Effects of constant and fluctuating incubation temperatures on hatching success and hatchling traits in the diamondback terrapin (Malaclemys terrapin) in the context of the warming climate

As global temperatures continue to rise, so too will the nest temperatures of many species of turtles. Yet for most turtle species, including the estuarine diamondback terrapin (Malaclemys terrapin), there is limited information on embryonic sensitivity to elevated temperature. We incubated eggs of M. terrapin at three, mean temperatures (31, 34, 37 °C) under two thermal exposure regimes (constant or semi-naturally fluctuating temperature) and measured hatching success, developmental rate, and hatchling size. Hatching success was 100% at 31 °C and 67% at 34 °C, respectively; at 37 °C, all eggs failed early in the incubation period. These values were unaffected by exposure regime. The modeled LT₅₀ (temperature that was lethal to 50% of the test population) was 34.0 °C in the constant and 34.2 °C in the fluctuating thermal regime, reflecting a steep decline in survival between 33 and 35 °C. Hatchlings having been incubated at a constant 34 °C hatched sooner than those incubated at 31 °C under either constant or fluctuating temperature. Hatchlings were smaller in straight carapace length (CL) and width after having been incubated at 34 °C compared to 31 °C. Larger (CL) hatchlings resulted from fluctuating temperature conditions relative to constant temperature conditions, regardless of mean temperature. Based upon recent temperatures in natural nests, the M. terrapin population studied here appears to possess resiliency to several degrees of elevated mean nest temperatures, beyond which, embryonic mortality will likely sharply increase. When considered within the mosaic of challenges that Maryland's M. terrapin face as the climate warms, including ongoing habitat losses due to sea level rise and impending thermal impacts on bioenergetics and offspring sex ratios, a future increase in embryonic mortality could be a critical factor for a population already experiencing ecological and physiological challenges due to climate change.     

Thermal performance curves under daily thermal fluctuation: A study in helmeted water toad tadpoles

Most research in physiological ecology has focused on the effects of mean changes in temperature under the classic “hot vs cold” acclimation treatment; however, current evidence suggests that an increment in both the mean and variance of temperature could act synergistically to amplify the negative effects of global temperature increase and how it would affect fitness and performance-related traits in ectothermic organisms. We assessed the effects of acclimation to daily variance of temperature on thermal performance curves of swimming speed in helmeted water toad tadpoles (Calyptocephalella gayi). Acclimation treatments were 20 °C ± 0.1 SD (constant) and 20 °C ± 1.5 SD (fluctuating). We draw two key findings: first, tadpoles exposed to daily temperature fluctuation had reduced maximal performance (Z_max), and flattened thermal performance curves, thus supporting the “vertical shift or faster-slower” hypothesis, and suggesting that overall swimming performance would be lower through an examination of temperatures under more realistic and ecologically-relevant fluctuating regimens; second, there was significant interindividual variation in performance traits by means of significant repeatability estimates.Our present results suggest that the widespread use of constant acclimation temperatures in laboratory experiments to estimate thermal performance curves (TPCs) may lead to an overestimation of actual organismal performance. We encourage the use of temperature fluctuation acclimation treatments to better understand the variability of physiological traits, which predict ecological and evolutionary responses to global change.     

Thermal sensitivity of a Neotropical amphibian (Engystomops pustulosus) and its vulnerability to climate change

