Extreme heat events and the vulnerability of endemic montane fishes to climate change

Identifying how close species live to their physiological thermal maxima is essential to understand historical warm‐edge elevational limits of montane faunas and forecast upslope shifts caused by future climate change. We used laboratory experiments to quantify the thermal tolerance and acclimation potential of four fishes (Notropis leuciodus, N. rubricroceus, Etheostoma rufilineatum, E. chlorobranchium) that are endemic to the southern Appalachian Mountains (USA), exhibit different historical elevational limits, and represent the two most species‐rich families in the region. All‐subsets selection of linear regression models using AIC_c indicated that species, acclimation temperature, collection location and month, and the interaction between species and acclimation temperature were important predictors of thermal maxima (T_max), which ranged from 28.5 to 37.2°C. Next, we implemented water temperature models and stochastic weather generation to characterize the magnitude and frequency of extreme heat events (T_extreme) under historical and future climate scenarios across 25 379 stream reaches in the upper Tennessee River system. Lastly, we used environmental niche models to compare warming tolerances (acclimation‐corrected T_max minus T_extreme) between historically occupied versus unoccupied reaches. Historical warming tolerances, ranging from +2.2 to +10.9°C, increased from low to high elevation and were positive for all species, suggesting that T_max does not drive warm‐edge (low elevation) range limits. Future warming tolerances were lower (?1.2 to +9.3°C) but remained positive for all species under the direst warming scenario except for a small proportion of reaches historically occupied by E. rufilineatum, indicating that T_max and acclimation potentials of southern Appalachian minnows and darters are adequate to survive future heat waves. We caution concluding that these species are invulnerable to 21st century warming because sublethal thermal physiology remains poorly understood. Integrating physiological sensitivity and warming exposure demonstrates a general and fine‐grained approach to assess climate change vulnerability for freshwater organisms across physiographically diverse riverscapes.     

Heat freezes niche evolution

Climate change is altering phenology and distributions of many species and further changes are projected. Can species physiologically adapt to climate warming? We analyse thermal tolerances of a large number of terrestrial ectotherm (n = 697), endotherm (n = 227) and plant (n = 1816) species worldwide, and show that tolerance to heat is largely conserved across lineages, while tolerance to cold varies between and within species. This pattern, previously documented for ectotherms, is apparent for this group and for endotherms and plants, challenging the longstanding view that physiological tolerances of species change continuously across climatic gradients. An alternative view is proposed in which the thermal component of climatic niches would overlap across species more than expected. We argue that hard physiological boundaries exist that constrain evolution of tolerances of terrestrial organisms to high temperatures. In contrast, evolution of tolerances to cold should be more frequent. One consequence of conservatism of upper thermal tolerances is that estimated niches for cold‐adapted species will tend to underestimate their upper thermal limits, thereby potentially inflating assessments of risk from climate change. In contrast, species whose climatic preferences are close to their upper thermal limits will unlikely evolve physiological tolerances to increased heat, thereby being predictably more affected by warming.     

Impacts of climate warming and habitat loss on extinctions at species' low-latitude range boundaries

Polewards expansions of species' distributions have been attributed to climate warming, but evidence for climate‐driven local extinctions at warm (low latitude/elevation) boundaries is equivocal. We surveyed the four species of butterflies that reach their southern limits in Britain. We visited 421 sites where the species had been recorded previously to determine whether recent extinctions were primarily due to climate or habitat changes. Coenonympha tullia had become extinct at 52% of study sites and all losses were associated with habitat degradation. Aricia artaxerxes was extinct from 50% of sites, with approximately one‐third to half of extinctions associated with climate‐related factors and the remainder with habitat loss. For Erebia aethiops (extinct from 24% of sites), approximately a quarter of the extinctions were associated with habitat and three‐quarters with climate. For Erebia epiphron, extinctions (37% of sites) were attributed mainly to climate with almost no habitat effects. For the three species affected by climate, range boundaries retracted 70–100 km northwards (A. artaxerxes, E. aethiops) and 130–150 m uphill (E. epiphron) in the sample of sites analysed. These shifts are consistent with estimated latitudinal and elevational temperature shifts of 88 km northwards and 98 m uphill over the 19‐year study period. These results suggest that the southern/warm range margins of some species are as sensitive to climate change as are northern/cool margins. Our data indicate that climate warming has been of comparable importance to habitat loss in driving local extinctions of northern species over the past few decades; future climate warming is likely to jeopardize the long‐term survival of many northern and mountain species.     

