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.

Habitat fragmentation and landscape change

This chapter begins by summarizing the conceptual approaches used to understand conservation in fragmented landscapes. We then examine the biophysical aspects of landscape change, and how such change affects species and communities, posing two main questions: (i) what are the implications for the patterns of occurrence of species and communities?; and (ii) how does landscape change affect processes that influence the distribution and viability of species and communities. The chapter concludes by identifying the kinds of actions that will enhance the conservation of biota in fragmented landscapes.

Integrating edge effects into studies of habitat fragmentation: a test using meiofauna in seagrass

Habitat fragmentation is thought to be an important process structuring landscapes in marine and estuarine environments, but effects on fauna are poorly understood, in part because of a focus on patchiness rather than fragmentation. Furthermore, despite concomitant increases in perimeter:area ratios with fragmentation, we have little understanding of how fauna change from patch edges to interiors during fragmentation. Densities of meiofauna were measured at different distances across the edges of four artificial seagrass treatments [continuous, fragmented, procedural control (to control for disturbance by fragmenting then restoring experimental plots), and patchy] 1 day, 1 week and 1 month after fragmentation. Experimental plots were established 1 week prior to fragmentation/disturbance. Samples were numerically dominated by harpacticoid copepods, densities of which were greater at the edge than 0.5 m into patches for continuous, procedural control and patchy treatments; densities were similar between the edge and 0.5 m in fragmented patches. For taxa that demonstrated edge effects, densities exhibited log-linear declines to 0.5 m into a patch with no differences observed between 0.5 m and 1 m into continuous treatments. In patchy treatments densities were similar at the internal and external edges for many taxa. The strong positive edge effect (higher densities at edge than interior) for taxa such as harpacticoid copepods implies some benefit of patchy landscapes. But the lack of edge effects during patch fragmentation itself demonstrates the importance of the mechanisms by which habitats become patchy.

Species- and sex-specific connectivity effects of habitat fragmentation in a suite of woodland birds

Loss of functional connectivity following habitat loss and fragmentation could drive species declines. A comprehensive understanding of fragmentation effects on functional connectivity of an ecological assemblage requires investigation of multiple species with different mobilities, at different spatial scales, for each sex, and in different landscapes. Based on published data on mobility and ecological responses to fragmentation of 10 woodland-dependent birds, and using simulation studies, we predicted that (1) fragmentation would impede dispersal and gene flow of eight "decliners" (species that disappear from suitable patches when landscape-level tree cover falls below species-specific thresholds), but not of two "tolerant" species (whose occurrence in suitable habitat patches is independent of landscape tree cover); and that fragmentation effects would be stronger (2) in the least mobile species, (3) in the more philopatric sex, and (4) in the more fragmented region. We tested these predictions by evaluating spatially explicit isolation-by-landscape-resistance models of gene flow in fragmented landscapes across a 50 x 170 km study area in central Victoria, Australia, using individual and population genetic distances. To account for sex-biased dispersal and potential scale- and configuration-specific effects, we fitted models specific to sex and geographic zones. As predicted, four of the least mobile decliners showed evidence of reduced genetic connectivity. The responses were strongly sex specific, but in opposite directions in the two most sedentary species. Both tolerant species and (unexpectedly) four of the more mobile decliners showed no reduction in gene flow. This is unlikely to be due to time lags because more mobile species develop genetic signatures of fragmentation faster than do less mobile ones. Weaker genetic effects were observed in the geographic zone with more aggregated vegetation, consistent with gene flow being unimpeded by landscape structure. Our results indicate that for all but the most sedentary species in our system, the movement of the more dispersive sex (females in most cases) maintains overall genetic connectivity across fragmented landscapes in the study area, despite some small-scale effects on the more philopatric sex for some species. Nevertheless, to improve population viability for the less mobile bird species, structural landscape connectivity must be increased.

