Warming and provenance limit tree recruitment across and beyond the elevation range of subalpine forest

Climate niche models project that subalpine forest ranges will extend upslope with climate warming. These projections assume that the climate suitable for adult trees will be adequate for forest regeneration, ignoring climate requirements for seedling recruitment, a potential demographic bottleneck. Moreover, local genetic adaptation is expected to facilitate range expansion, with tree populations at the upper forest edge providing the seed best adapted to the alpine. Here, we test these expectations using a novel combination of common gardens, seeded with two widely distributed subalpine conifers, and climate manipulations replicated at three elevations. Infrared heaters raised temperatures in heated plots, but raised temperatures more in the forest than at or above treeline because strong winds at high elevation reduced heating efficiency. Watering increased season‐average soil moisture similarly across sites. Contrary to expectations, warming reduced Engelmann spruce recruitment at and above treeline, as well as in the forest. Warming reduced limber pine first‐year recruitment in the forest, but had no net effect on fourth‐year recruitment at any site. Watering during the snow‐free season alleviated some negative effects of warming, indicating that warming exacerbated water limitations. Contrary to expectations of local adaptation, low‐elevation seeds of both species initially recruited more strongly than high‐elevation seeds across the elevation gradient, although the low‐provenance advantage diminished by the fourth year for Engelmann spruce, likely due to small sample sizes. High‐ and low‐elevation provenances responded similarly to warming across sites for Engelmann spruce, but differently for limber pine. In the context of increasing tree mortality, lower recruitment at all elevations with warming, combined with lower quality, high‐provenance seed being most available for colonizing the alpine, portends range contraction for Engelmann spruce. The lower sensitivity of limber pine to warming indicates a potential for this species to become more important in subalpine forest communities in the coming centuries.     

Tree demography suggests multiple directions and drivers for species range shifts in mountains of Northeastern United States

Climate change is expected to lead to upslope shifts in tree species distributions, but the evidence is mixed partly due to land‐use effects and individualistic species responses to climate. We examined how individual tree species demography varies along elevational climatic gradients across four states in the northeastern United States to determine whether species elevational distributions and their potential upslope (or downslope) shifts were controlled by climate, land‐use legacies (past logging), or soils. We characterized tree demography, microclimate, land‐use legacies, and soils at 83 sites stratified by elevation (~500 to ~1200 m above sea level) across 12 mountains containing the transition from northern hardwood to spruce‐fir forests. We modeled elevational distributions of tree species saplings and adults using logistic regression to test whether sapling distributions suggest ongoing species range expansion upslope (or contraction downslope) relative to adults, and we used linear mixed models to determine the extent to which climate, land use, and soil variables explain these distributions. Tree demography varied with elevation by species, suggesting a potential upslope shift only for American beech, downslope shifts for red spruce (more so in cool regions) and sugar maple, and no change with elevation for balsam fir. While soils had relatively minor effects, climate was the dominant predictor for most species and more so for saplings than adults of red spruce, sugar maple, yellow birch, cordate birch, and striped maple. On the other hand, logging legacies were positively associated with American beech, sugar maple, and yellow birch, and negatively with red spruce and balsam fir – generally more so for adults than saplings. All species exhibited individualistic rather than synchronous demographic responses to climate and land use, and the return of red spruce to lower elevations where past logging originally benefited northern hardwood species indicates that land use may mask species range shifts caused by changing climate.     

Leap frog in slow motion: Divergent responses of tree species and life stages to climatic warming in Great Basin subalpine forests

