Diurnal temperature variation impacts energetics but not reproductive effort across seasons in a temperate dung beetle.

Temperature varies on multiple timescales and ectotherms must adjust to these changes to survive. These adjustments may lead to energetic trade-offs between self-maintenance and reproductive investment. However, we know little about how diurnal and seasonal temperature changes impact energy allocation. Here we used a combination of empirical data and modeling of both thermoregulatory behaviors and body temperature to examine potential energetic trade-offs in the dung beetle Onthophagus taurus. Beginning in March 2020, universities and laboratories were officially closed due to the COVID-19 pandemic. We thus performed experiments at a private residence near Knoxville, Tennessee, USA, leveraging the heating, ventilation and air conditioning of the home to manipulate temperature and compare beetle responses to stable indoor temperatures versus variable outdoor temperatures. We collected O. taurus beetles in the early-, mid-, and late-breeding seasons to examine energetics and reproductive output in relation to diurnal and seasonal temperature fluctuations. We recorded the mass of field fresh beetles before and after a 24-h fast and used the resulting change in mass as a proxy for energetic costs of self-maintenance across seasons. To understand the impacts of diurnal fluctuations on energy allocation, we held beetles either indoors or outdoors for 14-day acclimation trials, fed them cow dung, and recorded mass change and reproductive output. Utilizing biophysical models, we integrated individual-level biophysical characteristics, microhabitat-specific performance, respirometry data, and thermoregulatory behaviors to predict temperature-induced changes to the allocation of energy toward survival and reproduction. During 24 h of outdoor fasting, we found that beetles experiencing reduced temperature variation lost more mass than those experiencing greater temperature variation, and this was not affected by season. By contrast, during the 14-day acclimation trials, we found that beetles experiencing reduced temperature variation (i.e., indoors) gained more mass than those experiencing greater temperature variation (i.e., outdoors). This effect may have been driven by shifts in the metabolism of the beetles during acclimation to increased temperature variation. Despite the negative relationship between temperature variation and energetic reserves, the only significant predictor of reproductive output was mean temperature. Taken together, we find that diurnal temperature fluctuations are important for driving energetics, but not reproductive output.

A trait-based framework for dung beetle functional ecology.

Traits are key for understanding the environmental responses and ecological roles of organisms. Trait approaches to functional ecology are well established for plants, whereas consistent frameworks for animal groups are less developed. Here we suggest a framework for the study of the functional ecology of animals from a trait-based response-effect approach, using dung beetles as model system. Dung beetles are a key group of decomposers that are important for many ecosystem processes. The lack of a trait-based framework tailored to this group has limited the use of traits in dung beetle functional ecology. We review which dung beetle traits respond to the environment and affect ecosystem processes, covering the wide range of spatial, temporal and biological scales at which they are involved. Dung beetles show trait-based responses to variation in temperature, water, soil properties, trophic resources, light, vegetation structure, competition, predation and parasitism. Dung beetles' influence on ecosystem processes includes trait-mediated effects on nutrient cycling, bioturbation, plant growth, seed dispersal, other dung-based organisms and parasite transmission, as well as some cases of pollination and predation. We identify 66 dung beetle traits that are either response or effect traits, or both, pertaining to six main categories: morphology, feeding, reproduction, physiology, activity and movement. Several traits pertain to more than one category, in particular dung relocation behaviour during nesting or feeding. We also identify 136 trait-response and 77 trait-effect relationships in dung beetles. No response to environmental stressors nor effect over ecological processes were related with traits of a single category. This highlights the interrelationship between the traits shaping body-plans, the multi-functionality of traits, and their role linking responses to the environment and effects on the ecosystem. Despite current developments in dung beetle functional ecology, many knowledge gaps remain, and there are biases towards certain traits, functions, taxonomic groups and regions. Our framework provides the foundations for the thorough development of trait-based dung beetle ecology. It also serves as an example framework for other taxa.

Experimental increases in temperature mean and variance alter reproductive behaviours in the dung beetle Phanaeus vindex.

