Viability and patch occupancy of a swamp rabbit metapopulation at the northern edge of its distribution

Swamp rabbits (Sylvilagus aquaticus) are state-endangered in Indiana, USA, and population decline has been attributed to habitat loss. We conducted pellet surveys as part of a long-term survey effort that has been conducted at approximate 10-year intervals over the last 40 years. We modeled patch occupancy and conducted a spatially-explicit population viability analysis (PVA). Although occupancy of individual patches varied over time, occupancy rate has been constant for the last 30 years, and Indiana swamp rabbits exist as a metapopulation that appears to be stable. Metapopulation dynamics were best characterized as being stationary, but area was an important factor in extinction rates; occupied patches (142 ± 37 ha) were significantly larger (P = 0.01) than unoccupied patches (79 ± 20 ha). We did not find strong support for models with colonization rates as a function of distance to neighboring patches, nor was distance to contiguous patches of habitat significantly different (P = 0.12) for occupied and unoccupied sites. Population viability analysis corroborated our findings based on occupancy modeling, and evaluation of the PVA model using occupancy data for the period 1985–2006 resulted in predictions that nearly matched our field observations (33% observed patch occupancy vs. 25% predicted patch occupancy). Population viability was most sensitive to reductions in survival and fecundity rates, but was otherwise robust to changes in parameters such as initial abundance and carrying capacity. Our findings provide novel insights into a poorly studied member of Sylvilagus and into species metapopulation dynamics at the edge of the range.     

A habitat-based population viability analysis for ocelots (Leopardus pardalis) in the United States

Strategies are needed to recover the ocelot Leopardus pardalis from the endangered species list. Recently, a population viability analysis (PVA) was developed which concluded that combinations of different recovery strategies were needed to effectively reduce ocelot extinction probability in the United States (US), with habitat protection and restoration identified as the most effective recovery scenario. We expanded this PVA model by incorporating landscape data to develop a more realistic habitat-based PVA for ocelots in southern Texas. We used RAMAS/gis software to conduct a habitat-based PVA by linking landscape data with a demographic metapopulation model. The primary goal of this study was to provide a model for evaluating ocelot recovery strategies in the US. Each model scenario was simulated 1000 times over 50 years and we defined extinction as one individual remaining. Using the RAMAS/gis program we identified 11 possible ocelot habitat patches (i.e., subpopulations) occurring in southern Texas. In addition, based on the habitat-based PVA model we found that combinations of different recovery strategies were needed to effectively reduce ocelot extinction probability in the US, with reducing road mortality the single most effective strategy. Short-term recovery strategies should include reducing ocelot road mortality, and translocation of ocelots into the US from northern Mexico. Long-term recovery strategies should include the restoration of habitat between and around existing ocelot habitat patches and the establishment of a dispersal corridor between ocelot breeding populations.     

An empirical evaluation of the area and isolation paradigm of metapopulation dynamics

Much of metapopulation theory assumes that the persistence of individual populations in a metapopulation, and persistence of the metapopulation as a whole, is best modeled by the area of habitat patches and their isolation. Estimates of isolation typically include a measure of geographic distance and a measure of either population size or patch area. This “area and isolation paradigm” assumes a functional relationship between the area of a patch and its extinction probability, and between isolation of a patch and its colonization probability. Although these assumptions are fundamental to use of incidence function models of metapopulation dynamics, the assumptions have been validated in only a small number of studies. We tested the ability of area and isolation to predict extinction and colonization patterns using multiple-year occupancy data for 10 species from three taxonomic groups (butterflies, amphibians, and birds). We examined 13 potential models of metapopulation dynamics. All models included four basic parameters: occupancy during the first year of the survey, probability of extinction, probability of colonization, and single-visit detection probability. In eight models, each parameter was either constant or time-dependent. Five models included a patch-level covariate of extinction probability (patch area or population size), colonization probability (connectivity, the inverse of isolation), or both. Extinction patterns generally were predicted more effectively as a function of local population size than as a function of patch area, a constant probability of extinction, or a time-dependent probability of extinction. In most cases, inclusion of connectivity as a patch-level covariate did not improve predictions of colonization patterns. We estimated single-visit detection probabilities for all species in our analyses, thus providing evidence-based guidelines for the refinement of future monitoring protocols.     

