Continuous versus binary representations of landscape heterogeneity in spatially-explicit models of mobile populations

How a landscape is represented is an important structural assumption in spatially-explicit simulation models. Simple models tend to specify just habitat and non-habitat (binary), while more complex models may use multiple levels or a continuum of habitat quality (continuous). How these different representations influence model projections is unclear. To assess the influence of landscape representation on population models, I developed a general, individual-based model with local dispersal and examined population persistence across binary and continuous landscapes varying in the amount and fragmentation of habitat. In binary and continuous landscapes habitat and non-habitat were assigned a unique mean suitability. In continuous landscapes, suitability of each individual site was then drawn from a normal distribution with fixed variance. Populations went extinct less often and abundances were higher in continuous landscapes. Production in habitat and non-habitat was higher in continuous landscapes, because the range of habitat suitability sampled by randomly dispersing individuals was higher than the overall mean habitat suitability. Increasing mortality, dispersal distance, and spatial heterogeneity all increased the discrepancy between continuous and binary landscapes. The effect of spatial structure on the probability of extinction was greater in binary landscapes. These results show that, under certain circumstances, model projections are influenced by how variation in suitability within a landscape is represented. Care should be taken to assess how a given species actually perceives the landscape when conducting population viability analyses or empirical validation of theory.     

Estimation of Core Terrestrial Habitat for Stream-Breeding Salamanders and Delineation of Riparian Buffers for Protection of Biodiversity

Abstract:  Many species of wildlife depend on riparian habitats for various life-history functions (e.g., breeding, foraging, overwintering). Although this unique habitat is critical for many species, delineations of riparian zones and buffers for various taxa are lacking. Typically when buffer zones are determined to mitigate edge effects, they are based on criteria that protect aquatic resources alone and do not consider impacts to wildlife and other terrestrial resources. Using two different survey methods (area-constrained daytime searches and nighttime visual encounter searches), we estimated core terrestrial habitat and buffer widths for stream-breeding salamanders in southern Appalachian streams from May to August 2004. A core terrestrial habitat of 27.0 m encompassed 95% of the salamander assemblage (four species of stream plethodontids), and an additional 50 m (to buffer edge effects) yielded a total buffer of 77.0 m. When each species of the assemblage was analyzed separately, the maximum core terrestrial habitat needed for the Blue Ridge two-lined salamander ( Eurycea wilderae ), a dominant member and the farthest-ranging species from the stream, was 42.6 m. Thus, we recommend an overall buffer width of 92.6 m in southern Appalachian streams. To protect stream amphibians and other wildlife dependent on riparian areas, land managers and policy makers must consider conserving more than aquatic resources alone. Developing core terrestrial habitat estimates and buffer zone widths for wildlife populations is a critical first step in the conservation of many semiaquatic organisms and protecting biodiversity.     

Tests of landscape influence: nest predation and brood parasitism in fragmented ecosystems

The effects of landscape fragmentation on nest predation and brood parasitism, the two primary causes of avian reproductive failure, have been difficult to generalize across landscapes, yet few studies have clearly considered the context and spatial scale of fragmentation. Working in two river systems fragmented by agricultural and rural-housing development, we tracked nesting success and brood parasitism in > 2500 bird nests in 38 patches of deciduous riparian woodland. Patches on both river systems were embedded in one of two local contexts (buffered from agriculture by coniferous forest, or adjacent to agriculture), but the abundance of agriculture and human habitation within 1 km of each patch was highly variable. We examined evidence for three models of landscape effects on nest predation based on (1) the relative importance of generalist agricultural nest predators, (2) predators associated with the natural habitats typically removed by agricultural development, or (3) an additive combination of these two predator communities. We found strong support for an additive predation model in which landscape features affect nest predation differently at different spatial scales. Riparian habitat with forest buffers had higher nest predation rates than sites adjacent to agriculture, but nest predation also increased with increasing agriculture in the larger landscape surrounding each site. These results suggest that predators living in remnant woodland buffers, as well as generalist nest predators associated with agriculture, affect nest predation rates, but they appear to respond at different spatial scales. Brood parasitism, in contrast, was unrelated to agricultural abundance on the landscape, but showed a strong nonlinear relationship with farm and house density, indicating a critical point at which increased human habitat causes increased brood parasitism. Accurate predictions regarding landscape effects on nest predation and brood parasitism will require an increased appreciation of the multiple scales at which landscape components influence predator and parasite behavior.     

