Synergisms among Habitat Fragmentation, Livestock Grazing, and Biotic Invasions in Southwestern Australia

Abstract: The agricultural development of southern Australia over the past 200 years has resulted in extensively fragmented systems, often with only small, isolated remnants of native vegetation remaining. Grazing by sheep and cattle has affected both the remaining fragments and the surrounding matrix, and non-native plant and animal species have had dramatic effects on the native biota. Invasive plant species have the potential to significantly alter ecosystem composition and functioning, and invasive animals, particularly rabbits ( Oryctolagus cuniculatus ), foxes (    Vulpes vulpes ) and cats (    Felis catus ) effectively alter habitat and drive native fauna to local extinction. These different influences often interact. For instance, smaller fragments are often more prone to plant invasion and are more likely to have been grazed in the past. Invasion of plant species is often linked with livestock grazing or rabbit invasion, and other higher-order interactions are also apparent. Classical fragmentation studies that concentrate on parameters such as habitat area and isolation but ignore changes in habitat condition brought about by livestock and invasive species are unlikely to yield meaningful results. Similarly, management of fragmented ecosystems must account for not only the spatial characteristics of the remaining habitat but also the importance of other influences, particularly those that impinge on fragments from the surrounding matrix.     

Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California

Abstract:   Livestock grazing represents a major human alteration of natural disturbance regimes in grasslands throughout the world, and its impacts on plant communities have been highly debated. We investigated the impact of cattle grazing on the California coastal prairie plant community with a focus on native annual forbs, a number of which are of conservation concern. In spring 2000 and 2001, we surveyed the vegetation community composition, vegetation structure, and soil chemical parameters at 25 paired grazed and ungrazed sites over a 670-km range of the ecosystem. Native annual forb species richness and cover were higher in grazed sites, and this effect was concomitant with decreased vegetation height and litter depth. Soil properties explained less of the variation. Exotic annual grass and forb cover were higher in grazed sites. Native grass cover and species richness did not differ in grazed and ungrazed sites, but cover and species richness of native perennial forbs were higher in ungrazed sites. Our results suggest that cattle grazing may be a valuable management tool with which to conserve native annual forbs in the ecosystem we studied but that grazing differentially affects the various life-history guilds. Therefore, land managers must focus on creating a matrix of disturbance regimes to maintain the suite of species native to these mesic grasslands. The results of this and other studies highlight the importance of considering the adaptation of vegetation communities to disturbance in making recommendations for grazing management.     

Meta-analysis of the differential effects of habitat fragmentation and degradation on plant genetic diversity

Genetic diversity is a key factor for population survival and evolution. However, anthropogenic habitat disturbance can erode it, making populations more prone to extinction. Aiming to assess the global effects of habitat disturbance on plant genetic variation, we conducted a meta-analysis based on 92 case studies obtained from published literature. We compared the effects of habitat fragmentation and degradation on plant allelic richness and gene diversity (equivalent to expected heterozygosity) and tested whether such changes are sensitive to different life-forms, life spans, mating systems, and commonness. Anthropogenic disturbance had a negative effect on allelic richness, but not on gene diversity. Habitat fragmentation had a negative effect on genetic variation, whereas habitat degradation had no effect. When we examined the individual effects in fragmented habitats, allelic richness and gene diversity decreased, but this decrease was strongly dependent on certain plant traits. Specifically, common long-lived trees and self-incompatible species were more susceptible to allelic richness loss. Conversely, gene diversity decreased in common short-lived species (herbs) with self-compatible reproduction. In a wider geographical context, tropical plant communities were more sensitive to allelic richness loss, whereas temperate plant communities were more sensitive to gene diversity loss. Our synthesis showed complex responses to habitat disturbance among plant species. In many cases, the absence of effects could be the result of the time elapsed since the disturbance event or reproductive systems favoring self-pollination, but attention must be paid to those plant species that are more susceptible to losing genetic diversity, and appropriate conservation should be actions taken.     

Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

Metacommunity organization of soil microorganisms depends on habitat defined by presence of Lobelia siphilitica plants

We tested regional-scale spatial patterns in soil microbial community composition for agreement with species sorting and dispersal limitation, two alternative mechanisms behind different models of metacommunity organization. Furthermore, we tested whether regional metacommunity organization depends on local habitat type. We sampled from sites across Ohio and West Virginia hosting populations of Lobelia siphilitica, and compared the metacommunity organization of soil microbial communities under L. siphilitica to those in adjacent areas at each site. In the absence of L. siphilitica, bacterial community composition across the region was consistent with species sorting. However, under L. siphilitica, bacterial community composition was consistent with dispersal limitation. Fungal community composition remained largely unexplained, although fungal communities under L. siphilitica were both significantly different in composition and less variable in composition than in adjacent areas. Our results show that communities in different local habitat types (e.g., in the presence or absence of a particular plant) may be structured on a regional scale by different processes, despite being separated by only centimeters at the local scale.     

