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1.
Mountain butterflies have evolved efficient thermoregulation strategies enabling their survival in marginal conditions with short flight season and unstable weather. Understanding the importance of their behavioural thermoregulation by habitat use can provide novel information for predicting the fate of alpine Lepidoptera and other insects under ongoing climate change. We studied the link between microhabitat use and thermoregulation in adults of seven species of a butterfly genus Erebia co-occurring in the Austrian Alps. We captured individuals in the field and measured their body temperature in relation to microhabitat and air temperature. We asked whether closely related species regulate their body temperature differently, and if so, what is the effect of behaviour, species traits and individual traits on body to air and body to microhabitat temperature differences. Co-occurring species differed in mean body temperature. These differences were driven by active microhabitat selection by individuals and also by species–specific habitat preferences. Species inhabiting grasslands and rocks utilised warmer microclimates to maintain higher body temperature than woodland species. Under low air temperatures, species of rocky habitats heated up more effectively than species of grasslands and woodlands which allowed them to stay active in colder weather. Species morphology and individual traits play rather minor roles in the thermoregulatory differences; although large species and young individuals maintained higher body temperature. We conclude that diverse microhabitat conditions at small spatial scales probably contribute to sympatric occurrence of closely related species with different thermal demands and that preserving heterogeneous conditions in alpine landscapes might mitigate detrimental consequences of predicted climate change.  相似文献   

2.
Populations at the warm range margins of the species distribution may be at the greatest risks of extinction from global warming unless they can tolerate extreme environmental conditions. Yet, some studies suggest that the thermal behavior of some lizard species is evolutionarily rigid. During two successive years, we compared the thermal biology of two populations of Liolaemus pictus living at the northern (warmer) and one population living at the southern (colder) range limits, thus spanning an 800 km latitudinal distance. Populations at the two range margins belong to two deeply divergent evolutionary clades. We quantified field body temperatures (Tb), laboratory preferred body temperatures (PBT), and used operative temperature data (Te) to calculate the effectiveness of thermoregulation (E). During one year in all populations, we further exposed half of the lizards to a cold or a hot acclimation treatment to test for plasticity in the thermal behavior. The environment at the southern range limit was characterized by cooler weather and lower Te. Despite that, females had higher Tb and both males and females had higher PBT in the southernmost population (or clade) than in the northernmost populations. Acclimation to cold conditions led to higher PBT in all populations suggesting that plastic responses to thermal conditions, instead of evolutionary history, may contribute to geographic variation. Lizards regulated moderately well their body temperature (E≈0.7): they avoided warm microhabitats in the northern range but capitalized on warm microhabitats in the southern range. We review literature data to show that Liolaemus species increase their thermoregulation efficiency in thermally challenging environments. Altogether, this indicates that habitats of low thermal quality generally select against thermoconformity in these lizards.  相似文献   

3.
Reptiles that live in cooler environments hibernate longer and, when active, limit daily activity times, allocate more time and energy toward thermoregulation, and consequently experience life-history constraints such as reduced fecundity and supra-annual reproductive cycles. This pattern becomes more extreme with increasing latitude and altitude. We compared the thermal biology of two populations of Liolaemus pictus argentinus living at two altitudes (771 and ∼1700 m asl). Environmental, microenvironmental, and operative temperatures were studied in order to describe the capture sites, sources of heat, and availability of microenvironments appropriate for thermoregulation. The body temperatures of L. p. argentinus at capture (Tb) and the preferred temperatures in the laboratory (Tp) were recorded and integrated with operative temperatures to calculate the effectiveness of thermoregulation. The high-altitude population was found to have a lower mean Tb (29 °C compared to 33 °C), while the Tp values for both populations were similar (36.7 °C). The analysis of operative temperatures and Tb in relation to Tp showed that L. p. argentinus behaves as a moderate thermoregulator at high altitude and as a poor thermoregulator at the low-altitude site probably due in part to the avoidance of predation risk.  相似文献   