Role of Light Availability and Dispersal in Exotic Plant Invasion along Roads and Streams in the H. J. Andrews Experimental Forest, Oregon

Abstract: We examined the roles of dispersal mechanism, a biological barrier; light availability, an environmental barrier; and level of disturbance, a physical barrier, in explaining the spatial patterns of exotic plant species along road and stream segments in a forest landscape in the western Cascade Range of Oregon (U.S.A). The presence or absence of 21 selected exotic plant species and light levels were observed along 0.3- to 1.0-km transects within four habitat types. Each habitat represented a different level of disturbance: high-use roads, low-use roads, abandoned roads, and streams in the H. J. Andrews Experimental Forest. Nearly 300 50 × 2–m sampling units were surveyed along five transects in each habitat type. We used ordination (nonmetric multidimensional scaling) and logistic regression to analyze data. All of the nearly 200 sampling units along roads with high and low levels of vehicle traffic contained at least one exotic plant species, and some contained as many as 14. Streams that were most recently disturbed by floods 20–30 years ago and abandoned spur roads with no traffic for 20–40 years also had numerous exotic species. Roads and streams apparently serve multiple functions that enhance exotic species invasion in this landscape: they act as corridors or agents for dispersal, provide suitable habitat, and contain reservoirs of propagules for future episodes of invasion. Species-specific dispersal mechanisms, habitat characteristics, and disturbance history each explain some, but not all, of the patterns of exotic species invasion observed in this study.     

Bird‐community responses to habitat creation in a long‐term,large‐scale natural experiment

Ecosystem function and resilience are compromised when habitats become fragmented due to land‐use change. This has led to national and international conservation strategies aimed at restoring habitat extent and improving functional connectivity (i.e., maintaining dispersal processes). However, biodiversity responses to landscape‐scale habitat creation and the relative importance of spatial and temporal scales are poorly understood, and there is disagreement over which conservation strategies should be prioritized. We used 160 years of historic post‐agricultural woodland creation as a natural experiment to evaluate biodiversity responses to habitat creation in a landscape context. Birds were surveyed in 101 secondary, broadleaf woodlands aged 10–160 years with ≥80% canopy cover and in landscapes with 0‐17% broadleaf woodland cover within 3000 m. We used piecewise structural equation modeling to examine the direct and indirect relationships between bird abundance and diversity, ecological continuity, patch characteristics, and landscape structure and quantified the relative conservation value of local and landscape scales for bird communities. Ecological continuity indirectly affected overall bird abundance and species richness through its effects on stand structure, but had a weaker influence (effect size near 0) on the abundance and diversity of species most closely associated with woodland habitats. This was probably because woodlands were rapidly colonized by woodland generalists in ≤10 years (minimum patch age) but were on average too young (median 50 years) to be colonized by woodland specialists. Local patch characteristics were relatively more important than landscape characteristics for bird communities. Based on our results, biodiversity responses to habitat creation depended on local‐ and landscape‐scale factors that interacted across time and space. We suggest that there is a need for further studies that focus on habitat creation in a landscape context and that knowledge gained from studies of habitat fragmentation and loss should be used to inform habitat creation with caution because the outcomes are not necessarily reciprocal.     

Terrestrial habitat selection and strong density-dependent mortality in recently metamorphosed amphibians

To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.     

Resource distributions among habitats determine solitary bee offspring production in a mosaic landscape.

Within mosaic landscapes, many organisms depend on attributes of the environment that operate over scales ranging from a single habitat patch to the entire landscape. One such attribute is resource distribution. Organisms' reliance on resources from within a local patch vs. those found among habitats throughout the landscape will depend on local habitat quality, patch quality, and landscape composition. The ability of individuals to move among complementary habitat types to obtain various resources may be a critical mechanism underlying the dynamics of animal populations and ultimately the level of biodiversity at different spatial scales. We examined the effects that local habitat type and landscape composition had on offspring production and survival of the solitary bee Osmia lignaria in an agri-natural landscape in California (U.S.A.). Female bees were placed on farms that did not use pesticides (organic farms), on farms that did use pesticides (conventional farms), or in seminatural riparian habitats. We identified pollens collected by bees nesting in different habitat types and matched these to pollens of flowering plants from throughout the landscape. These data enabled us to determine the importance of different plant species and habitat types in providing food for offspring, and how this importance changed with landscape and local nesting-site characteristics. We found that increasing isolation from natural habitat significantly decreased offspring production and survival for bees nesting at conventional farms, had weaker effects on bees in patches of seminatural habitat, and had little impact on those at organic farm sites. Pollen sampled from nests showed that females nesting in both farm and seminatural habitats relied on pollen from principally native plant species growing in seminatural habitat. Thus connectivity among habitats was critical for offspring production. Females nesting on organic farms were buffered to isolation effects by switching to floral resources growing at the farm site when seminatural areas were too distant. Overall local habitat conditions (farm management practices) can help bolster pollinators, but maintaining functional connectivity among habitats will likely be critical for persistence of pollinator populations as natural habitats are increasingly fragmented by human activities.     