In response to climate warming, subalpine treelines are expected to move up in elevation since treelines are generally controlled by growing season temperature. Where treeline is advancing, dispersal differences and early life stage environmental tolerances are likely to affect how species expand their ranges. Species with an establishment advantage will colonize newly available habitat first, potentially excluding species that have slower establishment rates. Using a network of plots across five mountain ranges, we described patterns of upslope elevational range shift for the two dominant Great Basin subalpine species, limber pine and Great Basin bristlecone pine. We found that the Great Basin treeline for these species is expanding upslope with a mean vertical elevation shift of 19.1 m since 1950, which is lower than what we might expect based on temperature increases alone. The largest advances were on limber pine‐dominated granitic soils, on west aspects, and at lower latitudes. Bristlecone pine juveniles establishing above treeline share some environmental associations with bristlecone adults. Limber pine above‐treeline juveniles, in contrast, are prevalent across environmental conditions and share few environmental associations with limber pine adults. Strikingly, limber pine is establishing above treeline throughout the region without regard to site characteristic such as soil type, slope, aspect, or soil texture. Although limber pine is often rare at treeline where it coexists with bristlecone pine, limber pine juveniles dominate above treeline even on calcareous soils that are core bristlecone pine habitat. Limber pine is successfully “leap‐frogging” over bristlecone pine, probably because of its strong dispersal advantage and broader tolerances for establishment. This early‐stage dominance indicates the potential for the species composition of treeline to change in response to climate change. More broadly, it shows how species differences in dispersal and establishment may result in future communities with very different specific composition.     

Plasticity in functional traits in the context of climate change: a case study of the subalpine forb Boechera stricta (Brassicaceae)

Environmental variation often induces shifts in functional traits, yet we know little about whether plasticity will reduce extinction risks under climate change. As climate change proceeds, phenotypic plasticity could enable species with limited dispersal capacity to persist in situ, and migrating populations of other species to establish in new sites at higher elevations or latitudes. Alternatively, climate change could induce maladaptive plasticity, reducing fitness, and potentially stalling adaptation and migration. Here, we quantified plasticity in life history, foliar morphology, and ecophysiology in Boechera stricta (Brassicaceae), a perennial forb native to the Rocky Mountains. In this region, warming winters are reducing snowpack and warming springs are advancing the timing of snow melt. We hypothesized that traits that were historically advantageous in hot and dry, low‐elevation locations will be favored at higher elevation sites due to climate change. To test this hypothesis, we quantified trait variation in natural populations across an elevational gradient. We then estimated plasticity and genetic variation in common gardens at two elevations. Finally, we tested whether climatic manipulations induce plasticity, with the prediction that plants exposed to early snow removal would resemble individuals from lower elevation populations. In natural populations, foliar morphology and ecophysiology varied with elevation in the predicted directions. In the common gardens, trait plasticity was generally concordant with phenotypic clines from the natural populations. Experimental snow removal advanced flowering phenology by 7 days, which is similar in magnitude to flowering time shifts over 2–3 decades of climate change. Therefore, snow manipulations in this system can be used to predict eco‐evolutionary responses to global change. Snow removal also altered foliar morphology, but in unexpected ways. Extensive plasticity could buffer against immediate fitness declines due to changing climates.     

Range dynamics of small mammals along an elevational gradient over an 80‐year interval

One expected response to observed global warming is an upslope shift of species elevational ranges. Here, we document changes in the elevational distributions of the small mammals within the Ruby Mountains in northeastern Nevada over an 80‐year interval. We quantified range shifts by comparing distributional records from recent comprehensive field surveys (2006–2008) to earlier surveys (1927–1929) conducted at identical and nearby locations. Collector field notes from the historical surveys provided detailed trapping records and locality information, and museum specimens enabled confirmation of species' identifications. To ensure that observed shifts in range did not result from sampling bias, we employed a binomial likelihood model (introduced here) using likelihood ratios to calculate confidence intervals around observed range limits. Climate data indicate increases in both precipitation and summer maximum temperature between sampling periods. Increases in winter minimum temperatures were only evident at mid to high elevations. Consistent with predictions of change associated with climate warming, we document upslope range shifts for only two mesic‐adapted species. In contrast, no xeric‐adapted species expanded their ranges upslope. Rather, they showed either static distributions over time or downslope contraction or expansion. We attribute these unexpected findings to widespread land‐use driven habitat change at lower elevations. Failure to account for land‐use induced changes in both baseline assessments and in predicting shifts in species distributions may provide misleading objectives for conservation policies and management practices.     