A species’ thermal sensitivity and its exposure to climate variation are key components in the prediction of its vulnerability to climate change. We tested the thermal sensitivity of a tropical amphibian that lives in a mild constant climate in which the thermal tolerance range is expected to closely match the experienced environmental temperature. The air temperature that this species is exposed to varies between 21.9 and 31.6°C with an annual mean of 27.2°C. We estimated the microhabitat water temperature variation under vegetation shade, which buffers the temperature by 1.8°C in relation to that of the air, and with open canopy, where the water was 1.9°C warmer than the air temperature. With broods of tadpoles split into five treatments (15°C, 21°C, 28°C, 31°C, and 33°C), we estimated the critical thermal maximum (CTMax) and critical thermal minimum (CTMin) after at least 7 days of acclimation. Both CTMax (42.3°C) and CTMin (11.8°C) were more extreme than the temperature range estimated for the field. We estimated the optimum temperature (T_o = 28.8°C) and the thermal performance breadth (range: 23.3–34.1°C) based on growth rate (g/day). The animals were able to acclimate more extensively to cold than to warm temperatures. These performance curve traits closely matched the air temperature. The estimated vulnerability varied according to the microhabitat prediction model used. The combination of tadpole data on thermal sensitivity and macro‐ and microhabitat variation provides a necessary framework to understand the effects of climate change on tropical amphibians.     

Effects of daily fluctuating temperatures on the Drosophila–Leptopilina boulardi parasitoid association

Koinobiont parasitoid insects, which maintain intimate and long-term relationships with their arthropod hosts, constitute an association of ectothermic organisms that is particularly sensitive to temperature variations. Because temperature shows pronounced natural daily fluctuations, we examined if experiments based on a constant temperature range can mask the real effects of the thermal regime on host-parasitoid interactions. The effects of two fluctuating thermal regimes on several developmental parameters of the Drosophila larval parasitoid Leptopilina boulardi were analyzed in this study. Regime 1 included a range of 16–23–16 °C and regime 2 included a range of 16–21–26–21–16 °C (mean temperature 20.1 °C) compared to a 20.1 °C constant temperature. Under an average temperature of 20.1 °C, which corresponds to a cold condition of L. boulardi development, we showed that the success of parasitism is significantly higher under a fluctuating temperature regime than at constant temperature. A fluctuating regime also correlated with a reduced development time of the parasitoids. In contrast, the thermal regime did not affect the ability of Drosophila to resist parasitoid infestation. Finally, we demonstrated that daily temperature fluctuation prevented the entry into diapause for this species, which is normally observed at a constant temperature of 21 °C. Overall, the results reveal that constant temperature experiments can produce misleading results, highlighting the need to study the thermal biology of organisms under fluctuating regimes that reflect natural conditions as closely as possible. This is particularly a major issue in host-parasitoid associations, which constitute a good model to understand the effect of climate warming on interacting species.     

Impact of fluctuating temperatures on development of the koinobiont endoparasitoid Venturia canescens

The effect of temperature on the biology of Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae) is well understood under constant temperature conditions, but less so under more natural, fluctuating conditions. Herein we studied the influence of fluctuating temperatures on biological parameters of V. canescens. Parasitized fifth-instar larvae of Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) were reared individually in incubators at six fluctuating temperature regimes (15–19.5 °C with a mean of 17.6 °C, 17.5–22.5 °C with a mean of 19.8 °C, 20–30 °C with a mean of 22.7 °C, 22.5–27.5 °C with a mean of 25 °C, 25.5-32.5 °C with a mean of 28.3 °C and 28.5–33 °C with a mean of 30 °C) until emergence and death of V. canescens adults. Developmental time from parasitism to adult eclosion, adult longevity and survival were recorded at each fluctuating temperature regime. In principle, developmental time decreased with an increase of the mean temperature of the fluctuating temperature regime. Upper and lower threshold temperatures for total development were estimated at 34.9 and 6.7 °C, respectively. Optimum temperature for development and thermal constant were 28.6 °C and 526.3 degree days, respectively. Adult longevity was also affected by fluctuating temperature, as it was significantly reduced at the highest mean temperature (7.0 days at 30 °C) compared to the lowest one (29.4 days at 17.6 °C). Survival was low at all tested fluctuating temperatures, apart from mean fluctuating temperature of 25 °C (37%). Understanding the thermal biology of V. canescens under more natural conditions is of critical importance in applied contexts. Thus, predictions of biological responses to fluctuating temperatures may be used in population forecasting models which potentially influence decision-making in IPM programs.     

