Patterns of spatiotemporal change in large mammal distribution and abundance in the southern Western Ghats, India

Large mammals face high risks of anthropogenic extinction owing to their larger body mass and associated life history traits. Recent worldwide mammal declines have highlighted the conservation importance of effective assessments of trends in distribution and abundance of species. Yet reliable data depicting the nature and extent of changes in population parameters is sparse, primarily due to logistical problems in covering large areas and difficulties in obtaining reliable information at large spatial scales, particularly over time. We used key informant surveys to generate detection histories for 18 species of large mammals (body mass > 2 kg) at two points in time (present and 30 years ago) in the Southern subregion of the Western Ghats global biodiversity hotspot. Multiple-season occupancy models were used to assess temporal trends in occupancy, detectability and vital rates of extinction and colonization for each species. Our results show significant declines in distribution for large carnivores, the Asian elephant and endemic ungulates and primates. There is a significant decline in detectability for 16 species, which suggests a decline in their abundance. These patterns of change in distribution and abundance repeat in our assessments of spatial variation in occupancy dynamics between the three contiguous forest complexes and two human-dominated landscapes into which the southern Western Ghats has been fragmented. Extinction rates are highest in the human-dominated landscapes. Declines in abundance for several species suggest the presence of extinction debts, which may soon be repaid with imminent range contractions and subsequent species extinctions unless immediate remedial conservation measures are taken. Detection/non-detection surveys of key informants used in an occupancy modeling framework provide potential for rapid conservation status assessments of multiple species across large spatial scales over time.     

Monitoring distributions using call surveys: estimating site occupancy, detection probabilities and inferring absence

Monitoring programs serve to track changes in the distribution and abundance of species. A major problem with most monitoring programs is that species detection is imperfect and some populations are inevitably missed. Therefore, in most monitoring programs the true distribution of a species will be underestimated. Here, we report a field test of the reliability and performance of a commonly used method to monitor the distribution of amphibians (anuran call surveys). We surveyed the distribution of four anuran species in western Switzerland, and estimated detection probabilities to account for imperfect species detection and used these estimates to adjust our estimate of site occupancy (i.e., distribution). Next, we assessed how detection probabilities were affected by weather and how site occupancy was affected by site specific covariates. For one species (Hyla arborea), call surveys proved efficient in determining the regional distribution with only few site visits because detection probabilities were relatively high. The call surveys apparently missed many populations of another common species (Bufo calamita) because detection probabilities were lower. Two other species (Bombina variegata and Alytes obstetricans) were uncommon and strong inference from the analysis is not possible. Thus, multispecies surveys may be inefficient for rare species. Estimates of detection probabilities were used to calculate how many site visits are necessary to infer the absence of a species with some predetermined statistical certainty. The implications of “false absences” are important in ecology as they are known to bias usual habitat suitability models and overestimate extinction/colonization events in metapopulations. Large-scale monitoring programs would benefit from the application of an estimation-based approach to monitoring the distribution of species.     

Brown hyaenas on roads: Estimating carnivore occupancy and abundance using spatially auto-correlated sign survey replicates

Carnivore survey protocols that properly address spatial sampling and detectability issues are seldom feasible at a landscape-scale. This limits knowledge of large-scale patterns in distribution, abundance and their underlying determinants, hindering conservation of globally threatened carnivore populations. Occupancy analysis of data from logistically efficient sign surveys along consecutive road segments (spatially auto-correlated replicates) offers a potential solution. We adapted and applied this newly-developed method over 62,979 km² of human-modified land in South Africa. Our aims were to (1) generate unbiased estimates of brown hyaena occupancy and abundance (2) investigate two suspected determinants of occupancy using a combination of biological and socio-economic sampling techniques, and (3) use simulations to evaluate the effort required for abundance and occupancy estimates with acceptable bias, precision and power. Brown hyaena occupancy was estimated at 0.748 (±SE 0.1), and estimated overall density in agricultural land (0.15/100 km², ±SE 0.08) was an order of magnitude lower than in protected areas. Positive attitudes to carnivores and presence of wildlife farms exerted strong positive effects on occupancy, so changes in these factors may well exert monotonic impacts on local metapopulation status. Producing reliable occupancy and abundance estimates would require ?6 replicates and ?12 replicates per site respectively. Detecting 50% and 30% declines in brown hyaena occupancy with adequate power would require five annual surveys at ?65 sites and ?125 sites respectively. Our results suggest that protocols based on spatially auto-correlated sign survey replicates could be used to monitor carnivore populations at large, and possibly even country-wide spatial scales.     