Butterfly community shifts over two centuries

Environmental changes strongly impact the distribution of species and subsequently the composition of species assemblages. Although most community ecology studies represent temporal snap shots, long‐term observations are rather rare. However, only such time series allow the identification of species composition shifts over several decades or even centuries. We analyzed changes in the species composition of a southeastern German butterfly and burnet moth community over nearly 2 centuries (1840–2013). We classified all species observed over this period according to their ecological tolerance, thereby assessing their degree of habitat specialisation. This classification was based on traits of the butterfly and burnet moth species and on their larval host plants. We collected data on temperature and precipitation for our study area over the same period. The number of species declined substantially from 1840 (117 species) to 2013 (71 species). The proportion of habitat specialists decreased, and most of these are currently endangered. In contrast, the proportion of habitat generalists increased. Species with restricted dispersal behavior and species in need of areas poor in soil nutrients had severe losses. Furthermore, our data indicated a decrease in species composition similarity between different decades over time. These data on species composition changes and the general trends of modifications may reflect effects from climate change and atmospheric nitrogen loads, as indicated by the ecological characteristics of host plant species and local changes in habitat configuration with increasing fragmentation. Our observation of major declines over time of currently threatened and protected species shows the importance of efficient conservation strategies.     

Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants

Abstract:  Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However, even small habitat patches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of the focal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.     

Interacting Effects of Landownership, Land Use, and Endangered Species on Conservation of Southwestern U.S. Rangelands

Abstract:  The contemporary southwestern United States is characterized by fire-adapted ecosystems; large numbers of federally listed threatened and endangered species; a patchwork of federal, state, and private landownership; and a long history of livestock grazing as the predominant land use. I compared eight sites in southern Arizona and New Mexico to assess the interacting effects of these characteristics on conservation practices and outcomes. There was widespread interest and private-sector leadership in restoring fire to southwestern rangelands, and there is a shortage of predictive scientific knowledge about the effects of fire and livestock grazing on threatened and endangered species. It was easier to restore fire to lands that were either privately owned or not grazed, in part because of obstacles created by threatened and endangered species on grazed public lands. Collaborative management facilitated conservation practices and outcomes, and periodic removal of livestock may be necessary for conservation, but permanent livestock exclusion may be counterproductive because of interactions with land-use and landownership patterns.     

Leaf quality, predators, and stochastic processes in the assembly of a diverse herbivore community

Ecological communities are structured by both deterministic, niche-based processes and stochastic processes such as dispersal. A pressing issue in ecology is to determine when and for which organisms each of these types of processes is important in community assembly. The roles of deterministic and stochastic processes have been studied for a variety of communities, but very few researchers have addressed their contribution to insect herbivore community structure. Insect herbivore niches are often described as largely shaped by the antagonistic pressures of predation and host plant defenses. However host plants are frequently discrete patches of habitat, and their spatial arrangement can affect herbivore dispersal patterns. We studied the roles of predation, host plant quality, and host spatial proximity for the assembly of a diverse insect herbivore community on Quercus alba (white oak) across two growing seasons. We examined abundances of feeding guilds to determine if ecologically similar species responded similarly to variation in niches. Most guilds responded similarly to leaf quality, preferring high-nitrogen, low-tannin host plants, particularly late in the growing season, while bird predation had little impact on herbivore abundance. The communities on the high-quality plants tended to be larger and, in some cases, have greater species richness. We analyzed community composition by correlating indices of community similarity with predator presence, leaf quality similarity, and host plant proximity. Birds did not affect community composition. Community similarity was significantly associated with distance between host plants and uncorrelated with leaf quality similarity. Thus although leaf quality significantly affected the total abundance of herbivores on a host plant, in some cases leading to increased species richness, dispersal limitation may weaken this relationship. The species composition of these communities may be driven by stochastic processes rather than variation in host plant characteristics or differential predation by insectivorous birds.     

Short-term and long-term facilitation of goose grazing by livestock in the Dutch Wadden Sea area

We studied the impact of livestock grazing on the distribution ofBranta bernicla bernicla (Dark-bellied Brent goose) in the Dutch Wadden Sea during spring. It was hypothesized that livestock facilitate short-term (within-season) grazing for geese as well as long-term (over years). Therefore we measured grazing pressure by geese in salt marsh and polder areas that were either grazed (spring-grazed) or ungrazed during spring (summer-grazed). Additionally, we carried out a preference experiment with captive geese to test the preference between spring-grazed and summer-grazed polder swards. We furthermore compared patterns of use by geese between long-term ungrazed and grazed salt marshes. In May, there is a difference in grazing pressure by geese between polder pastures that are grazed or ungrazed during spring. In this month, the ungrazed polder pastures are abandoned and the geese shift to either the grazed polder pastures or to the salt marsh. Vegetation in the polder that had been spring-grazed had a lower canopy height and a higher tiller density than summer-grazed vegetation. The captive geese in the preference experiment showed a clear preference for vegetation that had been spring-grazed by sheep over ungrazed vegetation. Goose grazing pressure was negatively correlated to canopy height, both on the polder and on the salt marsh. Within the plant communities dominated byFestuca rubra andPuccinellia maritima, marshes that were intensively grazed by livestock generally had higher grazing pressure by geese than long-term ungrazed or lightly grazed salt marshes.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species.     

