Non-climatic thermal adaptation: implications for species' responses to climate warming

There is considerable interest in understanding how ectothermic animals may physiologically and behaviourally buffer the effects of climate warming. Much less consideration is being given to how organisms might adapt to non-climatic heat sources in ways that could confound predictions for responses of species and communities to climate warming. Although adaptation to non-climatic heat sources (solar and geothermal) seems likely in some marine species, climate warming predictions for marine ectotherms are largely based on adaptation to climatically relevant heat sources (air or surface sea water temperature). Here, we show that non-climatic solar heating underlies thermal resistance adaptation in a rocky–eulittoral-fringe snail. Comparisons of the maximum temperatures of the air, the snail''s body and the rock substratum with solar irradiance and physiological performance show that the highest body temperature is primarily controlled by solar heating and re-radiation, and that the snail''s upper lethal temperature exceeds the highest climatically relevant regional air temperature by approximately 22°C. Non-climatic thermal adaptation probably features widely among marine and terrestrial ectotherms and because it could enable species to tolerate climatic rises in air temperature, it deserves more consideration in general and for inclusion into climate warming models.     

Variable effects of climate change on six species of North American birds

Many recent studies have shown that birds are advancing their laying date in response to long-term increases in spring temperatures. These studies have been conducted primarily in Europe and at local scales. If climate change is a large-scale phenomenon, then we should see responses at larger scales and in other regions. We examined the effects of long-term temperature change on the laying dates and clutch sizes of six ecologically diverse species of North American birds using 50 years of nest record data. As predicted, laying dates for most (four of six) species were earlier when spring temperatures were warmer. Over the long-term, laying dates advanced over time for two species (red-winged blackbirds, Agelaius phoeniceus and eastern bluebirds, Sialia sialis). Laying date of song sparrows (Melospiza melodia) also advanced with increasing temperature when the analysis was restricted to eastern populations. Neither laying date nor clutch sizes changed significantly over time in the remaining species (American coot, Fulica americana, killdeer, Charadrius vociferous, and American robin, Turdus migratorius), an unsurprising result given the lack of increase in temperatures over time at nest locations of these species. This study indicates that the relationship between climate change and breeding in birds is variable within and among species. In large-scale analyses of North American birds, four of seven species have shown advances in laying dates with increasing temperature (including song sparrows in the east). These variable responses within and among species highlight the need for more detailed studies across large spatial scales.     

Differential effects of past climate warming on mountain and flatland species distributions: a multispecies North American mammal assessment

Aim The magnitude of predicted range shifts during climate change is likely to be different for species living in mountainous environments compared with those living in flatland environments. The southern edges of ranges in mountain species may not shift northwards during warming as populations instead migrate up available elevational gradients; overall latitudinal range appears therefore to expand. In contrast, flatland species should shift range centroids northwards but not expand or contract their latitudinal range extent. These hypotheses were tested utilizing Late Pleistocene and modern occurrence data. Location North America. Methods The location and elevation of modern and Late Pleistocene species occurrences were collected from data bases for 26 species living in mountain or flatland environments. Regressions of elevation change over latitude, and southern and northern range edges were calculated for each species for modern and fossil data sets. A combination of regressions and anova s were used to test whether flatland species shift range edges and latitudinal extents more than mountain species do. Results Flatland species had significantly larger northward shifts at southern range edges than did mountain‐dwelling species from the Late Pleistocene to the present. There was also a significant negative correlation between the amount of change in the latitude of the southern edge of the range and the amount of elevational shifting from the Late Pleistocene to the present. Although significant, only c. 25% of the variance could be explained by this relationship. In addition, there was a weak indication that overall range expansion was less in flatland‐dwelling than in mountain‐dwelling species. Main conclusions The approach used here was to examine past species’ range responses to warming that occurred after the last ice ages as a means to better predict potential future responses to continued warming. The results confirm predictions of differential southern edge and overall range shifts for species occupying mountain and flatland regions in North America. The findings may be broadly applicable in other regions, thus allowing better modelling of future range and distribution related responses.     