Temperature profoundly impacts insect development, but plasticity of reproductive behaviours may mediate the impacts of temperature change on earlier life stages. Few studies have examined the potential for adult behavioural plasticity to buffer offspring from the warmer, more variable temperatures associated with climate change. We used a field manipulation to examine whether the dung beetle Phanaeus vindex alters breeding behaviours in response to temperature changes and whether behavioural shifts protect offspring from temperature changes. Dung beetles lay eggs inside brood balls made of dung that are buried underground. Brood ball depth impacts the temperatures offspring experience with consequences for development. We placed adult females in either control or greenhouse treatments that simultaneously increased temperature mean and variance. We found that females in greenhouse treatments produced more brood balls that were smaller and buried deeper than controls, suggesting brood ball number or burial depth may come at a cost to brood ball size, which can impact offspring nutrition. Despite being buried deeper, brood balls from the greenhouse treatment experienced warmer mean temperatures but similar amplitudes of temperature fluctuation relative to controls. Our findings suggest adult behaviours may partially buffer developing offspring from temperature changes.

The relative influence of history, climate, topography and vegetation structure on local animal richness varies among taxa and spatial grains.

Understanding the spatial scales at which environmental factors drive species richness patterns is a major challenge in ecology. Due to the trade-off between spatial grain and extent, studies tend to focus on a single spatial scale, and the effects of multiple environmental variables operating across spatial scales on the pattern of local species richness have rarely been investigated. Here, we related variation in local species richness of ground beetles, landbirds and small mammals to variation in vegetation structure and topography, regional climate, biome diversity and glaciation history for 27 sites across the USA at two different spatial grains. We studied the relative influence of broad-scale (landscape) environmental conditions using variables estimated at the site level (climate, productivity, biome diversity and glacial era ice cover) and fine-scale (local) environmental conditions using variables estimated at the plot level (topography and vegetation structure) to explain local species richness. We also examined whether plot-level factors scale up to drive continental scale richness patterns. We used Bayesian hierarchical models and quantified the amount of variance in observed richness that was explained by environmental factors at different spatial scales. For all three animal groups, our models explained much of the variation in local species richness (85%-89%), but site-level variables explained a greater proportion of richness variance than plot-level variables. Temperature was the most important site-level predictor for explaining variance in landbirds and ground beetles richness. Some aspects of vegetation structure were the main plot-level predictors of landbird richness. Environmental predictors generally had poor explanatory power for small mammal richness, while glacial era ice cover was the most important site-level predictor. Relationships between plot-level factors and richness varied greatly among geographical regions and spatial grains, and most relationships did not hold when predictors were scaled up to the continental scale. Our results suggest that the factors that determine richness may be highly dependent on spatial grain, geography, and animal group. We demonstrate that instead of artificially manipulating the resolution to study multiscale effects, a hierarchical approach that uses fine grain data at broad extents could help solve the issue of scale selection in environment-richness studies.

Dung beetles show metabolic plasticity as pupae and smaller adult body size in response to increased temperature mean and variance.

Though organisms may use thermal plasticity to cope with novel temperature regimes, our understanding of plastic responses is limited. Research on thermal plasticity has traditionally focused on the response of organisms to shifts in mean temperatures. However, increased temperature variation can have a greater impact on organismal performance than mean temperature alone. In addition, thermal plasticity studies are often designed to investigate plasticity in response to more extreme temperatures despite the fact that organisms make physiological adjustments to diurnal temperature fluctuations that they experience. Using pupae of the dung beetle Onthophagus taurus, we investigated the potential for plasticity in response to increasing temperature mean and variance using thermal regimes that were well within the species critical thermal limits. We reared 40 beetles from egg to pupae (n = 20) or adults (n = 20) at one of nine incubation treatments, including all combinations of three mean temperatures (22, 24, 26 °C) and three amplitudes of fluctuation (±2, ±4, ±8 °C). To measure thermal plasticity of pupae, we quantified CO2 production across a range of temperatures (i.e., 15, 20, 25, and 30 °C) for 20 beetles per treatment. The relationship between CO2 production and temperature provides an estimate of energetic costs at a given temperature (i.e., using the intercept) and thermal sensitivity (i.e., using the slope). We reared the remaining O. taurus in each treatment (n = 20) to adulthood and then recorded mass (g) to determine body size, a proxy for fitness. Pupae exhibited thermal plasticity in response to the additive and interactive effects of temperature mean and variance. Pupae reared in the warmest and most variable treatment (26 ± 8 °C) showed the greatest decrease in overall metabolism compared to all other treatments, and adult beetles from this treatment (26 ± 8 °C) were also significantly smaller than adult beetles from any other treatment. We found that both temperature mean and variance contributed to thermal plasticity of pupae and had consequences for adult body size, a trait related to dung beetle fitness. Importantly, the temperatures we used in our treatments are not extreme and are likely well below the critical thermal maxima of the species, demonstrating that organisms can make adjustments to temperatures they experience across diurnal or seasonal timescales.