Persistence and conservation of a consumer-resource metapopulation with local overexploitation of resources

The monophagous flightless weevil Hadramphus spinipennis causes the frequent local extinction of its host plant Aciphylla dieffenbachii through overexploitation. Both species are endemic and restricted to the Chatham Islands group (New Zealand). The weevil is classified as endangered and the plant as a threatened species. As this herbivore-plant system is locally unstable long-term persistence only appears possible via a metapopulation structure in which subpopulations are connected by dispersal. This paper investigates this hypothesis using a spatially explicit model of a consumer population whose resource is patchily distributed. The parameters are adapted to the H. spinipennis-A. dieffenbachii system. Our model includes local population dynamics and dispersal of the consumer, the destruction of host plant patches due to foraging and their regeneration. The results show that the key factor for long-term persistence is the short-range dispersal of the consumer with high mortality during dispersal. Only this highly inefficient dispersal prevents the synchronisation of local dynamics while ensuring sufficient colonisation of regenerated patches. We also show that although synchronisation through spatially correlated environmental fluctuations may be critical for long-term persistence, it cannot replace the strong effects of dispersal. Thus, in a consumer-resource metapopulation with deterministic local extinction, the strength and spatial scale of consumer dispersal in relation to the spatial and temporal scales of the resource patch dynamics (patch destruction and regeneration, number, size and distance of patches) play a vital role for long-term persistence. The consequence for conservation management is that such metapopulations could in fact profit rather than suffer from decreasing connectivity of resource patches.     

The relative importance of patch habitat quality and landscape attributes on a declining steppe-bird metapopulation

Metapopulation theory is one of the most popular approaches to identify the factors affecting the spatial and temporal dynamics of populations in fragmented habitat networks. Habitat quality, patch area and isolation are mainly focused on when analyzing distribution patterns in fragmented landscapes. The effects of landscape heterogeneity in the non-occupied matrix, however, have been largely neglected. Here, we determined the relative importance of patch quality and landscape attributes on the occurrence, density and extinction of the Dupont’s lark (Chersophilus duponti), an endangered steppe passerine whose habitat has been extremely reduced to highly isolated and fragmented patches embedded in a mainly unsuitable landscape matrix. Habitat patch quality, measured in terms of vegetation structure, grazing pressure, arthropod availability, predator abundance, and inter-specific competition, did not affect occurrence, density or extinction. At the landscape scale, however, the species’ occurrence was principally determined by the interactions among patch size, geographic isolation and landscape matrix. Isolation had the main independent contribution to explaining the probability of occurrence, followed by landscape matrix composition and patch size. The species’ density was negatively correlated to patch size, suggesting crowding effects in small fragments, while extinction events were exclusively related to isolation. Our findings suggest that landscape rather than local population characteristics are crucial in determining the patterns of distribution and abundance of non-equilibrium populations in highly fragmented habitat networks. Consequently, conservation measures for these species should simultaneously involve patch size, isolation and landscape matrix and apply to the entire metapopulation rather than to particular patches.     

Removing erosion control projects increases bank swallow (Riparia riparia) population viability modeled along the Sacramento River, California, USA

Spatially-explicit population viability analysis (PVA) is a powerful method for modeling the extinction risk of populations that show variation over space and time. It is especially effective for comparing relative effect of different management scenarios on population dynamics. Here, I present a habitat patch-based PVA for a population of the California state-listed threatened bank swallow (Riparia riparia) nesting along the Sacramento River. This model incorporates the effects of habitat availability and location, density dependence, site fidelity, and stochasticity in survival and fecundity. River bank habitat patches suitable for this species were delineated using a geographic information systems (GIS) model of river bank height and were used in a PVA scenario analysis to assess the effects of habitat restoration—that could occur by removing bank erosion control projects (bank revetment)—on population viability. Sensitivity analysis showed that the model estimated probability of quasi-extinction (dropping below 2000 breeding pairs) ranged from 0 to 0.8 depending on the input parameters, with juvenile survival causing the greatest variability. However, comparing changes to the probability of quasi-extinction between the restored habitat scenario and current conditions showed a consistent 40-60% decrease in probability of quasi-extinction across all parameter combinations. The results of this research reaffirm the need for continued protection of the bank swallow as a listed species indicate that the removal of bank erosion control projects would increase viability of this population.     

