Evaluating ecosystem protection and fragmentation of the world's major mountain regions.

Conserving mountains is important for protecting biodiversity because they have high beta diversity and endemicity, facilitate species movement, and provide numerous ecosystem benefits for people. Mountains are often thought to have lower levels of human modification and contain more protected area than surrounding lowlands. To examine this, we compared biogeographic attributes of the largest, contiguous, mountainous region on each continent. In each region, we generated detailed ecosystems based on Köppen-Geiger climate regions, ecoregions, and detailed landforms. We quantified anthropogenic fragmentation of these ecosystems based on human modification classes of large wild areas, shared lands, and cities and farms. Human modification for half the mountainous regions approached the global average, and fragmentation reduced the ecological integrity of mountain ecosystems up to 40%. Only one-third of the major mountainous regions currently meet the Kunming-Montreal Global Biodiversity Framework target of 30% coverage for all protected areas; furthermore, the vast majority of ecosystem types present in mountains were underrepresented in protected areas. By measuring ecological integrity and human-caused fragmentation with a detailed representation of mountain ecosystems, our approach facilitates tracking progress toward achieving conservation goals and better informs mountain conservation.

Evaluación de la protección y fragmentación ambiental de las principales regiones montañosas del mundo Resumen La conservación de las montañas es importante para proteger a la biodiversidad pues tienen una alta diversidad beta y endemismos, facilitan el movimiento y proporcionan numerosos beneficios ambientales para las personas. Con frecuencia creemos que las montañas tienen niveles más bajos de modificaciones humanas y que contienen más áreas protegidas que las tierras bajas que las rodean. Para evaluar lo anterior, hicimos una comparación entre los atributos biogeográficos de la región montañosa más grande y contigua en cada continente. En cada región generamos ecosistemas detallados con base en las regiones climáticas de Köppen­Geiger, ecorregiones y relieves detallados. Cuantificamos la fragmentación antropogénica de estos ecosistemas con base en las clases de modificación humana de las grandes áreas silvestres, tierras compartidas y ciudades y granjas. Las modificaciones humanas en la mitad de las regiones montañosas se aproximaron al promedio mundial, mientras que la fragmentación redujo la integridad ecológica de los ecosistemas montañosas hasta un 40%. Sólo un tercio de las principales regiones montañosas cumplen actualmente con el objetivo de 30% de cobertura para todas las áreas protegidas del Marco Mundial de Biodiversidad de Kunming­Montreal; además, la gran mayoría de los tipos de ecosistemas presentes en las montañas estaban subrepresentados dentro de las áreas protegidas. Con la medida de la integridad ecológica y la fragmentación antropogénica mediante una representación detallada de los ecosistemas montañosos, nuestra estrategia facilita el seguimiento del progreso hacia la obtención de los objetivos de conservación e informa de mejor manera a la conservación de las montañas.

Applying assessments of adaptive capacity to inform natural-resource management in a changing climate.

Adaptive capacity (AC)-the ability of a species to cope with or accommodate climate change-is a critical determinant of species vulnerability. Using information on species' AC in conservation planning is key to ensuring successful outcomes. We identified connections between a list of species' attributes (e.g., traits, population metrics, and behaviors) that were recently proposed for assessing species' AC and management actions that may enhance AC for species at risk of extinction. Management actions were identified based on evidence from the literature, a review of actions used in other climate adaptation guidance, and our collective experience in diverse fields of global-change ecology and climate adaptation. Selected management actions support the general AC pathways of persist in place or shift in space, in response to contemporary climate change. Some actions, such as genetic manipulations, can be used to directly alter the ability of species to cope with climate change, whereas other actions can indirectly enhance AC by addressing ecological or anthropogenic constraints on the expression of a species' innate abilities to adapt. Ours is the first synthesis of potential management actions directly linked to AC. Focusing on AC attributes helps improve understanding of how and why aspects of climate are affecting organisms, as well as the mechanisms by which management interventions affect a species' AC and climate change vulnerability. Adaptive-capacity-informed climate adaptation is needed to build connections among the causes of vulnerability, AC, and proposed management actions that can facilitate AC and reduce vulnerability in support of evolving conservation paradigms.

