Improving effectiveness of systematic conservation planning with density data

Systematic conservation planning aims to design networks of protected areas that meet conservation goals across large landscapes. The optimal design of these conservation networks is most frequently based on the modeled habitat suitability or probability of occurrence of species, despite evidence that model predictions may not be highly correlated with species density. We hypothesized that conservation networks designed using species density distributions more efficiently conserve populations of all species considered than networks designed using probability of occurrence models. To test this hypothesis, we used the Zonation conservation prioritization algorithm to evaluate conservation network designs based on probability of occurrence versus density models for 26 land bird species in the U.S. Pacific Northwest. We assessed the efficacy of each conservation network based on predicted species densities and predicted species diversity. High‐density model Zonation rankings protected more individuals per species when networks protected the highest priority 10‐40% of the landscape. Compared with density‐based models, the occurrence‐based models protected more individuals in the lowest 50% priority areas of the landscape. The 2 approaches conserved species diversity in similar ways: predicted diversity was higher in higher priority locations in both conservation networks. We conclude that both density and probability of occurrence models can be useful for setting conservation priorities but that density‐based models are best suited for identifying the highest priority areas. Developing methods to aggregate species count data from unrelated monitoring efforts and making these data widely available through ecoinformatics portals such as the Avian Knowledge Network will enable species count data to be more widely incorporated into systematic conservation planning efforts.     

Killing Tigers to Save Them: Fallacies of the Farming Argument

Abstract: The lucrative, illegal trade in tigers (Panthera tigris) remains a major conservation problem. Tiger farming has been proposed as a potential solution, with farmed tigers substituting for wild tigers. At first glance, this argument's logic seems simple: farming will increase the supply of tigers, prices will fall, and poaching will no longer be profitable. We contend, however, that this supply‐side argument relies on mistaken assumptions. First, tiger markets are imperfect, meaning they are dominated by a few producers who control price. Second, consumers prefer wild tigers to farmed tigers and therefore the two are not pure substitutes. In economic terms, products from wild tigers are luxury goods, commanding a price premium. Third, there is no evidence that farmed tigers can be produced or sold more cheaply than wild tigers. In sum, it is unlikely that farming will drive down the price of wild‐caught tigers or decrease profitability for tiger poachers. Rather, tiger farming is more likely to increase aggregate demand for tiger products and stimulate higher levels of poaching.     

Effects of national forest‐management regimes on unprotected forests of the Himalaya

Globally, deforestation continues, and although protected areas effectively protect forests, the majority of forests are not in protected areas. Thus, how effective are different management regimes to avoid deforestation in non‐protected forests? We sought to assess the effectiveness of different national forest‐management regimes to safeguard forests outside protected areas. We compared 2000–2014 deforestation rates across the temperate forests of 5 countries in the Himalaya (Bhutan, Nepal, China, India, and Myanmar) of which 13% are protected. We reviewed the literature to characterize forest management regimes in each country and conducted a quasi‐experimental analysis to measure differences in deforestation of unprotected forests among countries and states in India. Countries varied in both overarching forest‐management goals and specific tenure arrangements and policies for unprotected forests, from policies emphasizing economic development to those focused on forest conservation. Deforestation rates differed up to 1.4% between countries, even after accounting for local determinants of deforestation, such as human population density, market access, and topography. The highest deforestation rates were associated with forest policies aimed at maximizing profits and unstable tenure regimes. Deforestation in national forest‐management regimes that emphasized conservation and community management were relatively low. In India results were consistent with the national‐level results. We interpreted our results in the context of the broader literature on decentralized, community‐based natural resource management, and our findings emphasize that the type and quality of community‐based forestry programs and the degree to which they are oriented toward sustainable use rather than economic development are important for forest protection. Our cross‐national results are consistent with results from site‐ and regional‐scale studies that show forest‐management regimes that ensure stable land tenure and integrate local‐livelihood benefits with forest conservation result in the best forest outcomes.     