Habitat succession, hardwood encroachment and raccoons as limiting factors for Lower Keys marsh rabbits

The Lower Keys marsh rabbit (LKMR, Sylvilagus palustris hefneri), a marsh rabbit subspecies endemic to the Lower Keys, Florida was protected in 1990, however, populations continue to decline despite recovery efforts. We hypothesized on-going habitat loss and fragmentation due to succession and hardwood encroachment has lead to increased edge, reduced habitat quality, and increased activity by native raccoons (Procyon lotor). These factors reduce the suitability of patches in a later successional state, thus threatening LKMR recovery and metapopulation persistence. We surveyed 150 LKMR patches in 2008, tallying adult and juvenile rabbit pellets, estimating measures of habitat succession and quality (woody and herbaceous ground cover, distribution of herbaceous species) and recording raccoon activity (number of raccoon signs). We calculated patch edge (patch shape index) using ArcGIS. We evaluated the relationship between patch and habitat attributes and LKMR using regression analysis and model selection. We found both adult and juvenile LKMR pellet counts were lower in patches with higher shape indices and higher in patches with greater occurrence of bunchgrasses and forbs. We also found adult LKMR pellet counts were lower in patches with higher raccoon activity. Our results suggest patch edge, habitat succession and quality, and raccoons pose a threat to the persistence and recovery of LKMR populations. Recovery efforts should focus on reducing these trends through habitat management and raccoon removal implemented in carefully controlled experiments with proper monitoring. Measures of patch and habitat attributes important to LKMR should be incorporated into long-term metapopulation monitoring and used to evaluate recovery actions.     

Spatial population structure of the Cabrera vole in Mediterranean farmland: The relative role of patch and matrix effects

There is mounting evidence that both patch networks and the intervening matrix influence species persistence in fragmented landscapes, though the relative importance of each of these factors in determining spatial population structure remains poorly understood. This study examined this issue using a three-year data set on the distribution of Cabrera voles (Microtus cabrerae) in Mediterranean farmland. The spatial pattern appeared consistent with a metapopulation structure, as voles occupied discrete tall herb patches scattered across the agricultural landscape, where local extinctions and colonizations induced temporal changes in occupancy patterns. Patch dynamics determined deviations from classical metapopulation assumptions, with over half the extinctions resulting from agricultural disturbance or vegetation succession, and recolonizations often occurring after the recovery of suitable habitat conditions sometime after disturbance. Occupancy in undisturbed patches was more stable, with vole occurrence in one year strongly reflecting that in the previous year. Overall, occupancy increased with both patch size and connectivity, but the unique contribution of patch variables to explain variation in vole occurrence was far smaller than that of matrix attributes. Voles occurred more often in patches surrounded by natural pastures, while prevalence declined with increasing cover by shrubland, pine plantations, improved pastures and grazed cropland. It is hypothesised that unfavourable land uses may increase the effective isolation of habitat patches through increased predation risk of dispersing voles. Conservation of the Cabrera vole in Mediterranean farmland should thus strive to maintain lightly grazed fields surrounding well-connected networks of suitable habitat patches.     

Conservation implications of the distribution of genetic diversity at different scales: a case study using the marsh fritillary butterfly (Euphydryas aurinia)

In the UK, Euphydryas aurinia exists in fragmented habitat patches, and undergoes population fluctuations as a result of a larval parasitoid. Its range is declining in the UK and conservation is thought to require a landscape approach since populations spread over large areas in some years and contract to core breeding patches in others. We examined populations at a range of geographic scales using allozyme electrophoresis to look for evidence of gene flow and differences in genetic diversity among populations. Nationally, our F_ST value was 0.1542 but between population groups within the suspected colonisation range of the butterfly (ca. 20 km), F_ST values were not significantly different from zero. Genetic diversity in terms of number of alleles and heterozygosity was reasonably high in natural populations (H_e=0.267) but low in an introduced, isolated population. We infer that migration between closely spaced subpopulations (in a metapopulation) maintains a high genetic effective population size (large number of individuals in a population that contribute genes to the next generation) which offsets any local reductions in population numbers due to stochastic extinctions or parasitoid effects. We therefore conclude that effective conservation of the species must seek to provide networks of suitable habitat for groups of subpopulations, rather than maintaining habitat for isolated populations.     