Constant and fluctuating temperature acclimations have similar effects on phenotypic plasticity in springtails

Much interest exists in the extent to which constant versus fluctuating temperatures affect thermal performance traits and their phenotypic plasticity. Theory suggests that effects should vary with temperature, being especially pronounced at more extreme low (because of thermal respite) and high (because of Jensen's inequality) temperatures. Here we tested this idea by examining the effects of constant temperatures (10 to 30 °C in 5 °C increments) and fluctuating temperatures (means equal to the constant temperatures, but with fluctuations of ±5 °C) temperatures on the adult (F2) phenotypic plasticity of three thermal performance traits – critical thermal minimum (CT_min), critical thermal maximum (CT_max), and upper lethal temperature (ULT₅₀) in ten species of springtails (Collembola) from three families (Isotomidae 7 spp.; Entomobryidae 2 spp.; Onychiuridae 1 sp.). The lowest mean CT_min value recorded here was -3.56 ± 1.0 °C for Paristoma notabilis and the highest mean CT_max was 43.1 ± 0.8 °C for Hemisotoma thermophila. The Acclimation Response Ratio for CT_min was on average 0.12 °C/°C (range: 0.04 to 0.21 °C/°C), but was much lower for CT_max (mean: 0.017 °C/°C, range: -0.015 to 0.047 °C/°C) and lower also for ULT₅₀ (mean: 0.05 °C/°C, range: -0.007 to 0.14 °C/°C). Fluctuating versus constant temperatures typically had little effect on adult phenotypic plasticity, with effect sizes either no different from zero, or inconsistent in the direction of difference. Previous work assessing adult phenotypic plasticity of these thermal performance traits across a range of constant temperatures can thus be applied to a broader range of circumstances in springtails.     

Response of Development and Body Mass to Daily Temperature Fluctuations: a Study on <Emphasis Type="Italic">Tribolium castaneum</Emphasis>

Differences in thermal regimes are of paramount importance in insect development. However, experiments that examine trait development under constant temperature conditions may yield less evolutionarily relevant results than those that take naturally occurring temperature fluctuations into account. We investigated the effect of different temperature regimes (constant 30 °C, constant 35 °C, fluctuating with a daily mean of 30 °C, or fluctuating with a daily mean of 35 °C) on sex-specific development time and body mass in Tribolium castaneum. Using a half-sib breeding design, we also examined whether there is any evidence for genotype-by-environment interactions (GEI) for the studied traits. In response to fluctuating temperature regimes, beetles demonstrated reaction norm patterns in which thermal fluctuations influenced traits negatively above the species’ thermal optimum but had little to no effect close to the thermal optimum. Estimated heritabilities of development time were in general low and non-significant. In case of body mass of pupae and adults, despite significant genetic variance, we did not find any GEI due to crossing of reaction norms, both between temperatures and between variability treatments. We have observed a weak tendency towards higher heritabilities of adult and pupa body mass in optimal fluctuating thermal conditions. Thus, we have not found any biasing effect of stable thermal conditions as compared to fluctuating temperatures on the breeding values of heritable body-size traits. Contrary to this we have observed a strong population-wide effect of thermal fluctuations, indicated by the significant temperature-fluctuations interaction in both adult and pupa mass.     

Thermal niche predicts tolerance to habitat conversion in tropical amphibians and reptiles