Effects of an Exotic Plant Invasion on Native Understory Plants in a Tropical Dry Forest

Abstract: The dry forests of southern India, which are endangered tropical ecosystems and among the world's most important tiger (Panthera tigris) habitats, are extensively invaded by exotic plants. Yet, experimental studies exploring the impacts of these invasions on native plants in these forests are scarce. Consequently, little is known about associated implications for the long‐term conservation of tigers and other biodiversity in these habitats. I studied the impacts of the exotic plant Lantana camara on understory vegetation in a dry‐forest tiger habitat in southern India. I compared the richness, composition, and abundance of tree seedlings, herbs, and shrubs and the abundance of grass among plots in which Lantana was cleared or left standing. These plots were distributed across two blocks—livestock free and livestock grazed. Removal of Lantana had an immediate positive effect on herb–shrub richness in the livestock‐free block, but had no effect on that of tree seedlings in either livestock block. Tree‐seedling and herb–shrub composition differed significantly between Lantana treatment and livestock block, and Lantana removal significantly decreased survival of tree seedlings. Nevertheless, the absence of trees, in any stage between seedling and adult, indicates that Lantana may stall tree regeneration. Lantana removal decreased the abundance of all understory strata, probably because forage plants beneath Lantana are less accessible to herbivores, and plants in Lantana‐free open plots experienced greater herbivory. Reduced access to forage in invaded habitats could negatively affect ungulate populations and ultimately compromise the ability of these forests to sustain prey‐dependent large carnivores. Additional research focused on understanding and mitigating threats posed by exotic plants may be crucial to the long‐term protection of these forests as viable tiger habitats.     

Ecological‐Niche Modeling and Prioritization of Conservation‐Area Networks for Mexican Herpetofauna

Abstract: One of the most important tools in conservation biology is information on the geographic distribution of species and the variables determining those patterns. We used maximum‐entropy niche modeling to run distribution models for 222 amphibian and 371 reptile species (49% endemics and 27% threatened) for which we had 34,619 single geographic records. The planning region is in southeastern Mexico, is 20% of the country's area, includes 80% of the country's herpetofauna, and lacks an adequate protected‐area system. We used probabilistic data to build distribution models of herpetofauna for use in prioritizing conservation areas for three target groups (all species and threatened and endemic species). The accuracy of species‐distribution models was better for endemic and threatened species than it was for all species. Forty‐seven percent of the region has been deforested and additional conservation areas with 13.7% to 88.6% more native vegetation (76% to 96% of the areas are outside the current protected‐area system) are needed. There was overlap in 26 of the main selected areas in the conservation‐area network prioritized to preserve the target groups, and for all three target groups the proportion of vegetation types needed for their conservation was constant: 30% pine and oak forests, 22% tropical evergreen forest, 17% low deciduous forest, and 8% montane cloud forests. The fact that different groups of species require the same proportion of habitat types suggests that the pine and oak forests support the highest proportion of endemic and threatened species and should therefore be given priority over other types of vegetation for inclusion in the protected areas of southeastern Mexico.     

Dynamics of a low‐density tiger population in Southeast Asia in the context of improved law enforcement

Recovering small populations of threatened species is an important global conservation strategy. Monitoring the anticipated recovery, however, often relies on uncertain abundance indices rather than on rigorous demographic estimates. To counter the severe threat from poaching of wild tigers (Panthera tigris), the Government of Thailand established an intensive patrolling system in 2005 to protect and recover its largest source population in Huai Kha Khaeng Wildlife Sanctuary. Concurrently, we assessed the dynamics of this tiger population over the next 8 years with rigorous photographic capture‐recapture methods. From 2006 to 2012, we sampled across 624–1026 km² with 137–200 camera traps. Cameras deployed for 21,359 trap days yielded photographic records of 90 distinct individuals. We used closed model Bayesian spatial capture‐recapture methods to estimate tiger abundances annually. Abundance estimates were integrated with likelihood‐based open model analyses to estimate rates of annual and overall rates of survival, recruitment, and changes in abundance. Estimates of demographic parameters fluctuated widely: annual density ranged from 1.25 to 2.01 tigers/100 km², abundance from 35 to 58 tigers, survival from 79.6% to 95.5%, and annual recruitment from 0 to 25 tigers. The number of distinct individuals photographed demonstrates the value of photographic capture–recapture methods for assessments of population dynamics in rare and elusive species that are identifiable from natural markings. Possibly because of poaching pressure, overall tiger densities at Huai Kha Khaeng were 82–90% lower than in ecologically comparable sites in India. However, intensified patrolling after 2006 appeared to reduce poaching and was correlated with marginal improvement in tiger survival and recruitment. Our results suggest that population recovery of low‐density tiger populations may be slower than anticipated by current global strategies aimed at doubling the number of wild tigers in a decade.     

