Unraveling a century of global change impacts on winter bird distributions in the eastern United States.

One of the most pressing questions in ecology and conservation centers on disentangling the relative impacts of concurrent global change drivers, climate and land-use/land-cover (LULC), on biodiversity. Yet studies that evaluate the effects of both drivers on species' winter distributions remain scarce, hampering our ability to develop full-annual-cycle conservation strategies. Additionally, understanding how groups of species differentially respond to climate versus LULC change is vital for efforts to enhance bird community resilience to future environmental change. We analyzed long-term changes in winter occurrence of 89 species across nine bird groups over a 90-year period within the eastern United States using Audubon Christmas Bird Count (CBC) data. We estimated variation in occurrence probability of each group as a function of spatial and temporal variation in winter climate (minimum temperature, cumulative precipitation) and LULC (proportion of group-specific and anthropogenic habitats within CBC circle). We reveal that spatial variation in bird occurrence probability was consistently explained by climate across all nine species groups. Conversely, LULC change explained more than twice the temporal variation (i.e., decadal changes) in bird occurrence probability than climate change on average across groups. This pattern was largely driven by habitat-constrained species (e.g., grassland birds, waterbirds), whereas decadal changes in occurrence probabilities of habitat-unconstrained species (e.g., forest passerines, mixed habitat birds) were equally explained by both climate and LULC changes over the last century. We conclude that climate has generally governed the winter occurrence of avifauna in space and time, while LULC change has played a pivotal role in driving distributional dynamics of species with limited and declining habitat availability. Effective land management will be critical for improving species' resilience to climate change, especially during a season of relative resource scarcity and critical energetic trade-offs.

The effects of weather variability on patterns of genetic diversity in Tasmanian bettongs.

While the effects of climate (long-term, prevailing weather) on species abundance, range and genetic diversity have been widely studied, short-term, localized variations in atmospheric conditions (i.e., weather) can also rapidly alter species' geographical ranges and population sizes, but little is known about how they affect genetic diversity. We investigated the relationship between weather and range-wide genetic diversity in a marsupial, Bettongia gaimardi, using dynamic species distribution models (SDMs). Genetic diversity was lower in parts of the range where the weather-based SDM predicted high variability in probability of B. gaimardi occurrence during 1950-2009. This is probably an effect of lower population sizes and extinction-recolonization cycles in places with highly variable weather. Spatial variation in genetic diversity was also better predicted by mean probabilities of B. gaimardi occurrence from weather- than climate-based SDMs. Our results illustrate the importance of weather in driving population dynamics and species distributions on decadal timescales and thereby in affecting genetic diversity. Modelling the links between changing weather patterns, species distributions and genetic diversity will allow researchers to better forecast biological impacts of climate change.

Interrelated impacts of climate and land-use change on a widespread waterbird.

Together climate and land-use change play a crucial role in determining species distribution and abundance, but measuring the simultaneous impacts of these processes on current and future population trajectories is challenging due to time lags, interactive effects and data limitations. Most approaches that relate multiple global change drivers to population changes have been based on occurrence or count data alone. We leveraged three long-term (1995-2019) datasets to develop a coupled integrated population model-Bayesian population viability analysis (IPM-BPVA) to project future survival and reproductive success for common loons Gavia immer in northern Wisconsin, USA, by explicitly linking vital rates to changes in climate and land use. The winter North Atlantic Oscillation (NAO), a broad-scale climate index, immediately preceding the breeding season and annual changes in developed land cover within breeding areas both had strongly negative influences on adult survival. Local summer rainfall was negatively related to fecundity, though this relationship was mediated by a lagged interaction with the winter NAO, suggesting a compensatory population-level response to climate variability. We compared population viability under 12 future scenarios of annual land-use change, precipitation and NAO conditions. Under all scenarios, the loon population was expected to decline, yet the steepest declines were projected under positive NAO trends, as anticipated with ongoing climate change. Thus, loons breeding in the northern United States are likely to remain affected by climatic processes occurring thousands of miles away in the North Atlantic during the non-breeding period of the annual cycle. Our results reveal that climate and land-use changes are differentially contributing to loon population declines along the southern edge of their breeding range and will continue to do so despite natural compensatory responses. We also demonstrate that concurrent analysis of multiple data types facilitates deeper understanding of the ecological implications of anthropogenic-induced change occurring at multiple spatial scales. Our modelling approach can be used to project demographic responses of populations to varying environmental conditions while accounting for multiple sources of uncertainty, an increasingly pressing need in the face of unprecedented global change.

