Integrating presence-only and occupancy data to model habitat use for the northernmost population of jaguars.

Species distribution models (SDMs) have become an essential tool for the management and conservation of imperiled species. However, many at-risk species are rare and characterized by limited data on their spatial distribution and habitat relationships. This has led to the development of SDMs that integrate multiple types and sources of data to leverage more information and provide improved predictions of habitat associations. We developed a novel integrated species distribution model to predict habitat suitability for jaguars (Panthera onca) in the border region between northern Mexico and the southwestern USA. Our model combined presence-only and occupancy data to identify key environmental correlates, and we used model results to develop a probability of use map. We adopted a logistic regression modeling framework, which we found to be more straightforward and less computationally intensive to fit than Poisson point process-based models. Model results suggested that high terrain ruggedness and the presence of riparian vegetation were most strongly related to habitat use by jaguars in our study region. Our best model, on average, predicted that there is currently 25,463 km2 of usable habitat in our study region. The United States portion of the study region, which makes up 38.6% of the total area, contained 40.6% of the total usable habitat. Even though there have been few detections of jaguars in the southwestern USA in recent decades, our results suggest that protection of currently suitable habitats, along with increased conservation efforts, could significantly contribute to the recovery of jaguars in the USA.

The role of geography, environment, and genetic divergence on the distribution of pikas in the Himalaya.

Pikas (Ochotona Link, 1795) are high-altitude specialist species making them a useful bioindicator species to warming in high-altitude ecosystem. The Himalayan Mountains are an important part of their range, supporting approximately 23%-25% of total pika species worldwide, yet we lack basic information on the distribution patterns. We combine field-based surveys with genetics-based identification and phylogeny to identify differences in species-environment relationships. Further, we suggest putative evolutionary causes for the observed niche patterns. LOCATION: Himalayan high-altitude region. METHODS: We sampled 11 altitudinal transects (ranging from ~2,000 to 5,000 m) in the Himalaya to establish occurrence records. We collected 223 species records using genetic analyses to confirm species' identity (based on some invasive and mostly noninvasive biological samples). Niche and geographic overlap were estimated using kernel density estimates. RESULTS: Most pikas in the Himalaya span wide elevation ranges and exhibit extensive spatial overlap with other species. However, even in areas of high species diversity, we found species to have a distinct environmental niche. Despite apparent overlapping distributions at broad spatial scales, in our field surveys, we encountered few cases of co-occurrence of species in the sampled transects. Deeply diverged sister-species pair had the least environmental niche overlap despite having the highest geographic range overlap. In contrast, sister-species pair with shallow genetic divergence had a higher environmental niche overlap but was geographically isolated. We hypothesize that the extent of environmental niche divergence in pikas is a function of divergence time within the species complex. We assessed vulnerability of species to future climate change using environmental niche and geographic breadth sizes as a proxies. Our findings suggest that O. sikimaria may be the most vulnerable species. Ochotona roylii appears to have the most unique environmental niche space, with least niche overlap with other pika species from the study area.

Modeling spatial variation in density of golden eagle nest sites in the western United States.

In order to contribute to conservation planning efforts for golden eagles (Aquila chrysaetos) in the western U.S., we developed nest site models using >6,500 nest site locations throughout a >3,483,000 km2 area of the western U.S. We developed models for twelve discrete modeling regions, and estimated relative density of nest sites for each region. Cross-validation showed that, in general, models accurately estimated relative nest site densities within regions and sub-regions. Areas estimated to have the highest densities of breeding golden eagles had from 132-2,660 times greater densities compared to the lowest density areas. Observed nest site densities were very similar to those reported from published studies. Large extents of each modeling region consisted of low predicted nest site density, while a small percentage of each modeling region contained disproportionately high nest site density. For example, we estimated that areas with relative nest density values <0.3 represented from 62.8-97.8% ([Formula: see text] = 82.5%) of each modeling area, and those areas contained from 14.7-30.0% ([Formula: see text] = 22.1%) of the nest sites. In contrast, areas with relative nest density values >0.5 represented from 1.0-12.8% ([Formula: see text] = 6.3%) of modeling areas, and those areas contained from 47.7-66.9% ([Formula: see text] = 57.3%) of the nest sites. Our findings have direct application to: 1) large-scale conservation planning efforts, 2) risk analyses for land-use proposals such as recreational trails or wind power development, and 3) identifying mitigation areas to offset the impacts of human disturbance.

