Altitudinal and environmental variation in lifespan in the Copper butterfly Lycaena tityrus

1.  Variation in longevity within and between natural populations is widespread, and understanding the relative importance of environmental and genetic factors as well as their interactions in mediating such variation is crucial in longevity research.
2.  In this study lifespan of adult copper butterflies was examined in relation to altitude, temperature (20 and 27 °C), sex and adult feeding.
3.  As expected, longevity increased with decreasing temperature, and sucrose-fed butterflies had longer lifespans compared to water-fed and finally non-fed individuals. The impact of feeding, especially of having access to water or not, was larger at the higher compared to the lower temperature.
4.  Regarding altitudinal patterns, increased lifespan in high-altitude populations was largely restricted to beneficial feeding conditions, while under carbohydrate deprivation low-altitude animals lived longer, suggesting that low-altitude butterflies do better under food stress.
5.  Differences in longevity between sexes were small at 20 °C, while females lived substantially longer than males at the higher temperature. Consequently, females may be less susceptible to high temperature stress than males. Further, males suffered more from food stress than females, suggesting that females are generally more stress resistant than males.
6.  Using a full factorial design, this study demonstrates that variation in longevity is caused by several factors, and additionally by substantial interactive effects. Consequently, patterns of variation in longevity are complex, and one needs to be cautious when neglecting this source of variation, by focussing on individual factors only.     

Sexual differences in life-history traits in the butterfly Lycaena tityrus: a comparison between direct and diapause development

During direct development the butterfly Lycaena tityrus was previously found to display sex-related reaction norms in response to temperature. Based on selection for protandry in males and fecundity selection for larger females, males favoured early emergence over large size, leading to a dramatic weight loss at higher temperatures, whereas females maintained similar weights throughout. Because males were able to avoid a weight reduction relative to females in spite of their shorter development at lower temperatures, sexual size dimorphism existed at higher temperatures only. In the present paper we compare sexual differences in life-history traits in L. tityrus between direct and diapause development at 25 °C. We demonstrate that, regardless of developmental pathway, protandry persisted and relative sexual size dimorphism, with females being larger, remained unchanged. Although diapausing individuals were less time-constrained, allowing them to grow to considerably higher final weights in both sexes, males were not able to reduce their weight loss relative to females. This is explained by the pressure to gain a developmental advantage solely during post-diapause development, whereas direct developing males may spread the burden over the whole larval period. Our results highlight the importance of considering sexual differences in selective pressures, which may influence central life-history traits in manifold ways.     

Effects of rearing and induction temperature on the temporal dynamics of heat shock protein 70 expression in a butterfly

The temporal dynamics of heat shock protein 70 (HSP70) expression in response to longer‐term acclimation and rapid hardening in the butterfly Lycaena tityrus is investigated. After a 1‐h exposure to 1 °C or 37 °C, HSP70 is quickly up‐regulated within 1 h and down‐regulated within 2 h. The fast dynamic of HSP70 expression is in contrast to the patterns found in organisms inhabiting more stable thermal environments, and is interpreted as an adaptation to the large and rapid temperature variation experienced by flying ectotherms. HSP70 expression is higher in males than in females, as well as in animals reared at 27 °C than at 20 °C, although it is very similar across the high and low induction temperatures. Animals reared at the higher temperature, however, respond less strongly to high‐temperature stress.     

Evolutionary potential of Chamaecrista fasciculata in relation to climate change. I. Clinal patterns of selection along an environmental gradient in the great plains

Climate change will alter natural selection on native plant populations. Little information is available to predict how selection will change in the future and how populations will respond. Insight can be obtained by comparing selection regimes in current environments to selection regimes in environments similar to those predicted for the future. To mimic predicted temporal change in climate, three natural populations of the annual legume Chamaecrista fasciculata were sampled from a climate gradient in the Great Plains and progeny of formal crosses were reciprocally planted back into common gardens across this climate gradient. In each garden, native populations produced significantly more seed than the other populations, providing strong evidence of local adaptation. Phenotypic selection analysis conducted by site showed that plants with slower reproductive development, more leaves, and thicker leaves were favored in the most southern garden. Evidence of clinal variation in selection regimes was also found; selection coefficients were ordered according to the latitude of the common gardens. The adaptive value of native traits was indicated by selection toward the mean of local populations. Repeated clinal patterns in linear and nonlinear selection coefficients among populations and within and between sites were found. To the extent that temporal change in climate into the future will parallel the differences in selection across this spatial gradient, this study suggests that selection regimes will be displaced northward and different trait values will be favored in natural populations.     