4.
Behavioral thermoregulation is expected to be critical in determining the capacity of reptiles to respond to climate warming and how that response will vary with latitude. We used radio-telemetry to compare behavioral thermoregulation among ratsnake (Elaphe obsoleta) populations in Texas, Illinois, and Ontario, a latitudinal distance of >1500 km. Despite numerous specific differences among populations, overall the thermal ecology was surprisingly similar during the months that snakes in all three populations were active. Preferred temperatures varied only slightly across the snakes’ range, the extent of thermoregulation was similar, and by varying when during the day and season they thermoregulated, snakes in all three populations realized body temperatures within their preferred temperature range 15–20% of the time. The ability to use fine-scale behavioral thermoregulation (i.e., selective use of habitats and microclimates) to a similar extent and achieve similar outcomes across such a wide latitudinal and climatic gradient is made possible by large-scale differences in timing of activity (ratsnakes in Texas switch to nocturnal activity during summer, whereas in Illinois and Ontario activity is exclusively diurnal and hibernation lasts 5–7 months). Modeling indicated that a 3 °C increase in ambient temperature will generally improve thermal conditions for all three populations. Our empirical analyses suggest that the snakes’ ability to respond to climate warming will be determined more by their capacity to adjust when they are active than by changes in the extent of fine-scale behavioral thermoregulation. The ability to adjust timing of activity appears to make many snakes fundamentally different from lizards. As such, the consequences of climate warming may be very different for these two groups of reptiles.  相似文献   

5.
Summary We document activity field temperatures, daily activity patterns, and extent of thermoregulation in four species of Liolaemus lizards inhabiting at high altitude (above 3500 m) in the Andes of northern Chile. These four species have similar activity field temperature (Tb near 29°C) despite their being distributed at different altitudinal belts. However, conspicuous differences exist between higher-altitude (L. alticolor and L. jamesi) and lower-altitude (L. islugensis and L. ornatus) lizards regarding extent of thermoregulation and activity period. Some differences in morphology, behavior, and patterns of microhabitat occupancy are also apparent among these four species and are seemingly related to the thermal environment to which they are subjected. In comparison to eight low-altitude Liolaemus species in central Chile (Tb near 35°C) the four high-altitude species in northern Chile have lower activity field temperature. The latter is apparently due to the constraints imposed by the harsh Andean thermal environment, a hypothesis supported by the fact that high-altitude Liolaemus lizards under laboratory conditions demonstrate body temperatures that exceed by 5°C or more, those recorded in the field.  相似文献   

6.
Increasing evidence suggests that global warming significantly alters the range and phenology of plants and animals. Whereas thermophilous species usually benefit from rising temperatures, the living conditions of taxa adapted to cooler or continental climates are deteriorating. The woodland ringlet butterfly, Erebia medusa Fabricius (Lepidoptera: Nymphalidae) is one of the continental species that are supposed to be adversely affected by climate change, especially by rising winter temperatures. Here, we conduct an explorative study on the effects of low, moderate, and high winter temperatures on the pre‐adult and adult stages of E. medusa in a laboratory experiment. Compared to the two other temperature regimes, the warm winter treatment led to an earlier termination of diapause and higher larval weights at the end of the winter, but significantly lower survival rates. The after‐effects of the warm treatment included lower weight of the pupae and adult females, shorter forewings of adult males, and earlier emergence of both adult males and females. In natural environments, which are characterized by a much greater thermal variability and a much higher frequency of soil freeze‐thaw events compared to our experiment, the effects of rising winter temperatures might be stronger than in this study. Thus, we conclude that warmer winters pose a non‐negligible long‐term threat to E. medusa.  相似文献   