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.     

Using occupancy estimation to assess the effectiveness of a regional multiple-species conservation plan: Bats in the Pacific Northwest

Regional conservation plans are increasingly used to plan for and protect biodiversity at large spatial scales however the means of quantitatively evaluating their effectiveness are rarely specified. Multiple-species approaches, particular those which employ site-occupancy estimation, have been proposed as robust and efficient alternatives for assessing the status of wildlife populations over large spatial scales, but implemented examples are few. I used bats as a model to evaluate design considerations for the use of occupancy estimation to assess population status and habitat associations for eight species of bats covered under a regional conservation plan. Bats were one of the groups expected to benefit from a system of reserves for species associated with late-successional/old-growth (LSOG) habitat designated under the Northwest Forest Plan (NWFP). This study produced the first estimates for probabilities of occupancy and detection for bats at a regional scale. Overall probabilities of occupancy ranged from 0.586 to 0.783 and probabilities of detection ranged from 0.239 to 0.532 among the eight species modeled. Although point estimates of occupancy suggested association with NWFP habitat categories for some species, estimate precision was low. Models that assumed constant occupancy with respect to reserve- and LSOG-status were supported for most species. I used model-averaged estimates of occupancy and detection for each species to estimate survey effort necessary to meet precision targets. Occupancy estimation was best suited to species with the highest detection probabilities. Species that are rare or difficult to detect will require enhancement in survey methods or more intensive survey effort to produce meaningful estimates. Optimizing monitoring efforts to address multiple species requires tradeoffs among survey methods, levels of effort, and acceptable levels of precision.     

Dynamic multistate site occupancy models to evaluate hypotheses relevant to conservation of Golden Eagles in Denali National Park, Alaska

The recent development of multistate site occupancy models offers great opportunities to frame and solve decision problems for conservation that can be viewed in terms of site occupancy. These models have several characteristics (e.g., they account for detectability) that make them particularly well suited for addressing management and conservation problems. We applied multistate site occupancy models to evaluate hypotheses related to the conservation and management of Golden Eagles (Aquila chrysaetos) in Denali National Park, Alaska, and provided estimates of transition probabilities among three occupancy states for nesting areas (occupied with successful reproduction, occupied with unsuccessful reproduction, and unoccupied). Our estimation models included the effect of potential recreational activities (hikers) and environmental covariates such as a snowshoe hare (Lepus americanus) index on transition probabilities among the three occupancy states. Based on the most parsimonious model, support for the hypothesis of an effect of potential human disturbance on site occupancy dynamics was equivocal. There was some evidence that potential human disturbance negatively affected local colonization of territories, but there was no evidence of an effect on reproductive performance parameters. In addition, models that assume a positive relationship between the hare index and successful reproduction were well supported by the data. The statistical approach that we used is particularly useful to parameterize management models that can then be used to make optimal decisions related to the management of Golden Eagles in Denali. Although in our case we were particularly interested in managing recreational activities, we believe that such models should be useful to for a broad class of management and conservation problems.     

Conservation cornerstones: Capitalising on the endeavours of long-term monitoring projects

Ecological monitoring is widely used to measure change through time in ecosystems. The current extinction crisis has resulted in a wealth of monitoring programs focussed on tracking the status of threatened species, and the perceived importance of monitoring has seen it become the cornerstone of many biodiversity conservation programs. However, many monitoring programs fail to produce useful outcomes due to inherent flaws. Here we use a monitoring program from south-eastern Australia as a case study to illustrate the potential of such endeavours. The threatened carnivorous marsupial, the brush-tailed phascogale (Phascogale tapoatafa), has been monitored at various locations between 2000 and 2010. We present strong evidence for a decline in relative abundance during this period, and also describe relationships with environmental variables. These results provide insights likely to be valuable in guiding future management of the species. In the absence of the monitoring program, informed management would not be possible. While early detection of population declines is important, knowledge of the processes driving such declines is required for effective intervention. We argue that monitoring programs will be most effective as a tool for enhanced conservation management if they test specific hypotheses relating to changes in population trajectories. Greater emphasis should be placed on rigorous statistical analysis of monitoring datasets in order to capitalise on the resources devoted to monitoring activities. Many datasets are likely to exist for which careful analysis of results would have benefits for determining management directions.     