Global progress in incorporating climate adaptation into land protection for biodiversity since Aichi targets.

Climate adaptation strategies are being developed and implemented to protect biodiversity from the impacts of climate change. A well-established strategy involves the identification and addition of new areas for conservation, and most countries agreed in 2010 to expand the global protected area (PA) network to 17% by 2020 (Aichi Biodiversity Target 11). Although great efforts to expand the global PA network have been made, the potential of newly established PAs to conserve biodiversity under future climate change remains unclear at the global scale. Here, we conducted the first global-extent, country-level assessment of the contribution of PA network expansion toward three key land prioritization approaches for biodiversity persistence under climate change: protecting climate refugia, protecting abiotic diversity, and increasing connectivity. These approaches avoid uncertainties of biodiversity predictions under climate change as well as the issue of undescribed species. We found that 51% of the countries created new PAs in locations with lower mean climate velocity (representing better climate refugia) and 58% added PAs in areas with higher mean abiotic diversity compared to the available, non-human-dominated lands not chosen for protection. However, connectivity among PAs declined in 53% of the countries, indicating that many new PAs were located far from existing PAs. Lastly, we identified potential improvements for climate adaptation, showing that 94% of the countries have the opportunity to improve in executing one or more approaches to conserve biodiversity. Most countries (60%) were associated with multiple opportunities, highlighting the need for integrative strategies that target multiple land protection approaches. Our results demonstrate that a global improvement in the protection of climate refugia, abiotic diversity, and connectivity of reserves is needed to complement land protection informed by existing and projected species distributions. Our study also provides a framework for countries to prioritize land protection for climate adaptation using publicly available data.

Behavioural responses to warming differentially impact survival in introduced and native dung beetles.

Anthropogenic changes are often studied in isolation but may interact to affect biodiversity. For example, climate change could exacerbate the impacts of biological invasions if climate change differentially affects invasive and native species. Behavioural plasticity may mitigate some of the impacts of climate change, but species vary in their degree of behavioural plasticity. In particular, invasive species may have greater behavioural plasticity than native species since plasticity helps invasive species establish and spread in new environments. This plasticity could make invasives better able to cope with climate change. Here our goal was to examine whether reproductive behaviours and behavioural plasticity vary between an introduced and a native Onthophagus dung beetle species in response to warming temperatures and how differences in behaviour influence offspring survival. Using a repeated measures design, we exposed small colonies of introduced O. taurus and native O. hecate to three temperature treatments, including a control, low warming and high warming treatment, and then measured reproductive behaviours, including the number, size and burial depth of brood balls. We reared offspring in their brood balls in developmental temperatures that matched those of the brood ball burial depth to quantify survival. We found that the introduced O. taurus produced more brood balls and larger brood balls, and buried brood balls deeper than the native O. hecate in all treatments. However, the two species did not vary in the degree of behavioural plasticity in response to warming. Differences in reproductive behaviours did affect survival such that warming temperatures had a greater effect on survival of offspring of native O. hecate compared to introduced O. taurus. Overall, our results suggest that differences in behaviour between native and introduced species are one mechanism through which climate change may exacerbate negative impacts of biological invasions.

Different amplitudes of temperature fluctuation induce distinct transcriptomic and metabolomic responses in the dung beetle Phanaeus vindex.