Short- and long-term effects of habitat fragmentation differ but are predicted by response to the matrix

Habitat loss and fragmentation are major threats to biodiversity and ecosystem processes. Our current understanding of the impacts of habitat loss and fragmentation is based largely on studies that focus on either short-term or long-term responses. Short-term responses are often used to predict long-term responses and make management decisions. The lack of studies comparing short- and long-term responses to fragmentation means we do not adequately understand when and how well short-term responses can be extrapolated to predict long-term responses, and when or why they cannot. To address this gap, we used data from one of the world's longest-running fragmentation experiments, The Wog Wog Habitat Fragmentation Experiment. Using data for carabid beetles, we found that responses in the long term (more than 22 years post-fragmentation ~ 22 generations) often contrasted markedly with those in the short term (five years post-fragmentation). The total abundance of all carabids, species richness and the occurrence of six species declined in the short term in the fragments but increased over the long term. The occurrence of three species declined initially and continued to decline, whilst another species was positively affected initially but decreased in the long term. Species' responses to the matrix that surrounds the fragments strongly predicted both the direction (increase/decline in occurrence) and magnitude of their responses to fragmentation. Additionally, species' responses to the matrix were somewhat predicted by their preferences for different types of native habitat (open vs. shaded). Our study highlights the degree of the matrix's influence in fragmented landscapes, and how this influence can change over time. We urge caution in using short-term responses to forecast long-term responses in cases where the matrix a) impacts species' responses to fragmentation (by isolating them, creating new habitat or altering fragment habitat) and b) is likely to change through time. This article is protected by copyright. All rights reserved.

Predicting landscape-genetic consequences of habitat loss, fragmentation and mobility for multiple species of woodland birds

Inference concerning the impact of habitat fragmentation on dispersal and gene flow is a key theme in landscape genetics. Recently, the ability of established approaches to identify reliably the differential effects of landscape structure (e.g. land-cover composition, remnant vegetation configuration and extent) on the mobility of organisms has been questioned. More explicit methods of predicting and testing for such effects must move beyond post hoc explanations for single landscapes and species. Here, we document a process for making a priori predictions, using existing spatial and ecological data and expert opinion, of the effects of landscape structure on genetic structure of multiple species across replicated landscape blocks. We compare the results of two common methods for estimating the influence of landscape structure on effective distance: least-cost path analysis and isolation-by-resistance. We present a series of alternative models of genetic connectivity in the study area, represented by different landscape resistance surfaces for calculating effective distance, and identify appropriate null models. The process is applied to ten species of sympatric woodland-dependant birds. For each species, we rank a priori the expectation of fit of genetic response to the models according to the expected response of birds to loss of structural connectivity and landscape-scale tree-cover. These rankings (our hypotheses) are presented for testing with empirical genetic data in a subsequent contribution. We propose that this replicated landscape, multi-species approach offers a robust method for identifying the likely effects of landscape fragmentation on dispersal.

Fish responses to experimental fragmentation of seagrass habitat

Understanding the consequences of habitat fragmentation has come mostly from comparisons of patchy and continuous habitats. Because fragmentation is a process, it is most accurately studied by actively fragmenting large patches into multiple smaller patches. We fragmented artificial seagrass habitats and evaluated the impacts of fragmentation on fish abundance and species richness over time (1 day, 1 week, 1 month). Fish assemblages were compared among 4 treatments: control (single, continuous 9-m(2) patches); fragmented (single, continuous 9-m(2) patches fragmented to 4 discrete 1-m(2) patches); prefragmented/patchy (4 discrete 1-m(2) patches with the same arrangement as fragmented); and disturbance control (fragmented then immediately restored to continuous 9-m(2) patches). Patchy seagrass had lower species richness than actively fragmented seagrass (up to 39% fewer species after 1 week), but species richness in fragmented treatments was similar to controls. Total fish abundance did not vary among treatments and therefore was unaffected by fragmentation, patchiness, or disturbance caused during fragmentation. Patterns in species richness and abundance were consistent 1 day, 1 week, and 1 month after fragmentation. The expected decrease in fish abundance from reduced total seagrass area in fragmented and patchy seagrass appeared to be offset by greater fish density per unit area of seagrass. If fish prefer to live at edges, then the effects of seagrass habitat loss on fish abundance may have been offset by the increase (25%) in seagrass perimeter in fragmented and patchy treatments. Possibly there is some threshold of seagrass patch connectivity below which fish abundances cannot be maintained. The immediate responses of fish to experimental habitat fragmentation provided insights beyond those possible from comparisons of continuous and historically patchy habitat.

