Habitat fragmentation and species richness

In a recent article in this journal, Fahrig (2013, Journal of Biogeography, 40 , 1649–1663) concludes that variation in species richness among sampling sites can be explained by the amount of habitat in the ‘local landscape’ around the sites, while the spatial configuration of habitat within the landscape makes little difference. This conclusion may be valid for small spatial scales and when the total amount of habitat is large, but modelling and empirical studies demonstrate adverse demographic consequences of fragmentation when there is little habitat across large areas. Fragmentation effects are best tested with studies on individual species rather than on communities, as the latter typically consist of species with dissimilar habitat requirements. The total amount of habitat and the degree of fragmentation tend to be correlated, which poses another challenge for empirical studies. I conclude that fragmentation poses an extra threat to biodiversity, in addition to the threat posed by loss of habitat area.     

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.     

Habitat fragmentation may not matter to species diversity

Conservation biologists worry that fragmenting a bloc of natural habitat might reduce its species diversity. However, they also recognize the difficulty and importance of isolating the effect of fragmentation from that of simple loss of area. Using two different methods (species-area curve and Fisher's alpha index of diversity) to analyse the species diversities of plants, tenebrionid beetles and carabid beetles in a highly fragmented Mediterranean scrub landscape, we decoupled the effect of degree of fragmentation from that of area loss. In this system, fragmentation by itself seems not to have influenced the number of species. Our results, obtained at the scale of hectares, agree with similar results at island and continent scales.     

Habitat fragmentation: island v landscape perspectives on bird conservation

Fragments of habitat are often viewed as islands and are managed as such; however, habitat fragmentation includes a wide range of spatial patterns of environments that may occur on many spatial scales. Fragments exist in a complex landscape mosaic, and dynamics within a fragment are affected by external factors that vary as the mosaic structure changes. The simple analogy of fragments to islands, therefore, is unsatisfactory. Understanding how birds respond to these complexities of fragmentation requires mechanistic studies focused on habitat selection and movement behaviour. Conservation efforts must be based on viewing fragmentation as a range of conditions that occurs in a landscape mosaic, and management should be directed toward the mosaics rather than focusing solely on reserves.     

Habitat fragmentation and amphibian conservation: building a tool for assessing landscape matrix connectivity

We have attempted modelling biological connectivity for a toad population in a floodplain in the combined framework of population dynamics and landscape ecology. We took advantage of the GIS package for establishing friction maps for the focal species by attributing to each habitat type a resistance index. This approach made it possible to draw migration zones around each pond the area and the shape of which depended on landscape structure. We included mortality from road traffic by assigning a virtual population to each pond. The resulting map shows the potentiality of such an approach in the assistance of landscape management.     

Habitat fragmentation differentially shapes neutral and immune gene variation in a tropical bird species

Habitat fragmentation is a major cause of biodiversity loss, responsible for an alteration of intraspecific patterns of neutral genetic diversity and structure. Although neutral genetic variation can be informative for demographic inferences, it may be a poor predictor of adaptive genetic diversity and thus of the consequences of habitat fragmentation on selective evolutionary processes. In this context, we contrasted patterns of genetic diversity and structure of neutral loci (microsatellites) and immune genes (i.e., toll-like receptors) in an understorey bird species, the wedge-billed woodcreeper Glyphorynchus spirurus. The objectives were (1) to investigate forest fragmentation effects on population genetic diversity, (2) to disentangle the relative role of demography (genetic drift and migration) and selection, and (3) to assess whether immunogenetic patterns could be associated with variation of ectoparasite (i.e., ticks) pressures. Our results revealed an erosion of neutral genetic diversity and a substantial genetic differentiation among fragmented populations, resulting from a decrease in landscape connectivity and leading to the divergence of distinct genetic pools at a small spatial scale. Patterns of genetic diversity observed for TLR4 and TLR5 were concordant with neutral genetic patterns, whereas those observed for TLR3 and TLR21 were discordant. This result underlines that the dominant evolutionary force shaping immunogenetic diversity (genetic drift vs. selection) may be different depending on loci considered. Finally, tick prevalence was higher in fragmented environments. We discussed the hypothesis that pathogen selective pressures may contribute to maintain adaptive genetic diversity despite the negative demographic effect of habitat fragmentation on neutral genetic diversity.Subject terms: Tropical ecology, Genetic variation     