Most studies exploring molecular and physiological responses to temperature have focused on constant temperature treatments. To gain a better understanding of the impact of fluctuating temperatures, we investigated the effects of increased temperature variation on Phanaeus vindex dung beetles across levels of biological organization. Specifically, we hypothesized that increased temperature variation is energetically demanding. We predicted that thermal sensitivity of metabolic rate and energetic reserves would be reduced with increasing fluctuation. To test this, we examined the responses of dung beetles to constant (20°C), low fluctuation (20±5°C), or high fluctuation (20±12°C) temperature treatments using respirometry, assessment of energetic reserves and HPLC-MS-based metabolomics. We found no significant differences in metabolic rate or energetic reserves, suggesting increased fluctuations were not energetically demanding. To understand why there was no effect of increased amplitude of temperature fluctuation on energetics, we assembled and annotated a de novo transcriptome, finding non-overlapping transcriptomic and metabolomic responses of beetles exposed to different fluctuations. We found that 58 metabolites increased in abundance in both fluctuation treatments, but 15 only did so in response to high-amplitude fluctuations. We found that 120 transcripts were significantly upregulated following acclimation to any fluctuation, but 174 were upregulated only in beetles from the high-amplitude fluctuation treatment. Several differentially expressed transcripts were associated with post-translational modifications to histones that support a more open chromatin structure. Our results demonstrate that acclimation to different temperature fluctuations is distinct and may be supported by increasing transcriptional plasticity. Our results indicate for the first time that histone modifications may underlie rapid acclimation to temperature variation.

Life stages differ in plasticity to temperature fluctuations and uniquely contribute to adult phenotype in Onthophagus taurus dung beetles.

Adaptive thermal plasticity allows organisms to adjust their physiology to cope with fluctuating environments. However, thermal plasticity is rarely studied in response to thermal variability and is often measured in a single life stage. Plasticity in response to thermal variability likely differs from responses to constant temperature or acute stress. In addition, life stages likely differ in their plasticity, and responses in one stage may be affected by the experiences in a previous stage. Increasing the resolution with which we understand thermal plasticity in response to thermal variation across ontogeny is crucial to understanding how organisms cope with the thermal variation in their environment and to estimating the capacity of plasticity to mitigate costs of rapid environmental change. We wanted to know whether life stages differ in their capacity for thermal plasticity under temperature fluctuations. We reared Onthophagus taurus dung beetles in either low or high temperature fluctuation treatments and quantified thermal plasticity of metabolism of pupae and adults. We found that adults were thermally plastic and pupae were not. Next, we tested whether the plasticity observed in the adult life stage was affected by the thermal conditions during development. We again used low and high temperature fluctuation treatments and reared individuals in one condition through all egg to pupal stages. At eclosion, we switched half of the individuals in each treatment to the opposite fluctuation condition and, later, measured thermal plasticity of metabolism in adults. We found that temperature conditions experienced during the adult stage, but not egg to pupal stages, affect adult thermal plasticity. However, temperature fluctuations during development affect adult body size, suggesting that some aspects of the adult phenotype are decoupled from previous life stages and others are not. Our data demonstrate that life stages mount different responses to temperature variability and uniquely contribute to the adult phenotype. These findings emphasize the need to broadly integrate the life cycle into studies of phenotypic plasticity and physiology; doing so should enhance our ability to predict organismal responses to rapid global change and inform conservation efforts.

Panmixia across elevation in thermally sensitive Andean dung beetles.

Janzen's seasonality hypothesis predicts that organisms inhabiting environments with limited climatic variability will evolve a reduced thermal tolerance breadth compared with organisms experiencing greater climatic variability. In turn, narrow tolerance breadth may select against dispersal across strong temperature gradients, such as those found across elevation. This can result in narrow elevational ranges and generate a pattern of isolation by environment or neutral genetic differentiation correlated with environmental variables that are independent of geographic distance. We tested for signatures of isolation by environment across elevation using genome-wide SNP data from five species of Andean dung beetles (subfamily Scarabaeinae) with well-characterized, narrow thermal physiologies, and narrow elevational distributions. Contrary to our expectations, we found no evidence of population genetic structure associated with elevation and little signal of isolation by environment. Further, elevational ranges for four of five species appear to be at equilibrium and show no decay of genetic diversity at range limits. Taken together, these results suggest physiological constraints on dispersal may primarily operate outside of a stable realized niche and point to a lower bound on the spatial scale of local adaptation.

Fifty Years of Mountain Passes: A Perspective on Dan Janzen's Classic Article.

In 1967, Dan Janzen published "Why Mountain Passes Are Higher in the Tropics" in The American Naturalist. Janzen's seminal article has captured the attention of generations of biologists and continues to inspire theoretical and empirical work. The underlying assumptions and derived predictions are broadly synthetic and widely applicable. Consequently, Janzen's "seasonality hypothesis" has proven relevant to physiology, climate change, ecology, and evolution. To celebrate the fiftieth anniversary of this highly influential article, we highlight the past, present, and future of this work and include a unique historical perspective from Janzen himself.

