Climate mismatches with ectomycorrhizal fungi contribute to migration lag in North American tree range shifts.

Climate change will likely shift plant and microbial distributions, creating geographic mismatches between plant hosts and essential microbial symbionts (e.g., ectomycorrhizal fungi, EMF). The loss of historical interactions, or the gain of novel associations, can have important consequences for biodiversity, ecosystem processes, and plant migration potential, yet few analyses exist that measure where mycorrhizal symbioses could be lost or gained across landscapes. Here, we examine climate change impacts on tree-EMF codistributions at the continent scale. We built species distribution models for 400 EMF species and 50 tree species, integrating fungal sequencing data from North American forest ecosystems with tree species occurrence records and long-term forest inventory data. Our results show the following: 1) tree and EMF climate suitability to shift toward higher latitudes; 2) climate shifts increase the size of shared tree-EMF habitat overall, but 35% of tree-EMF pairs are at risk of declining habitat overlap; 3) climate mismatches between trees and EMF are projected to be greater at northern vs. southern boundaries; and 4) tree migration lag is correlated with lower richness of climatically suitable EMF partners. This work represents a concentrated effort to quantify the spatial extent and location of tree-EMF climate envelope mismatches. Our findings also support a biotic mechanism partially explaining the failure of northward tree species migrations with climate change: reduced diversity of co-occurring and climate-compatible EMF symbionts at higher latitudes. We highlight the conservation implications for identifying areas where tree and EMF responses to climate change may be highly divergent.

Climate change and population persistence in a hibernating marsupial.

Climate change has physiological consequences on organisms, ecosystems and human societies, surpassing the pace of organismal adaptation. Hibernating mammals are particularly vulnerable as winter survival is determined by short-term physiological changes triggered by temperature. In these animals, winter temperatures cannot surpass a certain threshold, above which hibernators arouse from torpor, increasing several fold their energy needs when food is unavailable. Here, we parameterized a numerical model predicting energy consumption in heterothermic species and modelled winter survival at different climate change scenarios. As a model species, we used the arboreal marsupial monito del monte (genus Dromiciops), which is recognized as one of the few South American hibernators. We modelled four climate change scenarios (from optimistic to pessimistic) based on IPCC projections, predicting that northern and coastal populations (Dromiciops bozinovici) will decline because the minimum number of cold days needed to survive the winter will not be attained. These populations are also the most affected by habitat fragmentation and changes in land use. Conversely, Andean and other highland populations, in cooler environments, are predicted to persist and thrive. Given the widespread presence of hibernating mammals around the world, models based on simple physiological parameters, such as this one, are becoming essential for predicting species responses to warming in the short term.

Ice age-driven range shifts of diploids and expanding autotetraploids of Biscutella laevigata within a conserved niche.

Early studies of the textbook mixed-ploidy system Biscutella laevigata highlighted diploids restricted to never-glaciated lowlands and tetraploids at high elevations across the European Alps, promoting the hypothesis that whole-genome duplication (WGD) is advantageous under environmental changes. Here we addressed long-held hypotheses on the role of hybridisation at the origin of the tetraploids, their single vs multiple origins, and whether a shift in climatic niche accompanied WGD. Climatic niche modelling together with spatial genetics and coalescent modelling based on ddRAD-seq genotyping of 17 diploid and 19 tetraploid populations was used to revisit the evolution of this species complex in space and time. Diploids differentiated into four genetic lineages corresponding to allopatric glacial refugia at the onset of the last ice age, whereas tetraploids displaying tetrasomic inheritance formed a uniform group that originated from southern diploids before the last glacial maximum. Derived from diploids occurring at high elevation, autotetraploids likely inherited their adaptation to high elevation rather than having evolved it through or after WGD. They further presented considerable postglacial expansion across the Alps and underwent admixture with diploids. Although the underpinnings of the successful expansion of autotetraploids remain elusive, differentiation in B. laevigata was chiefly driven by the glacial history of the Alps.

Insufficient and biased representation of species geographic responses to climate change.