Latitudinal variation of life-history traits of an exotic and a native impatiens species in Europe

Understanding the responses of invasive and native populations to environmental change is crucial for reliable predictions of invasions in the face of global change. While comparisons of responses across invasive species with different life histories have been performed before, comparing functional traits of congeneric native and invasive species may help to reveal driving factors associated with invasion. Here we compared morphological functional trait patterns of an invasive species (Impatiens parviflora) with its congeneric native species (I. noli-tangere) along an approximately 1600 km European latitudinal gradient from France (49°34′N) to Norway (63°40′N). Soil nitrogen was recorded during six weeks of the growing season, and light, soil moisture, and nutrient availability were estimated for each sampled population using community weighted means of indicator values for co-occurring species. Temperature data were gathered from nearby weather stations.Both the native and invasive species are taller at higher latitudes and this response is strongest in the invasive species. Seed mass and number of seeds per capsule increase in I. noli-tangere but decrease in I. parviflora towards higher latitudes. Surprisingly, plant height in the invasive I. parviflora decreases with increasing soil nitrogen availability. The latitudinal pattern in seed mass is positively related to temperature in I. noli-tangere and negatively in I. parviflora. Leaf area of both species decreases with increasing Ellenberg indicator values for nitrogen and light but increases with increasing soil moisture. Soil nitrogen concentrations and Ellenberg indicator values for nitrogen have significant positive (I. noli-tangere) and negative (I. parviflora) effects on the number of seeds per capsule. Our results show that the native I. noli-tangere has efficient reproduction at its range edge while the invasive I. parviflora shows a marked decrease in seed size and seed number per capsule. These patterns are unrelated to the growth and obtained size of the plants: even low soil nitrogen availability in the north seemed not to limit plant growth and size. Our results suggest that the invasive I. parviflora tends to become more invasive at lower latitudes by producing heavier seeds and more seeds per capsule.     

Latitudinal gradients in abundance, and the causes of rarity in the tropics: a test using Australian honeyeaters (Aves: Meliphagidae)

Several studies have uncovered interspecific latitudinal gradients in abundance (population density) such that tropical species tend to be, on average, less abundant than species at higher latitudes. The causes of this relationship remain poorly studied, in contrast to the relative wealth of literature examining the relationship to latitude of other variables such as range size and body mass. We used a cross-species phylogenetic comparative approach and a spatial approach to examine three potential determining factors (distribution, reproductive output and climate) that might explain why abundance correlates with latitude, using data from 54 species of honeyeaters (Meliphagidae) in woodland environments in eastern Australia. There is a strong positive correlation between mean abundance and latitude in these birds. Reproductive output (clutch size) was positively linked to both abundance and latitude, but partial correlation analysis revealed that clutch size is not related to abundance once the effects of latitude are removed. A subsequent multiple regression model that also considered range size, clutch size and body mass showed that latitude is the only strong predictor of abundance in honeyeaters. In the separate spatial analysis, the climatic variables that we considered (temperature, rainfall and seasonality) were all strongly linked to latitude, but none served as a better predictor of abundance than latitude per se, either individually or collectively. The most intriguing result of our analyses was that the cross-species latitudinal pattern in abundance was not evident within species. This suggests an intrinsic cause of the pattern of ‘rarity in the tropics’ in Australian honeyeaters. We suggest that evolutionary age may provide a key to understanding patterns of abundance in these birds.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Latitudinal variation in life-cycle characteristics of Potamogeton pectinatus L.: vegetative growth and asexual reproduction