Understanding Evolutionary Impacts of Seasonality: An Introduction to the Symposium.

Seasonality is a critically important aspect of environmental variability, and strongly shapes all aspects of life for organisms living in highly seasonal environments. Seasonality has played a key role in generating biodiversity, and has driven the evolution of extreme physiological adaptations and behaviors such as migration and hibernation. Fluctuating selection pressures on survival and fecundity between summer and winter provide a complex selective landscape, which can be met by a combination of three outcomes of adaptive evolution: genetic polymorphism, phenotypic plasticity, and bet-hedging. Here, we have identified four important research questions with the goal of advancing our understanding of evolutionary impacts of seasonality. First, we ask how characteristics of environments and species will determine which adaptive response occurs. Relevant characteristics include costs and limits of plasticity, predictability, and reliability of cues, and grain of environmental variation relative to generation time. A second important question is how phenological shifts will amplify or ameliorate selection on physiological hardiness. Shifts in phenology can preserve the thermal niche despite shifts in climate, but may fail to completely conserve the niche or may even expose life stages to conditions that cause mortality. Considering distinct environmental sensitivities of life history stages will be key to refining models that forecast susceptibility to climate change. Third, we must identify critical physiological phenotypes that underlie seasonal adaptation and work toward understanding the genetic architectures of these responses. These architectures are key for predicting evolutionary responses. Pleiotropic genes that regulate multiple responses to changing seasons may facilitate coordination among functionally related traits, or conversely may constrain the expression of optimal phenotypes. Finally, we must advance our understanding of how changes in seasonal fluctuations are impacting ecological interaction networks. We should move beyond simple dyadic interactions, such as predator prey dynamics, and understand how these interactions scale up to affect ecological interaction networks. As global climate change alters many aspects of seasonal variability, including extreme events and changes in mean conditions, organisms must respond appropriately or go extinct. The outcome of adaptation to seasonality will determine responses to climate change.

Biological Impacts of Thermal Extremes: Mechanisms and Costs of Functional Responses Matter.

Thermal performance curves enable physiological constraints to be incorporated in predictions of biological responses to shifts in mean temperature. But do thermal performance curves adequately capture the biological impacts of thermal extremes? Organisms incur physiological damage during exposure to extremes, and also mount active compensatory responses leading to acclimatization, both of which alter thermal performance curves and determine the impact that current and future extremes have on organismal performance and fitness. Thus, these sub-lethal responses to extreme temperatures potentially shape evolution of thermal performance curves. We applied a quantitative genetic model and found that beneficial acclimatization and cumulative damage alter the extent to which thermal performance curves evolve in response to thermal extremes. The impacts of extremes on the evolution of thermal performance curves are reduced if extremes cause substantial mortality or otherwise reduce fitness differences among individuals. Further empirical research will be required to understand how responses to extremes aggregate through time and vary across life stages and processes. Such research will enable incorporating passive and active responses to sub-lethal stress when predicting the impacts of thermal extremes.

Beyond the Mean: Biological Impacts of Cryptic Temperature Change.

Studies have typically used shifts in mean temperatures to make predictions about the biotic impacts of climate change. Though shifts in mean temperatures correlate with changes in phenology and distributions, other hidden, or cryptic, changes in temperature, such as temperature variation and extreme temperatures, could pose greater risks to species and ecological communities. Yet, these cryptic temperature changes have received relatively little attention because mean temperatures are readily available and the organism-appropriate temperature response is often elusive. An alternative to using mean temperatures is to view organisms as physiological filters of hourly temperature data. We explored three classes of physiological filters: (1) nonlinear thermal responses using performance curves of insect fitness, (2) cumulative thermal effects using degree-day models for corn emergence, and (3) threshold temperature effects using critical thermal maxima and minima for diverse ectotherms. For all three physiological filters, we determined the change in biological impacts of hourly temperature data from a standard reference period (1961-90) to a current period (2005-10). We then examined how well mean temperature changes during the same time period predicted the biotic impacts we determined from hourly temperature data. In all cases, mean temperature alone provided poor predictions of the impacts of climate change. These results suggest that incorporating high frequency temperature data can provide better predictions for how species will respond to temperature change.