Aplicación de Evaluaciones de la Capacidad Adaptativa para Informar la Gestión de Recursos Naturales en un Clima Cambiante Resumen La capacidad adaptativa (CA) - la habilidad que tiene una especie para sobrellevar o acomodarse al cambio climático - es una determinante crítica de la vulnerabilidad de una especie. El uso de la información sobre la CA de una especie dentro de la planeación de la conservación es de suma importancia para asegurar resultados exitosos. Identificamos las conexiones entre una lista de atributos de las especies (p. ej.: características, métricas poblacionales, comportamientos) que fueron propuestos recientemente para la evaluación de la CA de las especies y las acciones de gestión que pueden mejorar la CA para las especies que se encuentran en riesgo de extinción. Las acciones de gestión fueron identificadas con base en la evidencia de la literatura, una revisión de acciones usadas en otras guías de adaptación climática y nuestra experiencia colectiva en diferentes campos de la ecología del cambio global y la adaptación climática. Ciertas acciones de gestión respaldan las vías generales de CA de persistir en el lugar o cambiar en el espacio como respuesta al cambio climático contemporáneo. Algunas acciones, como la manipulación genética, pueden usarse para alterar directamente la habilidad que tienen las especies para sobrellevar el cambio climático, mientras que otras acciones pueden mejorar indirectamente la CA al combatir las restricciones ecológicas o antropogénicas que existen sobre la expresión de las habilidades innatas de una especie para adaptarse. Nuestra síntesis es la primera que aborda acciones potenciales de gestión conectadas directamente con la CA. Enfocarse en los atributos de la CA ayuda a mejorar el conocimiento sobre cómo y por qué los aspectos climáticos están afectando a los organismos, así como los mecanismos mediante los cuales las intervenciones de gestión afectan la CA y la vulnerabilidad al cambio climático de la especie. La adaptación climática orientada por la capacidad adaptativa es necesaria para establecer conexiones entre las causas de la vulnerabilidad, la CA y las acciones de gestión propuestas que pueden facilitar la CA y reducir la vulnerabilidad como apoyo a los paradigmas cambiantes de la conservación.

Is the grass always greener? Land surface phenology reveals differences in peak and season-long vegetation productivity responses to climate and management.

Vegetation phenology-the seasonal timing and duration of vegetative phases-is controlled by spatiotemporally variable contributions of climatic and environmental factors plus additional potential influence from human management. We used land surface phenology derived from the Advanced Very High Resolution Radiometer and climate data to examine variability in vegetation productivity and phenological dates from 1989 to 2014 in the U.S. Northwestern Plains, a region with notable spatial heterogeneity in climate, vegetation, and land use. We first analyzed interannual trends in six phenological measures as a baseline. We then demonstrated how including annual-resolution predictors can provide more nuanced insights into measures of phenology between plant communities and across the ecoregion. Across the study area, higher annual precipitation increased both peak and season-long productivity. In contrast, higher mean annual temperatures tended to increase peak productivity but for the majority of the study area decreased season-long productivity. Annual precipitation and temperature had strong explanatory power for productivity-related phenology measures but predicted date-based measures poorly. We found that relationships between climate and phenology varied across the region and among plant communities and that factors such as recovery from disturbance and anthropogenic management also contributed in certain regions. In sum, phenological measures did not respond ubiquitously nor covary in their responses. Nonclimatic dynamics can decouple phenology from climate; therefore, analyses including only interannual trends should not assume climate alone drives patterns. For example, models of areas exhibiting greening or browning should account for climate, anthropogenic influence, and natural disturbances. Investigating multiple aspects of phenology to describe growing-season dynamics provides a richer understanding of spatiotemporal patterns that can be used for predicting ecosystem responses to future climates and land-use change. Such understanding allows for clearer interpretation of results for conservation, wildlife, and land management.