Probable consequences of high female mortality for speckled warblers living in habitat remnants

The effects of habitat fragmentation on the Australian avifauna have been widespread with species richness and abundance declining with reduced remnant size and habitat quality and increased habitat isolation. The speckled warbler, Chthonicola sagittata is one species from the highly fragmented temperate woodlands of eastern Australia that has declined across its range and populations that remain appear to be patchily distributed in habitat remnants. Specific causes of decline are unknown but several aspects of its biology make the species particularly vulnerable to decline in fragmented landscapes. Here, we analyse survey data (presence/absence) of speckled warblers in a large sample of habitat remnants from three regions to identify patterns of occupancy. We explore the effects of patch size on extinction risk using population viability analyses (PVA) and detailed demographic data from a behavioural study of individuals in the Australian Capital Territory, south-eastern Australia. Patch size was a strong predictor of the persistence of speckled warblers in habitat remnants. High density populations had higher probabilities of persisting, and inclusion of an Allee effect during drought decreased the probability of persistence. In the absence of an Allee effect, only high density populations in patches greater than 300 ha and low density populations in patches greater than700 ha had more than an 80% probability of persisting over 100 years. The accelerating decrease in population persistence below approximately 200-400 ha suggests that small populations were particularly vulnerable to stochastic demographic and environmental events. Adult female mortality was the single most important factor in driving population extinction. Our PVA model predictions matched the survey data for the Australian Capital Territory region remarkably well, but failed to predict occupancy rates in remnants in other regions. Differences in occupancy patterns between regions may, however, have resulted from inbreeding depression. This study demonstrates both the strengths and limitations of PVA analysis. PVA can predict occupancy patterns with reasonable accuracy, given good demographic data, but data for one region cannot be used universally for all regions. We highlight the need for studies of demography in different regions to interpret regional patterns of occupancy and to identify mechanisms of decline in remnant habitat.     

Metapopulation dynamics and threatened species management: Why does the broad-toothed rat (Mastacomys fuscus) persist?

Many species that exist patchily across the landscape are declining due to incremental losses of their constituent sub-populations and increasing isolation of those that remain. For threatened species with such patchy metapopulation structure, it is particularly important to identify key habitat patches and understand what factors govern their occupancy so that their management can be targeted and effective. In this paper, we describe the spatial and temporal distribution of an endangered population of broad-toothed rats (Mastacomys fuscus) at Barrington Tops, New South Wales, and model its dynamics using metapopulation theory. The study population occurs patchily in swamps on the Barrington Plateau. Using faecal pellet searches, live-trapping and data from previous surveys, we identified 12 swamps where M. fuscus persists and 13 where there has been a history of colonisation and extinction. The species now appears to be entirely absent from seven of these latter swamps. Using logistic regression and model selection procedures, we found the strongest predictor of the presence of M. fuscus to be proximity to the nearest occupied swamp. Persistence declined strongly with swamp isolation, probably due to low success of individuals dispersing through the intervening habitat. These patterns support the interpretation that swamp patches at Barrington Tops contain a functioning but fragile metapopulation of M. fuscus. We predict continued loss of remaining sub-populations in peripheral swamps if current dynamics continue, and recommend research to identify the factors that are limiting dispersal and re-colonisation so that the species’ decline can be slowed and reversed.     

