Global patterns of protection of elevational gradients in mountain ranges.

Protected areas (PAs) that span elevational gradients enhance protection for taxonomic and phylogenetic diversity and facilitate species range shifts under climate change. We quantified the global protection of elevational gradients by analyzing the elevational distributions of 44,155 PAs in 1,010 mountain ranges using the highest resolution digital elevation models available. We show that, on average, mountain ranges in Africa and Asia have the lowest elevational protection, ranges in Europe and South America have intermediate elevational protection, and ranges in North America and Oceania have the highest elevational protection. We use the Convention on Biological Diversity's Aichi Target 11 to assess the proportion of elevational gradients meeting the 17% suggested minimum target and examine how different protection categories contribute to elevational protection. When considering only strict PAs [International Union for Conservation of Nature (IUCN) categories I-IV, n = 24,706], nearly 40% of ranges do not contain any PAs, roughly half fail to meet the 17% target at any elevation, and ∼75% fail to meet the target throughout ≥50% of the elevational gradient. Observed elevational protection is well below optimal, and frequently below a null model of elevational protection. Including less stringent PAs (IUCN categories V-VI and nondesignated PAs, n = 19,449) significantly enhances elevational protection for most continents, but several highly biodiverse ranges require new or expanded PAs to increase elevational protection. Ensuring conservation outcomes for PAs with lower IUCN designations as well as strategically placing PAs to better represent and connect elevational gradients will enhance ecological representation and facilitate species range shifts under climate change.

The relationship between tree mortality from a pine beetle epidemic and increased dissolved copper levels in the upper Big Thompson River, Colorado.

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.

Habitat availability and gene flow influence diverging local population trajectories under scenarios of climate change: a place-based approach.

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.

Exposure of U.S. National Parks to land use and climate change 1900-2100.

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.

Ring distributions leading to species formation: a global topographic analysis of geographic barriers associated with ring species.

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.

Birds track their Grinnellian niche through a century of 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.

A spectral three-dimensional color space model of tree crown health.

Protecting the future of forests in the United States and other countries depends in part on our ability to monitor and map forest health conditions in a timely fashion to facilitate management of emerging threats and disturbances over a multitude of spatial scales. Remote sensing data and technologies have contributed to our ability to meet these needs, but existing methods relying on supervised classification are often limited to specific areas by the availability of imagery or training data, as well as model transferability. Scaling up and operationalizing these methods for general broadscale monitoring and mapping may be promoted by using simple models that are easily trained and projected across space and time with widely available imagery. Here, we describe a new model that classifies high resolution (~1 m2) 3-band red, green, blue (RGB) imagery from a single point in time into one of four color classes corresponding to tree crown condition or health: green healthy crowns, red damaged or dying crowns, gray damaged or dead crowns, and shadowed crowns where the condition status is unknown. These Tree Crown Health (TCH) models trained on data from the United States (US) Department of Agriculture, National Agriculture Imagery Program (NAIP), for all 48 States in the contiguous US and spanning years 2012 to 2019, exhibited high measures of model performance and transferability when evaluated using randomly withheld testing data (n = 122 NAIP state x year combinations; median overall accuracy 0.89-0.90; median Kappa 0.85-0.86). We present examples of how TCH models can detect and map individual tree mortality resulting from a variety of nationally significant native and invasive forest insects and diseases in the US. We conclude with discussion of opportunities and challenges for extending and implementing TCH models in support of broadscale monitoring and mapping of forest health.

Predictors for reproductive isolation in a ring species complex following genetic and ecological divergence.

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.

Topography and human pressure in mountain ranges alter expected species responses to climate change.

Climate change is leading to widespread elevational shifts thought to increase species extinction risk in mountains. We integrate digital elevation models with a metric of human pressure to examine changes in the amount of intact land area available for species undergoing elevational range shifts in all major mountain ranges globally (n = 1010). Nearly 60% of mountainous area is under intense human pressure, predominantly at low elevations and mountain bases. Consequently, upslope range shifts generally resulted in modeled species at lower elevations expanding into areas of lower human pressure and, due to complex topography, encountering more intact land area relative to their starting position. Such gains were often attenuated at high elevations as land-use constraints diminished and topographic constraints increased. Integrating patterns of topography and human pressure is essential for accurate species vulnerability assessments under climate change, as priorities for protecting, connecting, and restoring mountain landscapes may otherwise be misguided.

Keeping pace with climate change in global terrestrial protected areas.

Protected areas (PAs) are essential to biodiversity conservation, but their static boundaries may undermine their potential for protecting species under climate change. We assessed how the climatic conditions within global terrestrial PAs may change over time. By 2070, protection is expected to decline in cold and warm climates and increase in cool and hot climates over a wide range of precipitation. Most countries are expected to fail to protect >90% of their available climate at current levels. The evenness of climatic representation under protection-not the amount of area protected-positively influenced the retention of climatic conditions under protection. On average, protection retention would increase by ~118% if countries doubled their climatic representativeness under protection or by ~102% if countries collectively reduced emissions in accordance with global targets. Therefore, alongside adoption of mitigation policies, adaptation policies that improve the complementarity of climatic conditions within PAs will help countries safeguard biodiversity.

Climate change adaptation benefits of potential conservation partnerships.