Neuroecology, chemical defense, and the keystone species concept

Neuroecology unifies principles from diverse disciplines, scaling from biophysical properties of nerve and muscle cells to community-wide impacts of trophic interactions. Here, these principles are used as a common fabric, woven from threads of chemosensory physiology, behavior, and population and community ecology. The "keystone species" concept, for example, is seminal in ecological theory. It defines a species whose impacts on communities are far greater than would be predicted from its relative abundance and biomass. Similarly, neurotoxins could function in keystone roles. They are rare within natural habitats but exert strong effects on species interactions at multiple trophic levels. Effects of two guanidine alkaloids, tetrodotoxin (TTX) and saxitoxin (STX), coalesce neurobiological and ecological perspectives. These molecules compose some of the most potent natural poisons ever described, and they are introduced into communities by one, or only a few, host species. Functioning as voltage-gated sodium channel blockers for nerve and muscle cells, TTX and STX serve in chemical defense. When borrowed by resistant consumer species, however, they are used either in chemical defense against higher order predators or for chemical communication as chemosensory excitants. Cascading effects of the compounds profoundly impact community-wide attributes, including species compositions and rates of material exchange. Thus, a diverse array of physiological traits, expressed differentially across many species, renders TTX and STX fully functional as keystone molecules, with vast ecological consequences at multiple trophic levels.     

Habitat fragmentation and the prevalence of parasites (Diptera,Streblidae) on three Phyllostomid bat species

Ectoparasitism in bats seems to be influenced strongly by the type of roost preferred by the hosts, and group size; however, the effect of habitat loss and fragmentation on the prevalence of ectoparasites in bats has scarcely been studied. In northeastern Yucatan, Mexico, we estimated the prevalence of infestation by Streblidae flies in three phyllostomid bat species with different roost preferences (caves, trees, or both) in two types of landscape matrices (tropical semi‐deciduous forest and man‐made pastures) that differed in area of forest cover and the number of forest fragments. Habitat fragmentation and the presence of a contrasting matrix may limit the availability of roosts (trees) and the movement of bats across the landscape. Accordingly, we hypothesized higher prevalence of Streblidae infestation in the pasture matrix and in the group of bats that roost in trees. Bat abundance was higher in the pasture matrix; however, the prevalence of infestation was significantly higher in the continuous forest matrix and in bats that roosted in caves. The prevalence of some species of Streblidae was affected by habitat fragmentation in species that roost in caves, such as Desmodus rotundus, as well as those using foliage and caves, such as Artibeus jamaicensis. Our results provide evidence that some species of Streblidae may respond differently to habitat fragmentation than their hosts, generating changes to bat‐ectoparasite interactions in fragmented areas. Environmental variations involving roosts, not evaluated in this study, may influence our results, since these factors affect ectoparasite abundance and reproduction.     

Disease,habitat fragmentation and conservation

Habitat loss and the resultant fragmentation of remaining habitat is the primary cause of loss of biological diversity. How do these processes affect the dynamics of parasites and pathogens? Hess has provided some important insights into this problem using metapopulation models for pathogens that exhibit ''S-I'' dynamics; for example, pathogens such as rabies in which the host population may be divided into susceptible and infected individuals. A major assumption of Hess''s models is that infected patches become extinct, rather than recovering and becoming resistant to future infections. In this paper, we build upon this framework in two different ways: first, we examine the consequences of including patches that are resistant to infection; second, we examine the consequences of including a second species of host that can act as a reservoir for the pathogen. Both of these effects are likely to be important from a conservation perspective. The results of both sets of analysis indicate that the benefits of corridors and other connections that allow species to disperse through the landscape far outweigh the possible risks of increased pathogen transmission. Even in the commonest case, where harmful pathogens are maintained by a common reservoir host, increased landscape connectance still allows greater coexistence and persistence of a threatened or endangered host.     

Habitat requirements of endangered species in a former coppice of high conservation value