Quantifying Ecological Integrity of Terrestrial Systems to Inform Management of Multiple-Use Public Lands in the United States.

The concept of ecological integrity has been applied widely to management of aquatic systems, but still is considered by many to be too vague and difficult to quantify to be useful for managing terrestrial systems, particularly across broad areas. Extensive public lands in the western United States are managed for diverse uses such as timber harvest, livestock grazing, energy development, and wildlife conservation, some of which may degrade ecological integrity. We propose a method for assessing ecological integrity on multiple-use lands that identifies the components of integrity and levels in the ecological hierarchy where the assessment will focus, and considers existing policies and management objectives. Both natural reference and societally desired environmental conditions are relevant comparison points. We applied the method to evaluate the ecological integrity of shrublands in Nevada, yielding an assessment based on six indicators of ecosystem structure, function, and composition, including resource- and stressor-based indicators measured at multiple scales. Results varied spatially and among indicators. Invasive plant cover and surface development were highest in shrublands in northwest and southeast Nevada. Departure from reference conditions of shrubland area, composition, patch size, and connectivity was highest in central and northern Nevada. Results may inform efforts to control invasive species and restore shrublands on federal lands in Nevada. We suggest that ecological integrity assessments for multiple-use lands be grounded in existing policies and monitoring programs, incorporate resource- and stressor-based metrics, rely on publicly available data collected at multiple spatial scales, and quantify both natural reference and societally desired resource conditions.

Conservation planning for species recovery under the Endangered Species Act: A case study with the Northern Spotted Owl.

The northern spotted owl (Strix occidentalis caurina) was listed as threatened under the U.S. Endangered Species Act (ESA) in 1990. We applied modern spatial conservation theory and models to evaluate several candidate critical habitat networks, and sought an efficient conservation solution that encompassed the highest value lands for spotted owl recovery rather than maximizing the total area of potential critical habitat. We created a map of relative habitat suitability, which served as input to the spatial conservation prioritization program Zonation. We used the spatially-explicit individual-based population model HexSim to estimate and compare simulated spotted owl population outcomes among a suite of candidate critical habitat networks that varied in size and spatial arrangement under alternative scenarios of future habitat suitability and barred owl (S. varia) effects. We evaluated simulated spotted owl population outcomes, including total population size, and extinction and quasi-extinction likelihoods for 108 combinations of candidate critical habitat networks by habitat change by barred owl scenarios, both range-wide and within 11 distinct portions of the owl's range. Barred owl encounter rates and the amount and suitability of habitat had substantial effects on simulated spotted owl populations. When barred owl encounter rates were high, changes in the amount and suitability of habitat had minimal impacts on population performance. Under lowered barred owl encounter rates, candidate critical habitat networks that included most existing high suitability habitat supported a high likelihood of long-term population persistence. Barred owls are currently the primary driving force behind poor population performance of NSOs; however, our models demonstrated that a sufficient area of high suitability habitat remains essential for recovery when effects of barred owls can be reduced. The modeling approach we employed is sufficiently flexible to incorporate new information about spotted owls as it becomes available and could likely be applied to conservation planning for other species.

Projecting species' vulnerability to climate change: Which uncertainty sources matter most and extrapolate best?