Interactive effects of rabbit grazing and environmental factors on plant species-richness on dunes of Norderney

On the German island Norderney rabbit grazing effects on vegetation, soil and microclimate were investigated with exclosures in different dune vegetation. Differences in species composition between grazed and ungrazed plots increased with grazing activity. A decline in growth height resulted in higher light availability and temperature on the grazed sites. One of the most visible effects was the decline in flowering plants. Most grazing effects varied between habitat types. Grey dunes were distinctly affected by rabbit grazing. In relation to the habitat type, life forms were differently affected: in closed vegetation, rabbit activities further open patches, resulting in more therophytes. In contrast, in open vegetation rabbit activities were too strong and following sand dynamics too high for many therophytes. In general, grazing diversified habitats, slightly supported the total species-richness, and enhanced the heterogeneity of species composition. Highest species-richness in total and of herbs was reached at intermediate levels of grazing pressure. The study is in line that highest species-diversity will be found at intermediate levels of disturbance and environmental stress and that grazing effects vary with the habitat type.     

Global Warming, Human Population Pressure, and Viability of the World's Smallest Butterfly

Abstract:  The effects of climate change and habitat destruction and their interaction are likely to be the greatest challenge to animal and plant conservation in the twenty-first century. We used the world's smallest butterfly, the Sinai baton blue ( Pseudophilotes sinaicus ), as an exemplar of how global warming and human population pressures may act together to cause species extinctions. We mapped the entire global range of this butterfly and obtained extensive data on the intensity of livestock grazing. As with an increasing number of species, it is confined to a network of small habitat patches and is threatened both by indirect human-induced factors (global warming) and by the direct activities of humans (in this case, livestock grazing and collection of medicinal plants). In the absence of global warming, grazing, and plant collection, our model suggested that the butterfly will persist for at least 200 years. Above a threshold intensity of global warming, the chance of extinction accelerated rapidly, implying that there may be an annual average temperature, specific to each endangered species, above which extinction becomes very much more likely. By contrast, there was no such threshold of grazing pressure—the chance of extinction increased steadily with increasing grazing. The impact of grazing, however, decreased with higher levels of year-to-year variation in habitat quality. The effect of global warming did not depend on the future level of grazing, suggesting that the impacts of global warming and grazing are additive. If the areas of habitat patches individually fall below certain prescribed levels, the butterfly is likely to go extinct. Two patches were very important for persistence: if either were lost the species would probably go extinct. Our results have implications for the conservation management of all species whose habitats are at risk because of the direct activities of humans and in the longer term because of climate change.     

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.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

Dispersal depression with habitat fragmentation in the bog fritillary butterfly

Habitat fragmentation is expected to impose strong selective pressures on dispersal rates. However, evolutionary responses of dispersal are not self-evident, since various selection pressures act in opposite directions. Here we disentangled the components of dispersal behavior in a metapopulation context using the Virtual Migration model, and we linked their variation to habitat fragmentation in the specialist butterfly Proclossiana eunomia. Our study provided a nearly unique opportunity to study how habitat fragmentation modifies dispersal at the landscape scale, as opposed to microlandscapes or simulation studies. Indeed, we studied the same species in four landscapes with various habitat fragmentation levels, in which large amounts of field data were collected and analyzed using similar methodologies. We showed the existence of quantitative variations in dispersal behavior correlated with increased fragmentation. Dispersal propensity from habitat patches (for a given patch size), and mortality during dispersal (for a given patch connectivity) were lower in more fragmented landscapes. We suggest that these were the consequences of two different evolutionary responses of dispersal behavior at the individual level: (1) when fragmentation increased, the reluctance of individuals to cross habitat patch boundaries also increased; (2) when individuals dispersed, they flew straighter in the matrix, which is the best strategy to improve dispersal success. Such evolutionary responses could generate complex nonlinear patterns of dispersal changes at the metapopulation level according to habitat fragmentation. Due to the small size and increased isolation of habitat patches in fragmented landscapes, overall emigration rate and mortality during dispersal remained high. As a consequence, successful dispersal at the metapopulation scale remained limited. Therefore, to what extent the selection of individuals with a lower dispersal propensity and a higher survival during dispersal is able to limit detrimental effects of habitat fragmentation on dispersal success is unknown, and any conclusion that metapopulations would compensate for them is flawed.     