Reproductive phenology of epiphytes in Monteverde, Costa Rica

Phenology of plants, or the timing of life cycle events, is important for understanding plant ecology, forest dynamics, and plant-animal interactions. In tropical forests, studies that document epiphyte reproductive phenology are relatively few because of the challenges of tracking plants that live in the canopy. Phenological patterns for 279 individuals of 7 epiphyte species were examined across 12 months in a tropical montane forest in Monteverde, Costa Rica. Epiphytes were located in one of two common tree species, Ficus tuerckheimii (Moraceae) or Ocotea tonduzii (Lauraceae). Flowering and fruiting (i.e., when ripe or unripe fruit is present on the plant) of study plants was recorded on monthly intervals, and phenology was examined as a function of the season at the study site (i.e., wet, transition, or dry season), and pollinator syndrome (bird-, or insect-pollinated) and seed dispersal syndrome (bird-, bat-, or wind-dispersed) of each plant. Though some epiphyte species flowered and fruited throughout the year, the majority showed significant seasonality in phenological events. Based on circular statistics, the timing of mean flowering of different epiphyte species varied, however, timing of mean fruiting for most species tended to occur during the wet season. Insect- and bird-pollinated species had peak flowering during the dry season and late wet season, respectively. Bird-dispersed fruits were present each month of the year with peaks from February to October and again in December. Wind-dispersed fruits were observed eight months of the year with a peak in the early wet season. The timing of epiphyte flowering coincided with flowering of large trees in the area. Epiphyte fruiting, however, is distinct from large tree fruiting. Our results demonstrate the seasonal nature of flowering and fruiting in individual epiphyte species while also highlighted the asynchronous nature of phenological events amongst the epiphyte community.

La fenología de las plantas, o el cronograma de eventos en el ciclo de vida, es importante para la comprensión de la ecología vegetal, la dinámica de los bosques y de las interacciones planta-animal. En los bosques tropicales, los estudios que documentan la fenología de las epífitas son relativamente pocos debido a los desafíos que representa darles seguimiento a las plantas que viven en el dosel. Aquí se presenta un reporte de los patrones fenológicos de 279 individuos de 7 especies de epífitas que abarca 12 meses en un bosque montano tropical en Monteverde, Costa Rica. Las epífitas se encuentran en una de las dos especies de árboles comunes, Ficus tuerckheimii (Moraceae) u Ocotea tonduzii (Lauraceae). Se registró la floración y fructificación (i.e. cuando la fruta madura o inmadura está presente en la planta) de las plantas de estudio en intervalos mensuales y examinó la fenología a través de las estaciones en el sitio de estudio (i.e. estación húmeda, transición, o seca) y el síndrome de polinizadores (ave- , o insecto-polinización ) y el síndrome de dispersión de semillas (ave-, murciélago-, o viento- dispersión) de cada planta. Aunque la mayoría de las especies de epífitas tuvieron flores y frutos durante todo el año, la mayoría mostró estacionalidad significativa en los eventos fenoló-gicos. Con base en estadísticas circulares, el tiempo de floración promedio de las especies de epífitas es variado, sin embargo, el momento de la fructificación promedio para la mayoría de las especies tiende a ocurrir durante la estación húmeda. Especies de insectos y aves de polinización tenían pico de floración durante la estación seca y la estación lluviosa tarde, respectivamente. Frutas dispersadas por aves estaban presentes todos los meses del año con picos de febrero a octubre y de nuevo en diciembre. Frutas dispersadas por el viento se observaron ocho meses del año con un pico en la temporada de lluvias temprana. El momento de la floración de las epífitas coincidió con la floración de árboles de gran tamaño en la zona. La fructificación de las epífitas, sin embargo, fue diferente de la fructificación de los árboles de gran tamaño. Nuestros resultados ponen de manifiesto el carácter estacional de la floración y fructificación de las especies epífitas individuales, además de destacar el carácter asincrónico de los eventos fenológicos entre toda la comunidad de epífitas.

Climate change creates rapid species turnover in montane communities.

Recent decades have seen substantial changes in patterns of biodiversity worldwide. Simultaneously, climate change is producing a widespread pattern of species' range shifts to higher latitudes and higher elevations, potentially creating novel assemblages as species shift at different rates. However, the direct link between species' turnover as a result of climate-induced range shifts has not yet been empirically evaluated. We measured rates of species turnover associated with species' range shifts in relatively undisturbed montane areas in Asia, Europe, North America, South America, and the Indo-Pacific. We show that species turnover is rapidly creating novel assemblages, and this can be explained by variable changes in species' range limits following warming. Across all the areas we analyzed, mean species' turnover was 12% per decade, which was nearly balanced between the loss of existing co-occurrences and the gain of novel co-occurrences. Turnover appears to be more rapid among ectothermic assemblages, and some evidence suggests tropical assemblages may be responding at more rapid rates than temperate assemblages.