Community science validates climate suitability projections from ecological niche modeling.

Climate change poses an intensifying threat to many bird species and projections of future climate suitability provide insight into how species may shift their distributions in response. Climate suitability is characterized using ecological niche models (ENMs), which correlate species occurrence data with current environmental covariates and project future distributions using the modeled relationships together with climate predictions. Despite their widespread adoption, ENMs rely on several assumptions that are rarely validated in situ and can be highly sensitive to modeling decisions, precluding their reliability in conservation decision-making. Using data from a novel, large-scale community science program, we developed dynamic occupancy models to validate near-term climate suitability projections for bluebirds and nuthatches in summer and winter. We estimated occupancy, colonization, and extinction dynamics across species' ranges in the United States in relation to projected climate suitability in the 2020s, and used a Gibbs variable selection approach to quantify evidence of species-climate relationships. We also included a Bird Conservation Region strata-level random effect to examine among-strata variation in occupancy that may be attributable to land-use and ecoregional differences. Across species and seasons, we found strong evidence that initial occupancy and colonization were positively related to 2020 climate suitability, illustrating an independent validation of projections from ENMs across a large geographic area. Random strata effects revealed that occupancy probabilities were generally higher than average in core areas and lower than average in peripheral areas of species' ranges, and served as a first step in identifying spatial patterns of occupancy from these community science data. Our findings lend much-needed support to the use of ENM projections for addressing questions about potential climate-induced changes in species' occupancy dynamics. More broadly, our work highlights the value of community scientist observations for ground-truthing projections from statistical models and for refining our understanding of the processes shaping species' distributions under a changing climate.

The pace of past climate change vs. potential bird distributions and land use in the United States.

Climate change may drastically alter patterns of species distributions and richness, but predicting future species patterns in occurrence is challenging. Significant shifts in distributions have already been observed, and understanding these recent changes can improve our understanding of potential future changes. We assessed how past climate change affected potential breeding distributions for landbird species in the conterminous United States. We quantified the bioclimatic velocity of potential breeding distributions, that is, the pace and direction of change for each species' suitable climate space over the past 60 years. We found that potential breeding distributions for landbirds have shifted substantially with an average velocity of 1.27 km yr(-1) , about double the pace of prior distribution shift estimates across terrestrial systems globally (0.61 km yr(-1) ). The direction of shifts was not uniform. The majority of species' distributions shifted west, northwest, and north. Multidirectional shifts suggest that changes in climate conditions beyond mean temperature were influencing distributional changes. Indeed, precipitation variables that were proxies for extreme conditions were important variables across all models. There were winners and losers in terms of the area of distributions; many species experienced contractions along west and east distribution edges, and expansions along northern distribution edges. Changes were also reflected in the potential species richness, with some regions potentially gaining species (Midwest, East) and other areas potentially losing species (Southwest). However, the degree to which changes in potential breeding distributions are manifested in actual species richness depends on landcover. Areas that have become increasingly suitable for breeding birds due to changing climate are often those attractive to humans for agriculture and development. This suggests that many areas might have supported more breeding bird species had the landscape not been altered. Our study illustrates that climate change is not only a future threat, but something birds are already experiencing.

Potential breeding distributions of U.S. birds predicted with both short-term variability and long-term average climate data.