Effects of Habitat Fragmentation on Effective Dispersal of Florida Scrub‐Jays

Abstract: Studies comparing dispersal in fragmented versus unfragmented landscapes show that habitat fragmentation alters the dispersal behavior of many species. We used two complementary approaches to explore Florida Scrub‐Jay (Aphelocoma c?rulescens) dispersal in relation to landscape fragmentation. First, we compared dispersal distances of color‐marked individuals in intensively monitored continuous and fragmented landscapes. Second, we estimated effective dispersal relative to the degree of fragmentation (as inferred from two landscape indexes: proportion of study site covered with Florida Scrub‐Jay habitat and mean distance to nearest habitat patch within each study site) by comparing genetic isolation‐by‐distance regressions among 13 study sites having a range of landscape structures. Among color‐banded individuals, dispersal distances were greater in fragmented versus continuous landscapes, a result consistent with other studies. Nevertheless, genetic analyses revealed that effective dispersal decreases as the proportion of habitat in the landscape decreases. These results suggest that although individual Florida Scrub‐Jays may disperse farther as fragmentation increases, those that do so are less successful as breeders than those that disperse short distances. Our study highlights the importance of combining observational data with genetic inferences when evaluating the complex biological and life‐history implications of dispersal.     

Value of Semi-Open Corridors for Simultaneously Connecting Open and Wooded Habitats: a Case Study with Ground Beetles

Abstract:  To counteract habitat fragmentation, the connectivity of a landscape should be enhanced. Corridors are thought to facilitate movement between disconnected patches of habitat, and linear strips of habitat connecting isolated patches are a popular type of corridor. On the other hand, the creation of new corridors can lead to fragmentation of the surrounding habitat. For example, heathland corridors connect patches of heathland and alternatively hedgerows connect patches of woodland. Nevertheless, these corridors themselves also break up previously connected patches of their surrounding habitat and in so doing fragment another type of habitat (heathland corridors fragment woodlands and woodland strips or hedgerows fragment heathlands). To overcome this challenge we propose the use of semi-open habitats (a mixture of heathland and woodland vegetation) as conservation corridors to enable dispersal of both stenotopic heathland and woodland species. We used two semi-open corridors with a mosaic of heathland and woody vegetation to investigate the efficiency of semi-open corridors for species dispersal and to assess whether these corridors might be a suitable approach for nature conservation. We conducted a mark-recapture study on three stenotopic flightless carabid beetles of heathlands and woodlands and took an inventory of all the carabid species in two semi-open corridors. Both methodological approaches showed simultaneous immigration of woodland and heathland species in the semi-open corridor. Detrended correspondence analysis showed a clear separation of the given habitats and affirmed that semi-open corridors are a good strategy for connecting woodlands and heathlands. The best means of creating and preserving semi-open corridors is probably through extensive grazing .     

Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20%, 22% and 50%, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53%, 38% and 17%. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characteized, byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition.     

Responses of Seed‐Dispersing Birds to Amount of Rainforest in the Landscape around Fragments

Habitat loss and fragmentation alter the composition of bird assemblages in rainforest. Because birds are major seed dispersers in rainforests, fragmentation‐induced changes to frugivorous bird assemblages are also likely to alter the ecological processes of seed dispersal and forest regeneration, but the specific nature of these changes is poorly understood. We assessed the influence of fragment size and landscape forest cover on the abundance, species composition, and functional properties of the avian seed disperser community in an extensively cleared, former rainforest landscape of subtropical Australia. Bird surveys of fixed time and area in 25 rainforest fragments (1–139 ha in size across a 1800 km² region) provided bird assemblage data which were coupled with prior knowledge of bird species’ particular roles in seed dispersal to give measurements of seven different attributes of the seed disperser assemblage. We used multimodel regression to assess how patch size and surrounding forest cover (within 200 m, 1000 m, and 5000 m radii) influenced variation in the abundance of individual bird species and of functional groups based on bird species’ responses to fragmentation and their roles in seed dispersal. Surrounding forest cover, specifically rainforest cover, generally had a greater effect on frugivorous bird assemblages than fragment size. Amount of rainforest cover within 200 m of fragments was the main factor positively associated with abundances of frugivorous birds that are both fragmentation sensitive and important seed dispersers. Our results suggest a high proportion of local rainforest cover is required for the persistence of seed‐dispersing birds and the maintenance of seed dispersal processes. Thus, even small rainforest fragments can function as important parts of habitat networks for seed‐dispersing birds, whether or not they are physically connected by vegetation. Respuestas de Aves Dispersoras de Semillas al Incremento de Selvas en el Paisaje Alrededor de Fragmentos