Fire and forest landscapes in the Georgia Piedmont: an assessment of spatial modeling assumptions

Landscape simulation models are widely used to study the behavior of ecological systems. As computing power has increased, these models have become more complex and incorporated more realistic spatial representations of landscape patterns and ecological processes. The goal of this research was to examine the sensitivity of simulated landscape patterns to fundamental spatial modeling assumptions. The LANDIS simulator was parameterized for forests of the Georgia Piedmont and used to model landscape-scale community dynamics at fire return intervals from 20 to 100 years. A base scenario incorporating localized seed dispersal along with landform-related variation in species establishment rates and disturbance regimes was contrasted with three alternative scenarios. The uniform habitat scenario applied the same set of species establishment coefficients across all landforms. The uniform dispersal scenario removed the effects of seed source abundance and pattern on species establishment. The uniform disturbance scenario assumed identical disturbance regimes on all landforms.At the shortest fire return intervals, fire severities were low and the stand age distribution was dominated by older forests. At longer fire return intervals, fire severities were high and the stand age distribution was skewed toward younger forests. Species composition generally followed a gradient from fire-resistant species at short fire return intervals to fire-sensitive species at longer fire return intervals. However, some species exhibited bimodal distributions with high abundances at both short and long fire return intervals. Landscape responses to fire were similar in the uniform habitat scenario and the base scenario. Communities were less sensitive to fire return interval and had more fire-sensitive species in the uniform dispersal scenario than in the base scenario. Species composition in the uniform disturbance scenario was similar to the base scenario for the longest fire-intervals, but was more sensitive to changes in the fire regime at shorter fire return intervals. In models of Piedmont forest landscapes, accurate spatial representations of dispersal and fire regime heterogeneity are essential for predicting landscape-scale species composition under changing fire regimes. In contrast, the precise spatial representation of species–habitat relationships may be considerably less important.     

Local extinction of grassland plants: the landscape matrix is more important than patch attributes

Past studies of local extinctions in fragmented habitats most often tested the influence of fragment size and isolation while ignoring how differences in the surrounding landscape matrix may govern extinction. We assessed how both the spatial attributes of remnant patches (area and isolation) and landscape factors (extent of urbanization and maximum inter-fire interval) influence the persistence of native plant species in grasslands in western Victoria, Australia. Persistence was determined in 2001 by resurveying 30 remnants first surveyed in the 1980s, and correlates of extinction were assessed using Bayesian logistic regression models. On average, 26% of populations of native species became locally extinct over two decades. Area and isolation had little effect on the probability of local extinction, but urbanization and longer maximum inter-fire intervals increased extinction risk. These findings suggest that the native grasslands studied are relatively insensitive to area- and isolation-based fragmentation effects and that short-term persistence of plant populations requires the maintenance of habitat quality. The latter is strongly influenced by the landscape matrix surrounding remnant patches through changes in fire regimes and increased exogenous disturbance.     

A spatially explicit model to predict future landscape composition of aspen woodlands under various management scenarios

Quaking aspen (Populus tremuloides) is declining across the western United States. Aspen habitats are among the most diverse plant communities in this region and loss of these habitats can result in shifts in biodiversity, productivity, and hydrology across a range of spatial scales. Western aspen occurs on the majority of sites seral to conifer species, and long-term maintenance of these aspen woodlands requires periodic fire. Over the past century, fire intervals, extents, and intensities have been insufficient to regenerate aspen stands at historic rates; however the effects of various fire regimes and management scenarios on aspen vegetation dynamics at broad spatial and temporal scales are unexplored. Here we use field data, remotely sensed data, and fire atlas information to develop a spatially explicit landscape simulation model to assess the effects of current and historic wildfire regimes and prescribed burning programs on landscape vegetation composition across two mountain ranges in the Owyhee Plateau, Idaho. Model outputs depict the future structural makeup and species composition of the landscape at selected time steps under simulated management scenarios. We found that under current fire regimes and in the absence of management activities, loss of seral aspen stands will continue to occur over the next two centuries. However, a return to historic fire regimes (burning 12–14% of the modeled landscape per decade) would maintain the majority of aspen stands in early and mid seral woodland stages and minimizes the loss of aspen. A fire rotation of 70–80 years was estimated for the historic fire regime while the current fire regime resulted in a fire rotation of 340–450 years, underscoring the fact that fire is currently lacking in the system. Implementation of prescribed burning programs, treating aspen and young conifer woodlands according to historic fire occurrence probabilities, are predicted to prevent conifer dominance and loss of aspen stands.     