Habitat conversion is a major driver of the biodiversity crisis, yet why some species undergo local extinction while others thrive under novel conditions remains unclear. We suggest that focusing on species' niches, rather than traits, may provide the predictive power needed to forecast biodiversity change. We first examine two Neotropical frog congeners with drastically different affinities to deforestation and document how thermal niche explains deforestation tolerance. The more deforestation‐tolerant species is associated with warmer macroclimates across Costa Rica, and warmer microclimates within landscapes. Further, in laboratory experiments, the more deforestation‐tolerant species has critical thermal limits, and a jumping performance optimum, shifted ~2 °C warmer than those of the more forest‐affiliated species, corresponding to the ~3 °C difference in daytime maximum temperature that these species experience between habitats. Crucially, neither species strictly specializes on either habitat – instead habitat use is governed by regional environmental temperature. Both species track temperature along an elevational gradient, and shift their habitat use from cooler forest at lower elevations to warmer deforested pastures upslope. To generalize these conclusions, we expand our analysis to the entire mid‐elevational herpetological community of southern Costa Rica. We assess the climatological affinities of 33 amphibian and reptile species, showing that across both taxonomic classes, thermal niche predicts presence in deforested habitat as well as or better than many commonly used traits. These data suggest that warm‐adapted species carry a significant survival advantage amidst the synergistic impacts of land‐use conversion and climate change.     

Changes to the elevational limits and extent of species ranges associated with climate change

The first expected symptoms of a climate change‐generated biodiversity crisis are range contractions and extinctions at lower elevational and latitudinal limits to species distributions. However, whilst range expansions at high elevations and latitudes have been widely documented, there has been surprisingly little evidence for contractions at warm margins. We show that lower elevational limits for 16 butterfly species in central Spain have risen on average by 212 m (± SE 60) in 30 years, accompanying a 1.3 °C rise (equivalent to c. 225 m) in mean annual temperature. These elevational shifts signify an average reduction in habitable area by one‐third, with losses of 50–80% projected for the coming century, given maintenance of the species thermal associations. The results suggest that many species have already suffered climate‐mediated habitat losses that may threaten their long‐term chances of survival.     

FLUCTUATING TEMPERATURE LEADS TO EVOLUTION OF THERMAL GENERALISM AND PREADAPTATION TO NOVEL ENVIRONMENTS

Environmental fluctuations can select for generalism, which is also hypothesized to increase organisms’ ability to invade novel environments. Here, we show that across a range of temperatures, opportunistic bacterial pathogen Serratia marcescens that evolved in fluctuating temperature (daily variation between 24°C and 38°C, mean 31°C) outperforms the strains that evolved in constant temperature (31°C). The growth advantage was also evident in novel environments in the presence of parasitic viruses and predatory protozoans, but less clear in the presence of stressful chemicals. Adaptation to fluctuating temperature also led to reduced virulence in Drosophila melanogaster host, which suggests that generalism can still be costly in terms of reduced fitness in other ecological contexts. While supporting the hypothesis that evolution of generalism is coupled with tolerance to several novel environments, our results also suggest that thermal fluctuations driven by the climate change could affect both species’ invasiveness and virulence.     

The influence of maternal thermal environments on reproductive traits and hatchling traits in a Lacertid lizard,Takydromus septentrionalis

The thermal environment can induce substantial variation in important life-history traits. Experimental manipulation of the thermal environment can help researchers determine the contribution of this factor to phenotypic variation in life-history traits. During the reproductive season, we kept female northern grass lizards, Takydromus septentrionalis (Lacertidae), in three temperature-controlled rooms (25, 28 and 32 °C) to measure the effect of the maternal thermal environment on reproductive traits. Maternal thermal environment remarkably affected reproductive frequency and thereby seasonal reproductive output, but had little effect on reproductive traits per clutch or hatchling traits. Females kept at 32 °C produced more clutches and thus had shorter clutch intervals than females from 28 to 25 °C. Clutch size, clutch mass, relative clutch mass, egg size and hatchling traits did not vary among the three treatments. The eggs produced by the females were incubated at 27 °C and the traits of hatchlings were measured. The result that egg (offspring) size was independent of maternal thermal environments is consistent with the prediction of the optimal egg size (offspring) theory. The eggs produced by low temperature females (28 and 25 °C) took longer time to complete their post-oviposition development than did eggs produced by high temperature females (32 °C). This suggests that the eggs from low temperatures might have been laid when the embryos were at relatively early stages. Therefore, maternal thermal environment prior to oviposition could affect post-oviposition development in T. septentrionalis.     