The geographic redistributions of species due to a rapidly changing climate are poised to perturb ecological communities and significantly impact ecosystems and human livelihoods. Effectively managing these biological impacts requires a thorough understanding of the patterns and processes of species geographic range shifts. While substantial recent redistributions have been identified and recognized to vary by taxon, region, and range geometry, there are large gaps and biases in the available evidence. Here, we use the largest compilation of geographic range change observations to date, comprised of 33,016 potential redistributions across 12,009 species, to formally assess within- and cross-species coverage and biases and to motivate future data collection. We find that species coverage varies strongly by taxon and underrepresents species at high and low latitudes. Within species, assessments of potential redistributions came from parts of their geographic range that were highly uneven and non-representative. For most species and taxa, studies were strongly biased toward the colder parts of species' distributions and thus significantly underrepresented populations that might get pushed beyond their maximum temperature limits. Coverage of potential leading and trailing geographic range edges under a changing climate was similarly uneven. Only 8% of studied species were assessed at both high and low latitude and elevation range edges, with most only covered at one edge. This suggests that substantial within-species biases exacerbate the considerable geographic and taxonomic among-species unevenness in evidence. Our results open the door for a more quantitative accounting for existing knowledge biases in climate change ecology and a more informed management and conservation. Our findings offer guidance for future data collection that better addresses information gaps and provides a more effective foundation for managing the biological impacts of climate change.

Can internal range structure predict range shifts?

Poleward and uphill range shifts are a common-but variable-response to climate change. We lack understanding regarding this interspecific variation; for example, functional traits show weak or mixed ability to predict range shifts. Characteristics of species' ranges may enhance prediction of range shifts. However, the explanatory power of many range characteristics-especially within-range abundance patterns-remains untested. Here, we introduce a hypothesis framework for predicting range-limit population trends and range shifts from the internal structure of the geographic range, specifically range edge hardness, defined as abundance within range edges relative to the whole range. The inertia hypothesis predicts that high edge abundance facilitates expansions along the leading range edge but creates inertia (either more individuals must disperse or perish) at the trailing range edge such that the trailing edge recedes slowly. In contrast, the limitation hypothesis suggests that hard range edges are the signature of strong limits (e.g. biotic interactions) that force faster contraction of the trailing edge but block expansions at the leading edge of the range. Using a long-term avian monitoring dataset from northern Minnesota, USA, we estimated population trends for 35 trailing-edge species and 18 leading-edge species and modelled their population trends as a function of range edge hardness derived from eBird data. We found limited evidence of associations between range edge hardness and range-limit population trends. Trailing-edge species with harder range edges were slightly more likely to be declining, demonstrating weak support for the limitation hypothesis. In contrast, leading-edge species with harder range edges were slightly more likely to be increasing, demonstrating weak support for the inertia hypothesis. These opposing results for the leading and trailing range edges might suggest that different mechanisms underpin range expansions and contractions, respectively. As data and state-of-the-art modelling efforts continue to proliferate, we will be ever better equipped to map abundance patterns within species' ranges, offering opportunities to anticipate range shifts through the lens of the geographic range.

Temperature-dependent dispersal and ectotherm species' distributions in a warming world.

Dispersal is a crucial component of species' responses to climate warming. Warming-induced changes in species' distributions are the outcome of how temperature affects dispersal at the individual level. Yet, there is little or no theory that considers the temperature dependence of dispersal when investigating the impacts of warming on species' distributions. Here I take a first step towards filling this key gap in our knowledge. I focus on ectotherms, species whose body temperature depends on the environmental temperature, not least because they constitute the majority of biodiversity on the planet. I develop a mathematical model of spatial population dynamics that explicitly incorporates mechanistic descriptions of ectotherm life history trait responses to temperature. A novel feature of this framework is the explicit temperature dependence of all phases of dispersal: emigration, transfer and settlement. I report three key findings. First, dispersal, regardless of whether it is random or temperature-dependent, allows both tropical and temperate ectotherms to track warming-induced changes in their thermal environments and to expand their distributions beyond the lower and upper thermal limits of their respective climate envelopes. In the absence of dispersal mortality, warming does not alter these new distributional limits. Second, an analysis based solely on trait response data predicts that tropical ectotherms should be able to expand their distributions polewards to a greater degree than temperate ectotherms. Analysis of the dynamical model confirms this prediction. Tropical ectotherms have an advantage when moving to cooler climates because they experience lower within-patch and dispersal mortality, and their higher thermal optima and maximal birth rates allow them to take advantage of the warmer parts of the year. Previous theory has shown that tropical ectotherms are more successful in invading and adapting the temperate climates than vice versa. This study provides the key missing piece, by showing how temperature-dependent dispersal could facilitate both invasion and adaptation. Third, dispersal mortality does not affect the poleward expansion of ectotherm distributions. But, it prevents both tropical and temperate ectotherms from maintaining sink populations in localities that are too warm to be viable in the absence of dispersal. Dispersal mortality also affects species' abundance patterns, causing a larger decline in abundance throughout the range when species disperse randomly rather than in response to thermal habitat suitability. In this way, dispersal mortality can facilitate the evolution of dispersal modes that maximize fitness in warmer thermal environments.