The range implications of lizard traits in changing environments

Aim Most predictions of species ranges are based on correlating current species localities to environmental conditions. These correlative models do not explicitly include a species' biology. In contrast, some mechanistic models link traits to energetics and population dynamics to predict species distributions. These models enable one to ask whether considering a species' biology is important for predicting its range. I implement mechanistic models to investigate how a species' morphology, physiology and life history influence its range. Location North America. Methods I compare the mechanistic model predictions with those of correlative models for eight species of North American lizards in both current environments and following a uniform 3 °C temperature warming. I then examine the implications of superimposing habitat and elevation requirements on constraints associated with environmental tolerances. Results In the mechanistic model, species with a narrower thermal range for activity are both predicted and observed to have more restricted distributions. Incorporating constraints on habitat and elevation further restricts species distributions beyond areas that are thermally suitable. While correlative models generally outperform mechanistic models at predicting current distributions, the performance of mechanistic models improves when incorporating additional factors. In response to a 3 °C temperature warming, the northward range shifts predicted by the mechanistic model vary between species according to trait differences and are of a greater extent than those predicted by correlative models. Main conclusions These findings highlight the importance of species traits for understanding the dynamics of species ranges in changing environments. The analysis demonstrates that mechanistic models may provide an important complement to correlative models for predicting range dynamics, which may underpredict climate‐induced range shifts.     

Asynchronous range shifts drive alpine plant–pollinator interactions and reduce plant fitness

Climate change is driving species' range shifts, which are in turn disrupting species interactions due to species‐specific differences in their abilities to migrate in response to climate. We evaluated the consequences of asynchronous range shifts in an alpine plant–pollinator community by transplanting replicated alpine meadow turfs downslope along an elevational gradient thereby introducing them to warmer climates and novel plant and pollinator communities. We asked how these novel plant–pollinator interactions affect plant reproduction. We found that pollinator communities differed substantially across the elevation/temperature gradient, suggesting that these plants will likely interact with different pollinator communities with warming climate. Contrary to the expectation that floral visitation would increase monotonically with warmer temperatures at lower elevations, visitation rate to the transplanted communities peaked under intermediate warming at midelevation sites. In contrast, visitation rate generally increased with temperature for the local, lower elevation plant communities surrounding the experimental alpine turfs. For two of three focal plant species in the transplanted high‐elevation community, reproduction declined at warmer sites. For these species, reproduction appears to be dependent on pollinator identity such that reduced reproduction may be attributable to decreased visitation from key pollinator species, such as bumble bees, at warmer sites. Reproduction in the third focal species appears to be primarily driven by overall pollinator visitation rate, regardless of pollinator identity. Taken together, the results suggest climate warming can indirectly affect plant reproduction via changes in plant–pollinator interactions. More broadly, the experiment provides a case study for predicting the outcome of novel species interactions formed under changing climates.     

Global variation in thermal tolerances and vulnerability of endotherms to climate change

The relationships among species'' physiological capacities and the geographical variation of ambient climate are of key importance to understanding the distribution of life on the Earth. Furthermore, predictions of how species will respond to climate change will profit from the explicit consideration of their physiological tolerances. The climatic variability hypothesis, which predicts that climatic tolerances are broader in more variable climates, provides an analytical framework for studying these relationships between physiology and biogeography. However, direct empirical support for the hypothesis is mostly lacking for endotherms, and few studies have tried to integrate physiological data into assessments of species'' climatic vulnerability at the global scale. Here, we test the climatic variability hypothesis for endotherms, with a comprehensive dataset on thermal tolerances derived from physiological experiments, and use these data to assess the vulnerability of species to projected climate change. We find the expected relationship between thermal tolerance and ambient climatic variability in birds, but not in mammals—a contrast possibly resulting from different adaptation strategies to ambient climate via behaviour, morphology or physiology. We show that currently most of the species are experiencing ambient temperatures well within their tolerance limits and that in the future many species may be able to tolerate projected temperature increases across significant proportions of their distributions. However, our findings also underline the high vulnerability of tropical regions to changes in temperature and other threats of anthropogenic global changes. Our study demonstrates that a better understanding of the interplay among species'' physiology and the geography of climate change will advance assessments of species'' vulnerability to climate change.     