Supporting the adaptive capacity of species through more effective knowledge exchange with conservation practitioners.

There is an imperative for conservation practitioners to help biodiversity adapt to accelerating environmental change. Evolutionary biologists are well-positioned to inform the development of evidence-based management strategies that support the adaptive capacity of species and ecosystems. Conservation practitioners increasingly accept that management practices must accommodate rapid environmental change, but harbour concerns about how to apply recommended changes to their management contexts. Given the interest from both conservation practitioners and evolutionary biologists in adjusting management practices, we believe there is an opportunity to accelerate the required changes by promoting closer collaboration between these two groups. We highlight how evolutionary biologists can harness lessons from other disciplines about how to foster effective knowledge exchange to make a substantive contribution to the development of effective conservation practices. These lessons include the following: (1) recognizing why practitioners do and do not use scientific evidence; (2) building an evidence base that will influence management decisions; (3) translating theory into a format that conservation practitioners can use to inform management practices; and (4) developing strategies for effective knowledge exchange. Although efforts will be required on both sides, we believe there are rewards for both practitioners and evolutionary biologists, not least of which is fostering practices to help support the long-term persistence of species.

Factors influencing distributional shifts and abundance at the range core of a climate-sensitive mammal.

Species are frequently responding to contemporary climate change by shifting to higher elevations and poleward to track suitable climate space. However, depending on local conditions and species' sensitivity, the nature of these shifts can be highly variable and difficult to predict. Here, we examine how the American pika (Ochotona princeps), a philopatric, montane lagomorph, responds to climatic gradients at three spatial scales. Using mixed-effects modeling in an information-theoretic approach, we evaluated a priori model suites regarding predictors of site occupancy, relative abundance, and elevational-range retraction across 760 talus patches, nested within 64 watersheds across the Northern Rocky Mountains of North America, during 2017-2020. The top environmental predictors differed across these response metrics. Warmer temperatures in summer and winter were associated with lower occupancy, lower relative abundances, and greater elevational retraction across watersheds. Occupancy was also strongly influenced by habitat patch size, but only when combined with climate metrics such as actual evapotranspiration. Using a second analytical approach, acute heat stress and summer precipitation best explained retraction residuals (i.e., the relative extent of retraction given the original elevational range of occupancy). Despite the study domain occurring near the species' geographic-range center, where populations might have higher abundances and be at lower risk of climate-related stress, 33.9% of patches showed evidence of recent extirpations. Pika-extirpated sites averaged 1.44℃ warmer in summer than did occupied sites. Additionally, the minimum elevation of pika occupancy has retracted upslope in 69% of watersheds (mean: 281 m). Our results emphasize the nuance associated with evaluating species' range dynamics in response to climate gradients, variability, and temperature exceedances, especially in regions where species occupy gradients of conditions that may constitute multiple range edges. Furthermore, this study highlights the importance of evaluating diverse drivers across response metrics to improve the predictive accuracy of widely used, correlative models.

Freezing in a warming climate: Marked declines of a subnivean hibernator after a snow drought.

Recent snow droughts associated with unusually warm winters are predicted to increase in frequency and affect species dependent upon snowpack for winter survival. Changes in populations of some cold-adapted species have been attributed to heat stress or indirect effects on habitat from unusually warm summers, but little is known about the importance of winter weather to population dynamics and how responses to snow drought vary among sympatric species. We evaluated changes in abundance of hoary marmots (Marmota caligata) over a period that included a year of record-low snowpack to identify mechanisms associated with weather and snowpack. To consider interspecies comparisons, our analysis used the same a priori model set as a concurrent study that evaluated responses of American pikas (Ochotona princeps) to weather and snowpack in the same study area of North Cascades National Park, Washington, USA. We hypothesized that marmot abundance reflected mechanisms related to heat stress, cold stress, cold exposure without an insulating snowpack, snowpack duration, atmospheric moisture, growing-season precipitation, or select combinations of these mechanisms. Changes in marmot abundances included a 74% decline from 2007 to 2016 and were best explained by an interaction of chronic dryness with exposure to acute cold without snowpack in winter. Physiological stress during hibernation from exposure to cold, dry air appeared to be the most likely mechanism of change in marmot abundance. Alternative mechanisms associated with changes to winter weather, including early emergence from hibernation or altered vegetation dynamics, had less support. A post hoc assessment of vegetative phenology and productivity did not support vegetation dynamics as a primary driver of marmot abundance across years. Although marmot and pika abundances were explained by strikingly similar models over periods of many years, details of the mechanisms involved likely differ between species because pika abundances increased in areas where marmots declined. Such differences may lead to diverging geographic distributions of these species as global change continues.