Two-Stage Bayesian Study Design for Species Occupancy Estimation

A problem of interest for ecology and conservation is that of determining the best allocation of survey effort in studies aimed at estimating the proportion of sites occupied by a species. Many species are difficult to detect and often remain undetected during surveys at sites where they are present. Hence, for the estimator of species occupancy to be unbiased, detectability needs to be taken into account. In such studies there is a trade-off between sampling more sites and expending more survey effort within each site. This design problem has not been addressed to date with an explicit consideration of the uncertainty in assumed parameter values. In this article we apply sequential and Bayesian design techniques and show how a simple two-stage design can significantly improve the efficiency of the study. We further investigate the optimal allocation of survey effort between the two study stages, given a prior distribution for the parameter values. We address this problem using asymptotic approximations and then explore how the results change when the sample size is small, considering second-order approximations and highlighting the value of simulations as a tool for study design. Given the efficiency gain, we recommend following the sequential design approach for species occupancy estimation. This article has supplementary material online.     

Grid-based monitoring methods for detecting population declines: Sensitivity to spatial scale and consequences of scale correction

A decrease in the area of occupancy of a species is often criterion by which a species is classified as threatened (i.e. IUCN Red List). However, it is unclear how to accurately measure change in area of occupancy. Area of occupancy is a measure of distribution and the spatial scale at which the distribution is measured will affect the ability to detect a decline. To overcome errors introduced by measuring distribution at different spatial scales, scale correction methods are often applied. It is known that scale correction reliably estimates area of occupancy; however, its suitability to estimate trends in area of occupancy has not been assessed. We investigate the effect of spatial scale and implementation of scale correction when estimating two different forms of decline in area of occupancy: spatially correlated and spatially uncorrelated declines. We explore these issues using simulations of three declining species and a grid-based monitoring method designed to detect the declines. Our results suggest that current grid-based methods are inadequate for detecting uncorrelated local extinctions, even if the total decline is substantial (e.g. 65% of the original range). We demonstrate that scale correction will lead to misleading conclusions in some situations. We suggest an alternative survey method to accurately estimate changes in area of occupancy over time. We provide guidelines for designing grid surveys to measure changes in area of occupancy to assess a species threat status using the IUCN Red List criterion.     

Needles in haystacks: Estimating detection probability and occupancy of rare and cryptic snakes

The species most in need of conservation or management are often also the most difficult to monitor, because of their rarity, secretive habits, or both. To combat these challenges, presence/absence (site occupancy) models can be used to track species occupancy at landscape scales. However, quantitative knowledge of detection probability (which is almost always <1) is required to reliably estimate site occupancy. Here, we present a case study that combines detection probabilities and site occupancy modeling to monitor a notoriously secretive guild of animals, North American aquatic snakes. Specifically, we use program PRESENCE to estimate detection probability (p) and probability of site occupancy (ψ) for seven snake species in relationship to site covariates, to understand the proximate and ultimate factors that influence habitat suitability. We were able to estimate p (3–46%) and ψ (12–96%) for each species and calculate the amount of unsuccessful effort necessary to declare absence of each species with statistical confidence (5–63 visits; 150–1890 trap-nights). We documented considerable interspecific variation in p and ψ; one species (Nerodia fasciata) was widespread and highly detectable, while another (Agkistrodon piscivorus) had low detectability despite its wide distribution. Five other species were secretive, or restricted to specific habitat types, or both, illustrating that complex and sometimes counterintuitive relationships exist between capture rate and occupancy. Incorporating p and ψ is essential to the success of large-scale monitoring programs for elusive species.     