Corridor Length and Patch Colonization by a Butterfly, Junonia coenia

Abstract: Corridors have been proposed to reduce isolation and increase population persistence in fragmented landscapes, yet little research has evaluated the types of landscapes in which corridors will be most effective. I tested the hypothesis that corridors increase patch colonization by a butterfly, Junonia coenia , regardless of the butterfly's initial distance from a patch. I chose J. coenia because it has been shown to move between patches preferentially through corridors. Individuals were released 16–192 m away from open experimental patches into adjacent open corridors or forest. Neither corridors nor distance had a significant effect on patch colonization, but there was a significant interaction between the presence or absence of corridors and distance. At small distances (16–64 m), J. coenia was more likely to colonize open patches when released within forest than within open corridors, most likely because J. coenia used corridors as habitat. Nevertheless, patch colonization by butterflies released within forest decreased rapidly as distance from patches increased, as predicted by a null model of random movement. Colonization did not change with distance in the corridor, and at long distances (128–192 m), butterflies released in corridors were twice as likely to colonize open patches as those released in forest. These results suggest that one critical factor, interpatch distance, may determine the relative effectiveness of corridors and other landscape configurations, such as stepping stones, in reducing isolation in fragmented landscapes. When distances between patches are short compared to an animal's movement ability, a stepping stone approach may most effectively promote dispersal. Alternatively, the conservation value of corridors is highest relative to other habitat configurations when longer distances separate patches in fragmented landscapes.     

Headwater riparian forest-floor invertebrate communities associated with alternative forest management practices.

Headwater streams and their riparian zones are a common, yet poorly understood, component of Pacific Northwest, USA, landscapes. We describe the ecological importance of headwater stream riparian zones as habitat for forest-floor invertebrate communities and assess how alternative management strategies for riparian zones may impact these communities. We compared community composition of forest-floor invertebrates at increasing distances along trans-riparian (stream edge to upslope) transects in mature forests, clearcuts, and riparian buffers of approximately 30-m width with upslope clearcuts. Invertebrates were collected using pitfall traps in five replicate blocks of three treatments each in the Willamette National Forest, Oregon, USA. We measured microclimate and microhabitat variables at pitfall locations. Despite strong elevation and block effects on community composition, community analyses revealed a distinct "riparian" invertebrate community within 1 m of the stream edge in mature forest treatments, which was strongly related to cool, humid microclimate conditions. Invertebrate community composition in buffer treatments was far more similar to that of mature forests than to clearcuts; a pattern mirrored by microclimate. These results suggest that, within our study sites, forest-floor invertebrate distributions are strongly associated with microclimate and that riparian buffers of approximately 30-m width do provide habitat for many riparian and forest species. Riparian reserves may serve as effective forest refugia and/or dispersal corridors for invertebrates and other taxa, and their incorporation into watershed management plans likely will contribute to meeting persistence and connectivity objectives.     

Appreciating Ecological Complexity: Habitat Contours as a Conceptual Landscape Model

Abstract:  Organisms respond to their surroundings at multiple spatial scales, and different organisms respond differently to the same environment. Existing landscape models, such as the "fragmentation model" (or patch-matrix-corridor model) and the "variegation model," can be limited in their ability to explain complex patterns for different species and across multiple scales. An alternative approach is to conceptualize landscapes as overlaid species-specific habitat contour maps. Key characteristics of this approach are that different species may respond differently to the same environmental conditions and at different spatial scales. Although similar approaches are being used in ecological modeling, there is much room for habitat contours as a useful conceptual tool. By providing an alternative view of landscapes, a contour model may stimulate more field investigations stratified on the basis of ecological variables other than human-defined patches and patch boundaries. A conceptual model of habitat contours may also help to communicate ecological complexity to land managers. Finally, by incorporating additional ecological complexity, a conceptual model based on habitat contours may help to bridge the perceived gap between pattern and process in landscape ecology. Habitat contours do not preclude the use of existing landscape models and should be seen as a complementary approach most suited to heterogeneous human-modified landscapes.     