Effects of fragmented habitat and fire on the distribution and ecology of the swamp antechinus (Antechinus minimus maritimus) in the Eastern Otways, Victoria

This study investigated the distribution, habitat and population dynamics of the swamp antechinus (Antechinus minimus maritimus) in the eastern Otway Ranges. The species has a restricted, disjunct distribution and has been recorded at 25 sites between 1969 and 1999. All sites were located within 7 km of the coast, occurred at altitudes up to 80 m above sea level and within 10 m of a gully. Analysis of landscape site variables identified sun index as being significant in determination of the probability of occurrence of A. minimus. The presence of A. minimus is negatively associated with sun index, occuring at sites that have a southerly aspect and gentle slope. A. minimus was located in a range of structural vegetation including Open Forest, Low Woodland, Shrubland and Hummock Grassland and a number of floristic groups, some characterised by high frequencies of sclerophyll shrubs, others by high frequencies of Pteridium esculentum, hummock grasses and herbaceous species. A. minimus occurs in fragmented, small populations with maximum population densities of 1.1–18 ha^–1. Populations at inland sites became extinct after the 1983 wildfire which burnt 41 000 ha. These sites have not been recolonised since, while on the coast the species did not re-establish until 1993–97. One population that is restricted to a narrow coastal strip of habitat is characterised by high levels of transient animals. The species is subject to extinction in the region due to habitat fragmentation, coastal developments and fire. Management actions to secure the present populations and ensure long-term survival of the species in the area are required and include implementation of appropriate fire regimes, prevention of habitat fragmentation, revegetation of habitat, and establishment of corridor habitat.

Estimating population size of the black-eared miner, with an assessment of landscape-scale habitat requirements

The decline of the Black-eared Miner Manorina melanotis has been caused primarily by habitat degradation and vegetation clearance. To better direct conservation actions for this species there was a need to assess habitat requirements on a regional-scale and to estimate the population size using quantitative methods. We used vegetation mapping and the current distribution of the Black-eared Miner to determine regional-scale habitat requirements. These findings were combined with the results of distance sampling to provide population estimates. The species is restricted to large tracts of intact mallee in the Murray Mallee of southeastern Australia that have not been burnt for at least 45 years. The density· of Black-eared Miners is highest in areas that are dominated by mallee- Triodia associations and have not been intensively grazed. The Bookmark Biosphere Reserve supports an estimated 501 (270-927, 95% CI) colonies, containing 3758 (2026-6954) phenotypically pure Black-eared Miners, 2255 (1 215-4170) hybrids and small numbers of Yellow-throated Miners Manorina flavigula. However, the effective population size is considerably smaller (390 Black-eared Miners '(21 0-726) and 234 hybrids (126-433)), due to a skewed adult sex ratio (1 female: 1.81 males) and complex social organization. A smaller population also persists in the Murray Sunset National Park containing 53 (32-85) Black-eared Miner/hybrid colonies. Both populations face a high risk of extinction from large-scale wildfire. The endangered status of the species under IUCN criteria remains warranted.

Genes and song : genetic and social connections in fragmented habitat in a woodland bird with limited dispersal

