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.

Modeling nonbreeding distributions of shorebirds and waterfowl in response to climate change.

To identify areas on the landscape that may contribute to a robust network of conservation areas, we modeled the probabilities of occurrence of several en route migratory shorebirds and wintering waterfowl in the southern Great Plains of North America, including responses to changing climate. We predominantly used data from the eBird citizen-science project to model probabilities of occurrence relative to land-use patterns, spatial distribution of wetlands, and climate. We projected models to potential future climate conditions using five representative general circulation models of the Coupled Model Intercomparison Project 5 (CMIP5). We used Random Forests to model probabilities of occurrence and compared the time periods 1981-2010 (hindcast) and 2041-2070 (forecast) in "model space." Projected changes in shorebird probabilities of occurrence varied with species-specific general distribution pattern, migration distance, and spatial extent. Species using the western and northern portion of the study area exhibited the greatest likelihoods of decline, whereas species with more easterly occurrences, mostly long-distance migrants, had the greatest projected increases in probability of occurrence. At an ecoregional extent, differences in probabilities of shorebird occurrence ranged from -0.015 to 0.045 when averaged across climate models, with the largest increases occurring early in migration. Spatial shifts are predicted for several shorebird species. Probabilities of occurrence of wintering Mallards and Northern Pintail are predicted to increase by 0.046 and 0.061, respectively, with northward shifts projected for both species. When incorporated into partner land management decision tools, results at ecoregional extents can be used to identify wetland complexes with the greatest potential to support birds in the nonbreeding season under a wide range of future climate scenarios.

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.

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.

Weather effects on avian breeding performance and implications of climate change.

The influence of recent climate change on the world's biota has manifested broadly, resulting in latitudinal range shifts, advancing dates of arrival of migrants and onset of breeding, and altered community relationships. Climate change elevates conservation concerns worldwide because it will likely exacerbate a broad range of identified threats to animal populations. In the past few decades, grassland birds have declined faster than other North American avifauna, largely due to habitat threats such as the intensification of agriculture. We examine the effects of local climatic variations on the breeding performance of a bird endemic to the shortgrass prairie, the Lark Bunting (Calamospiza melanocorys) and discuss the implications of our findings relative to future climate predictions. Clutch size, nest survival, and productivity all positively covaried with seasonal precipitation; yet relatively intense daily precipitation events temporarily depressed daily survival of nests. Nest survival was positively related to average temperatures during the breeding season. Declining summer precipitation may reduce the likelihood that Lark Buntings can maintain stable breeding populations in eastern Colorado although average temperature increases of up to 3 degrees C (within the range of this study) may ameliorate declines in survival expected with drier conditions. Historic climate variability in the Great Plains selects for a degree of vagility and opportunism rather than strong site fidelity and specific adaptation to local environments. These traits may lead to northerly shifts in distribution if climatic and habitat conditions become less favorable in the drying southern regions of the Great Plains. Distributional shifts in Lark Buntings could be constrained by future changes in land use, agricultural practices, or vegetative communities that result in further loss of shortgrass prairie habitats.

Potential misuse of avian density as a conservation metric.

Effective conservation metrics are needed to evaluate the success of management in a rapidly changing world. Reproductive rates and densities of breeding birds (as a surrogate for reproductive rate) have been used to indicate the quality of avian breeding habitat, but the underlying assumptions of these metrics rarely have been examined. When birds are attracted to breeding areas in part by the presence of conspecifics and when breeding in groups influences predation rates, the effectiveness of density and reproductive rate as indicators of habitat quality is reduced. It is beneficial to clearly distinguish between individual- and population-level processes when evaluating habitat quality. We use the term reproductive rate to refer to both levels and further distinguish among levels by using the terms per capita fecundity (number of female offspring per female per year, individual level) and population growth rate (the product of density and per capita fecundity, population level). We predicted how density and reproductive rate interact over time under density-independent and density-dependent scenarios, assuming the ideal free distribution model of how birds settle in breeding habitats. We predicted population density of small populations would be correlated positively with both per capita fecundity and population growth rate due to the Allee effect. For populations in the density-dependent growth phase, we predicted no relation between density and per capita fecundity (because individuals in all patches will equilibrate to the same success rate) and a positive relation between density and population growth rate. Several ecological theories collectively suggest that positive correlations between density and per capita fecundity would be difficult to detect. We constructed a decision tree to guide interpretation of positive, neutral, nonlinear, and negative relations between density and reproductive rates at individual and population levels.

Modeling post-fledging survival of Lark Buntings in response to ecological and biological factors.

We evaluated the influences of several ecological, biological, and methodological factors on post-fledging survival of a shortgrass prairie bird, the Lark Bunting (Calamospiza melanocorys). We estimated daily post-fledging survival (n = 206, 82 broods) using radiotelemetry and color bands to track fledglings. Daily survival probabilities were best explained by drought intensity, time in season (quadratic trend), ages < or = 3 d post-fledging, and rank given drought intensity. Drought intensity had a strong negative effect on survival. Rank was an important predictor of fledgling survival only during the severe drought of 2002 when the smallest fledglings had lower survival. Recently fledged young (ages < or = 3 d post-fledging) undergoing the transition from nest to surrounding habitat experienced markedly lower survival, demonstrating the vulnerable nature of this time period. Survival was greater in mid and late season than early season, corresponding to our assumptions of food availability. Neither mark type nor sex of attending parent influenced survival. The model-averaged product of the 22-d survival calculated using mean rank and median value of time in season was 0.360 +/- 0.08 in 2001 and 0.276 +/- 0.08 in 2002. Survival estimates that account for age, condition of young, ecological conditions, and other factors are important for parameterization of realistic population models. Biologists using population growth models to elucidate mechanisms of population declines should attempt to estimate species-specific of post-fledging survival rather than use generalized estimates.

Human Disturbance of an Avian Scavenging Guild.

In order to investigate the effects of human activities on relationships within foraging guilds, we examined inacanus dynamics of eagles, crows, and gulls scavenging on spawned salmon in the Pacific Northwest. We examined several hypotheses that postulate the asymmetric foraging relationships of the three guild members and that reveal the influence of competition and facilitation in these relationships. Spatial and temporal patterns of resource use by the three primary guild members varied with the presence and absence of human activity at experimental feeding stations. At control (undisturbed) stations, eagles preferred to feed >100 m from vegetative cover, whereas gulls fed <50 m from cover. At experimental (disturbed) stations, eagles rarely fed, and feeding activity by gulls increased at both near and far stations. Crows often fed on alternate food sources in fields adjacent to the river, especially when salmon carcasses were scarce, whereas eagles and gulls rarely did so. We also examined if and how the behavior of single guild members changes in the presence or absence of other guild members. In the absence of eagles, gulls and crows preferred stations far from cover, numbers of both increased at feeding stations, birds were distributed nearer to carcasses, and they fed more. We emphasize that guild theory lends important insights to our understanding of the effects of human disturbance on wildlife communities.