Modeling co-occurrence of northern spotted and barred owls: Accounting for detection probability differences

Barred owls (Strix varia) have recently expanded their range and now encompass the entire range of the northern spotted owl (Strix occidentalis caurina). This expansion has led to two important issues of concern for management of northern spotted owls: (1) possible competitive interactions between the two species that could contribute to population declines of northern spotted owls, and (2) possible changes in vocalization behavior and detection probabilities of northern spotted owls induced by presence of barred owls. We used a two-species occupancy model to investigate whether there was evidence of competitive exclusion between the two species at study locations in Oregon, USA. We simultaneously estimated detection probabilities for both species and determined if the presence of one species influenced the detection of the other species. Model selection results and associated parameter estimates provided no evidence that barred owls excluded spotted owls from territories. We found strong evidence that detection probabilities differed for the two species, with higher probabilities for northern spotted owls that are the object of current surveys. Non-detection of barred owls is very common in surveys for northern spotted owls, and detection of both owl species was negatively influenced by the presence of the congeneric species. Our results suggest that analyses directed at hypotheses of barred owl effects on demographic or occupancy vital rates of northern spotted owls need to deal adequately with imperfect and variable detection probabilities for both species.     

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.     

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.     

The use of historical collections to estimate population trends: A case study using Swedish longhorn beetles (Coleoptera: Cerambycidae)

Long term data to estimate population trends among species are generally lacking. However, Natural History Collections (NHCs) can provide such information, but may suffer from biases due to varying sampling effort. To analyze population trends and range-abundance dynamics of Swedish longhorn beetles (Coleoptera: Cerambycidae), we used collections of 108 species stretching over 100 years. We controlled for varying sampling effort by using the total number of database records as a reference for non-red-listed species. Because the general frequency of red-listed species increased over time, a separate estimate of sampling effort was used for that group. We observed large interspecific variation in population changes, from declines of 60% to several hundred percent increases. Most species showed stable or increasing ranges, whereas few seemed to decline in range. Among increasing species, rare species seemed to expand their range more than common species did, but this pattern was not observed in declining species. Historically, rare species did not seem to be at larger risk of local extinction, and population declines were mostly due to lower population density and not loss of sub-populations. We also evaluated the species’ declines under IUCN red-list criterion A, and four currently not red-listed species meet the suggested threshold for Near Threatened (NT). The results also suggested that species’ declines may be overlooked if estimated only from changes in species range.     

Assessment of cropping‐system impact on soil organic matter levels in short‐term field experiments

It has been suggested that short‐term field experiments are not suitable for the quantitative assessment of cropping‐systems impact on soil organic matter (SOM) levels in arable soils, as expectable temporal changes are very small compared to a large spatial variation of SOM background levels. However, applying an optimized sampling design based on repeated sampling in small plots, we were able to detect soil total carbon (STC) and nitrogen (STN) changes in the magnitude of ≈ 1% (STC) and ≈ 2% (STN) of background levels with only four replications, respectively. Gradually enlarging the sample size up to n = 24 did not considerably improve change detectability with STC, but with STN (n = 15 allowing for the dection of ≈ 1% change of background levels). The common calculation of minimum detectable differences (MDD) based on a state analysis of SOM levels instead of repeated measurements considerably underestimated change detectability.     

Patch-occupancy models indicate human activity as major determinant of forest elephant Loxodonta cyclotis seasonal distribution in an industrial corridor in Gabon

The importance of human activity and ecological features in influencing African forest elephant ranging behaviour was investigated in the Rabi-Ndogo corridor of the Gamba Complex of Protected Areas in southwest Gabon. Locations in a wide geographical area with a range of environmental variables were selected for patch-occupancy surveys using elephant dung to assess seasonal presence and absence of elephants. Patch-occupancy procedures allowed for covariate modelling evaluating hypotheses for both occupancy in relation to human activity and ecological features, and detection probability in relation to vegetation density. The best fitting models for old and fresh dung data sets indicate that (1) detection probability for elephant dung is negatively related to the relative density of the vegetation, and (2) human activity, such as presence and infrastructure, are more closely associated with elephant distribution patterns than are ecological features, such as the presence of wetlands and preferred fresh fruit. Our findings emphasize the sensitivity of elephants to human disturbance, in this case infrastructure development associated with gas and oil production. Patch-occupancy methodology offers a viable alternative to current transect protocols for monitoring programs with multiple covariates.     