Spatial uncertainty analysis of population models

This paper describes an approach for conducting spatial uncertainty analysis of spatial population models, and illustrates the ecological consequences of spatial uncertainty for landscapes with different properties. Spatial population models typically simulate birth, death, and migration on an input map that describes habitat. Typically, only a single “reference” map is available, but we can imagine that a collection of other, slightly different, maps could be drawn to represent a particular species’ habitat. As a first approximation, our approach assumes that spatial uncertainty (i.e., the variation among values assigned to a location by such a collection of maps) is constrained by characteristics of the reference map, regardless of how the map was produced. Our approach produces lower levels of uncertainty than alternative methods used in landscape ecology because we condition our alternative landscapes on local properties of the reference map. Simulated spatial uncertainty was higher near the borders of patches. Consequently, average uncertainty was highest for reference maps with equal proportions of suitable and unsuitable habitat, and no spatial autocorrelation. We used two population viability models to evaluate the ecological consequences of spatial uncertainty for landscapes with different properties. Spatial uncertainty produced larger variation among predictions of a spatially explicit model than those of a spatially implicit model. Spatially explicit model predictions of final female population size varied most among landscapes with enough clustered habitat to allow persistence. In contrast, predictions of population growth rate varied most among landscapes with only enough clustered habitat to support a small population, i.e., near a spatially mediated extinction threshold. We conclude that spatial uncertainty has the greatest effect on persistence when the amount and arrangement of suitable habitat are such that habitat capacity is near the minimum required for persistence.     

Scale Perspectives on Avian Diversity in Western Riparian Ecosystems

Conservation of riparian vegetation in western North America has, in part, emphasized providing habitats for a locally diverse avifauna. Site diversity, especially relative to the number of species present, is generally high within riparian avifaunas. Between-habitat diversity changes across a watershed, with riparian species assemblages differing most from upland assemblages at the highest and lowest elevations. This pattern can be attributed to enhanced avian movements within the riparian vegetation. The corridors for bird movements, in turn, facilitate faunal mixing on a broader scale, influencing regional diversity within landscapes. Riparian ecosystems are viewed as connectors of forests across fragmented landscapes. In western settings, however, they are highly linearized forests transecting watersheds between upland associations of high elevations and very different associations at lower elevations. Regionally, riparian vegetation represents linear islands that are internally both floristically and faunistically dynamic rather than mere bridges of homogeneous vegetation in landscape networks. The significance of riparian vegetation as habitat for western birds has been defined primarily at the local level. Conservation activities favoring site diversity are short-sighted, however, and could have severe consequences for unique elements of riparian avifaunas. Conservation actions must evaluate how local activities alter potential dispersal opportunities for ecological-generalist versus riparian-obligate species. Maintaining the character and integrity of riparian avifaunas requires planning from regional and continental perspectives.     

Landscape-level influences of terrestrial snake occupancy within the southeastern United States

Habitat loss and degradation are thought to be the primary drivers of species extirpations, but for many species we have little information regarding specific habitats that influence occupancy. Snakes are of conservation concern throughout North America, but effective management and conservation are hindered by a lack of basic natural history information and the small number of large-scale studies designed to assess general population trends. To address this information gap, we compiled detection/nondetection data for 13 large terrestrial species from 449 traps located across the southeastern United States, and we characterized the land cover surrounding each trap at multiple spatial scales (250-, 500-, and 1000-m buffers). We used occupancy modeling, while accounting for heterogeneity in detection probability, to identify habitat variables that were influential in determining the presence of a particular species. We evaluated 12 competing models for each species, representing various hypotheses pertaining to important habitat features for terrestrial snakes. Overall, considerable interspecific variation existed in important habitat variables and relevant spatial scales. For example, kingsnakes (Lampropeltis getula) were negatively associated with evergreen forests, whereas Louisiana pinesnake (Pituophis ruthveni) occupancy increased with increasing coverage of this forest type. Some species were positively associated with grassland and scrub/shrub (e.g., Slowinski's cornsnake, Elaphe slowinskii) whereas others, (e.g., copperhead, Agkistrodon contortrix, and eastern diamond-backed rattlesnake, Crotalus adamanteus) were positively associated with forested habitats. Although the species that we studied may persist in varied landscapes other than those we identified as important, our data were collected in relatively undeveloped areas. Thus, our findings may be relevant when generating conservation plans or restoration goals. Maintaining or restoring landscapes that are most consistent with the ancestral habitat preferences of terrestrial snake assemblages will require a diverse habitat matrix over large spatial scales.     