Optimal fire histories for biodiversity conservation

Fire is used as a management tool for biodiversity conservation worldwide. A common objective is to avoid population extinctions due to inappropriate fire regimes. However, in many ecosystems, it is unclear what mix of fire histories will achieve this goal. We determined the optimal fire history of a given area for biological conservation with a method that links tools from 3 fields of research: species distribution modeling, composite indices of biodiversity, and decision science. We based our case study on extensive field surveys of birds, reptiles, and mammals in fire‐prone semi‐arid Australia. First, we developed statistical models of species’ responses to fire history. Second, we determined the optimal allocation of successional states in a given area, based on the geometric mean of species relative abundance. Finally, we showed how conservation targets based on this index can be incorporated into a decision‐making framework for fire management. Pyrodiversity per se did not necessarily promote vertebrate biodiversity. Maximizing pyrodiversity by having an even allocation of successional states did not maximize the geometric mean abundance of bird species. Older vegetation was disproportionately important for the conservation of birds, reptiles, and small mammals. Because our method defines fire management objectives based on the habitat requirements of multiple species in the community, it could be used widely to maximize biodiversity in fire‐prone ecosystems. Historiales de Incendios Óptimos para la Conservación de la Biodiversidad     

Effects of landscape design of forest reserves on Saproxylic beetle diversity

Increasing the density of natural reserves in the forest landscape may provide conservation benefits for biodiversity within and beyond reserve borders. We used 2 French data sets on saproxylic beetles and landscape cover of forest reserves (LCFR) to test this hypothesis: national standardized data derived from 252 assessment plots in managed and reserve stands in 9 lowland and 5 highland forests and data from the lowland Rambouillet forest, a forested landscape where a pioneer conservation policy led to creation of a dense network of reserves. Abundance of rare and common saproxylic species and total saproxylic species richness were higher in forest reserves than in adjacent managed stands only in highland forests. In the lowland regional case study, as LCFR increased total species richness and common species abundance in reserves increased. In this case study, when there were two or more reserve patches, rare species abundance inside reserves was higher and common species richness in managed stands was higher than when there was a single large reserve. Spillover and habitat amount affected ecological processes underlying these landscape reserve effects. When LCFR positively affected species richness and abundance in reserves or managed stands, >12‐20% reserve cover led to the highest species diversity and abundance. This result is consistent with the target of 17% forested land area in reserves set at the Nagoya biodiversity summit in 2010. Therefore, to preserve biodiversity we recommend at least doubling the current proportion of forest reserves in European forested landscapes.     

Comparing modern and presettlement forest dynamics of a subboreal wilderness: does spruce budworm enhance fire risk?

Insect disturbance is often thought to increase fire risk through enhanced fuel loadings, particularly in coniferous forest ecosystems. Yet insect disturbances also affect successional pathways and landscape structure that interact with fire disturbances (and vice-versa) over longer time scales. We applied a landscape succession and disturbance model (LANDIS-II) to evaluate the relative strength of interactions between spruce budworm (Choristoneura fumiferana) outbreaks and fire disturbances in the Boundary Waters Canoe Area (BWCA) in northern Minnesota (USA). Disturbance interactions were evaluated for two different scenarios: presettlement forests and fire regimes vs. contemporary forests and fire regimes. Forest composition under the contemporary scenario trended toward mixtures of deciduous species (primarily Betula papyrifera and Populus spp.) and shade-tolerant conifers (Picea mariana, Abies balsamea, Thuja occidentalis), with disturbances dominated by a combination of budworm defoliation and high-severity fires. The presettlement scenario retained comparatively more "big pines" (i.e., Pinus strobus, P. resinosa) and tamarack (L. laricina), and experienced less budworm disturbance and a comparatively less-severe fire regime. Spruce budworm disturbance decreased area burned and fire severity under both scenarios when averaged across the entire 300-year simulations. Contrary to past research, area burned and fire severity during outbreak decades were each similar to that observed in non-outbreak decades. Our analyses suggest budworm disturbances within forests of the BWCA have a comparatively weak effect on long-term forest composition due to a combination of characteristics. These include strict host specificity, fine-scaled patchiness created by defoliation damage, and advance regeneration of its primary host, balsam fir (A. balsamea) that allows its host to persist despite repeated disturbances. Understanding the nature of the three-way interaction between budworm, fire, and composition has important ramifications for both fire mitigation strategies and ecosystem restoration initiatives. We conclude that budworm disturbance can partially mitigate long-term future fire risk by periodically reducing live ladder fuel within the mixed forest types of the BWCA but will do little to reverse the compositional trends caused in part by reduced fire rotations.     

