodiversity is challenging because these processes usually occur simultaneously. Additionally, their importance may be underestimated due to time-delayed extinction. In central Texas savannas, woody plant encroachment reduces the amount of habitat available to herbaceous species while fragmenting remaining habitat. We examined the relationships between present species richness and present and past habitat amount and fragmentation (measured as fractal dimension) using a series of aerial photographs taken over nearly 60 years. We show that woody plant encroachment, a common phenomenon in savannas worldwide, reduces the diversity of herbaceous vegetation through both habitat loss and fragmentation. Habitat loss has the strongest impact on species richness over short time spans and small spatial scales in these savannas. Habitat fragmentation, however, has the strongest impact over longer time spans and larger spatial scales and generates long-term extinction debts. We also demonstrate that examining habitat loss and habitat fragmentation across different time periods and at different spatial scales is essential for understanding their joint and individual effects on plant community composition.     

Breeding bird species diversity across gradients of land use from forest to agriculture in Europe

Loss, fragmentation and decreasing quality of habitats have been proposed as major threats to biodiversity world‐wide, but relatively little is known about biodiversity responses to multiple pressures, particularly at very large spatial scales. We evaluated the relative contributions of four landscape variables (habitat cover, diversity, fragmentation and productivity) in determining different components of avian diversity across Europe. We sampled breeding birds in multiple 1‐km² landscapes, from high forest cover to intensive agricultural land, in eight countries during 2001?2002. We predicted that the total diversity would peak at intermediate levels of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; forest and open‐habitat specialists would show threshold conditions along gradients of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; resident species would be more strongly impacted by forest cover and fragmentation than migratory species; and generalists and urban species would show weak responses. Measures of total diversity did not peak at intermediate levels of forest cover or fragmentation. Rarefaction‐standardized species richness decreased marginally and linearly with increasing forest cover and increased non‐linearly with productivity, whereas all measures increased linearly with increasing fragmentation and landscape diversity. Forest and open‐habitat specialists responded approximately linearly to forest cover and also weakly to habitat diversity, fragmentation and productivity. Generalists and urban species responded weakly to the landscape variables, but some groups responded non‐linearly to productivity and marginally to habitat diversity. Resident species were not consistently more sensitive than migratory species to any of the landscape variables. These findings are relevant to landscapes with relatively long histories of human land‐use, and they highlight that habitat loss, fragmentation and habitat‐type diversity must all be considered in land‐use planning and landscape modeling of avian communities.     

Dung beetle (Coleoptera: Scarabaeinae) persistence in Amazonian forest fragments and adjacent pastures: biogeographic implications for alpha and beta diversity

The aim of this study is to analyze the effects of habitat loss and forest replacement by cattle pasture on the alpha and beta diversity, abundance, biomass and species composition of dung beetles with different dispersal ability. Dung beetles were captured in 19 forest fragments and neighbouring pastures. Forest fragment area ranged from 3.7 to 4825 ha and in this study were grouped into four categories: small, medium, large and control forest. A total of 35,048 dung beetles representing 101 species were collected. Forest fragments had the highest richness with 81 species, followed by pasture with 58 species. Replacement of forest by pasture reduced species richness; however, due to the proximity and connectivity of these areas with Cerrado patches, pastures also had high species richness, but species composition was independent of adjacent fragments. Small fragments had lower abundance and species richness than our other habitat categories, even pastures. Our results highlight that proximity and connectivity with Cerrado areas influenced the patterns of alpha and beta diversity of dung beetles in fragments and pastures. We highlight that the ability to cross the pasture matrix is a strong adaptive trait for species living in human-modified landscapes. Consequently, species with these abilities are less susceptible to the effects of forest fragmentation and local extinction. Our results reinforce the importance of considering the biogeographic location and distribution pattern of species in forest fragmentation studies.     