Fluctuating environments impact thermal tolerance in an invasive insect species Bactrocera dorsalis (Diptera: Tephritidae)

The incidence and severity of environmental stressors associated with global climate change are increasing and insects frequently face variability in temperature and moisture regimes at variable spatio-temporal scales. Coincidental with this, is increased thermal and hydric stress on insects as warming increases vapour pressure deficit (VPD), the drying power of the air. While the effects of mean temperatures on fitness are widely documented, fluctuations in both temperature and relative humidity (RH) are largely unexplored. Here, we investigated the effects of dynamic temperature and RH fluctuations (around the mean [28°C; 65% RH]) on low and high thermal tolerance of laboratory-reared adult invasive Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), measured as critical thermal minima (CT_min), critical thermal maxima (CT_max), chill coma recovery time (CCRT) and heat knockdown time (HKDT). Our results show that increased environmental amplitude significantly influenced low and high temperature responses and varied across traits tested. The highest amplitude (δ12°C; 28% RH) compromised CT_min, CCRT and HKDT traits while enhancing CT_max. Similarly, acclimation to δ3°C; 7% RH compromised both low (CT_min and CCRT) and high (CT_max and HKDT) fitness traits. Variations in fitness reported here indicate significant roles of combined thermal and moisture fluctuations on B. dorsalis fitness suggesting caveats that are worthy considering when predicting species responses to climate change. These results are significant for B. dorsalis population phenology, management, quantifying vulnerability to climate variability and may help modelling future biogeographical patterns.     

Physiological costs of cold exposure on the parasitoid <Emphasis Type="Italic">Aphidius ervi</Emphasis>, without selection pressure and under constant or fluctuating temperatures

The aim of this study is to evaluate the physiological costs and the consequences on fitness of cold exposure on the parasitoid Aphidius ervi (Hymenoptera: Aphidiinae) under non-lethal conditions. We exposed 1-day-old mummies to different treatments: control at 20°C, 7°C constant, 7°C fluctuating (7°C for 22 h and 20°C for 2 h). Two performances of fitness were particularly affected after two weeks at 7°C constant: female longevity and the sex ratio of the progeny was male biased. In contrast, egg load at emergence, lifetime fecundity, frequency of asymmetric individuals in both sexes, and mating success were not significantly affected under all treatments. The effects of these treatments on trends such as utilisation of fat reserves and fertility of males and females are discussed.     

Similarities in butterfly emergence dates among populations suggest local adaptation to climate

Phenology shifts are the most widely cited examples of the biological impact of climate change, yet there are few assessments of potential effects on the fitness of individual organisms or the persistence of populations. Despite extensive evidence of climate‐driven advances in phenological events over recent decades, comparable patterns across species' geographic ranges have seldom been described. Even fewer studies have quantified concurrent spatial gradients and temporal trends between phenology and climate. Here we analyse a large data set (~129 000 phenology measures) over 37 years across the UK to provide the first phylogenetic comparative analysis of the relative roles of plasticity and local adaptation in generating spatial and temporal patterns in butterfly mean flight dates. Although populations of all species exhibit a plastic response to temperature, with adult emergence dates earlier in warmer years by an average of 6.4 days per °C, among‐population differences are significantly lower on average, at 4.3 days per °C. Emergence dates of most species are more synchronised over their geographic range than is predicted by their relationship between mean flight date and temperature over time, suggesting local adaptation. Biological traits of species only weakly explained the variation in differences between space‐temperature and time‐temperature phenological responses, suggesting that multiple mechanisms may operate to maintain local adaptation. As niche models assume constant relationships between occurrence and environmental conditions across a species' entire range, an important implication of the temperature‐mediated local adaptation detected here is that populations of insects are much more sensitive to future climate changes than current projections suggest.     