7.
Differences in body size are widely thought to allow closely related species to coexist in sympatry, but body size also varies as an adaptive response to climate. Here, we use a sister lineage approach to test the prediction that body size differences between closely related species of birds worldwide are greater for species whose ranges are sympatric rather than allopatric. We further test if body size differences among sympatric versus allopatric species vary with geography, evolutionary distance, and environmental temperatures. We find greater differences in size among sympatric compared with allopatric lineages, but only in closely related species that live where mean annual temperatures are above 25°C. These size differences in warm environments declined with the evolutionary distance between sister lineages. In species living in cooler regions, closely related allopatric and sympatric species did not differ significantly in size, suggesting either that colder temperatures constrain the evolutionary divergence of size in sympatry, or that the biotic selective pressures favoring size differences in sympatry are weaker in colder environments. Our results are consistent with suggestions by Wallace, Darwin, and Dobzhansky that climatic selective pressures are more important in cooler environments (e.g., high elevations and latitudes) whereas biotic selective pressures dominate in warm environments (e.g., lowland tropics).  相似文献   

8.
The habitat associations of individuals underpin the dynamics of species distributions. Broad‐scale gradients in climate can alter habitat associations across species’ geographic ranges, but topographic heterogeneity creates local microclimates which could generate variation in habitat use at finer spatial scales. We examined the selection of microhabitats for egg‐laying by populations of a thermally‐constrained butterfly, the skipper Hesperia comma, across 16 sites with different regional temperatures and topographic microclimates. Using models of thermal microclimate, we examined how the association between eggs and warm bare ground microhabitats varied with ambient temperature, and predicted bare ground associations in 287 existing H. comma populations, to investigate the relative impacts of regional temperatures and topographic microclimates on microhabitat use. Eggs were most strongly associated with bare ground in relatively cool sites, indicating climate‐driven changes in microhabitat use. The majority of temperature variation between study sites was attributable to topographic microclimates rather than regional temperature differences, such that changes in microhabitat associations occurred principally between north‐ and south‐facing slopes within the same region. Predicted microhabitat associations across the UK distribution of H. comma showed that, due to the large temperature differences generated by topography, most of the between‐population variation in microhabitat use occurs locally within 5 km grid squares, with a smaller proportion occurring at a regional level between 5 km squares. Our findings show how microclimatic variation generated by topography alters the habitat associations of populations at fine spatial scales, suggesting that microclimate‐driven changes in habitat suitability could shape species’ distribution dynamics and their responses to environmental change.  相似文献   

9.
Erebia aethiops is an aberrant lowland representative of a predominately arctic and alpine butterfly genus. We used behavioral records obtained during a mark-recapture study (MR) to extract information about its adult diurnal and within-season activity, and to compare its activity with previously studied mountain Erebia species. Similarly to them, maintenance activities (nectaring, basking) increased with season. Contrary to mountain relatives, E. aethiops nectars in morning hours and spends the hottest parts of the day in the shade. Therefore, high rather than low temperatures seem to be limiting E. aethiops activity. We also compared microhabitat preferences of sexes, finding that females preferred grassy patches, whereas males often entered scrub and sparse canopy forest. The study illustrates the utility of auxiliary MR data to analyze butterfly activity, and points to the necessity of finely structured habitat mosaics for E. aethiops, an open woodland species that retracted to abandoned grasslands and has become vulnerable in Central Europe.  相似文献   

10.
Biological impacts of climate warming are predicted to increase with latitude, paralleling increases in warming. However, the magnitude of impacts depends not only on the degree of warming but also on the number of species at risk, their physiological sensitivity to warming and their options for behavioural and physiological compensation. Lizards are useful for evaluating risks of warming because their thermal biology is well studied. We conducted macrophysiological analyses of diurnal lizards from diverse latitudes plus focal species analyses of Puerto Rican Anolis and Sphaerodactyus. Although tropical lowland lizards live in environments that are warm all year, macrophysiological analyses indicate that some tropical lineages (thermoconformers that live in forests) are active at low body temperature and are intolerant of warm temperatures. Focal species analyses show that some tropical forest lizards were already experiencing stressful body temperatures in summer when studied several decades ago. Simulations suggest that warming will not only further depress their physiological performance in summer, but will also enable warm-adapted, open-habitat competitors and predators to invade forests. Forest lizards are key components of tropical ecosystems, but appear vulnerable to the cascading physiological and ecological effects of climate warming, even though rates of tropical warming may be relatively low.  相似文献   