Ecological consequences of anomalies in atmospheric moisture and snowpack.

Although increased frequency of extreme-weather events is one of the most secure predictions associated with contemporary climate change, effects of such events on distribution and abundance of climate-sensitive species remain poorly understood. Montane ecosystems may be especially sensitive to extreme weather because of complex abiotic and biotic interactions that propagate from climate-driven reductions in snowpack. Snowpack not only protects subnivean biotas from extreme cold, but also influences forage availability through timing of melt-off and water availability. We related relative abundances of an alpine mammal, the American pika (Ochotona princeps), to measures of weather and snowpack dynamics over an 8-yr period that included before and after a year of record-low snowpack in Washington, USA. We sought to (1) quantify any change in pika abundance associated with the snowpack anomaly and (2) identify aspects of weather and snowpack that influenced abundance of pikas. Pikas showed a 1-yr lag response to the snowpack anomaly and exhibited marked declines in abundance at elevations below 1,400 m simultaneous with increased abundances at higher elevations. Atmospheric moisture, indexed by vapor pressure deficit (VPD), was especially important, evidenced by strong support for the top-ranked model that included the interaction of VPD with snowpack duration. Notably, our novel application of VPD from gridded climate data for analyses of animal abundances shows strong potential for improving species distribution models because VPD represents an important aspect of weather that influences the physiology and habitat of biota. Pikas were apparently affected by cold stress without snowpack at mid elevations, whereas changes to forage associated with snowpack and VPD were influential at high and low elevations. Our results reveal context dependency in pika responses to weather and illustrate how snow drought can lead to rapid change in the abundance of subnivean animals.

Change in dominance determines herbivore effects on plant biodiversity.

Herbivores alter plant biodiversity (species richness) in many of the world's ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis-that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally.

Adaptive population divergence and directional gene flow across steep elevational gradients in a climate-sensitive mammal.

The ecological effects of climate change have been shown in most major taxonomic groups; however, the evolutionary consequences are less well-documented. Adaptation to new climatic conditions offers a potential long-term mechanism for species to maintain viability in rapidly changing environments, but mammalian examples remain scarce. The American pika (Ochotona princeps) has been impacted by recent climate-associated extirpations and range-wide reductions in population sizes, establishing it as a sentinel mammalian species for climate change. To investigate evidence for local adaptation and reconstruct patterns of genomic diversity and gene flow across rapidly changing environments, we used a space-for-time design and restriction site-associated DNA sequencing to genotype American pikas along two steep elevational gradients at 30,966 SNPs and employed independent outlier detection methods that scanned for genotype-environment associations. We identified 338 outlier SNPs detected by two separate analyses and/or replicated in both transects, several of which were annotated to genes involved in metabolic function and oxygen transport. Additionally, we found evidence of directional gene flow primarily downslope from high-elevation populations, along with reduced gene flow at outlier loci. If this trend continues, elevational range contractions in American pikas will likely be from local extirpation rather than upward movement of low-elevation individuals; this, in turn, could limit the potential for adaptation within this landscape. These findings are of particular relevance for future conservation and management of American pikas and other elevationally restricted, thermally sensitive species.

Mechanistic variables can enhance predictive models of endotherm distributions: the American pika under current, past, and future climates.