Body size mediates trophic interaction strength of novel fish assemblages under climate change.

Ecological similarity plays an important role in biotic interactions. Increased body size similarity of competing species, for example, increases the strength of their biotic interactions. Body sizes of many exothermic species are forecast to be altered under global warming, mediating shifts in existing trophic interactions among species, in particular for species with different thermal niches. Temperate rocky reefs along the southeast coast of Australia are located in a climate warming hotspot and now house a mixture of temperate native fish species and poleward range-extending tropical fishes (vagrants), creating novel species assemblages. Here, we studied the relationship between body size similarity and trophic overlap between individual temperate native and tropical vagrant fishes. Dietary niche overlap between vagrant and native fish species increased as their body sizes converged, based on both stomach content composition (short-term diet), stable isotope analyses (integrated long-term diet) and similarity in consumed prey sizes. We conclude that the warming-induced faster growth rates of tropical range-extending fish species at their cool water ranges will continue to converge their body size towards and strengthen their degree of trophic interactions and dietary overlap with co-occurring native temperate species under increasing ocean warming. The strengthening of these novel competitive interactions is likely to drive changes to temperate food web structures and reshuffle existing species community structures.

Patterns and correlates of potential range shifts of bat species in China in the context of climate change.

Climate change may diminish biodiversity; thus, it is urgent to predict how species' ranges may shift in the future by integrating multiple factors involving more taxa. Bats are particularly sensitive to climate change due to their high surface-to-volume ratio. However, few studies have considered geographic variables associated with roost availability and even fewer have linked the distributions of bats to their thermoregulation and energy regulation traits. We used species distribution models to predict the potential distributions of 12 bat species in China under current and future greenhouse gas emission scenarios (SSP1-2.6 and SSP5-8.5) and examined factors that could affect species' range shifts, including climatic, geographic, habitat, and human activity variables and wing surface-to-mass ratio (S-MR). The results suggest that Ia io, Rhinolophus ferrumequinum, and Rhinolophus rex should be given the highest priority for conservation in future climate conservation strategies. Most species were predicted to move northward, except for I. io and R. rex, which moved southward. Temperature seasonality, distance to forest, and distance to karst or cave were the main environmental factors affecting the potential distributions of bats. We found significant relationships between S-MR and geographic distribution, current potential distribution, and future potential distribution in the 2050s. Our work highlights the importance of analyzing range shifts of species with multifactorial approaches, especially for species traits related to thermoregulation and energy regulation, to provide targeted conservation strategies.

Patrones y correlaciones de los cambios potenciales en la distribución de las especies de murciélago de China en el contexto del cambio climático Resumen El cambio climático puede disminuir la biodiversidad, por lo que es urgente pronosticar cómo puede cambiar en el futuro la distribución de las especies mediante la integración de múltiples factores que involucren a más taxones. Los murciélagos son particularmente sensibles al cambio climático debido a que tienen una gran proporción superficie­volumen. Sin embargo, hay pocos estudios que han considerado las variables asociadas con la disponibilidad de nidos y son todavía menos los que han conectado la distribución de los murciélagos con sus rasgos de termorregulación y regulación de energía. Usamos modelos de distribución de especies para pronosticar la distribución potencial de doce especies de murciélago en China bajo escenarios actuales y futuros de emisión de gases de efecto invernadero (SSP1­2.6 y SSP5­8.5) y analizamos los factores que podrían afectar el cambio en la distribución de las especies, incluyendo las variables climáticas, geográficas, de hábitat y de actividad humana y la proporción entre la superficie del ala y la masa (P S­M). Los resultados sugieren que Ia io, Rhinolophus ferrumequinum y R. rex deberían ser la mayor prioridad de conservación para las estrategias de conservación climáticas en el futuro. Pronosticamos que la mayoría de las especies se desplazarían al norte, a excepción de I. io y R. rex, que se desplazarían hacia el sur. Los principales factores que afectaron la distribución potencial de los murciélagos fueron la estacionalidad de la temperatura, la distancia al bosque y la distancia a la cueva o al karst. Encontramos una relación significativa entre la P S­M y la distribución geográfica, la distribución potencial actual y la distribución potencial para la década de 2050. Nuestra investigación destaca la importancia del análisis de los cambios de distribución de las especies con enfoques multifactoriales, especialmente para los rasgos de especie relacionados con la termorregulación y la regulación de energía, para proporcionar estrategias de conservación focalizadas.

