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.

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.

Variation in the impact of non-native seaweeds along gradients of habitat degradation: a meta-analysis and an experimental test

Biological invasions are acknowledged among the main drivers of global changes in biodiversity. Despite compelling evidence of species interactions being strongly regulated by environmental conditions, there is a dearth of studies investi-gating how the effects of non-native species vary among areas exposed to different anthropogenic pressures. Focusing on marine macroalgae, we performed a meta-analysis to test whether and how the direction and magnitude of their effects on resident communities and species varies in relation to cumulative anthropogenic impact levels. The relationship between human impact levels and non-native species impact intensity emerged only for a reduced subset of the response variables examined. Yet, there was a trend for the effects of non-native species on community biomass and abundance and on species abundance to become less negative at heavily impacted sites. By contrast, the magnitude of negative effects of seaweed on community evenness tended to increase with human impact levels. The hypothesis of decreasing severity of invader’ impacts along a gradient of habitat degradation was also tested experimentally at a regional scale by comparing the effects of the removal of non-native alga,
Caulerpa cylindracea, on resident assemblages among rocky reefs exposed to different anthropogenic pressures. Assemblages at urban and pristine site did not differ when invaded, but did so when C. cylindracea was removed. Our results suggest that, despite the generally weak relationship between human impacts levels and non-native species impacts, more negative impacts can be expected in less stressful environments (i.e. less degraded or pristine sites), where competitive interactions are presumably the driving force structuring resident communities. Implementing strategies for controlling the establishment of non-native seaweeds should be, thus, considered a priority for preserving biodiversity in relatively pristine areas. On the other hand, control of invaders at degraded sites could be warranted to lessen their role as propagule sources

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.

Plantations, not farmlands, cause biotic homogenisation of ground-active beetles in South-Eastern Australia

Following landscape change, species invasions and extinctions may lead to biotic homogenisation, resulting in increased taxonomic and functional similarity between previously distinct biotas. Biotic homogenisation is more likely to occur in landscapes where the matrix contrasts strongly with native vegetation patches. To test this, we examined the distribution of ground-active beetles in a landscape of remnant Eucalyptus open woodland patches where large areas of lower contrast matrix (farmland) are being transformed to high-contrast pine plantations in south-eastern Australia. We sampled beetles from 30 sites including six replicates of five categories; (1) remnants adjacent to farmland, (2) remnants adjacent to plantation, (3) farmland, (4) plantation, and, (5) remnants between pine plantation and farmland. Community composition in the pine matrix was similar to native patches embedded in pine (ANOSIM, Global R=. 0.49, P<. 0.000), which we suggest is due to biotic homogenisation. Remnant patches with edges of both farmland and pine plantation did not represent an intermediate community composition between patches surrounded by either matrix type, but rather a unique habitat with unique species. Farmland supported the greatest number of individuals (. F=. 9.049, df. =. 25, P<. 0.000) and species (. F=. 5.875, df. =. 25, P=. 0.002), even compared to native remnant patches. Our results suggest that matrix transformations can reduce species richness and homogenise within-patch populations. This may increase the risk of species declines in fragmented landscapes where plantations are not only replacing native vegetation patches, but also other matrix types that may better support biodiversity. Our findings are particularly concerning given expanding plantation establishment worldwide.

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.

The response of ground and bark foraging insectivorous birds across an urban–forest gradient

This paper assesses the response of four common species of forest dependant insectivorous birds to an urban–forest gradient. The presence or absence was recorded for each species in landscapes that varied in landscape and site level attributes. Landscapes were classified into three categories based on their level of urbanisation. Broad comparisons across the landscapes were used to determine species specific response to increasing levels of urbanisation. Site level attributes were modelled to predict the patch occupancy for each species in each of the landscape types. Two broad trends were identified: the superb fairy wren (Malurus cyaneus) and white-browed scrubwren (Sericornis frontalis) displayed a tolerance to urbanisation and the eastern yellowrobin (Eosaltrica australis) and white throated treecreeper (Cormobates leucophaeus) demonstrated a threshold response to urbanisation. The density of roads (−ve) and the extent of tree cover (+ve) in a landscape were highly correlated with the occurrence of urban sensitive species while at the site level the density of roads and density of rivers were the strongest contributors to their presence. The marked differences in the isolation and connectivity of patches where the threshold for urban sensitive species ceases are the likely contributors to their decline and sensitivity to suburban habitats. Conservation and management of urban sensitive species is largely dependant on the way urban development is managed. Of critical importance is careful planning in urban-fringe environments.

Social structure and landscape genetics of the endemic New Caledonian ant Leptomyrmex pallens Emery, 1883 (Hymenoptera: Formicidae: Dolichoderinae), in the context of fire-induced rainforest fragmentation

Habitat fragmentation is a major threat to biodiversity, as it can alter ecological processes at various spatial and trophic scales. At the species level, fragmentation leading to the isolation of populations can trigger reductions in genetic diversity, potentially having detrimental effects on population fitness, adaptability and ultimately population persistence. Leptomyrmex pallens is a widespread rainforest ant endemic to New Caledonia but now confined to habitat patches that have been fragmented by anthropogenic fire regimes over the last 200 years. We investigated the social structure of L. pallens in the Aoupinié region (c.a. 4900 ha), and assessed the impacts of habitat fragmentation on its population genetic structure. Allele frequencies at 13 polymorphic microsatellite loci were compared among 411 worker ants from 21 nests distributed across the region. High within-nest relatedness (r = 0.70 ± 0.02), and a single queen found in 38 % of the nests by pedigree analysis indicate that the species is monogynous to weakly polygynous. Estimates of gene flow and genetic structure across the region were subsequently determined using a combined dataset of single workers per nest and of unrelated foraging workers. These estimates coupled with a comprehensive landscape genetic analysis revealed no evidence of significant population structure or habitat effects, suggesting that the Aoupinié region harbours a single panmictic population. In contrast, analyses of mitochondrial DNA sequence data revealed a high degree of genetic structuring, indicating limited maternal gene flow and suggesting that gene flow among nests is driven primarily by winged males. Overall these findings suggest that fire-induced habitat fragmentation has had little impact on the population dynamics of L. pallens. Additional studies of less mobile species should therefore be conducted to gain further insights into fire related disturbances on the unique biodiversity and function of New Caledonian ecosystems.

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.