Elevational changes in the avian community of a Mesoamerican cloud forest park

Harboring many range‐restricted and specialized species, high elevation tropical cloud forests are diverse habitats represented in many protected areas. Despite this, many such areas receive little practical protection from deforestation and land conversion. Moreover, montane species may be more sensitive to climate change owing to various factors affecting community assembly across elevational gradients. Few studies have used annual monitoring to assess how biological communities in cloud forests may be shifting in response to habitat or climate change or assessed the efficacy of protected areas in buffering these effects. We analyzed avifaunal community trends in a 10‐yr dataset of constant‐effort bird point‐count data in a cloud forest national park in Honduras, Central America. We found that species richness and diversity increased at higher elevations, but decreased at lower elevations. Abundances of most dietary and forest‐dependency groups exhibited similar trends, and many key cloud forest species shifted upslope and/or increased in abundance. Taken together, our results suggest that the avian community is moving upslope and species composition is changing. Results for species richness and diversity were similar when only nondegraded transects were considered, suggesting the role of climate change as an important driver. At lower elevations, however, many species may be negatively affected by increased habitat degradation, favoring species with low forest dependency. Continued habitat conversion and climate change could push the cloud forest bird community further upslope, potentially resulting in increased competition, mortality, and even extirpation of some species. Increased protection is unlikely to mitigate the effects of climate change.     

Drought tolerance and growth in populations of a wide‐ranging tree species indicate climate change risks for the boreal north

Choosing drought‐tolerant planting stock in reforestation programs may help adapt forests to climate change. To inform such reforestation strategies, we test lodgepole pine (Pinus contorta Doug. ex Loud. var latifolia Englm.) population response to drought and infer potential benefits of a northward transfer of seeds from drier, southern environments. The objective is addressed by combining dendroecological growth analysis with long‐term genetic field trials. Over 500 trees originating from 23 populations across western North America were destructively sampled in three experimental sites in southern British Columbia, representing a climate warming scenario. Growth after 32 years from provenances transferred southward or northward over long distances was significantly lower than growth of local populations. All populations were affected by a severe natural drought event in 2002. The provenances from the most southern locations showed the highest drought tolerance but low productivity. Local provenances were productive and drought tolerant. Provenances from the boreal north showed lower productivity and less drought tolerance on southern test sites than all other sources, implying that maladaptation to drought may prevent boreal populations from taking full advantage of more favorable growing conditions under projected climate change.     

Drought stress limits the geographic ranges of two tree species via different physiological mechanisms

Range shifts are among the most ubiquitous ecological responses to anthropogenic climate change and have large consequences for ecosystems. Unfortunately, the ecophysiological forces that constrain range boundaries are poorly understood, making it difficult to mechanistically project range shifts. To explore the physiological mechanisms by which drought stress controls dry range boundaries in trees, we quantified elevational variation in drought tolerance and in drought avoidance‐related functional traits of a widespread gymnosperm (ponderosa pine – Pinus ponderosa) and angiosperm (trembling aspen – Populus tremuloides) tree species in the southwestern USA. Specifically, we quantified tree‐to‐tree variation in growth, water stress (predawn and midday xylem tension), drought avoidance traits (branch conductivity, leaf/needle size, tree height, leaf area‐to‐sapwood area ratio), and drought tolerance traits (xylem resistance to embolism, hydraulic safety margin, wood density) at the range margins and range center of each species. Although water stress increased and growth declined strongly at lower range margins of both species, ponderosa pine and aspen showed contrasting patterns of clinal trait variation. Trembling aspen increased its drought tolerance at its dry range edge by growing stronger but more carbon dense branch and leaf tissues, implying an increased cost of growth at its range boundary. By contrast, ponderosa pine showed little elevational variation in drought‐related traits but avoided drought stress at low elevations by limiting transpiration through stomatal closure, such that its dry range boundary is associated with limited carbon assimilation even in average climatic conditions. Thus, the same climatic factor (drought) may drive range boundaries through different physiological mechanisms – a result that has important implications for process‐based modeling approaches to tree biogeography. Further, we show that comparing intraspecific patterns of trait variation across ranges, something rarely done in a range‐limit context, helps elucidate a mechanistic understanding of range constraints.     

Linking population genetics and tree height growth models to predict impacts of climate change on forest production

Changes to forest growth models used widely in global change research and sustainable forest management are needed to account for expected climate change impacts. We provide a new approach that dynamically merges height–age functions prevalent in forest growth models with transfer functions prevalent in population adaptation research to better represent changes to forest productivity as climates gradually change. Our simulations with data from an extensive provenance test of lodgepole pine (Pinus contorta) in British Columbia, Canada, suggest that climate change will reduce production in lodgepole pine forests established today by at least 7–13% at the end of this century – considerably less than most predictions based solely on transfer or response functions, which do not integrate impacts as climate gradually changes. This work illustrates the need for forest productivity models to consider the changing climate in which a population is growing relative to the static climate of its origin. It also demonstrates the value of long‐term provenance trials in assessing the dynamic impact of climate change on forest productivity, and serves as an example of how provenance trials may be exploited in other forest productivity models or other research fields to assess plant responses to climate.     