The turnover of plant-frugivore interactions along plant range expansion: consequences for natural colonization processes.

Plant-animal mutualisms such as seed dispersal are key interactions for sustaining plant range shifts. It remains elusive whether the organization of interactions with seed dispersers is reconfigured along the expansion landscape template and, if so, whether its effects accelerate or slow colonization. Here we analyse plant-frugivore interactions in a scenario of rapid population expansion of a Mediterranean juniper. We combined network analyses with field surveys, sampling interactions between individual plants and frugivores by DNA-barcoding and phototrapping over two seasons. We assess the role of intrinsic and extrinsic intraspecific variability in shaping interactions and we estimate the individual plant contributions to the seed rain. The whole interaction network was highly structured, with a distinct set of modules including individual plants and frugivore species arranged concordantly along the expansion gradient. The modular configuration was partially shaped by individual neighbourhood context (density and fecundity) and phenotypic traits (cone size). Interaction reconfiguration resulted in a higher and more uneven propagule contribution, with most effective dispersers having a prominent role at the colonization front stand, where a distinct subset of early arriving plants dominated the seed rain. Our study offers new insights into the key role of mutualistic interactions in colonization scenarios by promoting fast plant expansion processes.

Observed forest trait velocities have not kept pace with hydraulic stress from climate change.

The extent to which future climate change will increase forest stress and the amount to which species and forest ecosystems can acclimate or adapt to increased stress is a major unknown. We used high-resolution maps of hydraulic traits representing the diversity in tree drought tolerance across the United States, a hydraulically enabled tree model, and forest inventory observations of demographic shifts to quantify the ability for within-species acclimation and between-species range shifts to mediate climate stress. We found that forests are likely to experience increases in both acute and chronic hydraulic stress with climate change. Based on current species distributions, regional hydraulic trait diversity was sufficient to buffer against increased stress in 88% of forested areas. However, observed trait velocities in 81% of forested areas are not keeping up with the rate required to ameliorate projected future stress without leaf area acclimation.

Sublethal consequences of ultraviolet radiation exposure on vertebrates: Synthesis through meta-analysis.

Ultraviolet radiation (UVR) from the sun is a natural daytime stressor for vertebrates in both terrestrial and aquatic ecosystems. UVR effects on the physiology of vertebrates manifest at the cellular level, but have bottom-up effects at the tissue level and on whole-animal performance and behaviours. Climate change and habitat loss (i.e. loss of shelter from UVR) could interact with and exacerbate the genotoxic and cytotoxic impacts of UVR on vertebrates. Therefore, it is important to understand the range and magnitude of effects that UVR can have on a diversity of physiological metrics, and how these may be shaped by taxa, life stage or geographical range in the major vertebrate groups. Using a meta-analytical approach, we used 895 observations from 47 different vertebrate species (fish, amphibian, reptile and bird), and 51 physiological metrics (i.e. cellular, tissue and whole-animal metrics), across 73 independent studies, to elucidate the general patterns of UVR effects on vertebrate physiology. We found that while UVR's impacts on vertebrates are generally negative, fish and amphibians were the most susceptible taxa, adult and larvae were the most susceptible life stages, and animals inhabiting temperate and tropical latitudes were the most susceptible to UVR stress. This information is critical to further our understanding of the adaptive capacity of vulnerable taxon to UVR stress, and the wide-spread sublethal physiological effects of UVR on vertebrates, such as DNA damage and cellular stress, which may translate up to impaired growth and locomotor performance. These impairments to individual fitness highlighted by our study may potentially cause disruptions at the ecosystem scale, especially if the effects of this pervasive diurnal stressor are exacerbated by climate change and reduced refuge due to habitat loss and degradation. Therefore, conservation of habitats that provide refuge to UVR stress will be critical to mitigate stress from this pervasive daytime stressor.