Species distribution models (SDMs) are commonly used to assess potential climate change impacts on biodiversity, but several critical methodological decisions are often made arbitrarily. We compare variability arising from these decisions to the uncertainty in future climate change itself. We also test whether certain choices offer improved skill for extrapolating to a changed climate and whether internal cross-validation skill indicates extrapolative skill. We compared projected vulnerability for 29 wetland-dependent bird species breeding in the climatically dynamic Prairie Pothole Region, USA. For each species we built 1,080 SDMs to represent a unique combination of: future climate, class of climate covariates, collinearity level, and thresholding procedure. We examined the variation in projected vulnerability attributed to each uncertainty source. To assess extrapolation skill under a changed climate, we compared model predictions with observations from historic drought years. Uncertainty in projected vulnerability was substantial, and the largest source was that of future climate change. Large uncertainty was also attributed to climate covariate class with hydrological covariates projecting half the range loss of bioclimatic covariates or other summaries of temperature and precipitation. We found that choices based on performance in cross-validation improved skill in extrapolation. Qualitative rankings were also highly uncertain. Given uncertainty in projected vulnerability and resulting uncertainty in rankings used for conservation prioritization, a number of considerations appear critical for using bioclimatic SDMs to inform climate change mitigation strategies. Our results emphasize explicitly selecting climate summaries that most closely represent processes likely to underlie ecological response to climate change. For example, hydrological covariates projected substantially reduced vulnerability, highlighting the importance of considering whether water availability may be a more proximal driver than precipitation. However, because cross-validation results were correlated with extrapolation results, the use of cross-validation performance metrics to guide modeling choices where knowledge is limited was supported.

Projected wetland densities under climate change: habitat loss but little geographic shift in conservation strategy.

Climate change poses major challenges for conservation and management because it alters the area, quality, and spatial distribution of habitat for natural populations. To assess species' vulnerability to climate change and target ongoing conservation investments, researchers and managers often consider the effects of projected changes in climate and land use on future habitat availability and quality and the uncertainty associated with these projections. Here, we draw on tools from hydrology and climate science to project the impact of climate change on the density of wetlands in the Prairie Pothole Region of the USA, a critical area for breeding waterfowl and other wetland-dependent species. We evaluate the potential for a trade-off in the value of conservation investments under current and future climatic conditions and consider the joint effects of climate and land use. We use an integrated set of hydrological and climatological projections that provide physically based measures of water balance under historical and projected future climatic conditions. In addition, we use historical projections derived from ten general circulation models (GCMs) as a baseline from which to assess climate change impacts, rather than historical climate data. This method isolates the impact of greenhouse gas emissions and ensures that modeling errors are incorporated into the baseline rather than attributed to climate change. Our work shows that, on average, densities of wetlands (here defined as wetland basins holding water) are projected to decline across the U.S. Prairie Pothole Region, but that GCMs differ in both the magnitude and the direction of projected impacts. However, we found little evidence for a shift in the locations expected to provide the highest wetland densities under current vs. projected climatic conditions. This result was robust to the inclusion of projected changes in land use under climate change. We suggest that targeting conservation towards wetland complexes containing both small and relatively large wetland basins, which is an ongoing conservation strategy, may also act to hedge against uncertainty in the effects of climate change.

Conserving tigers in working landscapes.

Tiger (Panthera tigris) conservation efforts in Asia are focused on protected areas embedded in human-dominated landscapes. A system of protected areas is an effective conservation strategy for many endangered species if the network is large enough to support stable metapopulations. The long-term conservation of tigers requires that the species be able to meet some of its life-history needs beyond the boundaries of small protected areas and within the working landscape, including multiple-use forests with logging and high human use. However, understanding of factors that promote or limit the occurrence of tigers in working landscapes is incomplete. We assessed the relative influence of protection status, prey occurrence, extent of grasslands, intensity of human use, and patch connectivity on tiger occurrence in the 5400 km(2) Central Terai Landscape of India, adjacent to Nepal. Two observer teams independently surveyed 1009 km of forest trails and water courses distributed across 60 166-km(2) cells. In each cell, the teams recorded detection of tiger signs along evenly spaced trail segments. We used occupancy models that permitted multiscale analysis of spatially correlated data to estimate cell-scale occupancy and segment-scale habitat use by tigers as a function of management and environmental covariates. Prey availability and habitat quality, rather than protected-area designation, influenced tiger occupancy. Tiger occupancy was low in some protected areas in India that were connected to extensive areas of tiger habitat in Nepal, which brings into question the efficacy of current protection and management strategies in both India and Nepal. At a finer spatial scale, tiger habitat use was high in trail segments associated with abundant prey and large grasslands, but it declined as human and livestock use increased. We speculate that riparian grasslands may provide tigers with critical refugia from human activity in the daytime and thereby promote tiger occurrence in some multiple-use forests. Restrictions on human-use in high-quality tiger habitat in multiple-use forests may complement existing protected areas and collectively promote the persistence of tiger populations in working landscapes.