Use of Inverse Spatial Conservation Prioritization to Avoid Biological Diversity Loss Outside Protected Areas

Globally expanding human land use sets constantly increasing pressure for maintenance of biological diversity and functioning ecosystems. To fight the decline of biological diversity, conservation science has broken ground with methods such as the operational model of systematic conservation planning (SCP), which focuses on design and on‐the‐ground implementation of conservation areas. The most commonly used method in SCP is reserve selection that focuses on the spatial design of reserve networks and their expansion. We expanded these methods by introducing another form of spatial allocation of conservation effort relevant for land‐use zoning at the landscape scale that avoids negative ecological effects of human land use outside protected areas. We call our method inverse spatial conservation prioritization. It can be used to identify areas suitable for economic development while simultaneously limiting total ecological and environmental effects of that development at the landscape level by identifying areas with highest economic but lowest ecological value. Our method is not based on a priori targets, and as such it is applicable to cases where the effects of land use on, for example, individual species or ecosystem types are relatively small and would not lead to violation of regional or national conservation targets. We applied our method to land‐use allocation to peat mining. Our method identified a combination of profitable production areas that provides the needed area for peat production while retaining most of the landscape‐level ecological value of the ecosystem. The results of this inverse spatial conservation prioritization are being used in land‐use zoning in the province of Central Finland.     

Applying network theory to prioritize multispecies habitat networks that are robust to climate and land‐use change

Designing connected landscapes is among the most widespread strategies for achieving biodiversity conservation targets. The challenge lies in simultaneously satisfying the connectivity needs of multiple species at multiple spatial scales under uncertain climate and land‐use change. To evaluate the contribution of remnant habitat fragments to the connectivity of regional habitat networks, we developed a method to integrate uncertainty in climate and land‐use change projections with the latest developments in network‐connectivity research and spatial, multipurpose conservation prioritization. We used land‐use change simulations to explore robustness of species’ habitat networks to alternative development scenarios. We applied our method to 14 vertebrate focal species of periurban Montreal, Canada. Accounting for connectivity in spatial prioritization strongly modified conservation priorities and the modified priorities were robust to uncertain climate change. Setting conservation priorities based on habitat quality and connectivity maintained a large proportion of the region's connectivity, despite anticipated habitat loss due to climate and land‐use change. The application of connectivity criteria alongside habitat‐quality criteria for protected‐area design was efficient with respect to the amount of area that needs protection and did not necessarily amplify trade‐offs among conservation criteria. Our approach and results are being applied in and around Montreal and are well suited to the design of ecological networks and green infrastructure for the conservation of biodiversity and ecosystem services in other regions, in particular regions around large cities, where connectivity is critically low.     

Response of Neotropical Bat Assemblages to Human Land Use

Neotropical bats are sensitive to human‐induced habitat changes, and some authors believe bats can be used as bioindicators. In the literature, however, the results are disparate. Some results show bat diversity deceases as disturbance increases, whereas others indicate no effect. Determining the general response patterns of bats when they encounter different degrees of human‐induced disturbance across the Neotropics would help to determine their usefulness as bioindicators. In a series of meta‐analyses, we compared the occurrence frequency of bat species between well‐preserved forests and human‐use areas. We obtained data through an extensive review of published peer‐reviewed articles, theses, and reports. The overall effect size indicated that human‐use areas harbored more bat species than well‐preserved forests. Different response patterns emerged when meta‐analyses were conducted separately by family, feeding habit, vegetation stratum, and conservation status. Our results suggest that bat assemblages display strong responses to forest loss and land‐use change and that the direction and magnitude of these responses depends on the bat group under study and the type of disturbance. Our results are consistent with the idea that bats are useful for assessing the effects of habitat changes in the Neotropics. However, with our meta‐analyses we could not detect fine differences in bat feeding habits, especially within Phyllostomidae, or elucidate the effect of landscape configuration. Respuesta de Ensambles de Murciélagos Neotropicales al Uso de Suelo por Humanos     

A Multispecies Framework for Landscape Conservation Planning

Abstract: Rapidly changing landscapes have spurred the need for quantitative methods for conservation assessment and planning that encompass large spatial extents. We devised and tested a multispecies framework for conservation planning to complement single‐species assessments and ecosystem‐level approaches. Our framework consisted of 4 elements: sampling to effectively estimate population parameters, measuring how human activity affects landscapes at multiple scales, analyzing the relation between landscape characteristics and individual species occurrences, and evaluating and comparing the responses of multiple species to landscape modification. We applied the approach to a community of terrestrial birds across 25,000 km² with a range of intensities of human development. Human modification of land cover, road density, and other elements of the landscape, measured at multiple spatial extents, had large effects on occupancy of the 67 species studied. Forest composition within 1 km of points had a strong effect on occupancy of many species and a range of negative, intermediate, and positive associations. Road density within 1 km of points, percent evergreen forest within 300 m, and distance from patch edge were also strongly associated with occupancy for many species. We used the occupancy results to group species into 11 guilds that shared patterns of association with landscape characteristics. Our multispecies approach to conservation planning allowed us to quantify the trade‐offs of different scenarios of land‐cover change in terms of species occupancy.     