The role of fragment area and isolation in the conservation of heathland species

Heathland area in the northwestern part of Belgium has been strongly reduced during the past 200 years, and the remaining heathland is forced back into several small and isolated relics. In this study, we investigated how the fragmentation of these heathlands affects the distribution patterns of heathland plant species. Furthermore, we tested whether differences in patch occupancy patterns could be explained in terms of life history traits related to dispersal capacity and persistence. Multivariate logistic regression models showed that the incidence of almost three quarters of the species was influenced by fragmentation. For the majority of these species, isolation was the most important factor determining their presence or absence in a heathland patch. Differences between the species in isolation-sensitivity could be attributed mainly to differences in seed bank characteristics, with species having a long living seed bank being less affected by isolation. In contrast, species having mechanisms facilitating long distance dispersal were as much affected by isolation as species lacking these mechanisms.Our results suggest that for the majority of the species extinction in a patch can be prevented by dispersal from neighbouring patches. Further isolation of the patches should therefore be prevented and connectivity between the patches needs to be assured. As almost none of the species is affected by a declining patch area, for most even small patches are important for their survival. Hence, conservation efforts should focus not only on large heathlands.     

Small mammals, habitat patches and PVA models: a field test of model predictive ability

The results are described of comparisons between actual values for patch occupancy for two species of Australian small mammals (Bush Rat Rattus fuscipes and Agile Antechinus Antechinus agilis) determined from field sampling and predictions of patch occupancy made using VORTEX, a generic simulation model for Population Viability Analysis (PVA). The work focussed on a fragmented forest in south-eastern Australia comprised of a network of 39 patches of native eucalypt forest surrounded by extensive stands of exotic softwood Radiata Pine (Pinus radiata) plantation. A range of modelling scenarios were completed in which four broad factors were varied: (1) inter-patch variation in habitat quality; (2) the pattern of inter-patch dispersal; (3) the rate of inter-patch dispersal; and (4) the population sink effects of the Radiata Pine matrix that surrounded the eucalypt patches. Model predictions were made for the total number of animals, the distribution of animal density among patches, the total number of occupied patches, and the probability of patch occupancy. Predictions were then compared with observed values for these same measures based on extensive field surveys of small mammals in the patch system. For most models for the Bush Rat, the predicted relative density of animals per patch correlated well with the values estimated from field surveys. Predictions of patch occupancy were not significantly different from the actual value for the number of occupied patches in half the models tested. The better models explained 10-16% of the log-likelihood of the probability of patch occupancy. While some of the models gave reasonable forecasts of the number of occupied patches, even in these cases, they had only moderate ability to predict which patches were occupied. Field surveys revealed there was no relationship between patch area and population density for the Agile Antechinus—an outcome correctly predicted by only a few models. Five of the 18 scenarios completed for the Agile Antechinus gave predicted numbers of occupied patches not significantly different from the observed number. In each of these five cases, large standard deviations around the mean predicted value meant uncertainty generated by the simulation model limited the predictive power of the PVA. Some of the models gave reasonable predictions for the number of occupied patches, but those models were unable to predict which ones were actually occupied. The results of our study suggest that key processes influencing which specific patches would be occupied were not modelled appropriately. High levels of variability and fecundity drive the population dynamics of the Bush Rat and Agile Antechinus, making the patch system unpredictable and difficult to model accurately. Despite the fact that both the Bush Rat and the Agile Antechinus are two of the most studied mammals in Australia, there are attributes of their biology that are presently poorly understood (which were not included in the VORTEX model), but which could strongly influence patch occupancy. For example, local landscape features may be important determinants of inter-patch movement and habitat utilisation in the patch system. Further empirical studies are needed to explore this aspect of small mammal biology.     

Potential factors controlling the population viability of newly introduced endangered marble trout populations

While several population viability analyses (PVAs) have been performed on anadromous salmonids, less attention has been given to stream-living salmonids. In this work, we explore the role of PVA as a tool in the recovery of threatened stream-living salmonid species. The analysis has been performed with reference to marble trout Salmo marmoratus, a salmonid with a limited geographic distribution and at risk of extinction due to hybridization with the non-native introduced brown trout. Demographic parameters, such as survival, fecundity and density-dependent patterns were estimated from an eight year on-going monitoring program of two translocated marble trout populations in pristine, previously fishless streams (Zakojska and Gorska) in the Soca and Idrijca river basins (Slovenia). To explore the importance of disturbance events such as floods on marble trout population dynamics, we performed a PVA under three scenarios: (1) occurrence of both severe and moderate floods; (2) occurrence of only moderate floods; (3) no flood events. Our analysis shows that population viability is threatened only by severe flood events, otherwise the two populations prove to be fairly stable with population abundance fluctuating around stream carrying capacity. A sensitivity analysis performed on model parameters highlighted that density-dependence in first-year survival and the magnitude of reduction in population size after a severe flood are the two most crucial parameters affecting population abundance and quasi-extinction probability, respectively. While only extreme floods can drive the population to extinction, the increase in juvenile survival when population abundance collapses after a major flood may allow the populations to quickly recover from few reproductive individuals back to stream carrying capacity.     