Protected areas not likely to serve as steppingstones for species undergoing climate-induced range shifts.

Species across the planet are shifting their ranges to track suitable climate conditions in response to climate change. Given that protected areas have higher quality habitat and often harbor higher levels of biodiversity compared to unprotected lands, it is often assumed that protected areas can serve as steppingstones for species undergoing climate-induced range shifts. However, there are several factors that may impede successful range shifts among protected areas, including the distance that must be traveled, unfavorable human land uses and climate conditions along potential movement routes, and lack of analogous climates. Through a species-agnostic lens, we evaluate these factors across the global terrestrial protected area network as measures of climate connectivity, which is defined as the ability of a landscape to facilitate or impede climate-induced movement. We found that over half of protected land area and two-thirds of the number of protected units across the globe are at risk of climate connectivity failure, casting doubt on whether many species can successfully undergo climate-induced range shifts among protected areas. Consequently, protected areas are unlikely to serve as steppingstones for a large number of species under a warming climate. As species disappear from protected areas without commensurate immigration of species suited to the emerging climate (due to climate connectivity failure), many protected areas may be left with a depauperate suite of species under climate change. Our findings are highly relevant given recent pledges to conserve 30% of the planet by 2030 (30 × 30), underscore the need for innovative land management strategies that allow for species range shifts, and suggest that assisted colonization may be necessary to promote species that are adapted to the emerging climate.

Migratory behavior and winter geography drive differential range shifts of eastern birds in response to recent climate change.

Over the past half century, migratory birds in North America have shown divergent population trends relative to resident species, with the former declining rapidly and the latter increasing. The role that climate change has played in these observed trends is not well understood, despite significant warming over this period. We used 43 y of monitoring data to fit dynamic species distribution models and quantify the rate of latitudinal range shifts in 32 species of birds native to eastern North America. Since the early 1970s, species that remain in North America throughout the year, including both resident and migratory species, appear to have responded to climate change through both colonization of suitable area at the northern leading edge of their breeding distributions and adaption in place at the southern trailing edges. Neotropical migrants, in contrast, have shown the opposite pattern: contraction at their southern trailing edges and no measurable shifts in their northern leading edges. As a result, the latitudinal distributions of temperate-wintering species have increased while the latitudinal distributions of neotropical migrants have decreased. These results raise important questions about the mechanisms that determine range boundaries of neotropical migrants and suggest that these species may be particularly vulnerable to future climate change. Our results highlight the potential importance of climate change during the nonbreeding season in constraining the response of migratory species to temperature changes at both the trailing and leading edges of their breeding distributions. Future research on the interactions between breeding and nonbreeding climate change is urgently needed.

Temporal dynamics of range expander and congeneric native plant responses during and after extreme drought events.

Climate change is causing range shifts of many species to higher latitudes and altitudes and increasing their exposure to extreme weather events. It has been shown that range-shifting plant species may perform differently in new soil than related natives; however, little is known about how extreme weather events affect range-expanding plants compared to related natives. In this study we used outdoor mesocosms to study how range-expanding plant species responded to extreme drought in live soil from a habitat in a new range with and without live soil from a habitat in the original range (Hungary). During summer drought, the shoot biomass of the range-expanding plant community declined. In spite of this, in the mixed community, range expanders produced more shoot biomass than congeneric natives. In mesocosms with a history of range expanders in the previous year, native plants produced less biomass. Plant legacy or soil origin effects did not change the response of natives or range expanders to summer drought. During rewetting, range expanders had less biomass than congeneric natives but higher drought resilience (survival) in soils from the new range where in the previous year native plant species had grown. The biomass patterns of the mixed plant communities were dominated by Centaurea spp.; however, not all plant species within the groups of natives and of range expanders showed the general pattern. Drought reduced the litter decomposition, microbial biomass, and abundances of bacterivorous, fungivorous, and carnivorous nematodes. Their abundances recovered during rewetting. There was less microbial and fungal biomass, and there were fewer fungivorous nematodes in soils from the original range where range expanders had grown in the previous year. We concluded that in mixed plant communities of range expanders and congeneric natives, range expanders performed better, under both ambient and drought conditions, than congeneric natives. However, when considering the responses of individual species, we observed variations among pairs of congenerics, so that under the present mixed-community conditions there was no uniformity in responses to drought of range expanders versus congeneric natives. Range-expanding plant species reduced soil fungal biomass and the numbers of soil fungivorous nematodes, suggesting that the effects of range-expanding plant species can trickle up in the soil food web.