Local and Landscape Correlates of Primate Distribution and Persistence in the Remnant Lowland Rainforests of the Upper Brahmaputra Valley,Northeastern India

Habitat fragmentation affects species distribution and abundance, and drives extinctions. Escalated tropical deforestation and fragmentation have confined many species populations to habitat remnants. How worthwhile is it to invest scarce resources in conserving habitat remnants within densely settled production landscapes? Are these fragments fated to lose species anyway? If not, do other ecological, anthropogenic, and species‐related factors mitigate the effect of fragmentation and offer conservation opportunities? We evaluated, using generalized linear models in an information‐theoretic framework, the effect of local‐ and landscape‐scale factors on the richness, abundance, distribution, and local extinction of 6 primate species in 42 lowland tropical rainforest fragments of the Upper Brahmaputra Valley, northeastern India. On average, the forest fragments lost at least one species in the last 30 years but retained half their original species complement. Species richness declined as proportion of habitat lost increased but was not significantly affected by fragment size and isolation. The occurrence of western hoolock gibbon (Hoolock hoolock) and capped langur (Trachypithecus pileatus) in fragments was inversely related to their isolation and loss of habitat, respectively. Fragment area determined stump‐tailed (Macaca arctoides) and northern pig‐tailed macaque occurrence (Macaca leonina). Assamese macaque (Macaca assamensis) distribution was affected negatively by illegal tree felling, and rhesus macaque (Macaca mulatta) abundance increased as habitat heterogeneity increased. Primate extinction in a fragment was primarily governed by the extent of divergence in its food tree species richness from that in contiguous forests. We suggest the conservation value of these fragments is high because collectively they retained the entire original species pool and individually retained half of it, even a century after fragmentation. Given the extensive habitat and species loss, however, these fragments urgently require protection and active ecological restoration to sustain this rich primate assemblage. Correlaciones Locales y de Paisaje de la Distribución y Persistencia de Primates en los Bosques Lluviosos Remanentes en el Valle del Alto Brahmaputra, Noreste de India     

Linking Management Effectiveness Indicators to Observed Effects of Protected Areas on Fire Occurrence in the Amazon Rainforest

Management‐effectiveness scores are used widely by donors and implementers of conservation projects to prioritize, track, and evaluate investments in protected areas. However, there is little evidence that these scores actually reflect the capacity of protected areas to deliver conservation outcomes. We examined the relation between indicators of management effectiveness in protected areas and the effectiveness of protected areas in reducing fire occurrence in the Amazon rainforest. We used data collected with the Management Effectiveness Tracking Tool (METT) scorecard, adopted by some of the world's largest conservation organizations to track management characteristics believed to be crucial for protected‐area effectiveness. We used the occurrence of forest fires from 2000 through 2010 as a measure of the effect of protected areas on undesired land‐cover change in the Amazon basin. We used matching to compare the estimated effect of protected areas with low versus high METT scores on fire occurrence. We also estimated effects of individual protected areas on fire occurrence and explored the relation between these effects and METT scores. The relations between METT scores and effects of protected areas on fire occurrence were weak. Protected areas with higher METT scores in 2005 did not seem to have performed better than protected areas with lower METT scores at reducing fire occurrence over the last 10 years. Further research into the relations between management‐effectiveness indicators and conservation outcomes in protected areas seems necessary, and our results show that the careful application of matching methods can be a suitable method for that purpose. Vinculación de Indicadores de Efectividad de Manejo con los Efectos Observados de la Ocurrencia de Fuego en Áreas Protegidas en la Amazonia     