Bet hedging in a warming ocean: predictability of maternal environment shapes offspring size variation in marine sticklebacks

Bet hedging at reproduction is expected to evolve when mothers are exposed to unpredictable cues for future environmental conditions, whereas transgenerational plasticity (TGP) should be favoured when cues reliably predict the environment offspring will experience. Since climate predictions forecast an increase in both temperature and climate variability, both TGP and bet hedging are likely to become important strategies to mediate climate change effects. Here, the potential to produce variably sized offspring in both warming and unpredictable environments was tested by investigating whether stickleback (Gasterosteus aculeatus) mothers adjusted mean offspring size and within‐clutch variation in offspring size in response to experimental manipulation of maternal thermal environment and predictability (alternating between ambient and elevated water temperatures). Reproductive output traits of F1 females were influenced by both temperature and environmental predictability. Mothers that developed at ambient temperature (17 °C) produced larger, but fewer eggs than mothers that developed at elevated temperature (21 °C), implying selection for different‐sized offspring in different environments. Mothers in unpredictable environments had smaller mean egg sizes and tended to have greater within‐female egg size variability, especially at 21 °C, suggesting that mothers may have dynamically modified the variance in offspring size to spread the risk of incorrectly predicting future environmental conditions. Both TGP and diversification influenced F2 offspring body size. F2 offspring reared at 21 °C had larger mean body sizes if their mother developed at 21 °C, but this TGP benefit was not present for offspring of 17 °C mothers reared at 17 °C, indicating that maternal TGP will be highly relevant for ocean warming scenarios in this system. Offspring of variable environment mothers were smaller but more variable in size than offspring from constant environment mothers, particularly at 21 °C. In summary, stickleback mothers may have used both TGP and diversified bet‐hedging strategies to cope with the dual stress of ocean warming and environmental uncertainty.     

Indirect selection of thermal tolerance during experimental evolution of Drosophila melanogaster

Natural selection alters the distribution of a trait in a population and indirectly alters the distribution of genetically correlated traits. Long‐standing models of thermal adaptation assume that trade‐offs exist between fitness at different temperatures; however, experimental evolution often fails to reveal such trade‐offs. Here, we show that adaptation to benign temperatures in experimental populations of Drosophila melanogaster resulted in correlated responses at the boundaries of the thermal niche. Specifically, adaptation to fluctuating temperatures (16–25°C) decreased tolerance of extreme heat. Surprisingly, flies adapted to a constant temperature of 25°C had greater cold tolerance than did flies adapted to other thermal conditions, including a constant temperature of 16°C. As our populations were never exposed to extreme temperatures during selection, divergence of thermal tolerance likely reflects indirect selection of standing genetic variation via linkage or pleiotropy. We found no relationship between heat and cold tolerances in these populations. Our results show that the thermal niche evolves by direct and indirect selection, in ways that are more complicated than assumed by theoretical models.     

Climate change is predicted to alter the current pest status of Globodera pallida and G. rostochiensis in the United Kingdom

The potato cyst nematodes Globodera pallida and G. rostochiensis are economically important plant pathogens causing losses to UK potato harvests estimated at £50 m/ year. Implications of climate change on their future pest status have not been fully considered. Here, we report growth of female G. pallida and G. rostochiensis over the range 15 to 25°C. Females per plant and their fecundity declined progressively with temperatures above 17.5°C for G. pallida, whilst females per plant were optimal between 17.5 and 22.5°C for G. rostochiensis. Relative reproductive success with temperature was confirmed on two potato cultivars infected with either species at 15, 22.5 and 25°C. The reduced reproductive success of G. pallida at 22.5°C relative to 15°C was also recorded for a further seven host cultivars studied. The differences in optimal temperatures for reproductive success may relate to known differences in the altitude of their regions of origin in the Andes. Exposure of G. pallida to a diurnal temperature stress for one week during female growth significantly suppressed subsequent growth for one week at 17.5°C but had no effect on G. rostochiensis. However, after two weeks of recovery, female size was not significantly different from that for the control treatment. Future soil temperatures were simulated for medium‐ and high‐emission scenarios and combined with nematode growth data to project future implications of climate change for the two species. Increased soil temperatures associated with climate change may reduce the pest status of G. pallida but benefit G. rostochiensis especially in the southern United Kingdom. We conclude that plant breeders may be able to exploit the thermal limits of G. pallida by developing potato cultivars able to grow under future warm summer conditions. Existing widely deployed resistance to G. rostochiensis is an important characteristic to retain for new potato cultivars.     