Lynx reintroductions in fragmented landscapes of Germany: Projects with a future or misunderstood wildlife conservation?

Eurasian lynx are slowly recovering in Germany after an absence of about 100 years, and additional reintroduction programs have been launched. However, suitable habitat is patchily distributed in Germany, and whether patches could host a viable population or contribute to the potential spread of lynx is uncertain. We combined demographic scenarios with a spatially explicit population simulation model to evaluate the viability and colonization success of lynx in the different patches, the aim being to conclude guidelines for reintroductions. The spatial basis of our model is a validated habitat model for the lynx in Germany. The dispersal module stems from a calibrated dispersal model, while the demographic module uses plausible published information on the lynx’ life history. The results indicate that (1) a viable population is possible, but that (2) source patches are not interconnected except along the German-Czech border, and that (3) from a demographic viewpoint at least 10 females and 5 males are required for a start that will develop into a viable population with an extinction probability of less than 5% in 50 years. The survival rate of resident adults was the most sensitive parameter, and the best management strategy for the success of reintroduction would be to reduce the mortality of residents in the source patches. Nevertheless, the extremely low probability of connectivity between suitable patches makes most of the reintroduction plans isolated efforts, and they are therefore questionable in the long run. With such a model, the suitability of the single habitat patches can be assessed and the most appropriate management scheme applied. This study shows that simulation models are useful tools for establishing the comparative effectiveness of reintroduction plans aimed at increasing the viability of the species.     

Wild-captive metapopulation viability analysis

We developed an interactive management model for wild and captive populations of the ploughshare tortoise or angonoka, Geochelone yniphora. Interactive management is based on the translocation of individuals between wild and captive populations to simulate a metapopulation. Demographic parameters of one captive and two wild populations of this rare tortoise were used to conduct a metapopulation viability analysis (MVA). The effectiveness of the conservation strategy proposed for this species was then evaluated by modifying the probability of extinction and growth of the metapopulation over a fixed period of time. Several alternative scenarios of interactive management were then tested and ranked in terms of their effect on the viability of the metapopulation. The model predicted that catastrophic events such as bush fires would likely have a negative effect on the future of remaining wild populations. However, the model also predicted that the use of captive-born offspring to establish additional wild populations would decrease the risk of extinction of the metapopulation as a whole. We believe that, when supported by sound knowledge of the demographic parameters of a species, the use of MVA as part of an interactive management program can be an effective conservation tool that allows assessment of the probable response of both captive and wild populations to different management alternatives. One of the most interesting aspects of this interactive management approach is the link between in situ and ex situ conservation.     

Viability of mountain caribou in British Columbia, Canada: Effects of habitat change and population density

Population viability analyses (PVA) are frequently employed to develop recovery plans and inform management of endangered species. Translating results from PVA into meaningful management recommendations often depends on an understanding of how population parameters change with environmental conditions as well as population density. The decline of mountain caribou (Rangifer tarandus caribou) in British Columbia, Canada, is believed to be caused by apparent competition with alternative prey species following changes to the forest age structure from timber harvest and wildfire. In addition, populations have been shown to decline at faster rates at low population density. To evaluate the potential effects of habitat change and population density on population persistence, we used stochastic projection models for 10 distinct populations varying in initial size from <10 to approximately 150 females. In an initial model, we used estimates of vital rates based on information sampled from >350 radiocollared caribou between 1984 and 2004. We then compared the results of the initial model to a set of models that evaluated the effects of habitat conditions and population density via their expected relationships to female adult survival. Assuming that vital rates remain constant over a 200-year time frame, only three populations have high probabilities (>0.95) of extinction. When models incorporate the declines in adult female survival know to occur with increasing proportions of young forest and declining population densities, all 10 populations are predicted to decline to extinction within <200 years. Based on our results, we suggest that PVA models that fail to incorporate the effects of changes in vital rates with habitat and population density may lead to overly optimistic assessments of the probability of population persistence in endangered species.     