Transformation of coppices to high forests has caused fundamental changes in site conditions and a decline of many species across Central Europe. Nevertheless, some formerly coppiced forests still harbour a number of the declining species and have become biodiversity hotspots in the changing landscape. We focused on the best-preserved remnant of formerly grazed and coppiced subcontinental oak forest in the Czech Republic – the Dúbrava forest near the town of Hodonín. To improve our understanding of the ecology of declining species, we studied local habitat requirements of vascular plants most endangered at the national level. We recorded vegetation composition and sampled important site variables in plots with the largest populations of endangered species and in additional plots placed randomly across all major forest habitats. We demonstrated that sites with endangered species have a highly uneven distribution in ecological space and that their species composition is often similar to open-canopy oak forests. Within this habitat, the endangered species are concentrated in places with a high light availability and high soil pH. Light-demanding species characteristic of subcontinental oak forests are the best indicators of these sites, while broadly distributed shade-tolerant and nutrient-demanding species avoid them. These results support the view that the occurrence of many endangered species in the Dúbrava forest is a legacy of the long history of traditional management that kept the canopies open. Light-demanding species are now threatened by ongoing successional changes. Therefore, active conservation measures are recommended, including opening up the canopies, early thinning of young stands, control of expansive and invasive species and understorey grazing or mowing.     

栖息地破碎化与鸟类生存

栖息地破碎化给野生动物带来的不良后果是全球生态学家和保护生物学家共同关心的问题.自从提出栖息地破碎化是导致生物多样性丧失的关键因素之一的论点后,近20年来,栖息地破碎化研究一直是生态学和保护生物学最活跃的前沿研究领域之一.栖息地破碎化是一动态过程,可在多尺度上发生并蕴涵着复杂的空间模式变化.栖息地破碎化对鸟类的生态学效应主要体现在面积效应、隔离效应和边缘效应等.这些效应影响着鸟类的分布、基因交流、种群动态、扩散行为、种间关系和生活史特征等,最终影响着鸟类的生存.介绍和总结了栖息地破碎化过程、研究的理论依据及栖息地破碎化对鸟类生存产生的诸多影响.     

Habitat fragmentation, variable edge effects, and the landscape-divergence hypothesis

Edge effects are major drivers of change in many fragmented landscapes, but are often highly variable in space and time. Here we assess variability in edge effects altering Amazon forest dynamics, plant community composition, invading species, and carbon storage, in the world's largest and longest-running experimental study of habitat fragmentation. Despite detailed knowledge of local landscape conditions, spatial variability in edge effects was only partially foreseeable: relatively predictable effects were caused by the differing proximity of plots to forest edge and varying matrix vegetation, but windstorms generated much random variability. Temporal variability in edge phenomena was also only partially predictable: forest dynamics varied somewhat with fragment age, but also fluctuated markedly over time, evidently because of sporadic droughts and windstorms. Given the acute sensitivity of habitat fragments to local landscape and weather dynamics, we predict that fragments within the same landscape will tend to converge in species composition, whereas those in different landscapes will diverge in composition. This 'landscape-divergence hypothesis', if generally valid, will have key implications for biodiversity-conservation strategies and for understanding the dynamics of fragmented ecosystems.     

Sex allocation theory aids species conservation

Supplementary feeding is often a key tool in the intensive management of captive and threatened species. Although it can increase such parameters as breeding frequency and individual survival, supplementary feeding may produce undesirable side effects that increase overall extinction risk. Recent attempts to increase breeding frequency and success in the kakapo Strigops habroptilus using supplementary feeding inadvertently resulted in highly male-biased chick sex ratios. Here, we describe how the inclusion of sex allocation theory has remedied this conservation dilemma. Our study is the first to manipulate chick sex ratios in an endangered species by altering maternal condition and highlights the importance of incorporating evolutionary theory into modern conservation practice.     

Invasive pythons,not anthropogenic stressors,explain the distribution of a keystone species

Untangling the causes of native species loss in human-modified systems is difficult and often controversial. Evaluating the impact of non-native species in these systems is particularly challenging, as additional human perturbations often precede or accompany introductions. One example is the ongoing debate over whether mammal declines within Everglades National Park (ENP) were caused by either the establishment of non-native Burmese pythons (Python molurus bivittatus) or the effects of other anthropogenic stressors. We examined the influence of both pythons and a host of alternative stressors—altered hydrology and habitat characteristics, mercury contamination and development—on the distribution of the marsh rabbit (Sylvilagus palustris), a once common mammal in ENP. Distance from the epicenter of the python invasion best explained marsh rabbit occurrence in suitable habitat patches, whereas none of the alternative stressors considered could explain marsh rabbit distribution. Estimates of the probability of marsh rabbit occurrence ranged from 0 at the python invasion epicenter to nearly 1.0 150 km from the invasion epicenter. These results support the hypothesis that invasive pythons shape the distribution of marsh rabbits in southern Florida. The loss of marsh rabbits and similar species will likely alter trophic interactions and ecosystem function within the Everglades, an internationally important hotspot of biodiversity. Further, our results suggest that non-native species can have profound impacts on mainland biodiversity.     