Assumption‐versus data‐based approaches to summarizing species’ ranges

For conservation decision making, species’ geographic distributions are mapped using various approaches. Some such efforts have downscaled versions of coarse‐resolution extent‐of‐occurrence maps to fine resolutions for conservation planning. We examined the quality of the extent‐of‐occurrence maps as range summaries and the utility of refining those maps into fine‐resolution distributional hypotheses. Extent‐of‐occurrence maps tend to be overly simple, omit many known and well‐documented populations, and likely frequently include many areas not holding populations. Refinement steps involve typological assumptions about habitat preferences and elevational ranges of species, which can introduce substantial error in estimates of species’ true areas of distribution. However, no model‐evaluation steps are taken to assess the predictive ability of these models, so model inaccuracies are not noticed. Whereas range summaries derived by these methods may be useful in coarse‐grained, global‐extent studies, their continued use in on‐the‐ground conservation applications at fine spatial resolutions is not advisable in light of reliance on assumptions, lack of real spatial resolution, and lack of testing. In contrast, data‐driven techniques that integrate primary data on biodiversity occurrence with remotely sensed data that summarize environmental dimensions (i.e., ecological niche modeling or species distribution modeling) offer data‐driven solutions based on a minimum of assumptions that can be evaluated and validated quantitatively to offer a well‐founded, widely accepted method for summarizing species’ distributional patterns for conservation applications.     

Using citizen science data to identify the sensitivity of species to human land use

Conservation practitioners must contend with an increasing array of threats that affect biodiversity. Citizen scientists can provide timely and expansive information for addressing these threats across large scales, but their data may contain sampling biases. We used randomization procedures to account for possible sampling biases in opportunistically reported citizen science data to identify species’ sensitivities to human land use. We analyzed 21,044 records of 143 native reptile and amphibian species reported to the Carolina Herp Atlas from North Carolina and South Carolina between 1 January 1990 and 12 July 2014. Sensitive species significantly associated with natural landscapes were 3.4 times more likely to be legally protected or treated as of conservation concern by state resource agencies than less sensitive species significantly associated with human‐dominated landscapes. Many of the species significantly associated with natural landscapes occurred primarily in habitats that had been nearly eradicated or otherwise altered in the Carolinas, including isolated wetlands, longleaf pine savannas, and Appalachian forests. Rare species with few reports were more likely to be associated with natural landscapes and 3.2 times more likely to be legally protected or treated as of conservation concern than species with at least 20 reported occurrences. Our results suggest that opportunistically reported citizen science data can be used to identify sensitive species and that species currently restricted primarily to natural landscapes are likely at greatest risk of decline from future losses of natural habitat. Our approach demonstrates the usefulness of citizen science data in prioritizing conservation and in helping practitioners address species declines and extinctions at large extents.     

Spatially Explicit Power Analyses for Occupancy‐Based Monitoring of Wolverine in the U.S. Rocky Mountains

Conservation scientists and resource managers often have to design monitoring programs for species that are rare or patchily distributed across large landscapes. Such programs are frequently expensive and seldom can be conducted by one entity. It is essential that a prospective power analysis be undertaken to ensure stated monitoring goals are feasible. We developed a spatially based simulation program that accounts for natural history, habitat use, and sampling scheme to investigate the power of monitoring protocols to detect trends in population abundance over time with occupancy‐based methods. We analyzed monitoring schemes with different sampling efforts for wolverine (Gulo gulo) populations in 2 areas of the U.S. Rocky Mountains. The relation between occupancy and abundance was nonlinear and depended on landscape, population size, and movement parameters. With current estimates for population size and detection probability in the northern U.S. Rockies, most sampling schemes were only able to detect large declines in abundance in the simulations (i.e., 50% decline over 10 years). For small populations reestablishing in the Southern Rockies, occupancy‐based methods had enough power to detect population trends only when populations were increasing dramatically (e.g., doubling or tripling in 10 years), regardless of sampling effort. In general, increasing the number of cells sampled or the per‐visit detection probability had a much greater effect on power than the number of visits conducted during a survey. Although our results are specific to wolverines, this approach could easily be adapted to other territorial species. Poder de Análisis Espacialmente Explícito para el Monitoreo Basado en Ocupación del Glotón (Gulo gulo) en las Montañas Rocallosas de Estados Unidos     

Value of protected areas to avian persistence across 20 years of climate and land-use change