Across latitudinal gradients, environmental conditions that influenceplant growth and reproduction largely change. Here we study clonal variation inlife-cycle characteristics of the cosmopolitan water plantPotamogetonpectinatus L. across a broad latitudinal range.Two consecutive experiments were performed under standardised laboratoryconditions (photoperiod, irradiance and temperature). In the first experimentweinvestigated asexual reproduction among fifteen clones, obtained from latitudesranging from 24 to 68° N. After 90 days of growth, high-latitude clonesproduced more but smaller tubers, while the aboveground biomass was lower ascompared to the clones obtained from low latitudes.In a second experiment we studied inherent differences in early growth,morphology and photosynthesis for eleven clones (obtained from the samelatitudinal range as in experiment 1). We found high among clonal variation formost measured variables, but the number of latitude-correlated traits waslimited. The only trait that correlated with latitude was the number of leavesper plant, which increased in clones from higher latitudes.Our results agree with the hypothesis of a latitude-correlated switch inlife-cycle strategy for this species. For northern clones this results in ashort life-cycle, with an early and high investment in tuber biomass, while forlow latitude clones the length of the life-cycle is prolonged, with a delayedreproduction and increased total plant biomass.     

Global richness patterns of venomous snakes reveal contrasting influences of ecology and history in two different clades

Recent studies addressing broad-scale species richness gradients have proposed two main primary drivers: contemporary climate and evolutionary processes (differential balance between speciation and extinction). Here, we analyze the global richness patterns of two venomous snake clades, Viperidae and Elapidae. We used ordinary least squares multiple regression (OLS) and partial regression analysis to investigate to what extent actual evapotranspiration (AET; summarizing current environmental conditions) and biogeographical regions (representing evolutionary effects) were associated with species richness. For viperids, AET explained 45.6% of the variance in richness whereas the effect of this variable for elapids was almost null (0.5%). On the other hand, biogeographic regions were the best predictors of elapid richness (56.5%), against its relatively small effect (25.9%) in viperid richness. Partial regressions also revealed similar patterns for independent effects of climate and history in both clades. However, the independent historical effect in Elapidae decreased from 45.2 to 17.8% when we excluded Australia from the analyses, indicating that the strong historical effect that had emerged for the global richness pattern was reflecting the historical process of elapid radiation into Australia. Even after excluding Australia, the historical signal in elapid richness in the rest of the globe was still significant and much higher than that observed in viperid richness at a global scale (2.7% after controlling for AET effects). Differences in the evolutionary age of these two clades can be invoked to explain these contrasting results, in that viperids probably had more time for diversification, generating richness responses to environmental gradients, whereas the pattern of distribution of elapid richness can be more directly interpreted in an evolutionary context. Moreover, these results show the importance of starting to adopt deconstructive approaches to species richness, since the driving factors of these patterns may vary from group to group according to their evolutionary history. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Implications of climate warming on seedling emergence and mortality of African savanna woody plants

Seedling emergence from scarified seeds and mortality of different seedling cohorts of five African savanna woody species (Acacia polyacantha, A. sieberana, Bauhinia thonningii, Dichrostachys cinerea and Ziziphus abyssinica) were studied under field conditions at a site in central Zambia. The study was conducted over a 4-year period, from 2003 to 2007. The objectives of the study were to determine climate factors that significantly influence seedling emergence rate and mortality in order to assess likely responses of the studied species to a warmer climate. Mean seedling emergence rate was 12% in D. cinerea and Z. abyssinica, 17% in B. thonningii, 47% in A. poyacantha and 62% in A. sieberana. Climate factors did not significantly affect seedling emergence in A. sieberana while temperature significantly influenced seedling emergence rate in the other species. Under a 1° warmer climate, seedling emergence rate was predicted to decline in A. polyacantha, B. thonningii and Z. abysssinica but is likely to increase slightly in D. cinerea. Time of seedling emergence during the wet season did not appear to affect seedling survival. Temperature also significantly influenced seedling mortality in all the studied species such that under a warmer climate, mortality was predicted to increase in A. sieberana and D. cinerea but decrease in A. polyacantha, B. thonningii and Z. abyssinica. As the studied species exhibited differential optimum temperature conditions for seedling emergence and seedling survival, they are likely to respond to climate warming in different but predictable ways. The results of the study are useful to forest management and development of climate change adaptation strategies in southern Africa.     