基于空间模拟退火算法的耕地质量布样及优化方法

耕地质量监测是保障耕地资源的永续利用,实现耕地产能提升、加强耕地资源的管理、保护、合理利用的重要措施,对实现持续粮食安全具有重要意义。该文提出了基于空间模拟退火算法的耕地质量布样优化方法,以空间模拟退火算法为基础生成一组最优样本,构成基础监测网络,在此基础上,通过多期耕地等级成果数据提取属性发生变化的分等因素和对应发生变化的区域,生成潜在变化区,并结合研究区实际情况辅以专家知识和异常监测点,对基础样本点进行增加、删除、替换等优化操作,生成最终监测样点。以北京市大兴区为例,最终确定布设55个监测样点,结果表明,该方法布设的样点在耕地质量预测方面的精度高于传统的随机抽样和分层抽样方法,能有效地预测县域耕地质量并监控耕地质量的变化情况。     

Modelling the effect of El Niño on the persistence of small populations: The Galápagos penguin as a case study

Small populations are vulnerable to long-term declines, even where short-term censuses indicate increasing trends in numbers. Census data for the Galápagos penguin (Spheniscus mendiculus) collected between 1970 and 2004 provide evidence that despite year-to-year population increases detected in most of the annual censuses, the strong El Niño events of 1982-83 and 1997-98 were followed by population declines of more than 60% from which the species has yet to recover. Such large declines raise concerns about the future viability of the species because the frequency and severity of El Niño events are predicted to increase. We used the simulation software VORTEX to evaluate the potential effects of El Niño on the risk of extinction of the Galápagos penguin population and its four constituent subpopulations. Weak and strong El Niño events were treated as catastrophes, with varying frequencies, which simulated past, current and future effects on the penguin population. The “Current El Niño” scenario, based on the frequency of El Niño events recorded in the Galápagos between 1965 and 2004, indicated an approximately 30% probability of extinction within the next 100 years for the penguin population. More ominously, the species may be at a greater risk if the frequency of strong El Niño episodes increases only marginally. A probability of extinction greater than 80% was predicted when the current frequency (5%) of strong El Niño events was doubled (to 10%). The probabilities of extinctions were higher for each subpopulation treated individually, ranging from 34% for Isabela and Fernandina, 64% for Bartolomé-Santiago to 78% for the smallest subpopulation on Floreana. Sensitivity analyses identified survival of penguins during El Niño events and sex ratio as influential parameters. The estimates of extinction risk may be conservative as other threats associated with increased human activities on the islands may further compromise species persistence.     

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.     

Assessing the extinction vulnerability of wood-inhabiting fungal species in fragmented northern Swedish boreal forests

Fragmentation of old-growth forests and greatly reduced amounts of coarse dead wood in managed forests threat the persistence of many saproxylic species in boreal Fennoscandia. Individual old-growth forest remnants may lose species over time as they pay off their extinction debt. We tested this by comparing the observed site occupancy of individual wood-inhabiting fungal species in isolated old-growth stands (i.e. woodland key habitats; WKHs) with statistical predictions of their occupancy assuming potential extinction debt had already been paid off. The occupancy of species was analysed in two sets of WKHs differing in time since isolation (i.e. recent and old isolates).Few species occurred more frequently than expected in WKHs. However, patterns across species and across all WKHs masked important differences among species in their risk of facing future extinction. The site occupancy decreased significantly between recent and old isolates for a group of annual, red-listed specialist fungal species, suggesting that an extinction debt in WKHs may exist among specific species confined to coarse dead wood and old-growth forest habitat. Generalist species that also occur in the surrounding matrix showed no negative trends, or actually increased in site occupancy, making future extinctions less likely. Thus, continuing loss of threatened species are likely if not preservation of WKHs are combined with other conservation efforts in managed forest landscapes. Natural forest landscapes may serve as important references when aiming to identify species in risk of future extinction but more detailed knowledge about the biology of the most vulnerable species is also required.