How climate constrains species' distributions through time and space is an important question in the context of conservation planning for climate change. Despite increasing awareness of the need to incorporate mechanism into species distribution models (SDMs), mechanistic modeling of endotherm distributions remains limited in this literature. Using the American pika (Ochotona princeps) as an example, we present a framework whereby mechanism can be incorporated into endotherm SDMs. Pika distribution has repeatedly been found to be constrained by warm temperatures, so we used Niche Mapper, a mechanistic heat-balance model, to convert macroclimate data to pika-specific surface activity time in summer across the western United States. We then explored the difference between using a macroclimate predictor (summer temperature) and using a mechanistic predictor (predicted surface activity time) in SDMs. Both approaches accurately predicted pika presences in current and past climate regimes. However, the activity models predicted 8-19% less habitat loss in response to annual temperature increases of ~3-5 °C predicted in the region by 2070, suggesting that pikas may be able to buffer some climate change effects through behavioral thermoregulation that can be captured by mechanistic modeling. Incorporating mechanism added value to the modeling by providing increased confidence in areas where different modeling approaches agreed and providing a range of outcomes in areas of disagreement. It also provided a more proximate variable relating animal distribution to climate, allowing investigations into how unique habitat characteristics and intraspecific phenotypic variation may allow pikas to exist in areas outside those predicted by generic SDMs. Only a small number of easily obtainable data are required to parameterize this mechanistic model for any endotherm, and its use can improve SDM predictions by explicitly modeling a widely applicable direct physiological effect: climate-imposed restrictions on activity. This more complete understanding is necessary to inform climate adaptation actions, management strategies, and conservation plans.

Predictors of Current and Longer-Term Patterns of Abundance of American Pikas (Ochotona princeps) across a Leading-Edge Protected Area.

American pikas (Ochotona princeps) have been heralded as indicators of montane-mammal response to contemporary climate change. Pikas no longer occupy the driest and lowest-elevation sites in numerous parts of their geographic range. Conversely, pikas have exhibited higher rates of occupancy and persistence in Rocky Mountain and Sierra Nevada montane 'mainlands'. Research and monitoring efforts on pikas across the western USA have collectively shown the nuance and complexity with which climate will often act on species in diverse topographic and climatic contexts. However, to date no studies have investigated habitat, distribution, and abundance of pikas across hundreds of sites within a remote wilderness area. Additionally, relatively little is known about whether climate acts most strongly on pikas through direct or indirect (e.g., vegetation-mediated) mechanisms. During 2007-2009, we collectively hiked >16,000 km throughout the 410,077-ha Glacier National Park, Montana, USA, in an effort to identify topographic, microrefugial, and vegetative characteristics predictive of pika abundance. We identified 411 apparently pika-suitable habitat patches with binoculars (in situ), and surveyed 314 of them for pika signs. Ranking of alternative logistic-regression models based on AICc scores revealed that short-term pika abundances were positively associated with intermediate elevations, greater cover of mosses, and taller forbs, and decreased each year, for a total decline of 68% during the three-year study; whereas longer-term abundances were associated only with static variables (longitude, elevation, gradient) and were lower on north-facing slopes. Earlier Julian date and time of day of the survey (i.e., midday vs. not) were associated with lower observed pika abundance. We recommend that wildlife monitoring account for this seasonal and diel variation when surveying pikas. Broad-scale information on status and abundance determinants of montane mammals, especially for remote protected areas, is crucial for land and wildlife-resource managers trying to anticipate mammalian responses to climate change.

Plastic pikas: Behavioural flexibility in low-elevation pikas (Ochotona princeps).