Understanding the processes leading to population declines in fragmented landscapes is essential for successful conservation management. However, isolating the influence of disparate processes, and dispersal in particular, is challenging. The Grey Shrike-thrush, Colluricincla harmonica, is a sedentary woodland-dependent songbird, with learned vocalizations whose incidence in suitable habitat patches falls disproportionally with decline in tree cover in the landscape. Although it has been suggested that gaps in tree cover might act as barriers to its dispersal, the species remains in many remnants of native vegetation in agricultural landscapes, suggesting that it may have responded to habitat removal and fragmentation by maintaining or even increasing dispersal distances. We quantified population connectivity of the Grey Shrike-thrush in a system fragmented over more than 120 years using genetic (microsatellites) and acoustic (song types) data. First, we tested for population genetic and acoustic structure at regional and local scales in search of barriers to dispersal or gene flow and signals of local spatial structuring indicative of restricted dispersal or localized acoustic similarity. Then we tested for effects of habitat loss and fragmentation on genetic and acoustic connectivity by fitting alternative models of mobility (isolation-by-distance [the null model] and reduced and increased movement models) across treeless vs. treed areas. Birds within 5 km of each other had more similar genotypes and song types than those farther away, suggesting that dispersal and song matching are limited in the region. Despite restricted dispersal detected for females (but not males), populations appeared to be connected by gene flow and displayed some cultural (acoustic) connectivity across the region. Fragmentation did not appear to impact greatly the dispersal of the Grey Shrike-thrush: none of the mobility models fit the genetic distances of males, whereas for females, an isolation-by-distance model could not be rejected in favor of the models of reduced or increased movement through treeless gaps. However, dissimilarities of the song types were more consistent with the model of reduced cultural connectivity through treeless areas, suggesting that fragmentation impedes song type sharing in the Grey Shrike-thrush. Our paper demonstrates that habitat fragmentation hinders important population processes in an Australian woodland bird even though its dispersal is not detectably impacted.

Preference for feeding at habitat edges declines among juvenile blue crabs as oyster reef patchiness increases and predation risk grows

Both habitat patchiness and behaviorally-mediated indirect effects (BMIEs; predator- induced changes in prey behavior that affect the prey's resources) are important in many food webs, but the relationships between these 2 factors have yet to be investigated. To explore effects of habitat patchiness and variation in perceived risk of predation on food-web dynamics, we conducted a factorial experiment in a model aquatic food chain of predator-prey-resource using 2 contrasting predators (adult blue crab Callinectes sapidus and toad fish Opsanus tau), juvenile blue crab as prey, and mussel Geukensia demissa as resource. Both predator presence and habitat patchiness influenced the prey's preference for consuming resources at patch edges instead of interiors. The preference of prey for consuming resources at habitat edges was 4 times stronger in continuous oyster reef habitat than in smaller habitat patches. This suggests that interior resources in continuous habitat experience a refuge from consumption, but this refuge is largely lost in patchy habitat. The mere presence of predators reduced the prey's preference for consuming resources at habitat edges. This BMIE was significant for the ambush predator (toadfish) and the treatment containing both predators, but not for the actively hunting predator (adult blue crab). We conclude that habitat patchiness and predator presence can jointly affect resource distribution by inducing shifts in prey foraging behavior, revealing a need to incorporate BMIEs into habitat fragmentation studies. This conclusion has broad and growing relevance as anthropogenic factors increasingly modify predator abundances and fragment coastal habitats. © Inter-Research 2012.

Ranging patterns of François' langur in a fragmented habitat in fusui reserve, Guangxi province, China

Studies of animal ranging patterns and the influencing ecological factors are useful for understanding the relationship between aspects of animal behavior and ecology. In a year-long study, we investigated the ranging behavior and other determining factors for a group of Francois' langur in an isolated habitat of approximately 25.7 ha in Fusui Reserve, China. The Francois' langur home range was estimated to be 15.3 ha, covering ∼60% of their total habitat. The mean yearly day range length estimate was 802.5 m (SD =295.5 m). Langurs changed sleeping sites approximately every 3 days, resulting in increases in the amount of grid cells used and the range length. Food availability of flowers and fruits were seasonal, whereas both mature and immature leaves of most trees were perennial. Ranging behavior was not significantly correlated with the availability of mature leaves, immature leaves, buds, fruits (ripe and unripe fruits) or seeds (p =0.05). These results suggested that variations in food type availability were not factors influencing ranging behavior for this langur group, whereas sleeping site changes, and probably predation avoidance, are factors that influence the ranging patterns of the langur group.