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Bark beetle outbreaks in the Rocky Mountains caused substantial tree mortality starting in the late 1990s, and continued into the 2000s, with the most severe mortality occurring from 2002 to 2012. Over the same time period, concentrations of dissolved copper in the Big Thompson River (BTR), Colorado, USA, increased significantly and are high enough to negatively affect aquatic life. We examined correlations between dissolved copper and tree mortality in the BTR. Two sites, one consisting of water from the western side of the continental divide and one consisting of water from the eastern side, demonstrated a positive relationship between percentage tree mortality and dissolved copper. The relationships were similar except that the best relationship occurred with a 3-year lag between tree mortality and subsequent dissolved copper levels at the eastern site and with a 5-year lag at the western site. The differential time lag is potentially the result of different levels of carbon in the soil in the watersheds associated with each site because carbon can affect copper mobility. Our results suggest that bark beetle-induced tree mortality may contribute significantly to dissolved copper levels in the BTR.

Ecological niche theory holds that species distributions are shaped by a large and complex suite of interacting factors. Species distribution models (SDMs) are increasingly used to describe species' niches and predict the effects of future environmental change, including climate change. Currently, SDMs often fail to capture the complexity of species' niches, resulting in predictions that are generally limited to climate-occupancy interactions. Here, we explore the potential impact of climate change on the American pika using a replicated place-based approach that incorporates climate, gene flow, habitat configuration, and microhabitat complexity into SDMs. Using contemporary presence-absence data from occupancy surveys, genetic data to infer connectivity between habitat patches, and 21 environmental niche variables, we built separate SDMs for pika populations inhabiting eight US National Park Service units representing the habitat and climatic breadth of the species across the western United States. We then predicted occurrence probability under current (1981-2010) and three future time periods (out to 2100). Occurrence probabilities and the relative importance of predictor variables varied widely among study areas, revealing important local-scale differences in the realized niche of the American pika. This variation resulted in diverse and - in some cases - highly divergent future potential occupancy patterns for pikas, ranging from complete extirpation in some study areas to stable occupancy patterns in others. Habitat composition and connectivity, which are rarely incorporated in SDM projections, were influential in predicting pika occupancy in all study areas and frequently outranked climate variables. Our findings illustrate the importance of a place-based approach to species distribution modeling that includes fine-scale factors when assessing current and future climate impacts on species' distributions, especially when predictions are intended to manage and conserve species of concern within individual protected areas.

Refugia have long been studied from paleontological and biogeographical perspectives to understand how populations persisted during past periods of unfavorable climate. Recently, researchers have applied the idea to contemporary landscapes to identify climate change refugia, here defined as areas relatively buffered from contemporary climate change over time that enable persistence of valued physical, ecological, and socio-cultural resources. We differentiate historical and contemporary views, and characterize physical and ecological processes that create and maintain climate change refugia. We then delineate how refugia can fit into existing decision support frameworks for climate adaptation and describe seven steps for managing them. Finally, we identify challenges and opportunities for operationalizing the concept of climate change refugia. Managing climate change refugia can be an important option for conservation in the face of ongoing climate change.

In the face of environmental change, species can evolve new physiological tolerances to cope with altered climatic conditions or move spatially to maintain existing physiological associations with particular climates that define each species' climatic niche. When environmental change occurs over short temporal and large spatial scales, vagile species are expected to move geographically by tracking their climatic niches through time. Here, we test for evidence of niche tracking in bird species of the Sierra Nevada mountains of California, focusing on 53 species resurveyed nearly a century apart at 82 sites on four elevational transects. Changes in climate and bird distributions resulted in focal species shifting their average climatological range over time. By comparing the directions of these shifts relative to the centroids of species' range-wide climatic niches, we found that 48 species (90.6%) tracked their climatic niche. Analysis of niche sensitivity on an independent set of occurrence data significantly predicted the temperature and precipitation gradients tracked by species. Furthermore, in 50 species (94.3%), site-specific occupancy models showed that the position of each site relative to the climatic niche centroid explained colonization and extinction probabilities better than a null model with constant probabilities. Combined, our results indicate that the factors limiting a bird species' range in the Sierra Nevada in the early 20th century also tended to drive changes in distribution over time, suggesting that climatic models derived from niche theory might be used successfully to forecast where and how to conserve species in the face of climate change.

Our ability to accurately forecast species' geographical responses to climate change requires knowledge of the proximate and ultimate drivers of their distribution. Here, we consider the ecophysiological and demographic determinants of the distribution of a partial migrant, the North American field sparrow, Spizella pusilla . From 1940 to 1963, the field sparrow extended its winter northern range margin 222 km polewards. Such expansion was coincident with not only a geographical expansion into suitable breeding habitats, but also a decrease in mean abundance across sites occupied during the winter surveys. Combined, these trends suggest that declining populations along the expansion front either stopped migrating or altered their autumn migration. The poleward expansion was not coincident with climatically induced decreases in peak metabolic energy demand, but it did track increases in ecosystem net primary productivity. After 1963, the species' lower lethal temperature prevented further poleward movement. These findings show how different ecophysiological constraints can interact to change migration and distribution in a demographically declining species.