Projected climate change impacts on the phylogenetic diversity of the world's terrestrial birds: more than species numbers.

Ongoing climate change is a major threat to biodiversity. As abiotic tolerances and dispersal abilities vary, species-specific responses have the potential to further amplify or ameliorate the ensuing impacts on species assemblages. Here, we investigate the effects of climate change on species distributions across non-marine birds, quantifying its projected impact on species richness (SR) as well as on different aspects of phylogenetic diversity globally. Going beyond previous work, we disentangle the potential impacts of species gains versus losses on assemblage-level phylogenetic diversity under climate change and compare the projected impacts to randomized assemblage changes. We show that beyond its effects on SR, climate change could have profound impacts on assemblage-level phylogenetic diversity and composition, which differ significantly from random changes and among regions. Though marked species losses are most frequent in tropical and subtropical areas in our projections, phylogenetic restructuring of species communities is likely to occur all across the globe. Furthermore, our results indicate that the most severe changes to the phylogenetic diversity of local assemblages are likely to be caused by species range shifts and local species gains rather than range reductions and extinctions. Our findings highlight the importance of considering diverse measures in climate impact assessments.

The untapped potential of macrofossils in ancient plant DNA research.

The rapid development of ancient DNA analysis in the last decades has induced a paradigm shift in ecology and evolution. Driven by a combination of breakthroughs in DNA isolation techniques, high-throughput sequencing, and bioinformatics, ancient genome-scale data for a rapidly growing variety of taxa are now available, allowing researchers to directly observe demographic and evolutionary processes over time. However, the vast majority of paleogenomic studies still focus on human or animal remains. In this article, we make the case for a vast untapped resource of ancient plant material that is ideally suited for paleogenomic analyses: plant remains, such as needles, leaves, wood, seeds, or fruits, that are deposited in natural archives, such as lake sediments, permafrost, or even ice caves. Such plant remains are commonly found in large numbers and in stratigraphic sequence through time and have so far been used primarily to reconstruct past local species presences and abundances. However, they are also unique repositories of genetic information with the potential to revolutionize the fields of ecology and evolution by directly studying microevolutionary processes over time. Here, we give an overview of the current state-of-the-art, address important challenges, and highlight new research avenues to inspire future research.

Ocean warming alters the distributional range, migratory timing, and spatial protections of an apex predator, the tiger shark (Galeocerdo cuvier).

Given climate change threats to ecosystems, it is critical to understand the responses of species to warming. This is especially important in the case of apex predators since they exhibit relatively high extinction risk, and changes to their distribution could impact predator-prey interactions that can initiate trophic cascades. Here we used a combined analysis of animal tracking, remotely sensed environmental data, habitat modeling, and capture data to evaluate the effects of climate variability and change on the distributional range and migratory phenology of an ectothermic apex predator, the tiger shark (Galeocerdo cuvier). Tiger sharks satellite tracked in the western North Atlantic between 2010 and 2019 revealed significant annual variability in the geographic extent and timing of their migrations to northern latitudes from ocean warming. Specifically, tiger shark migrations have extended farther poleward and arrival times to northern latitudes have occurred earlier in the year during periods with anomalously high sea-surface temperatures. A complementary analysis of nearly 40 years of tiger shark captures in the region revealed decadal-scale changes in the distribution and timing of shark captures in parallel with long-term ocean warming. Specifically, areas of highest catch densities have progressively increased poleward and catches have occurred earlier in the year off the North American shelf. During periods of anomalously high sea-surface temperatures, movements of tracked sharks shifted beyond spatial management zones that had been affording them protection from commercial fishing and bycatch. Taken together, these study results have implications for fisheries management, human-wildlife conflict, and ecosystem functioning.

Anna's hummingbird (Calypte anna) physiological response to novel thermal and hypoxic conditions at high elevations.