Contrasting growth forecasts across the geographical range of Scots pine due to altitudinal and latitudinal differences in climatic sensitivity

Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of a species’ geographical distribution, where differences in growth or population dynamics may result in range expansions or contractions. Understanding population responses to different climatic drivers along wide latitudinal and altitudinal gradients is necessary in order to gain a better understanding of plant responses to ongoing increases in global temperature and drought severity. We selected Scots pine (Pinus sylvestris L.) as a model species to explore growth responses to climatic variability (seasonal temperature and precipitation) over the last century through dendrochronological methods. We developed linear models based on age, climate and previous growth to forecast growth trends up to year 2100 using climatic predictions. Populations were located at the treeline across a latitudinal gradient covering the northern, central and southernmost populations and across an altitudinal gradient at the southern edge of the distribution (treeline, medium and lower elevations). Radial growth was maximal at medium altitude and treeline of the southernmost populations. Temperature was the main factor controlling growth variability along the gradients, although the timing and strength of climatic variables affecting growth shifted with latitude and altitude. Predictive models forecast a general increase in Scots pine growth at treeline across the latitudinal distribution, with southern populations increasing growth up to year 2050, when it stabilizes. The highest responsiveness appeared at central latitude, and moderate growth increase is projected at the northern limit. Contrastingly, the model forecasted growth declines at lowland‐southern populations, suggesting an upslope range displacement over the coming decades. Our results give insight into the geographical responses of tree species to climate change and demonstrate the importance of incorporating biogeographical variability into predictive models for an accurate prediction of species dynamics as climate changes.     

Islands in the sky or squeezed at the top? Ecological causes of elevational range limits in montane salamanders

Many animal and plant taxa reach their highest endemism and species richness in montane regions. The study of elevational range limits is central to understanding this widespread pattern and to predicting the responses of montane species to climate change. Yet, because large‐scale manipulations of the distributions of most species are difficult, the causes of species’ elevational range limits (e.g. competitive interactions, physiological specialization) are poorly understood. Here, we harness the power of new mechanistic approaches to dissect the factors that underlie the elevational replacement of two salamander species in the Appalachian Highlands. Our results challenge the long‐held idea that competitive interactions drive the lower elevational range limits of montane species and that physiological stress prevents low‐elevation species from expanding to high elevations. We show that physiological constraints drive the lower elevational range limit of the montane‐endemic species, Plethodon jordani. Conversely, we find that competition with P. jordani prevents the low‐elevation species, P. teyahalee, from expanding its range to include higher‐elevation habitats. These results are broadly consistent with the biogeography and behavior of other montane species, suggesting that similar mechanisms underlie patterns of elevational zonation across a variety of taxa and montane regions. To the extent that our findings are taxonomically and geographically widespread, these results challenge the idea that competitive release at species’ lower elevational range limits is driving the downslope range shifts exhibit by some montane taxa. Instead, our results raise the sobering possibility that even small changes in climate might cause erosion of the ranges of many high‐elevation species.     

An elevational shift in butterfly species richness and composition accompanying recent climate change

The geographic ranges of many species have shifted polewards and uphill in elevation associated with climate warming, leading to increases in species richness at high latitudes and elevations. However, few studies have addressed community‐level responses to climate change across the entire elevational gradients of mountain ranges, or at warm lower latitudes where ecological diversity is expected to decline. Here, we show uphill shifts in butterfly species richness and composition in the Sierra de Guadarrama (central Spain) between 1967–1973 and 2004–2005. Butterfly communities with comparable species compositions shifted uphill by 293 m (± SE 26), consistent with an upward shift of approximately 225 m in mean annual isotherms. Species richness had a humped relationship with elevation, but declined between surveys, particularly at low elevations. Changes to species richness and composition primarily reflect the loss from lower elevations of species whose regional distributions are restricted to the mountains. The few colonizations by specialist low‐elevation species failed to compensate for the loss of high‐elevation species, because there are few low‐elevation species in the region and the habitat requirements of some of these prevent them from colonizing the mountain range. As a result, we estimated a net decline in species richness in approximately 90% of the region, and increasing community domination by widespread species. The results suggest that climate warming, combined with habitat loss and other drivers of biological change, could lead to significant losses in ecological diversity in mountains and other regions where species encounter their lower latitudinal‐range margins.     