Key to understanding the implications of climate and land use change on biodiversity and natural resources is to incorporate the physiographic platform on which changes in ecological systems unfold. Here, we advance a detailed classification and high-resolution map of physiography, built by combining landforms and lithology (soil parent material) at multiple spatial scales. We used only relatively static abiotic variables (i.e., excluded climatic and biotic factors) to prevent confounding current ecological patterns and processes with enduring landscape features, and to make the physiographic classification more interpretable for climate adaptation planning. We generated novel spatial databases for 15 landform and 269 physiographic types across the conterminous United States of America. We examined their potential use by natural resource managers by placing them within a contemporary climate change adaptation framework, and found our physiographic databases could play key roles in four of seven general adaptation strategies. We also calculated correlations with common empirical measures of biodiversity to examine the degree to which the physiographic setting explains various aspects of current biodiversity patterns. Additionally, we evaluated the relationship between landform diversity and measures of climate change to explore how changes may unfold across a geophysical template. We found landform types are particularly sensitive to spatial scale, and so we recommend using high-resolution datasets when possible, as well as generating metrics using multiple neighborhood sizes to both minimize and characterize potential unknown biases. We illustrate how our work can inform current strategies for climate change adaptation. The analytical framework and classification of landforms and parent material are easily extendable to other geographies and may be used to promote climate change adaptation in other settings.

Many protected areas may not be adequately safeguarding biodiversity from human activities on surrounding lands and global change. The magnitude of such change agents and the sensitivity of ecosystems to these agents vary among protected areas. Thus, there is a need to assess vulnerability across networks of protected areas to determine those most at risk and to lay the basis for developing effective adaptation strategies. We conducted an assessment of exposure of U.S. National Parks to climate and land use change and consequences for vegetation communities. We first defined park protected-area centered ecosystems (PACEs) based on ecological principles. We then drew on existing land use, invasive species, climate, and biome data sets and models to quantify exposure of PACEs from 1900 through 2100. Most PACEs experienced substantial change over the 20th century (> 740% average increase in housing density since 1940, 13% of vascular plants are presently nonnative, temperature increase of 1 degree C/100 yr since 1895 in 80% of PACEs), and projections suggest that many of these trends will continue at similar or increasingly greater rates (255% increase in housing density by 2100, temperature increase of 2.5 degrees-4.5 degrees C/100 yr, 30% of PACE areas may lose their current biomes by 2030). In the coming century, housing densities are projected to increase in PACEs at about 82% of the rate of since 1940. The rate of climate warming in the coming century is projected to be 2.5-5.8 times higher than that measured in the past century. Underlying these averages, exposure of individual park PACEs to change agents differ in important ways. For example, parks such as Great Smoky Mountains exhibit high land use and low climate exposure, others such as Great Sand Dunes exhibit low land use and high climate exposure, and a few such as Point Reyes exhibit high exposure on both axes. The cumulative and synergistic effects of such changes in land use, invasives, and climate are expected to dramatically impact ecosystem function and biodiversity in national parks. These results are foundational to developing effective adaptation strategies and suggest policies to better safeguard parks under broad-scale environmental change.

Refugia have long been studied from paleontological and biogeographical perspectives to understand how populations persisted during past periods of unfavorable climate. Recently, researchers have applied the idea to contemporary landscapes to identify climate change refugia, here defined as areas relatively buffered from contemporary climate change over time that enable persistence of valued physical, ecological, and socio-cultural resources. We differentiate historical and contemporary views, and characterize physical and ecological processes that create and maintain climate change refugia. We then delineate how refugia can fit into existing decision support frameworks for climate adaptation and describe seven steps for managing them. Finally, we identify challenges and opportunities for operationalizing the concept of climate change refugia. Managing climate change refugia can be an important option for conservation in the face of ongoing climate change.