Wing morphology of the butterfly Coenonympha arcania in Europe: Traces of both historical isolation in glacial refugia and current adaptation

In this study, we examined the evolutionary outcome of and interplay between historic isolation and current selection pressures on traits more or less closely connected to fitness in the Pearly Heath butterfly (Coenonympha arcania) across its range in Europe. We hypothesized that a trait mean is more related to historic events if it has low connection to fitness, while a trait more closely connected with fitness is expected to have a mean that relates more to current selection pressures. In order to test this, we collected 322 butterflies from across the species range in Europe and measured five wing traits relating to size and color patterns. To infer a phylogeographic history for each individual, we sequenced a 594 bp fragment of the COI gene. The morphological data were then analyzed in relation to selected climatic variables and the history of individuals to disentangle which factors best correlated with morphological variation. The results supported our hypothesis in that wing sizes correlated with summer precipitation but not with its inferred location during the last glaciation. Eyespot position, on the other hand, correlated with the history of individuals but not with the analyzed climatic indicators. The sizes of the black spot and the white band, two traits that were expected to have intermediate selection pressure, were associated with both history and current conditions. Thus, this study illustrates the fascinating interplay between events and processes that lead to a specific evolutionary outcome.     

Trade-offs between baseline thermal tolerance and thermal tolerance plasticity are much less common than it appears

Thermal tolerance plasticity is a core mechanism by which organisms can mitigate the effects of climate change. As a result, there is a need to understand how variation in tolerance plasticity arises. The baseline tolerance/plasticity trade-off hypothesis (hereafter referred to as the trade-off hypothesis, TOH) has recently emerged as a potentially powerful explanation. The TOH posits that organisms with high baseline thermal tolerance have reduced thermal tolerance plasticity relative to those with low baseline tolerance. Many studies have found support for the TOH. However, this support must be regarded cautiously because the most common means of testing the TOH can yield spurious “trade-offs” due to regression to the mean. I acquired data for 25 previously published analyses that supported the TOH at the intraspecific level and reanalyzed them after applying a method that adjusts plasticity estimates for regression to the mean. Only six of the 25 analyses remained statistically significant after adjustment, and effect size and variance explained decreased in all cases. The few data sets in which support for the TOH was maintained after adjustment point to areas of future study, but are too few to make generalizations at this point. In sum, regression to the mean has led to a substantial overestimation of support for the TOH and must be accounted for in future tests of the hypothesis.     

Cold tolerance in relation to developmental and adult temperature in a butterfly

Abstract. Larvae of the butterfly Lycaena tityrus (Poda) are reared at 20 or 27 °C until adult eclosion, after which they are maintained at the same temperature or are transferred to the alternate temperature. The resulting adults are exposed to −20 °C for 8 min, returned to ambient temperature, and the recovery time to standing position is recorded. On the day of eclosion, individuals reared at 20 °C show 19% shorter recovery times than individuals reared at 27 °C. This effect of developmental temperature disappears when the same animals are tested 3 and 6 days later. However, adult temperature did not affect recovery time in these animals, presumably due to over-riding effects of previous cold shocks. This is suggested by another set of animals, not having experienced previous cold shocks, demonstrating recovery times that are 28% shorter in individuals maintained as adults for 3 days at 20 compared to 27 °C. Thus, L. tityrus appears to be capable of adapting to local temperatures.     