Life stage,not climate change,explains observed tree range shifts

Ongoing climate change is expected to shift tree species distribution and therefore affect forest biodiversity and ecosystem services. To assess and project tree distributional shifts, researchers may compare the distribution of juvenile and adult trees under the assumption that differences between tree life stages reflect distributional shifts triggered by climate change. However, the distribution of tree life stages could differ within the lifespan of trees, therefore, we hypothesize that currently observed distributional differences could represent shifts over ontogeny as opposed to climatically driven changes. Here, we test this hypothesis with data from 1435 plots resurveyed after more than three decades across the Western Carpathians. We compared seedling, sapling and adult distribution of 12 tree species along elevation, temperature and precipitation gradients. We analyzed (i) temporal shifts between the surveys and (ii) distributional differences between tree life stages within both surveys. Despite climate warming, tree species distribution of any life stage did not shift directionally upward along elevation between the surveys. Temporal elevational shifts were species specific and an order of magnitude lower than differences among tree life stages within the surveys. Our results show that the observed range shifts among tree life stages are more consistent with ontogenetic differences in the species' environmental requirements than with responses to recent climate change. The distribution of seedlings substantially differed from saplings and adults, while the distribution of saplings did not differ from adults, indicating a critical transition between seedling and sapling tree life stages. Future research has to take ontogenetic differences among life stages into account as we found that distributional differences recently observed worldwide may not reflect climate change but rather the different environmental requirements of tree life stages.     

Physical effects of habitat‐forming species override latitudinal trends in temperature

Latitudinal and elevational temperature gradients (LTG and ETG) play central roles in biogeographical theory, underpinning predictions of large‐scale patterns in organismal thermal stress, species' ranges and distributional responses to climate change. Yet an enormous fraction of Earth's taxa live exclusively in habitats where foundation species modify temperatures. We examine little‐explored implications of this widespread trend using a classic model system for understanding heat stresses – rocky intertidal shores. Through integrated field measurements and laboratory trials, we demonstrate that thermal buffering by centimetre‐thick mussel and seaweed beds eliminates differences in stress‐inducing high temperatures and associated mortality risk that would otherwise arise over 14° of latitude and ~ 1 m of shore elevation. These results reveal the extent to which physical effects of habitat‐formers can overwhelm broad‐scale thermal trends, suggesting a need to re‐evaluate climate change predictions for many species. Notably, inhabitant populations may exhibit deceptive resilience to warming until refuge‐forming taxa become imperiled.     

Wrong place,wrong time: climate change‐induced range shift across fragmented habitat causes maladaptation and declined population size in a modelled bird species

Many species are locally adapted to decreased habitat quality at their range margins, and therefore show genetic differences throughout their ranges. Under contemporary climate change, range shifts may affect evolutionary processes at the expanding range margin due to founder events. In addition, populations that are affected by such founder events will, in the course of time, become located in the range centre. Recent studies investigated evolutionary changes at the expanding range margin, but have not assessed eventual effects across the species' range. We explored the possible influence of range shift on the level of adaptation throughout the species' total range. For this we used a spatially explicit, individual‐based simulation model of a woodland bird, parameterized after the middle spotted woodpecker ( Dendrocopos medius) in fragmented habitat. We simulated its range under climate change, and incorporated genetic differences at a single locus that determined the individual's degree of adaptation to optimal temperature conditions. Generalist individuals had a large thermal tolerance, but relatively low overall fitness, whereas climate specialists had high fitness combined with a small thermal tolerance. In equilibrium, the populations in the range centre were comprised of the specialists, whereas the generalists dominated the margins. In contrast, under temperature increase, the generalist numbers increased at the expanding margin and eventually also occupied the centre of the shifting range, whereas the specialists were located in the retracting margins. This was caused by founder events and led to overall maladaptation of the species, which resulted in a reduced metapopulation size and thus impeded the species' persistence. We therefore found no evidence for a complementary effect of local adaptation and range shifts on species' survival. Instead, we showed that founder events can cause local maladaptation which can amplify throughout the species' range, and, as such, hamper the species' persistence under climate change.     