Vulnerability of breeding waterbirds to climate change in the Prairie Pothole Region, U.S.A.

The Prairie Pothole Region (PPR) of the north-central U.S. and south-central Canada contains millions of small prairie wetlands that provide critical habitat to many migrating and breeding waterbirds. Due to their small size and the relatively dry climate of the region, these wetlands are considered at high risk for negative climate change effects as temperatures increase. To estimate the potential impacts of climate change on breeding waterbirds, we predicted current and future distributions of species common in the PPR using species distribution models (SDMs). We created regional-scale SDMs for the U.S. PPR using Breeding Bird Survey occurrence records for 1971-2011 and wetland, upland, and climate variables. For each species, we predicted current distribution based on climate records for 1981-2000 and projected future distributions to climate scenarios for 2040-2049. Species were projected to, on average, lose almost half their current habitat (-46%). However, individual species projections varied widely, from +8% (Upland Sandpiper) to -100% (Wilson's Snipe). Variable importance ranks indicated that land cover (wetland and upland) variables were generally more important than climate variables in predicting species distributions. However, climate variables were relatively more important during a drought period. Projected distributions of species responses to climate change contracted within current areas of distribution rather than shifting. Given the large variation in species-level impacts, we suggest that climate change mitigation efforts focus on species projected to be the most vulnerable by enacting targeted wetland management, easement acquisition, and restoration efforts.

Efficient species-level monitoring at the landscape scale.

Monitoring the population trends of multiple animal species at a landscape scale is prohibitively expensive. However, advances in survey design, statistical methods, and the ability to estimate species presence on the basis of detection-nondetection data have greatly increased the feasibility of species-level monitoring. For example, recent advances in monitoring make use of detection-nondetection data that are relatively inexpensive to acquire, historical survey data, and new techniques in genetic evaluation. The ability to use indirect measures of presence for some species greatly increases monitoring efficiency and reduces survey costs. After adjusting for false absences, the proportion of sample units in a landscape where a species is detected (occupancy) is a logical state variable to monitor. Occupancy monitoring can be based on real-time observation of a species at a survey site or on evidence that the species was at the survey location sometime in the recent past. Temporal and spatial patterns in occupancy data are related to changes in animal abundance and provide insights into the probability of a species' persistence. However, even with the efficiencies gained when occupancy is the monitored state variable, the task of species-level monitoring remains daunting due to the large number of species. We propose that a small number of species be monitored on the basis of specific management objectives, their functional role in an ecosystem, their sensitivity to environmental changes likely to occur in the area, or their conservation importance.

Use of abundance of one species as a surrogate for abundance of others.

Indicator species concepts have a long history in conservation biology. Arguments in favor of these approaches generally stress expediency and assume efficacy. We tested the premise that the abundance patterns of one species can be used to infer those of other species. Our data consisted of 72,495 bird observations on 55 species across 1046 plots distributed across 30 sub basins. We analyzed abundance patterns at two spatial scales (plot and sub basin) and for empirical and a priori grouping. There were few significant indicator relationships at either scale or under either grouping rule, and those few we found did not explain a substantial portion of the abundance of other species. Coupled with the lack of proven efficacy for species surrogacy in the literature, our results indicate the utility of indicators and similar types of surrogate approaches must be demonstrated rather than assumed.