Forest Fragment and Breeding Habitat Characteristics Explain Frog Diversity and Abundance in Singapore

Habitat loss and fragmentation can have severe negative and irreversible effects on biodiversity. We investigated the effects of forest fragmentation on frog diversity in Singapore because of its high rates of deforestation and the demonstration that frogs are some of the most sensitive species to habitat degradation. We surveyed frog species in 12 forest fragments varying from 11 to 935 ha. We compared differences in species richness, abundance, and Shannon's index in relation to forest fragment size, connectivity (distance between fragments), and breeding habitat heterogeneity. A total of 20 species from 12 genera and five families were encountered in 12 fragments. Larger fragments and those closer to larger fragments had higher species richness. Abundance, however, was not correlated with forest area or connectivity, but we found fewer individual frogs in the larger fragments. We also found that breeding habitat heterogeneity best explained frog species diversity and abundance in forest fragments. Fragments with a high diversity of breeding habitats had more species. We found no evidence to suggest that abundance and diversity are strongly correlated, particularly in disturbed areas, but that breeding habitat heterogeneity is an under-appreciated factor that should be considered when prioritizing areas for anuran conservation. Enriching breeding habitat heterogeneity, creating corridors between fragments, and reforesting degraded areas are some of the most beneficial strategies for preserving urban frog biodiversity.     

Effects of forest fragmentation on species richness and composition of ground beetles (Coleoptera: Carabidae and Brachinidae) in urban landscapes

To clarify the effects of forest fragmentation in urban landscapes on the abundance, species richness, dominance, and species composition of ground beetles (Coleoptera: Carabidae and Brachinidae), we compared the beetles collected in 12 pitfall traps from April to July and from September to November between three continuous suburban forests and eight isolated urban forests (0.06–1.02 ha), most of which were in the precincts of shrines and temples in Hanshin District, Honshu, Japan. A total of 28 species and 4178 individuals of ground beetles were collected. Segregation of urban forests from continuous suburban forests has changed the species composition and resulted in the loss of some large‐sized forest species and the addition of some non‐forest species. Simpson's index of dominance (λ) also increased in the urban forests. The richness of forest species markedly decreased with the reduction in forest area but not with the distance from continuous forests, although the species richness of non‐forest species did not change with them. Also, species composition changed only with forest area. These findings indicate that continuous forests do not necessary serve as a “mainland” for urban forest species and that every urban habitat, however small in size, acts as a temporary reservoir of species. In comparison with populations of small‐sized species, populations of large‐sized forest species appeared to decline more readily during forest fragmentation.     

Islands,mainland, and terrestrial fragments: How isolation shapes plant diversity

The fragmentation of natural habitats is a major threat for biodiversity. However, the impact and spatial scale of natural isolation mechanisms leading to species loss, compared to anthropogenic fragmentation, are not clear, mainly due to differences between fragments and islands, such as matrix permeability. We studied a 500 km² Mediterranean region in France, including urban habitat fragments, continuous habitat, and continental‐shelf islands. On the basis of 295 floristic relevés, we built species–area relationships to compare isolation in fragments after urbanization, with continuous habitat and continental‐shelf islands. We assumed either no dispersal, infinite dispersal, or estimated intermediate levels of habitat reachability through graph theory. Isolation mechanisms occurred in fragments but with a lower strength than in near‐shore islands, and most importantly affected perennial plants. Annual plants were less affected, probably due to their smaller size and shorter life cycle. Isolation occurred at landscape level in fragments and at patch level in islands. The amount of reachable habitat (accounting for spatial configuration) explained local species richness in both systems, but the amount of habitat (no consideration of spatial configuration) was already a good predictor. These results suggest an important role of habitat amount around fragments in mitigating the isolation effects observed in near‐shore islands, and the importance of carefully considering different functional groups.     