Range-Wide Latitudinal and Elevational Temperature Gradients for the World's Terrestrial Birds: Implications under Global Climate Change

Species'' geographical distributions are tracking latitudinal and elevational surface temperature gradients under global climate change. To evaluate the opportunities to track these gradients across space, we provide a first baseline assessment of the steepness of these gradients for the world''s terrestrial birds. Within the breeding ranges of 9,014 bird species, we characterized the spatial gradients in temperature along latitude and elevation for all and a subset of bird species, respectively. We summarized these temperature gradients globally for threatened and non-threatened species and determined how their steepness varied based on species'' geography (range size, shape, and orientation) and projected changes in temperature under climate change. Elevational temperature gradients were steepest for species in Africa, western North and South America, and central Asia and shallowest in Australasia, insular IndoMalaya, and the Neotropical lowlands. Latitudinal temperature gradients were steepest for extratropical species, especially in the Northern Hemisphere. Threatened species had shallower elevational gradients whereas latitudinal gradients differed little between threatened and non-threatened species. The strength of elevational gradients was positively correlated with projected changes in temperature. For latitudinal gradients, this relationship only held for extratropical species. The strength of latitudinal gradients was better predicted by species'' geography, but primarily for extratropical species. Our findings suggest threatened species are associated with shallower elevational temperature gradients, whereas steep latitudinal gradients are most prevalent outside the tropics where fewer bird species occur year-round. Future modeling and mitigation efforts would benefit from the development of finer grain distributional data to ascertain how these gradients are structured within species'' ranges, how and why these gradients vary among species, and the capacity of species to utilize these gradients under climate change.     

The geographical range of British birds expands during 15 years of warming

Capsule Abundance monitoring data suggest that the short-term response of breeding birds to recent warming in Great Britain has been range expansion, caused by poleward shifts of leading range margins and no significant shifts of trailing range margins.

Aims To quantify latitudinal and elevational shifts of breeding bird populations in Great Britain and test for differential shifts in range margins during a period of warming (1994–2009).

Methods We modelled the population density of 80 species as a smooth function of latitude, longitude, elevation and year. Reference points on the distribution curve were used to describe latitudinal and elevational shifts.

Results Across species, poleward shifts in the leading range margin were greater than in the range-centre. The trailing range margin was largely static, providing evidence for significant range expansion. The magnitude of latitudinal range shift lagged behind the equivalent shift in temperature, suggesting that species may be accumulating a climatic debt. There was no evidence for consistent elevational shifts.

Conclusion Contrary to the generally expected long-term consequences of climate change of range contraction, we show that the short-term response to recent warming has been range expansion. This suggests the mechanisms of short-term and long-term consequences of climate change may differ.     

The endemic kelp Lessonia corrugata is being pushed above its thermal limits in an ocean warming hotspot

Kelps are in global decline due to climate change, which includes ocean warming. To identify vulnerable species, we need to identify their tolerances to increasing temperatures and determine whether tolerances are altered by co-occurring drivers such as inorganic nutrient levels. This is particularly important for those species with restricted distributions, which may already be experiencing thermal stress. To identify thermal tolerance of the range-restricted kelp Lessonia corrugata, we conducted a laboratory experiment on juvenile sporophytes to measure performance (growth, photosynthesis) across its thermal range (4–22°C). We determined the upper thermal limit for growth and photosynthesis to be ~22–23°C, with a thermal optimum of ~16°C. To determine if elevated inorganic nitrogen availability could enhance thermal tolerance, we compared the performance of juveniles under low (4.5 μmol · d⁻¹) and high (90 μmol · d⁻¹) nitrate conditions at and above the thermal optimum (16–23.5°C). Nitrate enrichment did not enhance thermal performance at temperatures above the optimum but did lead to elevated growth rates at the thermal optimum. Our results indicate L. corrugata is likely to be extremely susceptible to moderate ocean warming and marine heatwaves. Peak sea surface temperatures during summer in eastern and northeastern Tasmania can reach up to 20–21°C, and climate projections suggest that L. corrugata's thermal limit will be regularly exceeded by 2050 as southeastern Australia is a global ocean-warming hotspot. By identifying the upper thermal limit of L. corrugata, we have taken a critical step in predicting the future of the species in a warming climate.     