Patch dynamics and the timing of colonization-abandonment events by male Kirtland’s Warblers in an early succession habitat

Habitat colonization and abandonment affects the distribution of a species in space and time, ultimately influencing the duration of time habitat is used and the total area of habitat occupied in any given year. Both aspects have important implications to long-term conservation planning. The importance of patch isolation and area to colonization-extinction events is well studied, but little information exists on how changing regional landscape structure and population dynamics influences the variability in the timing of patch colonization and abandonment events. We used 26 years of Kirtland’s Warbler (Dendroica kirtlandii) population data taken during a habitat restoration program (1979-2004) across its historical breeding range to examine the influence of patch attributes and temporal large-scale processes, specifically the rate of habitat turnover and fraction of occupied patches, on the year-to-year timing of patch colonization and abandonment since patch origin. We found the timing of patch colonization and abandonment was influenced by patch and large-scale regional factors. In this system, larger patches were typically colonized earlier (i.e., at a younger age) and abandoned later than smaller patches. Isolated patches (i.e., patches farther from another occupied patch) were generally colonized later and abandoned earlier. Patch habitat type affected colonization and abandonment; colonization occurred at similar patch ages between plantation and wildfire areas (9 and 8.5 years, respectively), but plantations were abandoned at earlier ages (13.9 years) than wildfire areas (16.4 years) resulting in shorter use. As the fraction of occupied patches increased, patches were colonized and abandoned at earlier ages. Patches were abandoned at older ages when the influx of new habitat patches was at low and high rates. Our results provide empirical support for the temporal influence of patch dynamics (i.e., patch destruction, creation, and succession) on local colonization and extinction processes that help explain large-scale patterns of habitat occupancy. Results highlight the need for practitioners to consider the timing of habitat restoration as well as total amount and spatial arrangement of habitat to sustain populations.     

The influence of social behaviour, dispersal and landscape fragmentation on population structure in a sedentary bird

White-browed Babblers Pomatostomus superciliosus lived in groups of up to 13 birds in the highly fragmented landscape of the WA wheatbelt. Contacts between these groups and sexual differences in dispersal behaviour interacted with the landscape mosaic at a number of spatial scales to produce a hierarchically structured population with four levels of organization: (1) groups, which were the basic breeding unit; (2) social neighbourhoods, where group interactions were frequent, and male dispersal and female post-natal and breeding dispersal occurred; (3) local population neighbourhoods, which contained social neighbourhoods between which female natal dispersal was frequent; and (4) metapopulations, which contained local population neighbourhoods between which dispersal was infrequent. The boundaries of these structural units, with the exception of the group, were not discrete and were influenced by the structure of the landscape they occupied.Interactions between groups occupying different patches were rare, and the frequency of group interactions was lower in small patches. Male dispersal was restricted to groups within the same patch or in patches less than 1 km apart. Therefore, decreasing patch size and increasing patch isolation resulted in smaller social neighbourhoods. Males generally dispersed to smaller groups and these dispersals may have enhanced the productivity of these groups by increasing their size. Therefore, habitat loss and fragmentation are likely to disrupt social neighbourhoods resulting in lower levels of social interaction and reduced productivity.The size and configuration of local populations were dependent on female natal dispersal, which in turn depended on landscape connectivity. White-browed Babblers used remnant vegetation in preference to other landscape elements when dispersing, but were not dependent solely on corridors. The permeability to dispersal of the boundaries between remnants and agricultural vegetation was dependent on patch configuration. Changes in boundary permeability were found to alter connectivity between habitat patches in a complex and asymmetric manner. Therefore, it is essential to consider landscape connectivity in a spatially explicit context for species that use some elements of the landscape mosaic in preference to others when dispersing.Habitat loss and fragmentation impose a complex set of changes, at a number of different scales, to processes that affect aspects of a species' life history. In order to manage species in fragmented agricultural landscapes it is necessary to understand the hierarchical structure of their populations, and how processes affect the different organizational levels within this structure.