Non-equilibrium succession dynamics indicate continued northern migration of lodgepole pine

Because species affect ecosystem functioning, understanding migration processes is a key component of predicting future ecosystem responses to climate change. This study provides evidence of range expansion under current climatic conditions of an indigenous species with strong ecosystem effects. Surveys of stands along the northern distribution limit of lodgepole pine (Pinus contorta var. latifolia) in central Yukon Territory, Canada showed consistent increases in pine dominance following fire. These patterns differed strongly from those observed at sites where pine has been present for several thousand years. Differences in species thinning rates are unlikely to account for the observed increases in pine dominance. Rates of pine regeneration at its range limits were equivalent to those of spruce, indicating a capacity for rapid local population expansion. The study also found no evidence of strong climatic limitation of pine population growth at the northern distribution limit. We interpret these data as evidence of current pine expansion at its range limits and conclude that the northern distribution of lodgepole pine is not in equilibrium with current climate. This study has implications for our ability to predict vegetation response to climate change when populations may lag in their response to climate.     

Can community structure track sea‐level rise? Stress and competitive controls in tidal wetlands

Climate change impacts, such as accelerated sea‐level rise, will affect stress gradients, yet impacts on competition/stress tolerance trade‐offs and shifts in distributions are unclear. Ecosystems with strong stress gradients, such as estuaries, allow for space‐for‐time substitutions of stress factors and can give insight into future climate‐related shifts in both resource and nonresource stresses. We tested the stress gradient hypothesis and examined the effect of increased inundation stress and biotic interactions on growth and survival of two congeneric wetland sedges, Schoenoplectus acutus and Schoenoplectus americanus. We simulated sea‐level rise across existing marsh elevations and those not currently found to reflect potential future sea‐level rise conditions in two tidal wetlands differing in salinity. Plants were grown individually and together at five tidal elevations, the lowest simulating an 80‐cm increase in sea level, and harvested to assess differences in biomass after one growing season. Inundation time, salinity, sulfides, and redox potential were measured concurrently. As predicted, increasing inundation reduced biomass of the species commonly found at higher marsh elevations, with little effect on the species found along channel margins. The presence of neighbors reduced total biomass of both species, particularly at the highest elevation; facilitation did not occur at any elevation. Contrary to predictions, we documented the competitive superiority of the stress tolerator under increased inundation, which was not predicted by the stress gradient hypothesis. Multifactor manipulation experiments addressing plant response to accelerated climate change are integral to creating a more realistic, valuable, and needed assessment of potential ecosystem response. Our results point to the important and unpredicted synergies between physical stressors, which are predicted to increase in intensity with climate change, and competitive forces on biomass as stresses increase.     

Significance of summer fog and overcast for drought stress and ecological functioning of coastal California endemic plant species

Aim Fog drip is a crucial water source for plants in many ecosystems, including a number of global biodiversity hotspots. In California, dozens of rare, drought‐sensitive plant species are endemic to coastal areas where the dominant summer moisture source is fog. Low clouds that provide water to these semi‐arid ecosystems through fog drip can also sharply reduce evaporative water losses by providing shade. We quantified the relative hydrological importance of cloud shading vs. fog drip. We then examined how both factors influence the range dynamics of an apparently fog‐dependent plant species spanning a small‐scale cloud gradient. Location The study area is on Santa Cruz Island off the coast of southern California. It is near the southern range limit of bishop pine (Pinus muricata D. Don), a tree endemic to the coasts of California and Baja, Mexico. Methods We measured climate across a pine stand along a 7 km, coastal–inland elevation transect. Short‐term (1–5 years) monitoring and remote sensing data revealed strong climatic gradients driven primarily by cloud cover. Long‐term (102 years) effects of these gradients were estimated using a water balance model. Results We found that shade from persistent low clouds near the coast reduced annual drought stress by 22–40% compared with clearer conditions further inland. Fog drip at higher elevations provided sufficient extra water to reduce annual drought stress by 20–36%. Sites located at both high elevation and nearer the coast were subject to both effects. Together, these effects reduced average annual drought stress by 56% and dramatically reduced the frequency of severe drought over the last century. At lower elevation (without appreciable fog drip) and also near the inland edge of the stand (with less cloud shading) severe droughts episodically kill most pine recruits, thereby limiting the local range of this species. Main conclusions Persistent cloud shading can influence hydrology as much as fog drip in cloud‐affected ecosystems. Understanding the patterns of both cloud shading and fog drip and their respective impacts on ecosystem water budgets is necessary to fully understand past species range shifts and to anticipate future climate change‐induced range shifts in fog‐dependent ecosystems.     