Effects of Fragmentation and Sea-Level Changes upon Frog Communities of Land-Bridge Islands off the Southeastern Coast of Brazil

We investigate the composition of anuran communities of land-bridge islands off the southeastern coast of Brazil. These islands provide natural long-term experiments on the effects of fragmentation in the Brazilian Atlantic Forest (BAF). We hypothesize that Pleistocene sea-level changes, in combination with other abiotic variables such as area and habitat diversity, has affected anuran species richness and community composition on these islands. Data from the literature and collections databases were used to produce species lists for eight land-bridge islands and for the mainland adjacent to the islands. We assess the effects of area, number of breeding habitats and distance to the mainland upon anuran species richness on land-bridge islands. Additionally we use nestedness analysis to quantify the extent to which the species on smaller and less habitat-diverse islands correspond to subsets of those on larger and more diverse ones. We found that area has both direct and indirect effects on anuran species richness on land-bridge islands, irrespective of distance to the mainland. However, on islands with comparable sizes, differences in species richness can be attributed to the number and quality of breeding habitats. Anuran communities on these islands display a nested pattern, possibly caused by selective extinction related to habitat loss. Common lowland pond-breeders were conspicuous by their absence. In the BAF, the conservation of fragments with a high diversity of breeding habitats could compensate for the generally negative effect of small area upon species richness. We suggest that sea-level changes have an important role in shaping composition of anuran species on coastal communities.     

Effects of small‐scale grassland fragmentation and frequent mowing on population density and species diversity of orthopterans: a long‐term study

Abstract 1. Habitat fragmentation is considered one of the major threats to invertebrate diversity in semi‐natural grassland. However, the effects of habitat fragmentation through mowing on the rich insect fauna of these grasslands have not been sufficiently investigated and experiments are especially rare. 2. We studied the impact of small‐scale grassland fragmentation on orthopterans over 7 years in an experiment which allowed us to additionally investigate the effect of frequent mowing on Orthoptera communities. 3. Overall, Orthoptera density and species richness increased over time. This was likely a result of increased small‐scale habitat heterogeneity and the provision of a short‐turf habitat suitable for xerophilous species. The fragmentation affected orthopteran density and species composition but not species richness whose response lagged behind the changes in abundance. 4. Responses differed between suborders. Ensifera density was higher in fragment than in control plots. Caelifera density did not differ between fragment and control plots. The mown matrix was an unsuitable habitat for most of the species, particularly within the Ensifera. 5. Our experiment shows that even small‐scale fragmentation can affect Orthopteran communities and that the effects became more pronounced over time. As the mown matrix was unsuitable for many Ensifera species, they may go locally extinct when large areas are mown simultaneously.     

Comparing bird assemblages in large and small fragments of the Atlantic Forest hotspots

The Atlantic Forest (AF) is one of the five most threatened and megadiverse world hotspots. It is arguably the most devastated and highly threatened ecosystem on the planet. The vast scope of habitat loss and extreme fragmentation in the AF hotspots has left intact very few extensive and continuous forested fragments. We compared bird assemblages between small (<100 ha) and large (>6,000 ha) forest remnants, in one of the largest AF remnants in Argentina. We performed 84 point-counts of birds in four large fragments (LF) and 67 points in 25 small fragments (SF). We recorded 4,527 bird individuals belonging to 173 species; 2,632 belonging to 153 species in LF and 1,897 in 124 species in SF. Small fragments suffered a significant loss of bird richness, mainly forest dependent species, but the birds abundance did not decrease, due to an increase in abundance of forest independent and semi-dependent bird species (edge and non forest species) that benefit from forest fragmentation. The bird guilds of frugivores, undestory, terrestrial and midstory insectivores, nectarivores and raptors, and the endemic species of AF were area sensitive, decreasing significantly in richness and abundance in the SF. Terrestrial granivores were the only guild positively affected by forest fragmentation, containing mainly edge species, which forage in open areas or borders including crops. Our first observations on fragmentation effects on bird assemblages in the southernmost Argentinean Atlantic Forests did not validate the hypothesis on pre-adaptation to human disturbances in the bird communities of AF. On the contrary, we observed that forest dependent, endemic and several sensitive bird guilds were strongly affected by fragmentation, putting in evidence the vulnerability to the fragmentation process and the necessity to conserve large remnants to avoid reduction of the high biodiversity of AF birds.     