Behaviour is an important mechanism for accommodating rapid environmental changes. Understanding a species' capacity for behavioural plasticity is therefore a key, but understudied, aspect of developing tractable conservation and management plans under climate-change scenarios. Here, we quantified behavioural differences between American pikas (Ochotona princeps) living in an atypical, low-elevation habitat versus those living in a more-typical, alpine habitat. With respect to foraging strategy, low-elevation pikas spent more time consuming vegetation and less time caching food for winter, compared to high-elevation pikas. Low-elevation pikas were also far more likely to be detected in forested microhabitats off the talus than their high-elevation counterparts at midday. Finally, pikas living in the atypical habitat had smaller home range sizes compared to those in typical habitat or any previously published home ranges for this species. Our findings indicate that behavioural plasticity likely allows pikas to accommodate atypical conditions in this low-elevation habitat, and that they may rely on critical habitat factors such as suitable microclimate refugia to behaviourally thermoregulate. Together, these results suggest that behavioural adjustments are one important mechanism by which pikas can persist outside of their previously appreciated dietary and thermal niches.

Assessing the components of adaptive capacity to improve conservation and management efforts under global change.

Natural-resource managers and other conservation practitioners are under unprecedented pressure to categorize and quantify the vulnerability of natural systems based on assessment of the exposure, sensitivity, and adaptive capacity of species to climate change. Despite the urgent need for these assessments, neither the theoretical basis of adaptive capacity nor the practical issues underlying its quantification has been articulated in a manner that is directly applicable to natural-resource management. Both are critical for researchers, managers, and other conservation practitioners to develop reliable strategies for assessing adaptive capacity. Drawing from principles of classical and contemporary research and examples from terrestrial, marine, plant, and animal systems, we examined broadly the theory behind the concept of adaptive capacity. We then considered how interdisciplinary, trait- and triage-based approaches encompassing the oft-overlooked interactions among components of adaptive capacity can be used to identify species and populations likely to have higher (or lower) adaptive capacity. We identified the challenges and value of such endeavors and argue for a concerted interdisciplinary research approach that combines ecology, ecological genetics, and eco-physiology to reflect the interacting components of adaptive capacity. We aimed to provide a basis for constructive discussion between natural-resource managers and researchers, discussions urgently needed to identify research directions that will deliver answers to real-world questions facing resource managers, other conservation practitioners, and policy makers. Directing research to both seek general patterns and identify ways to facilitate adaptive capacity of key species and populations within species, will enable conservation ecologists and resource managers to maximize returns on research and management investment and arrive at novel and dynamic management and policy decisions.

Successes and challenges from formation to implementation of eleven broad-extent conservation programs.

Integration of conservation partnerships across geographic, biological, and administrative boundaries is increasingly relevant because drivers of change, such as climate shifts, transcend these boundaries. We explored successes and challenges of established conservation programs that span multiple watersheds and consider both social and ecological concerns. We asked representatives from a diverse set of 11 broad-extent conservation partnerships in 29 countries 17 questions that pertained to launching and maintaining partnerships for broad-extent conservation, specifying ultimate management objectives, and implementation and learning. Partnerships invested more funds in implementing conservation actions than any other aspect of conservation, and a program's context (geographic extent, United States vs. other countries, developed vs. developing nation) appeared to substantially affect program approach. Despite early successes of these organizations and benefits of broad-extent conservation, specific challenges related to uncertainties in scaling up information and to coordination in the face of diverse partner governance structures, conflicting objectives, and vast uncertainties regarding future system dynamics hindered long-term success, as demonstrated by the focal organizations. Engaging stakeholders, developing conservation measures, and implementing adaptive management were dominant challenges. To inform future research on broad-extent conservation, we considered several challenges when we developed detailed questions, such as what qualities of broad-extent partnerships ensure they complement, integrate, and strengthen, rather than replace, local conservation efforts and which adaptive management processes yield actionable conservation strategies that account explicitly for dynamics and uncertainties regarding multiscale governance, environmental conditions, and knowledge of the system?

Understanding relationships among abundance, extirpation, and climate at ecoregional scales.