Quantifying the multi-scale response of avifauna to prescribed fire experiments in the southwest United States.

Landscape-scale disturbance events, including ecological restoration and fuel reduction activities, can modify habitat and affect relationships between species and their environment. To reduce the risk of uncharacteristic stand-replacing fires in the southwestern United States, land managers are implementing restoration and fuels treatments (e.g., mechanical thinning, prescribed fire) in progressively larger stands of dry, lower elevation ponderosa pine (Pinus ponderosa) forest. We used a Before-After/Control-Impact experimental design to quantify the multi-scale response of avifauna to large (approximately 250-400 ha) prescribed fire treatments on four sites in Arizona and New Mexico dominated by ponderosa pine. Using distance sampling and an information-theoretic approach, we estimated changes in density for 14 bird species detected before (May-June 2002-2003) and after (May-June 2004-2005) prescribed fire treatments. We observed few site-level differences in pre- and posttreatment density, and no species responded strongly to treatment on all four sites. Point-level spatial models of individual species response to treatment, habitat variables, and fire severity revealed ecological relationships that were more easily interpreted. At this scale, pretreatment forest structure and patch characteristics were important predictors of posttreatment differences in bird species density. Five species (Pygmy Nuthatch [Sitta pygmaea], Western Bluebird [Sialia mexicana], Steller's Jay [Cyanocitta stelleri], American Robin [Turdus migratorius], and Hairy Woodpecker [Picoides villosus]) exhibited a strong treatment response, and two of these species (American Robin and Hairy Woodpecker) could be associated with meaningful fire severity response functions. The avifaunal response patterns that we observed were not always consistent with those reported by more common studies of wildland fire events. Our results suggest that, in the short-term, the distribution and abundance of common members of the breeding bird community in Southwestern ponderosa pine forests appear to be tolerant of low- to moderate-intensity prescribed fire treatments at multiple spatial scales and across multiple geographic locations.

Conservation of the northern spotted owl under the Northwest Forest Plan.

Development of the Northwest Forest Plan (NWFP) was motivated by concerns about the over-harvest of late-seral forests and the effects of intensive forest management on the long-term viability of the Northern Spotted Owl (Strix occidentalis caurina). Following several years of intense political and legal debates, the final NWFP was approved in 1994. Even though the plan evolved with a broad ecosystem perspective, it remained anchored in the Spotted Owl reserve design proposed in 1990. Based on a criterion of stable or increasing populations, a decade later it remains unclear whether the enactment of the NWFP has improved the conservation status of Spotted Owls. The results of intensive monitoring of several Spotted Owl populations for over a decade suggest a continuing range-wide decline even though rates of timber harvest have declined dramatically on federal lands. The cause of the decline is difficult to determine because the research needed to establish cause and effect relations has not been done. One plausible hypothesis is that the owl's life history greatly constrains its rate of population growth even when habitat is no longer limiting. Since enactment of the NWFP, new threats have arisen, including the movement of Barred Owls (S. varia) into the range of the Spotted Owl, political pressure to increase levels of timber harvest, and recent changes to forest laws that eliminate the requirement to assess the viability of wildlife populations on U.S. Department of Agriculture Forest Service lands. At this time is appears that Spotted Owl conservation rests critically on continued implementation of the protections afforded by the NWFP and the U.S. Endangered Species Act.

Predicting abundance of desert riparian birds: validation and calibration of the Effective Area Model.