11.
W. U. Blanckenhorn 《Oecologia》1997,109(3):342-352
 Field phenologies of high- (ca. 1500 m) and low- (ca. 500 m) altitude populations of the two most common European species of dung flies, Scathophaga stercoraria and Sepsis cynipsea, differ quite markedly due to differences in climate. To differentiate genetic adaptation due to natural selection and phenotypic plasticity, I compared standard life history characters of pairs of high- and low-altitude populations from three disjunctive sites in Switzerland in a laboratory experiment. The F1 rearing environment did not affect any of the variables of the F2 generation with which all experiments were conducted; hence, there were no carry-over or maternal effects. In Sc. stercoraria, high-altitude individuals were smaller but laid larger eggs; the latter may be advantageous in the more extreme (i.e. more variable and less predictable) high-altitude climate. Higher rearing temperature strongly decreased development time, body size and the size difference between males and females (males are larger), produced female-biased sex ratios and led to suboptimal adult emergence rates. Several of these variables also varied among the three sites, producing some interactions complicating the patterns. In Se. cynipsea, high-altitude females were marginally smaller, less long-lived and laid fewer clutches. Higher rearing temperature strongly decreased development time and body size but tended to increase the size difference between males and females (males are smaller); it also increased clutch size but decreased physiological longevity. Again, interpretation is complicated by variation across sites and some significant interactions. Overall, genetic adaptation to high-altitude conditions appears weak, probably prevented by substantial gene flow, and may be swamped by the effects of other geographic variables among populations. In contrast, phenotypic plasticity is extensive. This may be due to selection of flexible, multi-purpose genotypes. The results suggest that differences in season length between high- and low-altitude locations alone do not explain well the patterns of variation in phenology and body size. Received: 21 March 1996 / Accepted: 1 September 1996  相似文献   

12.
The geographic variation of external characters of the satyrid Erebia euryale (Esper, [1805]), distributed in the north of European Russia, was analyzed. One subspecies, E. euryale euryaloides Tengström, 1869, was shown to inhabit the northern Russian Plain and the Urals. The subspecies is characterized by a gradual decrease in body size and reduction of the wing pattern in adults towards the north and a distinct geographic segregation of the plain and mountain populations into two races by the color of the postdiscal band in females. The plain populations, in which females have silver-white bands, are referred to as the “taiga” race, and the Ural populations with golden-yellow bands, as the “flaveoides” race, according to the names of the subspecific taxa previously described. A probable historical scenario of the distribution of E. euryale and the origin of geographic isolation of its races are considered.  相似文献   

13.
Responses to climate change are particularly complicated in species that engage in symbioses, as the niche of one partner may be modified by that of the other. We explored thermal traits in gut symbionts of honeybees and bumblebees, which are vulnerable to rising temperatures. In vitro assays of symbiont strains isolated from 16 host species revealed variation in thermal niches. Strains from bumblebees tended to be less heat-tolerant than those from honeybees, possibly due to bumblebees maintaining cooler nests or inhabiting cooler climates. Overall, however, bee symbionts grew at temperatures up to 44°C and withstood temperatures up to 52°C, at or above the upper thermal limits of their hosts. While heat-tolerant, most strains of the symbiont Snodgrassella grew relatively slowly below 35°C, perhaps because of adaptation to the elevated body temperatures that bees maintain through thermoregulation. In a gnotobiotic bumblebee experiment, Snodgrassella was unable to consistently colonize bees reared at 29°C under conditions that limit thermoregulation. Thus, host thermoregulatory behaviour appears important in creating a warm microenvironment for symbiont establishment. Bee–microbiome–temperature interactions could affect host health and pollination services, and inform research on the thermal biology of other specialized gut symbionts.  相似文献   