Habitat fragmentation disrupts the demography of a widespread native mammal

Fragmentation theory predicts that population persistence should be positively correlated with the size of habitat fragments. The patterns of occurrence of many species are consistent with this prediction, but the demographic processes that determine how species respond to fragmentation are poorly understood. In addition, habitat quality may interact with fragment size as an influence on demographic performance. We investigated these predictions for the native bush rat Rattus fuscipes by testing the following hypotheses: 1) population performance (i.e. viability as determined by various demographic parameters) is positively correlated with fragment size; and 2) population performance is positively correlated with habitat quality. Populations of R. fuscipes were censused in two large (>49 ha) and eight small (<2.5 ha) forest fragments in an agricultural region of southeastern Australia. Fragments with high and low quality habitat were included in each size category. Fragment size influenced multiple aspects of population demography; populations in large fragments had higher densities, older age structures, received more potential immigrants, and were more likely to recruit adults than those in small fragments. Reproductive patterns were more predictable in large fragments. Habitat quality per se had less marked effects; adult females were heavier and subadults more prevalent in fragments with high quality habitat. However, high quality habitat enhanced population performance in small fragments more so than in large ones. Despite being widespread in the study area, R. fuscipes populations are profoundly impacted by habitat fragmentation, with population performance declining with fragment size. Studies based on patterns of species occurrence should be interpreted with caution as they may mask critical processes occurring at the population level. For a thorough understanding of the effects of habitat fragmentation, population‐level studies are required.     

The Habitat Amount Hypothesis implies negative effects of habitat fragmentation on species richness

The habitat amount hypothesis (HAH) predicts that species richness in a habitat site increases with the amount of habitat in the ‘local landscape’ defined by an appropriate distance around the site, with no distinct effects of the size of the habitat patch in which the site is located. It has been stated that a consequence of the HAH, if supported, would be that it is unnecessary to consider habitat configuration to predict or manage biodiversity patterns, and that conservation strategies should focus on habitat amount regardless of fragmentation. Here, I assume that the HAH holds and apply the HAH predictions to all habitat sites over entire landscapes that have the same amount of habitat but differ in habitat configuration. By doing so, I show that the HAH actually implies clearly negative effects of habitat fragmentation, and of other spatial configuration changes, on species richness in all or many of the habitat sites in the landscape, and that these habitat configuration effects are distinct from those of habitat amount in the landscape. I further show that, contrary to current interpretations, the HAH is compatible with a steeper slope of the species–area relationship for fragmented than for continuous habitat, and with higher species richness for a single large patch than for several small patches with the same total area (SLOSS). This suggests the need to revise the ways in which the HAH has been interpreted and can be actually tested. The misinterpretation of the HAH has arisen from confounding and overlooking the differences in the spatial scales involved: the individual habitat site at which the HAH gives predictions, the local landscape around an individual site and the landscapes or regions (with multiple habitat sites and different local landscapes) that need to be analysed and managed. The HAH has been erroneously viewed as negating or diminishing the relevance of fragmentation effects, while it actually supports the importance of habitat configuration for biodiversity. I conclude that, even in the cases where the HAH holds, habitat fragmentation and configuration are important for understanding and managing species distributions in the landscape.     

Habitat fragmentation and population extinction of birds

It has not been established that a major cause of extinction in birds or any other taxa is failure of metapopulation dynamics: the collapse of a network of ephemeral but discrete populations as movement between them becomes increasingly infrequent. The few data on who goes where and who mates with whom suggest that most species are structured as either a single large population or a small set of source populations and a larger set of sinks. The extinction of the latter is irrelevant to the persistence of the species. However, regional decline of a species in the face of habitat destruction and fragmentation can mimic a failure of metapopulation dynamics, because distinct aggregations of individuals will disappear much as they would if populations in an interacting network were eliminated one by one. Any species with highly restricted range is at great risk of extinction from spatially localized forces, such as cyclones or deforestation. Restricted range rather than inherent weakness is the main reason that so many island species have gone extinct or are endangered. Species with small populations in contact with much larger heterospecific ones with which they are interfertile are threatened with extinction by hybridization. Finally, the disappearance of a species from a site may be due to subtle habitat change, even if this observation seems superficially consistent with some general population theory, such as the dynamic equilibrium theory of island biogeography. Current theory is an inadequate substitute for intensive field studies as a means to address the conservation problems of individual species.