Contemporary climate change alters the pace and drivers of extinction

Contemporary climate change is expected to affect the distributions of most species, but the nature, tempo, and mechanics of contemporary range shifts are still largely speculative. Here, we use fine‐scale distributional records developed over the past Century, combined with spatially comprehensive microclimatic data, to demonstrate a dramatic shift in the range of a climate‐sensitive mammal and to infer the increasingly important role of climate in local extinctions of this species across a 38.2 million‐ha area. Changes in the distribution of the American pika (Ochotona princeps) throughout the Great Basin ecoregion were characterized using records from 1898–2008, revealing a nearly five‐fold increase in the rate of local extinction and an 11‐fold increase in the rate of upslope range retraction during the last ten years, compared with during the 20th Century. Four of ten local pika extinctions have occurred since 1999, and across this ecoregion the low‐elevation range boundary for this species is now moving upslope at an average rate of about 145 m per decade. The rapid, ecoregional range shift of this small, talus‐dwelling species stands in remarkable contrast with the dynamics and determinants of endangerment previously observed for most terrestrial species, and to earlier extinction determinants for O. princeps in this region. Further investigation of widely distributed species will clarify rates at which biotic response to environmental change is occurring, and how factors driving that change are evolving in different portions of the earth.     

Seed dispersers help plants to escape global warming

Plants are shifting their ranges towards higher elevations in response to global warming, yet such shifts are occurring at a rate slower than is needed to keep pace with a rapidly changing climate. There is, however, an almost complete lack of knowledge on seed dispersal across altitude, a key process to understand what constrains climate‐driven range shifts. Here, we report the first direct empirical evidence on altitudinal seed dispersal mediated by two common frugivorous mammals: the red fox Vulpes vulpes and the pine marten Martes martes. We conducted a three‐year (bait‐marking) experiment in a mountainous region of Spain. We offered experimental fruits containing colour‐coded seed mimics at feeding stations that simulated source trees. The colour codes allowed us to identify the exact origin of seed mimics found later in mammal scats. Nearly half (47%) of the dispersal events occurred towards higher elevations, despite only ca 25% of the study area being above the average altitude of the feeding stations (1344 m). Seeds dispersed uphill gained an average of 106 m (median = 111 m) and a maximum of 288 m, greatly exceeding the estimated requirements to escape warming (35.4 m per decade). Yet, foxes mediated much more uphill seed dispersal than martens (57% and 26% of dispersal events, respectively), which can be explained by between‐disperser differences in home range size and habitat specificity. Dispersers with larger home ranges move farther and potentially disperse more seeds to higher altitudes, while habitat generalism is necessary to transport seeds above vegetation belts delimiting contrasting habitat types. We discuss how both traits (home range size and habitat specificity) can be used to infer altitudinal seed dispersal across disperser species and mountainous landscapes.     