Background In the mid 20th century, Ernst Mayr and Theodosius Dobzhansky championed the significance of circular overlaps or ring species as the perfect demonstration of speciation, yet in the over 50 years since, only a handful of such taxa are known. We developed a topographic model to evaluate whether the geographic barriers that favor processes leading to ring species are common or rare, and to predict where other candidate ring barriers might be found. Results Of the 952,147 geographic barriers identified on the planet, only about 1% are topographically similar to barriers associated with known ring taxa, with most of the likely candidates occurring in under-studied parts of the world (for example, marine environments, tropical latitudes). Predicted barriers separate into two distinct categories: (i) single cohesive barriers (< 50,000 km2), associated with taxa that differentiate at smaller spatial scales (salamander: Ensatina eschscholtzii; tree: Acacia karroo); and (ii) composite barriers - formed by groups of barriers (each 184,000 to 1.7 million km2) in close geographic proximity (totaling 1.9 to 2.3 million km2) - associated with taxa that differentiate at larger spatial scales (birds: Phylloscopus trochiloides and Larus (sp. argentatus and fuscus)). When evaluated globally, we find a large number of cohesive barriers that are topographically similar to those associated with known ring taxa. Yet, compared to cohesive barriers, an order of magnitude fewer composite barriers are similar to those that favor ring divergence in species with higher dispersal. Conclusions While these findings confirm that the topographic conditions that favor evolutionary processes leading to ring speciation are, in fact, rare, they also suggest that many understudied natural systems could provide valuable demonstrations of continuous divergence towards the formation of new species. Distinct advantages of the model are that it (i) requires no a priori information on the relative importance of features that define barriers, (ii) can be replicated using any kind of continuously distributed environmental variable, and (iii) generates spatially explicit hypotheses of geographic species formation. The methods developed here - combined with study of the geographical ecology and genetics of taxa in their environments - should enable recognition of ring species phenomena throughout the world.

The ability of species to respond to novel future climates is determined in part by their physiological capacity to tolerate climate change and the degree to which they have reached and continue to maintain distributional equilibrium with the environment. While broad-scale correlative climatic measurements of a species’ niche are often described as estimating the fundamental niche, it is unclear how well these occupied portions actually approximate the fundamental niche per se, versus the fundamental niche that exists in environmental space, and what fitness values bounding the niche are necessary to maintain distributional equilibrium. Here, we investigate these questions by comparing physiological and correlative estimates of the thermal niche in the introduced North American house sparrow (Passer domesticus). Our results indicate that occupied portions of the fundamental niche derived from temperature correlations closely approximate the centroid of the existing fundamental niche calculated on a fitness threshold of 50% population mortality. Using these niche measures, a 75-year time series analysis (1930–2004) further shows that: (i) existing fundamental and occupied niche centroids did not undergo directional change, (ii) interannual changes in the two niche centroids were correlated, (iii) temperatures in North America moved through niche space in a net centripetal fashion, and consequently, (iv) most areas throughout the range of the house sparrow tracked the existing fundamental niche centroid with respect to at least one temperature gradient. Following introduction to a new continent, the house sparrow rapidly tracked its thermal niche and established continent-wide distributional equilibrium with respect to major temperature gradients. These dynamics were mediated in large part by the species’ broad thermal physiological tolerances, high dispersal potential, competitive advantage in human-dominated landscapes, and climatically induced changes to the realized environmental space. Such insights may be used to conceptualize mechanistic climatic niche models in birds and other taxa.

Background Reproductive isolation (RI) is widely accepted as an important "check point" in the diversification process, since it defines irreversible evolutionary trajectories. Much less consensus exists about the processes that might drive RI. Here, we employ a formal quantitative analysis of genetic interactions at several stages of divergence within the ring species complex Ensatina eschscholtzii in order to assess the relative contribution of genetic and ecological divergence for the development of RI. Results By augmenting previous genetic datasets and adding new ecological data, we quantify levels of genetic and ecological divergence between populations and test how they correlate with a restriction of genetic admixture upon secondary contact. Our results indicate that the isolated effect of ecological divergence between parental populations does not result in reproductively isolated taxa, even when genetic transitions between parental taxa are narrow. Instead, processes associated with overall genetic divergence are the best predictors of reproductive isolation, and when parental taxa diverge in nuclear markers we observe a complete cessation of hybridization, even to sympatric occurrence of distinct evolutionary lineages. Although every parental population has diverged in mitochondrial DNA, its degree of divergence does not predict the extent of RI. Conclusions These results show that in Ensatina, the evolutionary outcomes of ecological divergence differ from those of genetic divergence. While evident properties of taxa may emerge via ecological divergence, such as adaptation to local environment, RI is likely to be a byproduct of processes that contribute to overall genetic divergence, such as time in geographic isolation, rather than being a direct outcome of local adaptation.

Niche theory is central to understanding how species respond geographically to climate change. It defines a species' realized niche in a biological community, its fundamental niche as determined by physiology, and its potential niche--the fundamental niche in a given environment or geographic space. However, most predictions of the effects of climate change on species' distributions are limited to correlative models of the realized niche, which assume that species are in distributional equilibrium with respect to the variables or gradients included in the model. Here, I present a mechanistic niche model that measures species' responses to major seasonal temperature gradients that interact with the physiology of the organism. I then use lethal physiological temperatures to parameterize the model for bird species in North and South America and show that most focal bird species are not in direct physiological equilibrium with the gradients. Results also show that most focal bird species possess broad thermal tolerances encompassing novel climates that could become available with climate change. I conclude with discussion of how mechanistic niche models may be used to (i) gain insights into the processes that cause species to respond to climate change and (ii) build more accurate correlative distribution models in birds and other species.