Climate conditions, such as temperature or precipitation, averaged over several decades strongly affect species distributions, as evidenced by experimental results and a plethora of models demonstrating statistical relations between species occurrences and long-term climate averages. However, long-term averages can conceal climate changes that have occurred in recent decades and may not capture actual species occurrence well because the distributions of species, especially at the edges of their range, are typically dynamic and may respond strongly to short-term climate variability. Our goal here was to test whether bird occurrence models can be predicted by either covariates based on short-term climate variability or on long-term climate averages. We parameterized species distribution models (SDMs) based on either short-term variability or long-term average climate covariates for 320 bird species in the conterminous USA and tested whether any life-history trait-based guilds were particularly sensitive to short-term conditions. Models including short-term climate variability performed well based on their cross-validated area-under-the-curve AUC score (0.85), as did models based on long-term climate averages (0.84). Similarly, both models performed well compared to independent presence/absence data from the North American Breeding Bird Survey (independent AUC of 0.89 and 0.90, respectively). However, models based on short-term variability covariates more accurately classified true absences for most species (73% of true absences classified within the lowest quarter of environmental suitability vs. 68%). In addition, they have the advantage that they can reveal the dynamic relationship between species and their environment because they capture the spatial fluctuations of species potential breeding distributions. With this information, we can identify which species and guilds are sensitive to climate variability, identify sites of high conservation value where climate variability is low, and assess how species' potential distributions may have already shifted due recent climate change. However, long-term climate averages require less data and processing time and may be more readily available for some areas of interest. Where data on short-term climate variability are not available, long-term climate information is a sufficient predictor of species distributions in many cases. However, short-term climate variability data may provide information not captured with long-term climate data for use in SDMs.

Conservation hotspots for marine turtle nesting in the United States based on coastal development.

Coastal areas provide nesting habitat for marine turtles that is critical for the persistence of their populations. However, many coastal areas are highly affected by coastal development, which affects the reproductive success of marine turtles. Knowing the extent to which nesting areas are exposed to these threats is essential to guide management initiatives. This information is particularly important for coastal areas with both high nesting density and dense human development, a combination that is common in the United States. We assessed the extent to which nesting areas of the loggerhead (Caretta caretta), the green (Chelonia mydas), the Kemp's ridley (Lepidochelys kempii), and leatherback turtles (Dermochelys coriacea) in the continental United States are exposed to coastal development and identified conservation hotspots that currently have high reproductive importance and either face high exposure to coastal development (needing intervention), or have low exposure to coastal development, and are good candidates for continued and future protection. Night-time light, housing, and population density were used as proxies for coastal development and human disturbance. About 81.6% of nesting areas were exposed to housing and human population, and 97.8% were exposed to light pollution. Further, most (>65%) of the very high- and high-density nesting areas for each species/subpopulation, except for the Kemp's ridley, were exposed to coastal development. Forty-nine nesting sites were selected as conservation hotspots; of those high-density nesting sites, 49% were sites with no/low exposure to coastal development and the other 51% were exposed to high-density coastal development. Conservation strategies need to account for ~66.8% of all marine turtle nesting areas being on private land and for nesting sites being exposed to large numbers of seasonal residents.

The rise of novelty in ecosystems.

Rapid and ongoing change creates novelty in ecosystems everywhere, both when comparing contemporary systems to their historical baselines, and predicted future systems to the present. However, the level of novelty varies greatly among places. Here we propose a formal and quantifiable definition of abiotic and biotic novelty in ecosystems, map abiotic novelty globally, and discuss the implications of novelty for the science of ecology and for biodiversity conservation. We define novelty as the degree of dissimilarity of a system, measured in one or more dimensions relative to a reference baseline, usually defined as either the present or a time window in the past. In this conceptualization, novelty varies in degree, it is multidimensional, can be measured, and requires a temporal and spatial reference. This definition moves beyond prior categorical definitions of novel ecosystems, and does not include human agency, self-perpetuation, or irreversibility as criteria. Our global assessment of novelty was based on abiotic factors (temperature, precipitation, and nitrogen deposition) plus human population, and shows that there are already large areas with high novelty today relative to the early 20th century, and that there will even be more such areas by 2050. Interestingly, the places that are most novel are often not the places where absolute changes are largest; highlighting that novelty is inherently different from change. For the ecological sciences, highly novel ecosystems present new opportunities to test ecological theories, but also challenge the predictive ability of ecological models and their validation. For biodiversity conservation, increasing novelty presents some opportunities, but largely challenges. Conservation action is necessary along the entire continuum of novelty, by redoubling efforts to protect areas where novelty is low, identifying conservation opportunities where novelty is high, developing flexible yet strong regulations and policies, and establishing long-term experiments to test management approaches. Meeting the challenge of novelty will require advances in the science of ecology, and new and creative. conservation approaches.