14.
Rhododendron przewalskii is an important dwarf shrub species in alpine environments of western Sichuan that offers a unique opportunity to expand current dendrochronological networks into extreme environments beyond the survival limit of trees. Our objectives in this study are to evaluate the dendroclimatological potentials of R. przewalskii and determine the major limiting climate factor for the species’ growth rings. We sampled 25 cross-sections of R. przewalskii at an elevation of 4050 m, about 150 m above treeline, on Zhegu Mountain of Miyaluo in western Sichuan of southeastern Tibetan Plateau. R. przewalskii has well-defined growth rings such that most stem sections could be cross-dated. The resulting 61-year long standard chronology (A.D. 1949–2009) was derived from 38 series from 19 cross-sections. Response analysis revealed that radial growth of the Zhegu Mountain R. przewalskii is significantly and negatively correlated to late winter temperature (January–February), mainly driven by maximum temperatures. This correlation indicates that colder daytime temperatures during late-winter lead to improved growth the following growing season. Minimum winter temperatures do not appear important for radial growth of this population. R. przewalskii ring widths are also strongly correlated to late-winter maximum temperatures over the southeastern Tibetan Plateau region. April maximum and July minimum temperatures are positively and significantly correlated to radial growth suggesting that warm April days and warm July nights promote current-season radial growth. In contrast, radial growth is only negatively correlated to April precipitation, indicating that wet soil conditions inhibit total radial increment. The main differences of R. przewalskii compared to tree species living at high-altitude nearby regions is that R. przewalskii has a weaker positive growth response to summer temperature and winter minimum temperature, but a much stronger negative response to winter temperature. Due to the strong climatic signal recorded in the growth curves of R. przewalskii, this dwarf shrub should be useful for climate–growth studies in alpine regions where no forests are present.  相似文献   

15.
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.  相似文献   

16.
Respiration for maintenance and growth ofReynoutria japonica ecotypes from altitudes of 700 and 2420 m on Mt Fuji were measured in two controlled thermal conditions. The maintenance respiration of the high-altitude ecotype at both 15 and 25°C was significantly (1.7-fold) higher than that of the low-altitude ecotype, whereas growth respiration was independent of both ecotype and temperature. The temperature coefficient (Q10) of the maintenance respiration was about 1.9 in both ecotypes. The results show that there is ecotypic differentiation in the performance of maintenance respiration. It is suggested that the high maintenance respiration of the high-altitudeR. japonica ecotype is advantageous in severe upland environments but disadvantageous in a warm lowland climate in terms of carbon economy.  相似文献   

17.
Many ectotherms effectively reduce their exposure to low or high environmental temperatures using behavioral thermoregulation. In terrestrial ectotherms, thermoregulatory strategies range from accurate thermoregulation to thermoconformity according to the costs and limits of thermoregulation, while in aquatic taxa the quantification of behavioral thermoregulation have received limited attention. We examined thermoregulation in two sympatric newt species, Ichthyosaura alpestris and Lissotriton vulgaris, exposed to elevated water temperatures under semi-natural conditions. According to a recent theory, we predicted that species for which elevated water temperatures pose a lower thermal quality habitat, would thermoregulate more effectively than species in thermally benign conditions. In the laboratory thermal gradient, L. vulgaris maintained higher body temperatures than I. alpestris. Semi-natural thermal conditions provided better thermal quality of habitat for L. vulgaris than for I. alpestris. Thermoregulatory indices indicated that I. alpestris actively thermoregulated its body temperature, whereas L. vulgaris remained passive to the thermal heterogeneity of aquatic environment. In the face of elevated water temperatures, sympatric newt species employed disparate thermoregulatory strategies according to the species-specific quality of the thermal habitat. Both strategies reduced newt exposure to suboptimal water temperatures with the same accuracy but with or without the costs of thermoregulation. The quantification of behavioral thermoregulation proves to be an important conceptual and methodological tool for thermal ecology studies not only in terrestrial but also in aquatic ectotherms.  相似文献   