Many species have not tracked their thermal niches upslope as predicted by climate change, potentially because higher elevations are associated with abiotic challenges beyond temperature. To better predict whether organisms can continue to move upslope with rising temperatures, we need to understand their physiological performance when subjected to novel high-elevation conditions. Here, we captured Anna's hummingbirds - a species expanding their elevational distribution in concordance with rising temperatures - from across their current elevational distribution and tested their physiological response to novel abiotic conditions. First, at a central aviary within their current elevational range, we measured hovering metabolic rate to assess their response to oxygen conditions and torpor use to assess their response to thermal conditions. Second, we transported the hummingbirds to a location 1200 m above their current elevational range limit to test for an acute response to novel oxygen and thermal conditions. Hummingbirds exhibited lower hovering metabolic rates above their current elevational range limit, suggesting lower oxygen availability may reduce performance after an acute exposure. Alternatively, hummingbirds showed a facultative response to thermal conditions by using torpor more frequently and for longer. Finally, post-experimental dissection found that hummingbirds originating from higher elevations within their range had larger hearts, a potential plastic response to hypoxic environments. Overall, our results suggest lower oxygen availability and low air pressure may be difficult challenges to overcome for hummingbirds shifting upslope as a consequence of rising temperatures, especially if there is little to no long-term acclimatization. Future studies should investigate how chronic exposure and acclimatization to novel conditions, as opposed to acute experiments, may result in alternative outcomes that help organisms better respond to abiotic challenges associated with climate-induced range shifts.

Air temperatures over-predict changes to stream fish assemblages with climate warming compared with water temperatures.

Studies predicting how the distribution of aquatic organisms will shift with climate change often use projected increases in air temperature or water temperature. However, the assumed correlations between water temperature change and air temperature change can be problematic, especially for mountainous, high elevation streams. Using stream fish assemblage data from 1,442 surveys across a mountain-plains gradient (Wyoming, USA; 1990-2018), we compared the responsiveness of thermal guilds, native status groups, and assemblage structure to projected climate warming from generalized air temperature models and stream-specific water temperature models. Air temperature models consistently predicted greater range shift differences between warm-water and cold-water species, with air temperatures predicting greater increases in occurrence and greater range expansions for warm-water species. The "over-prediction" of warm-water species expansions resulted in air temperature models predicting higher rates of novel species combinations, greater increases in local species richness, and higher magnitudes of biotic homogenization compared with water temperature models. Despite differences in model predictions for warm-water species, both air and water temperature models predicted that three cold-water species would exhibit similar decreases in occurrence (decline of 1.0% and 1.8% of sites per 1°C warming, respectively) and similar range contractions (16.6 and 21.5 m elevation loss per 1°C warming, respectively). The "over-prediction" for warm-water species is partially attributable to water temperatures warming at slower rates than air temperatures because local, stream-scale factors (e.g., riparian cover, groundwater inputs) buffer high elevation streams from rising air temperatures. Our study provides the first comparison of how inferences about climate-induced biotic change at the species- and assemblage-levels differ when modeling with generalized air temperatures versus stream-specific water temperatures. We recommend that future studies use stream-specific water temperature models, especially for mountainous, high elevation streams, to avoid the "over-prediction" of biotic changes observed from air temperature variables.

"No-regrets" pathways for navigating climate change: planning for connectivity with land use, topography, and climate.

As both plant and animal species shift their ranges in response to a changing climate, maintaining connectivity between present habitat and suitable habitat in the future will become increasingly important to ensure lasting protection for biodiversity. Because the temporal period commensurate with planning for mid-century change is multi-generational for most species, connectivity designed to facilitate climate adaptation requires pathways with 'stepping-stones' between current and future habitat. These areas should have habitats suitable not only for dispersal, but for all aspects of species lifecycles. We integrated present-day land use, topographic diversity, and projections of shifting climate regimes into a single connectivity modeling approach to identify pathways for mid-century shifts in species ranges. Using Omniscape we identified climate linkages, or areas important for climate change-driven movement, as the areas with more current flow than would be expected in the absence of climate considerations. This approach identified connectivity potential between natural lands in the present climate and natural lands with future analogous climate following topo-climatically diverse routes. We then translated the model output into a strategic framework to improve interpretation and to facilitate a more direct connection with conservation action. Across modified landscapes, pathways important to climate-driven movement were highly coincident with the last remaining present-day linkages, reinforcing their importance. Across unfragmented lands, the presence of climate-adapted pathways helped inform the prioritization of conservation actions in areas where multiple connectivity options still exist. Many climate linkages follow major watercourses along elevational gradients, highlighting the importance of protecting or managing for these natural linear pathways that provide movement routes for climate adaptation. By integrating enduring landscape features with climate projections and present-day land uses, our approach reveals "no-regrets" pathways to plan for a connected landscape in an uncertain future.