Assessing Risks to Biodiversity from Future Landscape Change

We examined the impacts of possible future land development patterns on the biodiversity of a landscape. Our landscape data included a remote sensing derived map of the current habitat of the study area and six maps of future habitat distributions resulting from different land development scenarios. Our species data included lists of all bird, mammal, reptile, and amphibian species in the study area, their habitat associations, and area requirements for each. We estimated the area requirements using home ranges, sampled population densities, or genetic area requirements that incorporate dispersal distances. Our measures of biodiversity were species richness and habitat abundance. We calculated habitat abundance in two ways. First, we computed the total habitat area for each species in each landscape. Second, we calculated the number of habitat units for each species in each landscape by dividing the size of each habitat patch in the landscape by the area requirement and summing over all patches. Species richness was based on presence of habitat. Species became extinct in the landscape if they had no habitat area or no habitat units, respectively. We then computed ratios of habitat abundance in each future landscape to habitat abundance in the present for each species. We also computed the ratio of future to present species richness. We then calculated summary statistics across all species. Species richness changed little from present to future. There were distinctly greater risks to habitat abundance in landscapes that extrapolated from present trends or zoning patterns, however, as opposed to landscapes in which land development activities followed more constrained patterns. These results were stable when tested using Monte Carlo simulations and sensitivity tests on the area requirements. We conclude that this methodology can begin to discriminate the effects of potential changes in land development on vertebrate biodiversity.     

Modeling mammal response to fire based on species’ traits

Fire has shaped ecological communities worldwide for millennia, but impacts of fire on individual species are often poorly understood. We performed a meta-analysis to predict which traits, habitat, or study variables and fire characteristics affect how mammal species respond to fire. We modeled effect sizes of measures of population abundance or occupancy as a function of various combinations of these traits and variables with phylogenetic least squares regression. Nine of 115 modeled species (7.83%) returned statistically significant effect sizes, suggesting most mammals are resilient to fire. The top-ranked model predicted a negative impact of fire on species with lower reproductive rates, regardless of fire type (estimate = –0.68), a positive impact of burrowing in prescribed fires (estimate = 1.46) but not wildfires, and a positive impact of average fire return interval for wildfires (estimate = 0.93) but not prescribed fires. If a species’ International Union for Conservation of Nature Red List assessment includes fire as a known or possible threat, the species was predicted to respond negatively to wildfire relative to prescribed fire (estimate = –2.84). These findings provide evidence of experts’ abilities to predict whether fire is a threat to a mammal species and the ability of managers to meet the needs of fire-threatened species through prescribed fire. Where empirical data are lacking, our methods provide a basis for predicting mammal responses to fire and thus can guide conservation actions or interventions in species or communities.     

Quantifying Habitat Requirements of Tree-Living Species in Fragmented Boreal Forests with Bayesian Methods

Abstract:  Quantitative conservation objectives require detailed consideration of the habitat requirements of target species. Tree-living bryophytes, lichens, and fungi are a critical and declining biodiversity component of boreal forests. To understand their requirements, Bayesian methods were used to analyze the relationships between the occurrence of individual species and habitat factors at the tree and the stand scale in a naturally fragmented boreal forest landscape. The importance of unexplained between-stand variation in occurrence of species was estimated, and the ability of derived models to predict species' occurrence was tested. The occurrence of species was affected by quality of individual trees. Furthermore, the relationships between occurrence of species at the tree level and size and shape of stands indicated edge effects, implying that some species were restricted to interior habitats of large, regular stands. Yet for the habitat factors studied, requirements of many species appeared similar. Species occurrence also varied between stands; most of the seemingly suitable trees in some stands were unoccupied. The models captured most variation in species occurrence at tree level. They also successfully accounted for between-stand variation in species occurrence, thus providing realistic simulations of stand-level occupancy of species. Important unexplained between-stand variation in species occurrence warns against a simplified view that only local habitat factors influence species' occurrence. Apparently, similar stands will host populations of different sizes due to historical, spatial, and stochastic factors. Thus, habitat suitability cannot be assessed simply by population sizes, and stands lacking a species may still provide suitable habitat and merit protection.     