Latitudinal variation in the fruiting phenology of a bird-dispersed plant (shape Crataegus monogyna) in Western Europe

A number of fleshy-fruit-bearing plants of temperate regions are dispersed by migratory frugivorous birds. It has been suggested that the more southerly populations of such species should produce ripe fruits later than more northerly populations, to ensure that fruit is available when the birds arrive. I will call this the adaptive delay hypothesis. To test this hypothesis, I monitored fruiting phenology of Crataegus monogyna Jacq. at ten sites (in all of which fruit consumption is very largely by redwings, Turdus iliacus, and fieldfares, Turdus pilaris) between northern Spain (42° N) and northern Scotland (59° N). There was no negative correlation between latitude and date of fruit ripening (earliest recorded date on which ripe fruits had appeared, or earliest recorded date by which all fruits had ripened). My results thus argue against the adaptive delay hypothesis.     

The elevation history of the Tibetan Plateau and its implications for the Asian monsoon

The determination of the evolving palaeoaltitude of the Tibetan Plateau since the India-Eurasia collision underpins our understanding of how orography in central Asia affects the intensity of the monsoon and hence global climate change. Palaeoaltitudes, however, cannot be measured directly and need to be inferred from proxy observations that are usually model-dependent. Differing tectonic models for the behaviour of the lithosphere during continental collision have contrasting implications for the elevation of the plateau. However, two techniques recently employed for determining palaeo-elevation are independent of tectonic models, the first involving the variation with altitude of oxygen isotopes in precipitation and the second involving the change of leaf morphology with moist static energy of the atmosphere.Elevation studies have focused on southern Tibet, largely due to the relative ease of access to the region. There is a remarkable unanimity amongst the diverse techniques applied that the altitude of the southern plateau has not significantly changed since at least the mid Miocene (ca. 15 Ma) arguing for an onset of the monsoon system during or before the early Miocene. A range of tectonic studies suggest that the northern and eastern parts of the plateau are younger geomorphological features, but there are few quantitative constraints of the timing of elevation from these regions of Tibet. Since both the elevation and the surface area of the plateau impact on atmospheric circulation, palaeoaltitude studies need to be extended to chart the increasing areas of elevated land surface through time.     

Gravity and the evolution of cardiopulmonary morphology in snakes

Physiological investigations of snakes have established the importance of heart position and pulmonary structure in contexts of gravity effects on blood circulation. Here we investigate morphological correlates of cardiopulmonary physiology in contexts related to ecology, behavior and evolution. We analyze data for heart position and length of vascular lung in 154 species of snakes that exhibit a broad range of characteristic behaviors and habitat associations. We construct a composite phylogeny for these species, and we codify gravitational stress according to species habitat and behavior. We use conventional regression and phylogenetically independent contrasts to evaluate whether trait diversity is correlated with gravitational habitat related to evolutionary transitions within the composite tree topology. We demonstrate that snake species living in arboreal habitats, or which express strongly climbing behaviors, possess relatively short blood columns between the heart and the head, as well as relatively short vascular lungs, compared to terrestrial species. Aquatic species, which experience little or no gravity stress in water, show the reverse — significantly longer heart-head distance and longer vascular lungs. These phylogenetic differences complement the results of physiological studies and are reflected in multiple habitat transitions during the evolutionary histories of these snake lineages, providing strong evidence that heart-to-head distance and length of vascular lung are co-adaptive cardiopulmonary features of snakes.     