Modeled distribution shifts of North American birds over four decades based on suitable climate alone do not predict observed shifts.

As climate change alters the global environment, it is critical to understand the relationship between shifting climate suitability and species distributions. Key questions include whether observed changes in population abundance are aligned with the velocity and direction of shifts predicted by climate suitability models and if the responses are consistent among species with similar ecological traits. We examined the direction and velocity of the observed abundance-based distribution centroids compared with the model-predicted bioclimatic distribution centroids of 250 bird species across the United States from 1969 to 2011. We hypothesized that there is a significant positive correlation in both direction and velocity between the observed and the modeled shifts. We then tested five additional hypotheses that predicted differential shifting velocity based on ecological adaptability and climate change exposure. Contrary to our hypotheses, we found large differences between the observed and modeled shifts among all studied bird species and within specific ecological guilds. However, temperate migrants and habitat generalist species tended to have higher velocity of observed shifts than other species. Neotropical migratory and wetland birds also had significantly different observed velocities than their counterparts, which may be due to their climate change exposure. The velocity based on modeled bioclimatic suitability did not exhibit significant differences among most guilds. Boreal forest birds were the only guild with significantly faster modeled-shifts than the other groups, suggesting an elevated conservation risk for high latitude and altitude species. The highly idiosyncratic species responses to climate and the mismatch between shifts in modeled and observed distribution centroids highlight the challenge of predicting species distribution change based solely on climate suitability and the importance of non-climatic factors traits in shaping species distributions.

Projected changes in bird assemblages due to climate change in a Canadian system of protected areas.

National parks often serve as a cornerstone for a country's species and ecosystem conservation efforts. However, despite the protection these sites afford, climate change is expected to drive a substantial change in their bird assemblages. We used species distribution models to predict the change in environmental suitability (i.e., how well environmental conditions explain the presence of a species) of 49 Canadian national parks during summer and winter for 434 bird species under a 2°C warming scenario, anticipated to occur in Canada around the mid-21st century. We compared these to existing species distributions in the 2010s, and classified suitability projections for each species at each park as potential extirpation, worsening, stable, improving, or potential colonisation. Across all parks, and both seasons, 70% of the projections indicate change, including a 25% turnover in summer assemblages and 30% turnover in winter assemblages. The majority of parks are projected to have increases in species richness and functional traits in winter, compared to a mix of increases and decreases in both in summer. However, some changes are expected to vary by region, such as Arctic region parks being likely to experience the most potential colonisation, while some of the Mixedwood Plains and Atlantic Maritime region parks may experience the greatest turnover and potential extirpation in summer if management actions are not taken to mitigate some of these losses. Although uncertainty exists around the precise rate and impacts of climate change, our results indicate that conservation practices that assume stationarity of environmental conditions will become untenable. We propose general guidance to help managers adapt their conservation actions to consider the potentially substantive changes in bird assemblages that are projected, including managing for persistence and change.

Testing the Role of Climate Change in Species Decline: Is the Eastern Quoll a Victim of a Change in the Weather?

To conserve a declining species we first need to diagnose the causes of decline. This is one of the most challenging tasks faced by conservation practitioners. In this study, we used temporally explicit species distribution models (SDMs) to test whether shifting weather can explain the recent decline of a marsupial carnivore, the eastern quoll (Dasyurus viverrinus). We developed an SDM using weather variables matched to occurrence records of the eastern quoll over the last 60 years, and used the model to reconstruct variation through time in the distribution of climatically suitable range for the species. The weather model produced a meaningful prediction of the known distribution of the species. Abundance of quolls, indexed by transect counts, was positively related to the modelled area of suitable habitat between 1990 and 2004. In particular, a sharp decline in abundance from 2001 to 2003 coincided with a sustained period of unsuitable weather over much of the species' distribution. Since 2004, abundance has not recovered despite a return to suitable weather conditions, and abundance and area of suitable habitat have been uncorrelated. We suggest that fluctuations in weather account for the species' recent decline, but other unrelated factors have suppressed recovery.