Reliable prediction of the effects of landscape change on species abundance is critical to land managers who must make frequent, rapid decisions with long-term consequences. However, due to inherent temporal and spatial variability in ecological systems, previous attempts to predict species abundance in novel locations and/or time frames have been largely unsuccessful. The Effective Area Model (EAM) uses change in habitat composition and geometry coupled with response of animals to habitat edges to predict change in species abundance at a landscape scale. Our research goals were to validate EAM abundance predictions in new locations and to develop a calibration framework that enables absolute abundance predictions in novel regions or time frames. For model validation, we compared the EAM to a null model excluding edge effects in terms of accurate prediction of species abundance. The EAM outperformed the null model for 83.3% of species (N=12) for which it was possible to discern a difference when considering 50 validation sites. Likewise, the EAM outperformed the null model when considering subsets of validation sites categorized on the basis of four variables (isolation, presence of water, region, and focal habitat). Additionally, we explored a framework for producing calibrated models to decrease prediction error given inherent temporal and spatial variability in abundance. We calibrated the EAM to new locations using linear regression between observed and predicted abundance with and without additional habitat covariates. We found that model adjustments for unexplained variability in time and space, as well as variability that can be explained by incorporating additional covariates, improved EAM predictions. Calibrated EAM abundance estimates with additional site-level variables explained a significant amount of variability (P < 0.05) in observed abundance for 17 of 20 species, with R2 values >25% for 12 species, >48% for six species, and >60% for four species when considering all predictive models. The calibration framework described in this paper can be used to predict absolute abundance in sites different from those in which data were collected if the target population of sites to which one would like to statistically infer is sampled in a probabilistic way.

Post-fledging survival of northern goshawks: the importance of prey abundance, weather, and dispersal.

Effective wildlife conservation strategies require an understanding of how fluctuating environmental conditions affect sensitive life stages. As part of a long-term study, we examined post-fledging and post-independence survival of 89 radio-marked juvenile Northern Goshawks (Accipiter gentilis) produced from 48 nests in northern Arizona, USA, during 1998-2001. Information-theoretic methods were used to examine within- and among-year variation in survival relative to environmental (prey abundance, weather), territory (hatching date, brood size), and individual (gender, body mass) sources of variation. The results support age- and cohort-specific differences in survival that were best explained by behaviors occurring at distinct stages of juvenile development, annual changes in the density of primary bird and mammal prey species, and gender-related differences in body mass. Survival between fledging and independence increased linearly with age and varied among annual cohorts of radio-marked juveniles from 0.81 (95% CI = 0.60-0.93) to 1.00 (95% CI = 0.95-1.00) in association with annual differences in prey density; the slope coefficient for the additive effect of prey density on survival was 1.12 (95% CI = 0.06-2.19). Survival declined to 0.71 (95% CI = 0.60-0.93) shortly after juveniles initiated dispersal (weeks 8-12 post-fledging) and moved to more open habitats at lower elevations. Survival was not closely associated with weather or territory-level parameters. A comparison of the predictions of environmental-, territory-, and individual-based models of survival demonstrated that food availability was the primary factor limiting juvenile survival. This finding indicates that forest management prescriptions designed to support abundant prey populations while providing forest structural conditions that allow goshawks to access their prey within breeding areas should benefit juvenile survival.

Integrating Scientific Methods with Habitat Conservation Planning: Reserve Design for Northern Spotted Owls.

To meet the requirements of Congressional legislation mandating the production of a "scientifically credible" conservation strategy for the threatened Northern Spotted Owl (Strix occidentalis caurina), the Interagency Spotted Owl Scientific Committee employed scientific methods to design a habitat reserve system. Information on the current and historical distributions of the owl and its habitats was reviewed in light of economic, political, and legal constraints; results were used to develop a preliminary reserve system of habitat "polygons." A map representing these polygons and their attendant properties served as a set of hypotheses that were tested. Statistical analyses of empirical data, predictions from ecological theory, predictions from population dynamics models, and inferences drawn from studies of related species were used to test properties of the preliminary map, including the number and sizes of habitat conservation areas (HCAs), their distribution, configuration, and spacing, and the nature of the landscape matrix between HCAs. Conclusions that failed to confirm specific map properties were used to refine the reserve system, a process that continued iteratively until all relevant data had been examined and all map properties had been tested. This conservation planning process has proven to be credible, repeatable, and scientifically defendable, and should serve as a model for wildlife management, endangered species recovery, and national forest planning.