Parental overwintering history affects the responses of Thlaspi arvense to warming winters in the North

The overwintering conditions of northern plants are expected to change substantially due to global warming. For perennial plants, winter warming may increase the risk of frost damage if the plants start dehardening prematurely. On the other hand, evergreen plants may remain photosynthetically active and thereby benefit from milder winters. The positive and negative effects of mild winters on annual plants remain, however, largely unknown. We postulated that summer annuals may be susceptible to frost damage if the seeds germinate during a mild spell in winter. Winter annuals may utilize a warm period for photosynthesis. These questions were addressed in two consecutive experiments in which pot-grown individuals of Thlaspi arvense that overwintered in the field were exposed to an elevated temperature for 8 days in growth chambers in mid-winter. No premature germination was observed in summer annuals. However, in accordance with our hypothesis, winter annuals started photosynthesising very rapidly upon exposure to elevated temperature. The winter warming treatment affected neither the total number of seeds produced nor the mean seed weight. These seeds, possessing divergent parental overwintering histories, were used as starting material for the second experiment. Seeds originating from both summer and winter annual plants germinated both in the autumn and in the following spring. We observed a major parental effect associated with the winter warming treatment. The warm spell experienced by the mother plant (either as a winter annual rosette or as a summer annual seed) reduced the proportion of autumn germination in the next generation. Only 43% of the seeds of summer annuals possessing a parental warming history germinated before the winter, whereas the germination percentage of seeds with no previous winter warming history was 71%. In the case of seeds collected from winter annual plants, 4% of the seeds germinated in autumn if the mother plants experienced the warming treatment during the previous winter, whereas the corresponding value was 37% if the mother plants did not experience warming. Our results show that summer and winter annual individuals show diverse responses to warm spells in winter. Since the responses are not limited only to the generation that actually experiences the warm spell, but also appear in their offspring, long-term studies consisting of several generations are called for.     

Implications of climate change for North American wood warblers (Parulidae)

Since 1912, Neotropical–Nearctic migrant birds may now have up to 20 fewer days to travel between Southern Illinois, where spring is arriving later, and Northern Minnesota, where spring is arriving earlier, to exploit optimal habitat conditions (expanding leaves and caterpillar activity) for refueling and breeding. As case studies of the effect of climate change on bird migration, I analyzed two long‐term data sets of arrival times for eight species of northern breeding migratory wood warblers (Parulidae) gathered over a 100 year period in east‐central Illinois (IL, USA) and a 40 year period in western Minnesota (MN, USA). Six (IL) and seven (MN) of the wood warbler species showed no significant tendency to migrate earlier in response to earlier springs in their breeding range. These results suggest that climate change may force many species of long‐distance migratory songbirds to become uncoupled in the spring from their food resources that are driven by temperature.     

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.     

气候变暖对长白山主要树种的潜在影响

应用LINKAGES模型对长白山自然保护区内主要树种在各斑块类型中对气候变化的潜在响应进行了模拟,模拟时选择了目前和未来变暖2种气候条件,对于目前气候状态,模型使用目前气象参数;而对于未来变暖气候,则按温度增加5℃,降水无明显变化作为模拟假设,温度的增加假定各月都相同,即各月均增加5度,模拟结果表明,对于高山岳桦林,气温变暖后岳桦依然扮演重要角色,但落叶松,云极,冷杉等目前这一林带的伴生树种,在气温上升后,其生物量均有较大辐度的增加,部分占据目前岳桦的位置,即目前下部的云冷杉林带有上移的趋势;对于亚高山云冷杉林,其优势种云杉和冷杉在气温变暖后,生物量有较大幅度的增加,落叶松虽有增加的趋势但幅度较小,即云杉和冷杉在未来气温变暖后依然是这一林带的优势种,但生长会加快,阔叶红松林的主要建群种在气温升高后,其生物量只有较小的增加,其它主要伴生种的生物量随气温上升的增加趋势非常相似,表明阔叶红松林在未来气候变暖情况下仍将维持目前的结构状态。     

Empirical methodology for predicting changes in species range extension and richness associated with climate warming

Species richness and extent of geographical range for North American freshwater amphipods and crayfish are strongly correlated with mean annual absolute, or range of, temperature, respectively. Such spatially based relationships are shown to be useful for predicting the biological consequences arising through scenarios of temporal climate warming.     