Recent research on mountain-dwelling species has illustrated changes in species distributional patterns in response to climate change. Abundance of a species will likely provide an earlier warning indicator of change than will occupancy, yet relationships between abundance and climatic factors have received less attention. We tested whether predictors of counts of American pikas (Ochotona princeps) during surveys from the Great Basin region in 1994-1999 and 2003-2008 differed between the two periods. Additionally, we tested whether various modeled aspects of ecohydrology better predicted relative density than did average annual precipitation, and whether risk of site-wide extirpation predicted subsequent population counts of pikas. We observed several patterns of change in pika abundance at range edges that likely constitute early warnings of distributional shifts. Predictors of pika abundance differed strongly between the survey periods, as did pika extirpation patterns previously reported from this region. Additionally, maximum snowpack and growing-season precipitation resulted in better-supported models than those using average annual precipitation, and constituted two of the top three predictors of pika density in the 2000s surveys (affecting pikas perhaps via vegetation). Unexpectedly, we found that extirpation risk positively predicted subsequent population size. Our results emphasize the need to clarify mechanisms underlying biotic responses to recent climate change at organism-relevant scales, to inform management and conservation strategies for species of concern.

Testing alternative models of climate-mediated extirpations.

Biotic responses to climate change will vary among taxa and across latitudes, elevational gradients, and degrees of insularity. However, due to factors such as phenotypic plasticity, ecotypic variation, and evolved tolerance to thermal stress, it remains poorly understood whether losses should be greatest in populations experiencing the greatest climatic change or living in places where the prevailing climate is closest to the edge of the species' bioclimatic envelope (e.g., at the hottest, driest sites). Research on American pikas (Ochotona princeps) in montane areas of the Great Basin during 1994-1999 suggested that 20th-century population extirpations were predicted by a combination of biogeographic, anthropogenic, and especially climatic factors. Surveys during 2005-2007 documented additional extirpations and within-site shifts of pika distributions at remaining sites. To evaluate the evidence in support of alternative hypotheses involving effects of thermal stress on pikas, we placed temperature sensors at 156 locations within pika habitats in the vicinity of 25 sites with historical records of pikas in the Basin. We related these time series of sensor data to data on ambient temperature from weather stations within the Historical Climate Network. We then used these highly correlated relationships, combined with long-term data from the same weather stations, to hindcast temperatures within pika habitats from 1945 through 2006. To explain patterns of loss, we posited three alternative classes of direct thermal stress: (1) acute cold stress (number of days below a threshold temperature); (2) acute heat stress (number of days above a threshold temperature); and (3) chronic heat stress (average summer temperature). Climate change was defined as change in our thermal metrics between two 31-yr periods: 1945-1975 and 1976-2006. We found that patterns of persistence were well predicted by metrics of climate. Our best models suggest some effects of climate change; however, recent and long-term metrics of chronic heat stress and acute cold stress, neither previously recognized as sources of stress for pikas, were some of the best predictors of pika persistence. Results illustrate that extremely rapid distributional shifts can be explained by climatic influences and have implications for conservation topics such as reintroductions and early-warning indicators.

Taxonomic considerations in listing subspecies under the U.S. Endangered Species Act.

The U.S. Endangered Species Act (ESA) allows listing of subspecies and other groupings below the rank of species. This provides the U.S. Fish and Wildlife Service and the National Marine Fisheries Service with a means to target the most critical unit in need of conservation. Although roughly one-quarter of listed taxa are subspecies, these management agencies are hindered by uncertainties about taxonomic standards during listing or delisting activities. In a review of taxonomic publications and societies, we found few subspecies lists and none that stated standardized criteria for determining subspecific taxa. Lack of criteria is attributed to a centuries-old debate over species and subspecies concepts. Nevertheless, the critical need to resolve this debate for ESA listings led us to propose that minimal biological criteria to define disjunct subspecies (legally or taxonomically) should include the discreteness and significance criteria of distinct population segments (as defined under the ESA). Our subspecies criteria are in stark contrast to that proposed by supporters of the phylogenetic species concept and provide a clear distinction between species and subspecies. Efforts to eliminate or reduce ambiguity associated with subspecies-level classifications will assist with ESA listing decisions. Thus, we urge professional taxonomic societies to publish and periodically update peer-reviewed species and subspecies lists. This effort must be paralleled throughout the world for efficient taxonomic conservation to take place.