A Framework for Conceptualizing Human Effects on Landscapes and Its Relevance to Management and Research Models

Abstract: The concept of habitat fragmentation is limited in its ability to describe the range of possible landscape configurations created by a variety of disturbances. This limitation is especially problematic in landscapes where human use of the habitat matrix occurs at multiple levels and where habitat modification may be a more important consideration than a simple binary classification of habitat versus nonhabitat. We propose a synthesizing scheme that places intact, variegated, fragmented, and relictual landscape states on a continuum, depending on the degree of habitat destruction. At a second level, the scheme considers the patterns of habitat modification that are imposed on remaining habitats. Management for conservation involves halting and sometimes reversing the trends of habitat destruction and modification. Conservation strategies will differ according to the state of alteration of the landscape, but all strategies include some consideration of the degree of modification of the matrix in determining habitat viability. It is convenient for biologists to assess landscape alteration state in terms of the persistence of large structural elements such as trees. Because animal species use habitats differently, however, they also experience the landscape differently. A landscape considered structurally fragmented by humans may be functionally variegated to other species. Therefore, it is necessary to consider the extent to which the entire landscape, including the matrix, is accessible and utilized by organisms with different spatial scales of resource use.     

Managing Boreal Forest Landscapes for Flying Squirrels

Abstract: Flying squirrel (Pteromys volans) populations have declined severely during the past few decades, and the species has become a focal species in forest management and the conservation debate in Finland. We compared landscape structure around known flying squirrel home ranges with randomly chosen forest sites to determine which landscape patterns characterize the areas occupied by the species in northern Finland. We sought to identify the key characteristics of the landscape that support the remaining flying squirrel populations. We analyzed landscape structure within circular areas with 1- and 3-km radii around 63 forest sites occupied by flying squirrels, and around 96 random sites. We applied stepwise analysis of the landscape structure where landscapes were built up step-by-step by adding patch types in order of their suitability for the flying squirrel. The land-use and forest-resource data for the analysis were derived from multisource national forest inventory and imported to a geographical information system. Landscape patch types were divided into three suitability categories: breeding habitat (mixed spruce-deciduous forests); dispersal habitat ( pine and young forests); and unsuitable habitat ( young sapling stands, open habitats, water). Flying squirrel landscapes contained more suitable breeding habitat patches and were better connected by dispersal habitats than random landscapes. Our results suggest that for the persistence of the flying squirrel, forest managers should 1) maintain a deciduous mixture, particularly in spruce-dominated forests; 2) maintain physical connectivity between optimal breeding habitats; and 3) impose coarse-grained structures on northeastern Finnish landscapes at current levels of habitat availability.     

Grizzly bear habitat selection is scale dependent.

The purpose of our study is to show how ecologists' interpretation of habitat selection by grizzly bears (Ursus arctos) is altered by the scale of observation and also how management questions would be best addressed using predetermined scales of analysis. Using resource selection functions (RSF) we examined how variation in the spatial extent of availability affected our interpretation of habitat selection by grizzly bears inhabiting mountain and plateau landscapes. We estimated separate models for females and males using three spatial extents: within the study area, within the home range, and within predetermined movement buffers. We employed two methods for evaluating the effects of scale on our RSF designs. First, we chose a priori six candidate models, estimated at each scale, and ranked them using Akaike Information Criteria. Using this method, results changed among scales for males but not for females. For female bears, models that included the full suite of covariates predicted habitat use best at each scale. For male bears that resided in the mountains, models based on forest successional stages ranked highest at the study-wide and home range extents, whereas models containing covariates based on terrain features ranked highest at the buffer extent. For male bears on the plateau, each scale estimated a different highest-ranked model. Second, we examined differences among model coefficients across the three scales for one candidate model. We found that both the magnitude and direction of coefficients were dependent upon the scale examined; results varied between landscapes, scales, and sexes. Greenness, reflecting lush green vegetation, was a strong predictor of the presence of female bears in both landscapes and males that resided in the mountains. Male bears on the plateau were the only animals to select areas that exposed them to a high risk of mortality by humans. Our results show that grizzly bear habitat selection is scale dependent. Further, the selection of resources can be dependent upon the availability of a particular vegetation type on the landscape. From a management perspective, decisions should be based on a hierarchical process of habitat selection, recognizing that selection patterns vary across scales.     