Defining and Evaluating the Umbrella Species Concept for Conserving and Restoring Landscape Connectivity

Conserving or restoring landscape connectivity between patches of breeding habitat is a common strategy to protect threatened species from habitat fragmentation. By managing connectivity for some species, usually charismatic vertebrates, it is often assumed that these species will serve as conservation umbrellas for other species. We tested this assumption by developing a quantitative method to measure overlap in dispersal habitat of 3 threatened species—a bird (the umbrella), a butterfly, and a frog—inhabiting the same fragmented landscape. Dispersal habitat was determined with Circuitscape, which was parameterized with movement data collected for each species. Despite differences in natural history and breeding habitat, we found substantial overlap in the spatial distributions of areas important for dispersal of this suite of taxa. However, the intuitive umbrella species (the bird) did not have the highest overlap with other species in terms of the areas that supported connectivity. Nevertheless, we contend that when there are no irreconcilable differences between the dispersal habitats of species that cohabitate on the landscape, managing for umbrella species can help conserve or restore connectivity simultaneously for multiple threatened species with different habitat requirements. Definición y Evaluación del Concepto de Especie Paraguas para Conservar y Restaurar la Conectividad de Paisajes     

Use of linkage mapping and centrality analysis across habitat gradients to conserve connectivity of gray wolf populations in western North America

Centrality metrics evaluate paths between all possible pairwise combinations of sites on a landscape to rank the contribution of each site to facilitating ecological flows across the network of sites. Computational advances now allow application of centrality metrics to landscapes represented as continuous gradients of habitat quality. This avoids the binary classification of landscapes into patch and matrix required by patch-based graph analyses of connectivity. It also avoids the focus on delineating paths between individual pairs of core areas characteristic of most corridor- or linkage-mapping methods of connectivity analysis. Conservation of regional habitat connectivity has the potential to facilitate recovery of the gray wolf (Canis lupus), a species currently recolonizing portions of its historic range in the western United States. We applied 3 contrasting linkage-mapping methods (shortest path, current flow, and minimum-cost-maximum-flow) to spatial data representing wolf habitat to analyze connectivity between wolf populations in central Idaho and Yellowstone National Park (Wyoming). We then applied 3 analogous betweenness centrality metrics to analyze connectivity of wolf habitat throughout the northwestern United States and southwestern Canada to determine where it might be possible to facilitate range expansion and interpopulation dispersal. We developed software to facilitate application of centrality metrics. Shortest-path betweenness centrality identified a minimal network of linkages analogous to those identified by least-cost-path corridor mapping. Current flow and minimum-cost-maximum-flow betweenness centrality identified diffuse networks that included alternative linkages, which will allow greater flexibility in planning. Minimum-cost-maximum-flow betweenness centrality, by integrating both land cost and habitat capacity, allows connectivity to be considered within planning processes that seek to maximize species protection at minimum cost. Centrality analysis is relevant to conservation and landscape genetics at a range of spatial extents, but it may be most broadly applicable within single- and multispecies planning efforts to conserve regional habitat connectivity.     

Toward Meaningful Snag-Management Guidelines for Postfire Salvage Logging in North American Conifer Forests

Abstract:  The bird species in western North America that are most restricted to, and therefore most dependent on, severely burned conifer forests during the first years following a fire event depend heavily on the abundant standing snags for perch sites, nest sites, and food resources. Thus, it is critical to develop and apply appropriate snag-management guidelines to implement postfire timber harvest operations in the same locations. Unfortunately, existing guidelines designed for green-tree forests cannot be applied to postfire salvage sales because the snag needs of snag-dependent species in burned forests are not at all similar to the snag needs of snag-dependent species in green-tree forests. Birds in burned forests have very different snag-retention needs from those cavity-nesting bird species that have served as the focus for the development of existing snag-management guidelines. Specifically, many postfire specialists use standing dead trees not only for nesting purposes but for feeding purposes as well. Woodpeckers, in particular, specialize on wood-boring beetle larvae that are superabundant in fire-killed trees for several years following severe fire. Species such as the Black-backed Woodpecker ( Picoides arcticus ) are nearly restricted in their habitat distribution to severely burned forests. Moreover, existing postfire salvage-logging studies reveal that most postfire specialist species are completely absent from burned forests that have been (even partially) salvage logged. I call for the long-overdue development and use of more meaningful snag-retention guidelines for postfire specialists, and I note that the biology of the most fire-dependent bird species suggests that even a cursory attempt to meet their snag needs would preclude postfire salvage logging in those severely burned conifer forests wherein the maintenance of biological diversity is deemed important.     