Projected effects of climate change on tick phenology and fitness of pathogens transmitted by the North American tick Ixodes scapularis

Ixodes scapularis is the principal tick vector of the Lyme borreliosis agent Borrelia burgdorferi and other tick-borne zoonoses in northeastern North America. The degree of seasonal synchrony of nymphal and larval ticks may be important in influencing the basic reproductive number of the pathogens transmitted by I. scapularis. Because the seasonal phenology of tick vectors is partly controlled by ambient temperature, climate and climate change could shape the population biology of tick-borne pathogens. We used projected monthly normal temperatures, obtained from the second version of the Canadian Coupled Global Climate Model (CGCM2) under emissions scenario A2 of the Intergovernmental Panel on Climate Change for a site in southern Ontario, Canada, to simulate the phenology of I. scapularis in a mathematical model. The simulated seasonal abundance of ticks then determined transmission of three candidate pathogens amongst a population of white-footed mice (Peromyscus leucopus) using a susceptible-infected-recovered (SIR) model. Fitness of the different pathogens, in terms of resilience to changes in tick and rodent mortality, minima for infection duration, transmission efficiency and particularly any additional mortality of rodents specifically associated with infection, varied according to the seasonal pattern of immature tick activity, which was different under the temperature conditions projected for the 2020s, 2050s and 2080s. In each case, pathogens that were long-lived, highly transmissible and had little impact on rodent mortality rates were the fittest. However, under the seasonal tick activity patterns projected for the 2020s and 2050s, the fitness of pathogens that are shorter-lived, less efficiently transmitted, and more pathogenic to their natural hosts, increased. Therefore, climate change may affect the frequency and distribution of I. scapularis-borne pathogens and alter their evolutionary trajectories.     

Geographic variation in body size: the effects of ambient temperature and precipitation

Latitudinal trends in body size have been explained as a response to temperature- or water-related factors, which are predictors of primary production. We used the first principal component calculated from three body parameters (weight, body length and the greatest length of the skull) of a sample of mammals from Israel and Sinai to determine those species that vary in size geographically, and whether such variation is related to annual rainfall, average minimum January temperature and average maximum August temperature. We used a conservative approach to discern the effects of precipitation and temperature by applying sequential regression. Variable priorities were assigned according to their bivariate correlation with body size, except for rainfall and its interactions that entered into the model last. Eleven species (Acomys cahirinus, Apodemus mystacinus, Canis lupus, Crocidura suaveolens, Gerbillus dasyurus, Hyaena hyaena, Lepus capensis, Meles meles, Meriones tristrami, Rousettus aegyptius and Vulpes vulpes) of the 17 species examined varied in size geographically. In five of them, rainfall was positively related to body size, while in one species it was negatively related to it. Contrary to the prediction of Bergmann’s rule, mean minimum January temperature was positively related to body size in five species and negatively related to body size in two species (C. suaveolens and G. dasyurus). As predicted by Bergmann’s rule, maximum June temperature was negatively related to body size in three species, and positively so in one (L. capensis). Primary production, particularly in desert and semi-desert areas, is determined mainly by precipitation. The above results indicate that, in our sample, primary production has an important effect on body size of several species of mammals. This is evident from the considerable proportion of the variability in body size explained by rain. However, low ambient temperatures may slow down and even inhibit photosynthesis. Hence, the observed positive relationships between average minimum January temperature and body size in four of the six species influenced by rain further support this conclusion.     

Potential impact of climate warming on the distribution of the Golden-striped salamander, Chioglossa lusitanica, on the Iberian Peninsula

The increase in the concentration of greenhouse gases in the atmosphere is expected to impact the world's climate on a time-scale of just decades. We simulated the potential impact of climate warming on the range of the Iberian endemic Golden-striped salamander, Chioglossa lusitanica, by extrapolating present-day GIS-based distribution models. The results indicated a significant decrease in the distribution of the species for 2050 and 2080, with losses ranging from 19 to 35% in Portugal and from 17 to 22% in Spain. The models also predict the fragmentation of the species range, in a pattern mirroring past distributions inferred from the spatial analysis of genetic data.