Establishing protected areas, where human activities and land cover changes are restricted, is among the most widely used strategies for biodiversity conservation. This practice is based on the assumption that protected areas buffer species from processes that drive extinction. However, protected areas can maintain biodiversity in the face of climate change and subsequent shifts in distributions have been questioned. We evaluated the degree to which protected areas influenced colonization and extinction patterns of 97 avian species over 20 years in the northeastern United States. We fitted single-visit dynamic occupancy models to data from Breeding Bird Atlases to quantify the magnitude of the effect of drivers of local colonization and extinction (e.g., climate, land cover, and amount of protected area) in heterogeneous landscapes that varied in the amount of area under protection. Colonization and extinction probabilities improved as the amount of protected area increased, but these effects were conditional on landscape context and species characteristics. In this forest-dominated region, benefits of additional land protection were greatest when both forest cover in a grid square and amount of protected area in neighboring grid squares were low. Effects did not vary with species’ migratory habit or conservation status. Increasing the amounts of land protection benefitted the range margins species but not the core range species. The greatest improvements in colonization and extinction rates accrued for forest birds relative to open-habitat or generalist species. Overall, protected areas stemmed extinction more than they promoted colonization. Our results indicate that land protection remains a viable conservation strategy despite changing habitat and climate, as protected areas both reduce the risk of local extinction and facilitate movement into new areas. Our findings suggest conservation in the face of climate change favors creation of new protected areas over enlarging existing ones as the optimal strategy to reduce extinction and provide stepping stones for the greatest number of species.     

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.     

Paying the Extinction Debt in Southern Wisconsin Forest Understories

Abstract: The lack of long‐term baseline data restricts the ability to measure changes in biological diversity directly and to determine its cause. This hampers conservation efforts and limits testing of basic tenets of ecology and conservation biology. We used a historical baseline survey to track shifts in the abundance and distribution of 296 native understory species across 82 sites over 55 years in the fragmented forests of southern Wisconsin. We resurveyed stands first surveyed in the early 1950s to evaluate the influence of patch size and surrounding land cover on shifts in native plant richness and heterogeneity and to evaluate changes in the relative importance of local site conditions versus the surrounding landscape context as drivers of community composition and structure. Larger forests and those with more surrounding forest cover lost fewer species, were more likely to recruit new species, and had lower rates of homogenization than smaller forests in more fragmented landscapes. Nearby urbanization further reduced both alpha and beta understory diversity. Similarly, understory composition depended strongly on local site conditions in the original survey but only weakly reflected the surrounding landscape composition. By 2005, however, the relative importance of these factors had reversed such that the surrounding landscape structure is now a much better predictor of understory composition than are local site conditions. Collectively, these results strongly support the idea that larger intact habitat patches and landscapes better sustain native species diversity and demonstrate that humans play an increasingly important role in driving patterns of native species diversity and community composition.     

Occupancy Patterns of Regionally Declining Grassland Sparrow Populations in a Forested Pennsylvania Landscape

Organisms can be affected by processes in the surrounding landscape outside the boundary of habitat areas and by local vegetation characteristics. There is substantial interest in understanding how these processes affect populations of grassland birds, which have experienced substantial population declines. Much of our knowledge regarding patterns of occupancy and density stem from prairie systems, whereas relatively little is known regarding how occurrence and abundance of grassland birds vary in reclaimed surface mine grasslands. Using distance sampling and single‐season occupancy models, we investigated how the occupancy probability of Grasshopper (Ammodramus savannarum) and Henslow's Sparrows (A. henslowii) on 61 surface mine grasslands (1591 ha) in Pennsylvania changed from 2002 through 2011 in response to landscape, grassland, and local vegetation characteristics . A subset (n = 23; 784 ha) of those grasslands were surveyed in 2002, and we estimated changes in sparrow density and vegetation across 10 years. Grasshopper and Henslow's Sparrow populations declined 72% and 49%, respectively from 2002 to 2011, whereas overall woody vegetation density increased 2.6 fold. Henslow's Sparrows avoided grasslands with perimeter–area ratios ≥0.141 km/ha and woody shrub densities ≥0.04 shrubs/m². Both species occupied grasslands ≤13 ha, but occupancy probability declined with increasing grassland perimeter–area ratio and woody shrub density. Grassland size, proximity to nearest neighboring grassland ( = 0.2 km), and surrounding landscape composition at 0.5, 1.5, and 3.0 km were not parsimonious predictors of occupancy probability for either species. Our results suggest that reclaimed surface mine grasslands, without management intervention, are ephemeral habitats for Grasshopper and Henslow's Sparrows. Given the forecasted decline in surface coal production for Pennsylvania, it is likely that both species will continue to decline in our study region for the foreseeable future. Patrones de Ocupación de Poblaciones Regionalmente Declinantes de Gorriones de Pastizales en un Paisaje Boscoso de Pennsylvania