Predicting potential responses to future climate in an alpine ungulate: interspecific interactions exceed climate effects

The altitudinal shifts of many montane populations are lagging behind climate change. Understanding habitual, daily behavioural rhythms, and their climatic and environmental influences, could shed light on the constraints on long‐term upslope range‐shifts. In addition, behavioural rhythms can be affected by interspecific interactions, which can ameliorate or exacerbate climate‐driven effects on ecology. Here, we investigate the relative influences of ambient temperature and an interaction with domestic sheep (Ovis aries) on the altitude use and activity budgets of a mountain ungulate, the Alpine chamois (Rupicapra rupicapra). Chamois moved upslope when it was hotter but this effect was modest compared to that of the presence of sheep, to which they reacted by moving 89–103 m upslope, into an entirely novel altitudinal range. Across the European Alps, a range‐shift of this magnitude corresponds to a 46% decrease in the availability of suitable foraging habitat. This highlights the importance of understanding how factors such as competition and disturbance shape a given species’ realised niche when predicting potential future responses to change. Furthermore, it exposes the potential for manipulations of species interactions to ameliorate the impacts of climate change, in this case by the careful management of livestock. Such manipulations could be particularly appropriate for species where competition or disturbance already strongly restricts their available niche. Our results also reveal the potential role of behavioural flexibility in responses to climate change. Chamois reduced their activity when it was warmer, which could explain their modest altitudinal migrations. Considering this behavioural flexibility, our model predicts a small 15–30 m upslope shift by 2100 in response to climate change, less than 4% of the altitudinal shift that would be predicted using a traditional species distribution model‐type approach (SDM), which assumes that species’ behaviour remains unchanged as climate changes. Behavioural modifications could strongly affect how species respond to a changing climate.     

Upslope Range Shifts of Andean Dung Beetles in Response to Deforestation: Compounding and Confounding Effects of Microclimatic Change

Warmer, and sometimes drier, conditions associated with global climate change are driving many species to shift poleward and/or upslope. I hypothesized that microclimatic changes related to deforestation cause similar shifts for forest species persisting within degraded landscapes. This appears to be the first study to examine this novel hypothesis. I examined elevational distributions of dung beetle communities along parallel intact and disturbed elevational gradients from 290 to 3450 m asl in the Andes of southeastern Peru. Deforested sites were consistently warmer and drier than forested sites. To maintain the same ambient temperature as in forest, species in a deforested landscape would need to shift on average 489±59 m upslope. Dung beetle species showed a mean upslope range shift of 132±64 m (maximum=743 m) in the deforested landscape. Eight species occurred farther upslope in the degraded landscape, while none shifted downslope. In addition to upper range limit expansions, six species shifting upslope also showed range contractions or population declines at their lower range boundary. High elevation and disturbance‐tolerant species did not show range shifts. These findings suggest that land‐use change may both confound and compound the influence of global climate change on biodiversity. Synergies between habitat degradation and climate change could more than double previous range shift projections for this century, leading to unexpectedly rapid changes in biodiversity, especially for sensitive organisms such as tropical insects. On the other hand, range shifts caused by habitat degradation may be mistakenly attributed to global climate change. Abstract in Spanish is available in the online version of this article.     

Upward elevation and northwest range shifts for alpine Meconopsis species in the Himalaya–Hengduan Mountains region

Climate change may impact the distribution of species by shifting their ranges to higher elevations or higher latitudes. The impacts on alpine plant species may be particularly profound due to a potential lack of availability of future suitable habitat. To identify how alpine species have responded to climate change during the past century as well as to predict how they may react to possible global climate change scenarios in the future, we investigate the climatic responses of seven species of Meconopsis, a representative genus endemic in the alpine meadow and subnival region of the Himalaya–Hengduan Mountains. We analyzed past elevational shifts, as well as projected shifts in longitude, latitude, elevation, and range size using historical specimen records and species distribution modeling under optimistic (RCP 4.5) and pessimistic (RCP 8.5) scenarios across three general circulation models for 2070. Our results indicate that across all seven species, there has been an upward shift in mean elevation of 302.3 m between the pre‐1970s (1922–1969) and the post‐1970s (1970–2016). The model predictions suggest that the future suitable climate space will continue to shift upwards in elevation (as well as northwards and westwards) by 2070. While for most of the analyzed species, the area of suitable climate space is predicted to expand under the optimistic emission scenario, the area contracts, or, at best, shows little change under the pessimistic scenario. Species such as M. punicea, which already occupy high latitudes, are consistently predicted to experience a contraction of suitable climate space across all the models by 2070 and may consequently deserve particular attention by conservation strategies. Collectively, our results suggest that the alpine high‐latitude species analyzed here have already been significantly impacted by climate change and that these trends may continue over the coming decades.