18.
Climate warming is likely to shift the range margins of species poleward, but fine‐scale temperature differences near the ground (microclimates) may modify these range shifts. For example, cold‐adapted species may survive in microrefugia when the climate gets warmer. However, it is still largely unknown to what extent cold microclimates govern the local persistence of populations at their warm range margin. We located 99 microrefugia, defined as sites with edge populations of 12 widespread boreal forest understory species (vascular plants, mosses, liverworts and lichens) in an area of ca. 24,000 km2 along the species' southern range margin in central Sweden. Within each population, a logger measured temperature eight times per day during one full year. Using univariate and multivariate analyses, we examined the differences of the populations' microclimates with the mean and range of microclimates in the landscape, and identified the typical climate, vegetation and topographic features of these habitats. Comparison sites were drawn from another logger data set (n = 110), and from high‐resolution microclimate maps. The microrefugia were mainly places characterized by lower summer and autumn maximum temperatures, late snow melt dates and high climate stability. Microrefugia also had higher forest basal area and lower solar radiation in spring and autumn than the landscape average. Although there were common trends across northern species in how microrefugia differed from the landscape average, there were also interspecific differences and some species contributed more than others to the overall results. Our findings provide biologically meaningful criteria to locate and spatially predict potential climate microrefugia in the boreal forest. This opens up the opportunity to protect valuable sites, and adapt forest management, for example, by keeping old‐growth forests at topographically shaded sites. These measures may help to mitigate the loss of genetic and species diversity caused by rear‐edge contractions in a warmer climate.  相似文献   

19.
Summary Seedlings representative of Verbascum thapsus L. populations from thermally diverse habitats were grown under uniform, controlled conditions. The plants were used to obtain temperature response curves for net photosynthesis over a range of 15–40°C. In general, all experimental plants exhibited similar rates of net photosynthesis at 20, 25, 30, and 35°C. Plants representative of cool habitat populations (high-latidude and high-altitude) had greatest rates of net photosynthesis at the lower temperatures and much lower rates at 40°C. Plants representative of warm habitat populations (low-latitude and low-altitude) exhibited rates of net photosynthesis at 40°C which were nearly twice those of plants representative of cool habitat populations. Carbon dioxide transfer resistances are discussed with reference to plant control of photosynthesis at different temperatures. Patterns of photosynthesis and resistance response among plants representative of different habitats suggest ecotypic variation has occurred only to a very limited extent. Therefore, the patterns exhibited by experimental plants suggest that Verbascum thapsus' success in a number of diverse sites is related to the ability of all members of the species to photosynthesize over a broad range of temperatures.  相似文献   

20.
The consequences of climate change for biogeographic range dynamics depend on the spatial scales at which climate influences focal species directly and indirectly via biotic interactions. An overlooked question concerns the extent to which microclimates modify specialist biotic interactions, with emergent properties for communities and range dynamics. Here, we use an in-field experiment to assess egg-laying behaviour of a range-expanding herbivore across a range of natural microclimatic conditions. We show that variation in microclimate, resource condition and individual fecundity can generate differences in egg-laying rates of almost two orders of magnitude in an exemplar species, the brown argus butterfly (Aricia agestis). This within-site variation in fecundity dwarfs variation resulting from differences in average ambient temperatures among populations. Although higher temperatures did not reduce female selection for host plants in good condition, the thermal sensitivities of egg-laying behaviours have the potential to accelerate climate-driven range expansion by increasing egg-laying encounters with novel hosts in increasingly suitable microclimates. Understanding the sensitivity of specialist biotic interactions to microclimatic variation is, therefore, critical to predict the outcomes of climate change across species'' geographical ranges, and the resilience of ecological communities.  相似文献   

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