Restoration of Landscape Structure Altered by Fire Suppression

There is increasing interest in applying landscape ecological research to the management of wildlands, particularly regarding the negative effects of fragmentation and the benefits of corridors. Patch-producing large disturbances, such as fires and floods, produce a spatial mosaic structure in landscapes to which many species are sensitive. Management of the spatial structure of the patch mosaic has seldom been an explicit concern, however, in part because of insufficient knowledge about bow this spatial structure is affected by alterations in the disturbance regime. Yet the patch mosaic structure of many landscapes has been altered by disturbance control (such as fire suppression), and there is substantial interest in restoring natural disturbance regimes in some wildland landscapes. It has been proposed that, in landscapes subjected to decades of fire suppression, simple reinstatement of the natural fire regime may lead to adverse effects because fuel buildup during fire suppression may result in unusually large fires. It has also been proposed that the use of small prescribed fires may be an effective approach to restoration of landscapes subjected to fire suppression. Here I use a spatial GIS-based simulation model to analyze the effects of reinstating a natural fire regime in the Boundary Waters Canoe Area, Minnesota, after 82 years of fire suppression. The simulation experiment suggests that suppression can be expected to significantly alter landscape structure, but landscape structure can generally be restored within 50–75 years by reinstating the natural fire regime. Unusually large fires would probably hasten the restoration of landscape structure, while small prescribed fires will not restore the landscape but instead will produce further alteration.     

Validating graph-based connectivity models with independent presence–absence and genetic data sets

Habitat connectivity is a key objective of current conservation policies and is commonly modeled by landscape graphs (i.e., sets of habitat patches [nodes] connected by potential dispersal paths [links]). These graphs are often built based on expert opinion or species distribution models (SDMs) and therefore lack empirical validation from data more closely reflecting functional connectivity. Accordingly, we tested whether landscape graphs reflect how habitat connectivity influences gene flow, which is one of the main ecoevolutionary processes. To that purpose, we modeled the habitat network of a forest bird (plumbeous warbler [Setophaga plumbea]) on Guadeloupe with graphs based on expert opinion, Jacobs’ specialization indices, and an SDM. We used genetic data (712 birds from 27 populations) to compute local genetic indices and pairwise genetic distances. Finally, we assessed the relationships between genetic distances or indices and cost distances or connectivity metrics with maximum-likelihood population-effects distance models and Spearman correlations between metrics. Overall, the landscape graphs reliably reflected the influence of connectivity on population genetic structure; validation R² was up to 0.30 and correlation coefficients were up to 0.71. Yet, the relationship among graph ecological relevance, data requirements, and construction and analysis methods was not straightforward because the graph based on the most complex construction method (species distribution modeling) sometimes had less ecological relevance than the others. Cross-validation methods and sensitivity analyzes allowed us to make the advantages and limitations of each construction method spatially explicit. We confirmed the relevance of landscape graphs for conservation modeling but recommend a case-specific consideration of the cost-effectiveness of their construction methods. We hope the replication of independent validation approaches across species and landscapes will strengthen the ecological relevance of connectivity models.     

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.     

Simulating Historical Variability in the Amount of Old Forests in the Oregon Coast Range

Abstract: We developed the landscape age-class demographics simulator ( LADS) to model historical variability in the amount of old-growth and late-successional forest in the Oregon Coast Range over the past 3,000 years. The model simulated temporal and spatial patterns of forest fires along with the resulting fluctuations in the distribution of forest age classes across the landscape. Parameters describing historical fire regimes were derived from data from a number of existing dendroecological and paleoecological studies. Our results indicated that the historical age-class distribution was highly variable and that variability increased with decreasing landscape size. Simulated old-growth percentages were generally between 25% and 75% at the province scale (2,250,000 ha) and never fell below 5%. In comparison, old-growth percentages varied from 0 to 100% at the late-successional reserve scale (40,000 ha). Province-scale estimates of current old-growth (5%) and late-successional forest (11%) in the Oregon Coast Range were lower than expected under the simulated historical fire regime, even when potential errors in our parameter estimates were considered. These uncertainties do, however, limit our ability to precisely define ranges of historical variability. Our results suggest that in areas where historical disturbance regimes were characterized by large, infrequent fires, management of forest age classes based on a range of historical variability may be feasible only at relatively large spatial scales. Comprehensive landscape management strategies will need to consider other factors besides the percentage of old forests on the landscape, including the spatial pattern of stands and the rates and pathways of landscape change.     