Habitat fragmentation,tree diversity,and plant invasion interact to structure forest caterpillar communities

Habitat fragmentation and invasive species are two of the most prominent threats to terrestrial ecosystems. Few studies have examined how these factors interact to influence the diversity of natural communities, particularly primary consumers. Here, we examined the effects of forest fragmentation and invasion of exotic honeysuckle (Lonicera maackii, Caprifoliaceae) on the abundance and diversity of the dominant forest herbivores: woody plant-feeding Lepidoptera. We systematically surveyed understory caterpillars along transects in 19 forest fragments over multiple years in southwestern Ohio and evaluated how fragment area, isolation, tree diversity, invasion by honeysuckle and interactions among these factors influence species richness, diversity and abundance. We found strong seasonal variation in caterpillar communities, which responded differently to fragmentation and invasion. Abundance and richness increased with fragment area, but these effects were mitigated by high levels of honeysuckle, tree diversity, landscape forest cover, and large recent changes in area. Honeysuckle infestation was generally associated with decreased caterpillar abundance and diversity, but these effects were strongly dependent on other fragment traits. Effects of honeysuckle on abundance were moderated when fragment area, landscape forest cover and tree diversity were high. In contrast, negative effects of honeysuckle invasion on caterpillar diversity were most pronounced in fragments with high tree diversity and large recent increases in area. Our results illustrate the complex interdependencies of habitat fragmentation, plant diversity and plant invasion in their effects on primary consumers and emphasize the need to consider these processes in concert to understand the consequences of anthropogenic habitat change for biodiversity.     

Effects of habitat fragmentation and disturbance on howler monkeys: a review

We examined the literature on the effects of habitat fragmentation and disturbance on howler monkeys (genus Alouatta) to (1) identify different threats that may affect howlers in fragmented landscapes; (2) review specific predictions developed in fragmentation theory and (3) identify the empirical evidence supporting these predictions. Although howlers are known for their ability to persist in both conserved and disturbed conditions, we found evidence that they are negatively affected by high levels of habitat loss, fragmentation and degradation. Patch size appears to be the main factor constraining populations in fragmented habitats, probably because patch size is positively related to food availability, and negatively related to anthropogenic pressures, physiological stress and parasite loads. Patch isolation is not a strong predictor of either patch occupancy or population size in howlers, a result that may be related to the ability of howlers to move among forest patches. Thus, we propose that it is probable that habitat loss has larger consistent negative effects on howler populations than habitat fragmentation per se. In general, food availability decreases with patch size, not only due to habitat loss, but also because the density of big trees, plant species richness and howlers' home range size are lower in smaller patches, where howlers' population densities are commonly higher. However, it is unclear which vegetation attributes have the biggest influence on howler populations. Similarly, our knowledge is still limited concerning the effects of postfragmentation threats (e.g. hunting and logging) on howlers living in forest patches, and how several endogenous threats (e.g. genetic diversity, physiological stress, and parasitism) affect the distribution, population structure and persistence of howlers. More long‐term studies with comparable methods are necessary to quantify some of the patterns discussed in this review, and determine through meta‐analyses whether there are significant inter‐specific differences in species' responses to habitat loss and fragmentation. Am. J. Primatol. 72:1–16, 2010. © 2009 Wiley‐Liss, Inc.     

Habitat fragmentation and species diversity in competitive communities

Habitat loss is one of the key drivers of the ongoing decline of biodiversity. However, ecologists still argue about how fragmentation of habitat (independent of habitat loss) affects species richness. The recently proposed habitat amount hypothesis posits that species richness only depends on the total amount of habitat in a local landscape. In contrast, empirical studies report contrasting patterns: some find positive and others negative effects of fragmentation per se on species richness. To explain this apparent disparity, we devise a stochastic, spatially explicit model of competitive species communities in heterogeneous habitats. The model shows that habitat loss and fragmentation have complex effects on species diversity in competitive communities. When the total amount of habitat is large, fragmentation per se tends to increase species diversity, but if the total amount of habitat is small, the situation is reversed: fragmentation per se decreases species diversity.