We evaluate the world terrestrial network of protected areas (PAs) for its partnership potential in responding to climate change. That is, if a PA engaged in collaborative, trans-boundary management of species, by investing in conservation partnerships with neighboring areas, what climate change adaptation benefits might accrue? We consider core tenets of conservation biology related to protecting large areas with high environmental heterogeneity and low climate change velocity and ask how a series of biodiversity adaptation indicators change across spatial scales encompassing potential PA and non-PA partners. Less than 1% of current world terrestrial PAs equal or exceed the size of established and successful conservation partnerships. Partnering at this scale would increase the biodiversity adaptation indicators by factors up to two orders of magnitude, compared to a null model in which each PA is isolated. Most partnership area surrounding PAs is comprised of non-PAs (70%), indicating the importance of looking beyond the current network of PAs when promoting climate change adaptation. Given monumental challenges with PA-based species conservation in the face of climate change, partnerships provide a logical and achievable strategy for helping areas adapt. Our findings identify where strategic partnering efforts in highly vulnerable areas of the world may prove critical in safeguarding biodiversity.

Correction: Managing Climate Change Refugia for Climate Adaptation.

[This corrects the article DOI: 10.1371/journal.pone.0159909.].

Managing Climate Change Refugia for Climate Adaptation.

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.

Ecologically-Relevant Maps of Landforms and Physiographic Diversity for Climate Adaptation Planning.

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.

Protected Area Tourism in a Changing Climate: Will Visitation at US National Parks Warm Up or Overheat?

Climate change will affect not only natural and cultural resources within protected areas but also tourism and visitation patterns. The U.S. National Park Service systematically collects data regarding its 270+ million annual recreation visits, and therefore provides an opportunity to examine how human visitation may respond to climate change from the tropics to the polar regions. To assess the relationship between climate and park visitation, we evaluated historical monthly mean air temperature and visitation data (1979-2013) at 340 parks and projected potential future visitation (2041-2060) based on two warming-climate scenarios and two visitation-growth scenarios. For the entire park system a third-order polynomial temperature model explained 69% of the variation in historical visitation trends. Visitation generally increased with increasing average monthly temperature, but decreased strongly with temperatures > 25°C. Linear to polynomial monthly temperature models also explained historical visitation at individual parks (R2 0.12-0.99, mean = 0.79, median = 0.87). Future visitation at almost all parks (95%) may change based on historical temperature, historical visitation, and future temperature projections. Warming-mediated increases in potential visitation are projected for most months in most parks (67-77% of months; range across future scenarios), resulting in future increases in total annual visits across the park system (8-23%) and expansion of the visitation season at individual parks (13-31 days). Although very warm months at some parks may see decreases in future visitation, this potential change represents a relatively small proportion of visitation across the national park system. A changing climate is likely to have cascading and complex effects on protected area visitation, management, and local economies. Results suggest that protected areas and neighboring communities that develop adaptation strategies for these changes may be able to both capitalize on opportunities and minimize detriment related to changing visitation.

Climate exposure of US national parks in a new era of change.

US national parks are challenged by climate and other forms of broad-scale environmental change that operate beyond administrative boundaries and in some instances are occurring at especially rapid rates. Here, we evaluate the climate change exposure of 289 natural resource parks administered by the US National Park Service (NPS), and ask which are presently (past 10 to 30 years) experiencing extreme (<5th percentile or >95th percentile) climates relative to their 1901-2012 historical range of variability (HRV). We consider parks in a landscape context (including surrounding 30 km) and evaluate both mean and inter-annual variation in 25 biologically relevant climate variables related to temperature, precipitation, frost and wet day frequencies, vapor pressure, cloud cover, and seasonality. We also consider sensitivity of findings to the moving time window of analysis (10, 20, and 30 year windows). Results show that parks are overwhelmingly at the extreme warm end of historical temperature distributions and this is true for several variables (e.g., annual mean temperature, minimum temperature of the coldest month, mean temperature of the warmest quarter). Precipitation and other moisture patterns are geographically more heterogeneous across parks and show greater variation among variables. Across climate variables, recent inter-annual variation is generally well within the range of variability observed since 1901. Moving window size has a measureable effect on these estimates, but parks with extreme climates also tend to exhibit low sensitivity to the time window of analysis. We highlight particular parks that illustrate different extremes and may facilitate understanding responses of park resources to ongoing climate change. We conclude with discussion of how results relate to anticipated future changes in climate, as well as how they can inform NPS and neighboring land management and planning in a new era of change.

Forecasting distributional responses of limber pine to climate change at management-relevant scales in Rocky Mountain National Park.

Resource managers at parks and other protected areas are increasingly expected to factor climate change explicitly into their decision making frameworks. However, most protected areas are small relative to the geographic ranges of species being managed, so forecasts need to consider local adaptation and community dynamics that are correlated with climate and affect distributions inside protected area boundaries. Additionally, niche theory suggests that species' physiological capacities to respond to climate change may be underestimated when forecasts fail to consider the full breadth of climates occupied by the species rangewide. Here, using correlative species distribution models that contrast estimates of climatic sensitivity inferred from the two spatial extents, we quantify the response of limber pine (Pinus flexilis) to climate change in Rocky Mountain National Park (Colorado, USA). Models are trained locally within the park where limber pine is the community dominant tree species, a distinct structural-compositional vegetation class of interest to managers, and also rangewide, as suggested by niche theory. Model forecasts through 2100 under two representative concentration pathways (RCP 4.5 and 8.5 W/m(2)) show that the distribution of limber pine in the park is expected to move upslope in elevation, but changes in total and core patch area remain highly uncertain. Most of this uncertainty is biological, as magnitudes of projected change are considerably more variable between the two spatial extents used in model training than they are between RCPs, and novel future climates only affect local model predictions associated with RCP 8.5 after 2091. Combined, these results illustrate the importance of accounting for unknowns in species' climatic sensitivities when forecasting distributional scenarios that are used to inform management decisions. We discuss how our results for limber pine may be interpreted in the context of climate change vulnerability and used to help guide adaptive management.

Niche tracking and rapid establishment of distributional equilibrium in the house sparrow show potential responsiveness of species to climate change.

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.

A mechanistic niche model for measuring species' distributional responses to seasonal temperature gradients.

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.