Intraspecific Variation of Reproductive Traits in a Mediterranean Lizard: Clutch, Population, and Lineage Effects

Widely distributed terrestrial ectotherms from the southern European peninsulas show patterns of subdivision (related to isolation in temperate refugia) that allow us to test the relative importance of phylogeographic lineage, population of origin and familial effects as sources of variation for life-history traits. We collected gravid females from 15 geographically separated populations of the lacertid lizard Psammodromus algirus, a widely distributed species with well differentiated eastern and western lineages. We incubated eggs under two treatments of constant (28°C) and fluctuating (28 ± 4°C) temperature, and we examined clutch, population, and lineage effects on several traits of females, eggs, and hatchlings. Incubation time was mainly explained by differences between lineages, but it was also influenced by population and female effects. Within each lineage, incubation was shorter at cooler and wetter sites, and for a given climate it was shorter for eastern than for western populations, suggesting that countergradient variation has evolved independently in the two lineages. Female size, clutch size, and relative fecundity were primarily influenced by inter-population differences, a pattern that seemed attributable to environmental differences in productivity, because mean female size was positively correlated with a gradient of increasing precipitation and decreasing temperature. Clutch size was mainly, but not entirely, dependent on female SVL, suggesting both a proximate effect of local conditions and intrinsic differences among populations. Females from drier and warmer sites produced larger hatchlings. Mean egg mass was mainly determined by familial effects. Eggs incubated at a constant temperature hatched earlier than did their siblings incubated at fluctuating temperatures, a fact that could be explained by considering that in Mediterranean environments developmental rate might increase at a lower speed above average incubation temperature than it does decrease below it.     

The effect of acclimation temperature on thermal activity thresholds in polar terrestrial invertebrates

In the Maritime Antarctic and High Arctic, soil microhabitat temperatures throughout the year typically range between ?10 and +5 °C. However, on occasion, they can exceed 20 °C, and these instances are likely to increase and intensify as a result of climate warming. Remaining active under both cool and warm conditions is therefore important for polar terrestrial invertebrates if they are to forage, reproduce and maximise their fitness. In the current study, lower and upper thermal activity thresholds were investigated in the polar Collembola, Megaphorura arctica and Cryptopygus antarcticus, and the mite, Alaskozetes antarcticus. Specifically, the effect of acclimation on these traits was explored. Sub-zero activity was exhibited in all three species, at temperatures as low as ?4.6 °C in A. antarcticus. At high temperatures, all three species had capacity for activity above 30 °C and were most active at 25 °C. This indicates a comparable spread of temperatures across which activity can occur to that seen in temperate and tropical species, but with the activity window shifted towards lower temperatures. In all three species following one month acclimation at ?2 °C, chill coma (=the temperature at which movement and activity cease) and the critical thermal minimum (=low temperature at which coordination is no longer shown) occurred at lower temperatures than for individuals maintained at +4 °C (except for the CTmin of M. arctica). Individuals acclimated at +9 °C conversely showed little change in their chill coma or CTmin. A similar trend was demonstrated for the heat coma and critical thermal maximum (CTmax) of all species. Following one month at ?2 °C, the heat coma and CTmax were reduced as compared with +4 °C reared individuals, whereas the heat coma and CTmax of individuals acclimated at +9 °C showed little adjustment. The data obtained suggest these invertebrates are able to take maximum advantage of the short growing season and have some capacity, in spite of limited plasticity at high temperatures, to cope with climate change.