Effects of Weed-Management Burning on Reptile Assemblages in Australian Tropical Savannas

Abstract: Fire is frequently used for land management purposes and may be crucial for effective control of invasive non‐native plants. Nevertheless, fire modifies environments and may affect nontarget native biodiversity, which can cause conflicts for conservation managers. Native Australian reptiles avoid habitat invaded by the alien plant rubber vine (Cryptostegia grandiflora) and may be susceptible to the impacts of burning, a situation that provides a model system in which to examine possible conservation trade‐offs between managing invasive plants and maintaining native biodiversity. We used replicated, experimental fire treatments (unburned, dry‐season burned, and wet‐season burned) in 2 habitats (riparian and adjacent open woodland) to examine the short‐ (within 12 months of fire) and longer‐term (within 3 years of fire) changes of reptile assemblages in response to wet‐ and dry‐season burning for weed management in tropical savannas of northern Australia. Within 12 months of fire, abundances of the skink Carlia munda (Scincidae) were higher in the burned sites, but overall reptile composition was structured by habitat type rather than by effects of burning. Within 3 years of a fire, the effects of fire were evident. Reptiles, especially the gecko Heteronotia binoei (Gekkonidae), were least abundant in dry‐season burned sites; litter‐associated species, including Carlia pectoralis (Scincidae), were rarely observed in burned habitat; and there were fewer species in the wet‐season burned sites. Reptile abundance was associated with vegetation structure, which suggests that fire‐induced changes detrimentally altered the availability of resources for some reptiles, particularly leaf‐litter species. Invasive alien plants, such as rubber vine, have severe effects on native biodiversity, and control of such species is a fundamental land management objective. Nevertheless, fire management of invasive alien plants may adversely affect native biodiversity, creating a conservation conundrum. In such scenarios, land managers will need to identify the most desired conservation goal and consider the consequences for native biota.     

Spatial occurrence of a habitat-tracking saproxylic beetle inhabiting a managed forest landscape.

Because of the dynamic nature of many managed habitats, proper evaluation of conservation efforts calls for models that take into account both spatial and temporal habitat dynamics. We develop a metapopulation model for successional-type systems, in which habitat quality changes over time in a predictable fashion. The occupancy and recruitment of the predatory saproxylic (dependent on dead wood) beetle Harminius undulatus was studied in a managed boreal forest landscape, covering 24,449 ha, in central Sweden. In a first step, we analyzed the beetle's occupancy pattern in relation to stand characteristics, and the amounts of present and past habitat in the surrounding landscape. Managed forest is suitable habitat when > or =60 years old, and immediately after cutting, but not between the ages of 10 and 60 years. The observed occupancy of H. undulatus was positively correlated with the stand's age as habitat. We used a metapopulation model to predict the current probability of occurrence in each forest stand, given the spatiotemporal distribution of suitable forest stands during the last 50 years. Metapopulation parameters were estimated by matching predicted spatial distributions with observed spatial distributions. The model predicted observed spatial distributions better than a similar model that assumed constant habitat quality of each forest stand. Thus, metapopulation models for successional-type systems, such as dead wood dependent organisms in managed forest landscapes, should include habitat dynamics. An estimated 82% of the landscape-wide recruitment took place in managed stands, which covered 87% of the forest area, in comparison with 18% in unmanaged stands, which covered 13% of the forest area. Among the managed stand types, > or =60-year-old stands and 3-7-year-old clear-cuttings contributed to 79% of the total recruitment while 8-59-year-old stands only contributed 3%. The results suggest the following guidelines to improve conditions for H. undulatus and other species with similar habitat requirements: (1) the proportion of the landscape constituted by younger stands should not be allowed to grow too large, (2) the rotation period of managed stands should not be allowed to be too short, and (3) dead wood should be retained and created at final cutting.     

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.