The impact of seasonality in temperature on thermal tolerance and elevational range size.

Environmental temperature variation can influence physiology, biogeography, and life history, with large consequences for ecology, evolution, and the impacts of climate change. Based on the seasonality hypothesis, greater annual temperature variation at high latitudes should result in greater thermal tolerance and, consequently, larger elevational ranges in temperate compared to tropical species. Despite the mechanistic nature of this hypothesis, most research has used latitude as a proxy for seasonality, failing to directly examine the impact of temperature variation on physiology and range size. We used phylogenetically matched beetles from locations spanning 60 degrees of latitude to explore links between seasonality, physiology and elevational range. Thermal tolerance increased with seasonality across all beetle groups, but realized seasonality (temperature variation restricted to the months species are active) was a better predictor of thermal tolerance than was annual seasonality. Additionally, beetles with greater thermal tolerance had larger elevational ranges. Our results support a mechanistic framework linking variation in realized temperature to physiology and distributions.

On a collision course: competition and dispersal differences create no-analogue communities and cause extinctions during climate change.

Most climate change predictions omit species interactions and interspecific variation in dispersal. Here, we develop a model of multiple competing species along a warming climatic gradient that includes temperature-dependent competition, differences in niche breadth and interspecific differences in dispersal ability. Competition and dispersal differences decreased diversity and produced so-called 'no-analogue' communities, defined as a novel combination of species that does not currently co-occur. Climate change altered community richness the most when species had narrow niches, when mean community-wide dispersal rates were low and when species differed in dispersal abilities. With high interspecific dispersal variance, the best dispersers tracked climate change, out-competed slower dispersers and caused their extinction. Overall, competition slowed the advance of colonists into newly suitable habitats, creating lags in climate tracking. We predict that climate change will most threaten communities of species that have narrow niches (e.g. tropics), vary in dispersal (most communities) and compete strongly. Current forecasts probably underestimate climate change impacts on biodiversity by neglecting competition and dispersal differences.

Climate change and community disassembly: impacts of warming on tropical and temperate montane community structure.

Both tropical and temperate species are responding to global warming through range shifts, but our understanding of the consequences of these shifts for whole communities is limited. Here, we use current elevational range data for six taxonomic groups spanning 90° in latitude to examine the potential impacts of climate-driven range shifts on community change, or 'disassembly', across latitude. Elevational ranges are smaller at low latitudes for most groups and, as a consequence, tropical communities appear to be more sensitive to temperature increases compared with temperate communities. Under site-specific temperature projections, we generally found greater community disassembly in tropical compared with temperate communities, although this varied by dispersal assumptions. Mountain height can impact the amount of community disassembly, with greater change occurring on smaller mountains. Finally, projected community disassembly was higher for ectotherms than endotherms, although the variation among ectotherms was greater than the variation separating endotherms and ectotherms.

Impacts of climate warming on terrestrial ectotherms across latitude.

The impact of anthropogenic climate change on terrestrial organisms is often predicted to increase with latitude, in parallel with the rate of warming. Yet the biological impact of rising temperatures also depends on the physiological sensitivity of organisms to temperature change. We integrate empirical fitness curves describing the thermal tolerance of terrestrial insects from around the world with the projected geographic distribution of climate change for the next century to estimate the direct impact of warming on insect fitness across latitude. The results show that warming in the tropics, although relatively small in magnitude, is likely to have the most deleterious consequences because tropical insects are relatively sensitive to temperature change and are currently living very close to their optimal temperature. In contrast, species at higher latitudes have broader thermal tolerance and are living in climates that are currently cooler than their physiological optima, so that warming may even enhance their fitness. Available thermal tolerance data for several vertebrate taxa exhibit similar patterns, suggesting that these results are general for terrestrial ectotherms. Our analyses imply that, in the absence of ameliorating factors such as migration and adaptation, the greatest extinction risks from global warming may be in the tropics, where biological diversity is also greatest.