Habitat Split as a Cause of Local Population Declines of Amphibians with Aquatic Larvae

Abstract:  Most amphibian species have biphasic life histories and undergo an ontogenetic shift from aquatic to terrestrial habitats. In deforested landscapes, streams and forest fragments are frequently disjunct, jeopardizing the life cycle of forest-associated amphibians with aquatic larvae. We tested the impact of habitat split—defined as human-induced disconnection between habitats used by different life-history stages of a species—on four forest-associated amphibian species in a severely fragmented landscape of the Brazilian Atlantic Forest. We surveyed amphibians in forest fragments with and without streams (referred to as wet and dry fragments, respectively), including the adjacent grass-field matrix. Our comparison of capture rates in dry fragments and nearby streams in the matrix allowed us to evaluate the number of individuals that engaged in high-risk migrations through nonforested habitats. Adult amphibians moved from dry fragments to matrix streams at the beginning of the rainy season, reproduced, and returned at the end of the breeding period. Juveniles of the year moved to dry fragments along with adults. These risky reproductive migrations through nonforested habitats that expose individuals to dehydration, predation, and other hazards may cause population declines in dry fragments. Indeed, capture rates were significantly lower in dry fragments compared with wet fragments. Declining amphibians would strongly benefit from investments in the conservation and restoration of riparian vegetation and corridors linking breeding and nonbreeding areas.     

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.     

Dispersal ecology of deadwood organisms and connectivity conservation

Limited knowledge of dispersal for most organisms hampers effective connectivity conservation in fragmented landscapes. In forest ecosystems, deadwood‐dependent organisms (i.e., saproxylics) are negatively affected by forest management and degradation globally. We reviewed empirically established dispersal ecology of saproxylic insects and fungi. We focused on direct studies (e.g., mark‐recapture, radiotelemetry), field experiments, and population genetic analyses. We found 2 somewhat opposite results. Based on direct methods and experiments, dispersal is limited to within a few kilometers, whereas genetic studies showed little genetic structure over tens of kilometers, which indicates long‐distance dispersal. The extent of direct dispersal studies and field experiments was small and thus these studies could not have detected long‐distance dispersal. Particularly for fungi, more studies at management‐relevant scales (1–10 km) are needed. Genetic researchers used outdated markers, investigated few loci, and faced the inherent difficulties of inferring dispersal from genetic population structure. Although there were systematic and species‐specific differences in dispersal ability (fungi are better dispersers than insects), it seems that for both groups colonization and establishment, not dispersal per se, are limiting their occurrence at management‐relevant scales. Because most studies were on forest landscapes in Europe, particularly the boreal region, more data are needed from nonforested landscapes in which fragmentation effects are likely to be more pronounced. Given the potential for long‐distance dispersal and the logical necessity of habitat area being a more fundamental landscape attribute than the spatial arrangement of habitat patches (i.e., connectivity sensu strict), retaining high‐quality deadwood habitat is more important for saproxylic insects and fungi than explicit connectivity conservation in many cases.     

Metapopulation Extinction Risk under Spatially Autocorrelated Disturbance

Abstract:  Recent extinction models generally show that spatial aggregation of habitat reduces overall extinction risk because sites emptied by local extinction are more rapidly recolonized. We extended such an investigation to include spatial structure in the disturbance regime. A spatially explicit metapopulation model was developed with a wide range of dispersal distances. The degree of aggregation of both habitat and disturbance pattern could be varied from a random distribution, through the intermediate case of a fractal distribution, all the way to complete aggregation (single block). Increasing spatial aggregation of disturbance generally increased extinction risk. The relative risk faced by populations in different landscapes varied greatly, depending on the disturbance regime. With random disturbance, the spatial aggregation of habitat reduced extinction risk, as in earlier studies. Where disturbance was spatially autocorrelated, however, this advantage was eliminated or reversed because populations in aggregated habitats are at risk of mass extinction from coarse-scale disturbance events. The effects of spatial patterns on extinction risk tended to be reduced by long-distance dispersal. Given the high levels of spatial correlation in natural and anthropogenic disturbance processes, population vulnerability may be greatly underestimated both by classical (nonspatial) models and by those that consider spatial structure in habitat alone.