Illuminating geographical patterns in species' range shifts

Species' range shifts in response to ongoing climate change have been widely documented, but although complex spatial patterns in species' responses are expected to be common, comprehensive comparisons of species' ranges over time have undergone little investigation. Here, we outline a modeling framework based on historical and current species distribution records for disentangling different drivers (i.e. climatic vs. nonclimatic) and assessing distinct facets (i.e. colonization, extirpation, persistence, and lags) of species' range shifts. We used extensive monitoring data for stream fish assemblages throughout France to assess range shifts for 32 fish species between an initial period (1980–1992) and a contemporary one (2003–2009). Our results provide strong evidence that the responses of individual species varied considerably and exhibited complex mosaics of spatial rearrangements. By dissociating range shifts in climatically suitable and unsuitable habitats, we demonstrated that patterns in climate‐driven colonization and extirpation were less marked than those attributed to nonclimatic drivers, although this situation could rapidly shift in the near future. We also found evidence that range shifts could be related to some species' traits and that the traits involved varied depending on the facet of range shift considered. The persistence of populations in climatically unsuitable areas was greater for short‐lived species, whereas the extent of the lag behind climate change was greater for long‐lived, restricted‐range, and low‐elevation species. We further demonstrated that nonclimatic extirpations were primarily related to the size of the species' range, whereas climate‐driven extirpations were better explained by thermal tolerance. Thus, the proposed framework demonstrated its potential for markedly improving our understanding of the key processes involved in range shifting and also offers a template for informing management decisions. Conservation strategies would greatly benefit from identifying both the geographical patterns and the species' traits associated with complex modifications of species' distributions in response to global changes.     

Coping with the cold: minimum temperatures and thermal tolerances dominate the ecology of mountain ants

1. Ants (Hymenoptera: Formicidae) are often cited as highly thermophilic and this has led to a range of studies investigating their thermal tolerances. It is unknown, however, if the geographic distribution of ant thermal tolerance conforms to the two major macropyhsiological rules that have been found in other taxa: Janzen's and Brett's rules. In addition, there is a paucity of data on how the lower thermal tolerances of ants are able to influence behaviour. 2. These two knowledge gaps were addressed here by sampling ants across a 1500 m elevational gradient in southern Africa and estimating the upper (CTmax) and lower (CTmin) thermal tolerances of 31 and 28 species, respectively. Ant abundances and soil temperatures were also recorded across the gradient over 6 years. 3. It was found that the average CTmin of the ants declined with elevation along with environmental temperatures. It was also found that the correlation between abundance and local temperature depended on the ant species' CTmin. The activity of species with a low CTmin was not constrained by temperature, whereas those with a high CTmin were limited by low temperatures. 4. For the first time, evidence is provided here that the thermal tolerances of ants are consistent with two major macrophysiological rules: Brett's rule and Janzen's rule. A mechanistic link between physiology, behaviour and the environment is also shown, which highlights that the ability of ants to deal with the cold may be a key, but often overlooked, factor allowing multiple ant species to succeed within an environment.     

Vanishing islands in the sky? A comparison of correlation- and mechanism-based forecasts of range dynamics for montane salamanders under climate change

Forecasting the effects of climate change on species and populations is a fundamental goal of conservation biology, especially for montane endemics which seemingly are under the greatest threat of extinction given their association with cool, high elevation habitats. Species distribution models (also known as niche models) predict where on the landscape there is suitable habitat for a species of interest. Correlative niche modeling, the most commonly employed approach to predict species' distributions, relies on correlations between species' localities and current environmental data. This type of model could spuriously forecast less future suitable habitat because species' current distributions may not adequately represent their thermal tolerance, and future climate conditions may not be analogous to current conditions. We compared the predicted distributions for three montane species of Plethodon salamanders in the southern Appalachian Mountains of North America using a correlative modeling approach and a mechanistic model. The mechanistic model incorporates species-specific physiology, morphology and behavior to predict an annual energy budget on the landscape. Both modeling approaches performed well at predicting the species' current distributions and predicted that all species could persist in habitats at higher elevation through 2085. The mechanistic model predicted more future suitable habitat than the correlative model. We attribute these differences to the mechanistic approach being able to model shifts in key range-limiting biological processes (changes in surface activity time and energy costs) that the correlative approach cannot. Choice of global circulation model (GCM) contributed significantly to distribution predictions, with a tenfold difference in future suitability based on GCM, indicating that GCM variability should be either directly included in models of species distributions or, indirectly, through the use of multi-model ensemble averages. Our results indicate that correlative models are over-predicting habitat loss for montane species, suggesting a critical need to incorporate mechanisms into forecasts of species' range dynamics.     

Breeding success at the range margin of a desert species: implications for a climate‐induced elevational shift

If the breeding range of a species is limited by biotic or abiotic environmental factors that depress breeding success at the range margin, then range expansion is expected when those limiting factors are alleviated. Over a three‐year period, we measured breeding success of a desert species, black‐throated sparrow Amphispiza bilineata, along a steep elevation gradient between the Peninsular Mountains and Colorado Desert (San Diego County, California) that is undergoing a warming trend. We compared breeding success at geographically marginal locations (higher‐elevation chaparral sites) to more central locations (lower‐elevation desert scrub sites) only a short distance apart. Breeding success was measured at the nest level, territory level, and population level. At each level measured, breeding success tended to be greater at higher‐elevation chaparral sites at the distribution margin compared to lower‐elevation sites where the bird was more common. Black‐throated sparrows had 100% reproductive failure at lower‐elevation sites during the two driest years of our study (2006–2007), but did relatively well at higher‐elevation sites. Only in a wetter year (2008) was breeding success improved at lower‐elevation sites. Surprisingly, there was no evidence of an upward elevational shift in distribution over a 26‐year period despite a clear warming trend and drier conditions. Greater territory density at lower‐elevation sites with reproductive failure during dry years suggests the possibility of an ecological trap in this system, which could prevent or delay climate‐induced range shifts. A common presumption has been that desert species will undergo relatively mild negative impacts due to a warming climate, but it is possible that some desert species are already at or near their temperature and aridity tolerance limits within their current range and shifts may not always be possible.     

Thermal specialization across large geographical scales predicts the resilience of mangrove crab populations to global warming

The broad prediction that ectotherms will be more vulnerable to climate change in the tropics than in temperate regions includes assumptions about centre/edge population effects that can only be tested by within‐species comparisons across wide latitudinal gradients. Here, we investigated the thermal vulnerability of two mangrove crab species, comparing populations at the centre (Kenya) and edge (South Africa) of their distributions. At the same time, we investigated the role of respiratory mode (water‐ versus air‐breathing) in determining the thermal tolerance in amphibious organisms. To do this, we compared the vulnerability to acute temperature fluctuations of two sympatric species with two different lifestyle adaptations: the free living Perisesarma guttatum and the burrowing Uca urvillei, both pivotal to the ecosystem functioning of mangroves. The results revealed the air‐breathing U. urvillei to be a thermal generalist with much higher thermal tolerances than P. guttatum. Importantly, however, we found that, while U. urvillei showed little difference between edge and centre populations, P. guttatum showed adaptation to local conditions. Equatorial populations had elevated tolerances to acute heat stress and mechanisms of partial thermoregulation, which make them less vulnerable to global warming than temperate conspecifics. The results reveal both the importance of respiratory mode to thermal tolerance and the unexpected potential for low latitude populations/species to endure a warming climate. The results also contribute to a conceptual model on the latitudinal thermal tolerance of these key species. This highlights the need for an integrated population‐level approach to predict the consequences of climate change.     

Compositional shifts in Costa Rican forests due to climate‐driven species migrations

Species are predicted to shift their distributions upslope or poleward in response to global warming. This prediction is supported by a growing number of studies documenting species migrations in temperate systems but remains poorly tested for tropical species, and especially for tropical plant species. We analyzed changes in tree species composition in a network of 10 annually censused 1‐ha plots spanning an altitudinal gradient of 70–2800 m elevation in Costa Rica. Specifically, we combined plot data with herbarium records (accessed through GBIF) to test if the plots' community temperature scores (CTS, average thermal mean of constituent species weighted by basal area) have increased over the past decade as is predicted by climate‐driven species migrations. In addition, we quantified the contributions of stem growth, recruitment, and mortality to the observed patterns. Supporting our a priori hypothesis of upward species migrations, we found that there have been consistent directional shifts in the composition of the plots, such that the relative abundance of lowland species, and hence CTS, increased in 90% of plots. The rate of the observed compositional shifts corresponds to a mean thermal migration rate (TMR) of 0.0065 °C yr^?1 (95% CI = 0.0005–0.0132 °C yr^?1). While the overall TMR is slower than predicted based on concurrent regional warming of 0.0167 °C yr^?1, migrations were on pace with warming in 4 of the 10 plots. The observed shifts in composition were driven primarily by mortality events (i.e., the disproportionate death of highland vs. lowland species), suggesting that individuals of many tropical tree species will not be able to tolerate future warming and thus their persistence in the face of climate change will depend on successful migrations. Unfortunately, in Costa Rica and elsewhere, land area inevitably decreases at higher elevations; hence, even species that are able to migrate successfully will face heightened risks of extinction.     

Butterfly phenology in Mediterranean mountains using space‐for‐time substitution

Inferring species' responses to climate change in the absence of long‐term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14‐day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23‐day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8‐day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space‐for‐time substitutions, understanding species‐specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.