The behavioural and physiological strategies of bird and reptile embryos in response to unpredictable variation in nest temperature

Temperature profoundly affects the rate and trajectory of embryonic development, and thermal extremes can be fatal. In viviparous species, maternal behaviour and physiology can buffer the embryo from thermal fluctuations; but in oviparous animals (like most reptiles and all birds), an embryo is likely to encounter unpredictable periods when incubation temperatures are unfavourable. Thus, we might expect natural selection to have favoured traits that enable embryos to maintain development despite those fluctuations. Our review of recent research identifies three main routes that embryos use in this way. Extreme temperatures (i) can be avoided (e.g. by accelerating hatching, by moving within the egg, by cooling the egg by enhanced rates of evaporation, or by hysteresis in rates of heating versus cooling); (ii) can be tolerated (e.g. by entering diapause, by producing heat‐shock proteins, or by changing oxygen use); or (iii) the embryo can adjust its physiology and/or developmental trajectory in ways that reduce the fitness penalties of unfavourable thermal conditions (e.g. by acclimating, by exploiting brief windows of favourable conditions, or by producing the hatchling phenotype best suited to those incubation conditions). Embryos are not simply passive victims of ambient conditions. Like free‐living stages of the life cycle, embryos exhibit behavioural and physiological plasticity that enables them to deal with unpredictable abiotic challenges.     

Environmental and genetic control of cold tolerance in the Glanville fritillary butterfly

Thermal tolerance has a major effect on individual fitness and species distributions and can be determined by genetic variation and phenotypic plasticity. We investigate the effects of developmental and adult thermal conditions on cold tolerance, measured as chill coma recovery (CCR) time, during the early and late adult stage in the Glanville fritillary butterfly. We also investigate the genetic basis of cold tolerance by associating CCR variation with polymorphisms in candidate genes that have a known role in insect physiology. Our results demonstrate that a cooler developmental temperature leads to reduced cold tolerance in the early adult stage, whereas cooler conditions during the adult stage lead to increased cold tolerance. This suggests that adult acclimation, but not developmental plasticity, of adult cold tolerance is adaptive. This could be explained by the ecological conditions the Glanville fritillary experiences in the field, where temperature during early summer, but not spring, is predictive of thermal conditions during the butterfly's flight season. In addition, an amino acid polymorphism (Ala‐Glu) in the gene flightin, which has a known function in insect flight and locomotion, was associated with CCR. These amino acids have distinct biochemical properties and may thus affect protein function and/or structure. To our knowledge, our study is the first to link genetic variation in flightin to cold tolerance, or thermal adaptation in general.     

Life-history plasticity in the butterfly Lycaena hippothoe: local adaptations and trade-offs

Life-history theory predicts some cost to be associated with short development time, the most frequently assumed being small adult size. Alternatively, insects may increase developmental rates and grow fast to a larger size. Seasonal environments should select for phenotypic plasticity in growth and development, based on the need to complete development up to the diapausing stage before the onset of unfavourable season. Nevertheless, there must be some limit beyond which a compensation for a shorter development cannot be achieved. By comparing three geographically isolated populations of Lycaena hippothoe in common environments we show that in the Hungarian population development time seems to be traded off against size at maturity. This population is the only bivoltine one within this principally monovoltine species. Thus, realization of an additional generation per year, achieved through largely reduced development times, appears to carry the cost of substantially lower adult weights compared with other populations. In contrast, differences in development time in two monovoltine populations were not accompanied by a trade-off between development time and size. These results suggest that clear trade-offs are restricted to stressful situations, when compensation by an increase in growth rates is no longer feasible. We suggest the particularly short development time in the Hungarian population (facilitating a second generation), as well as the shorter development in an alpine (short vegetation period) compared with a western German population, to be adaptations to local climatic conditions. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 173–185.     

Increasing ocean temperature reduces the metabolic performance and swimming ability of coral reef damselfishes

Tropical coral reef teleosts are exclusively ectotherms and their capacity for physical and physiological performance is therefore directly influenced by ambient temperature. This study examined the effect of increased water temperature to 3 °C above ambient on the swimming and metabolic performance of 10 species of damselfishes (Pomacentridae) representing evolutionary lineages from two subfamilies and four genera. Five distinct performance measures were tested: (a) maximum swimming speed (U_crit), (b) gait‐transition speed (the speed at which they change from strictly pectoral to pectoral‐and‐caudal swimming, U_p?c), (c) maximum aerobic metabolic rate (MO_2?MAX), (d) resting metabolic rate (MO_2?REST), and (e) aerobic scope (ratio of MO_2?MAX to MO_2?REST, A_SC). Relative to the control (29 °C), increased temperature (32 °C) had a significant negative effect across all performance measures examined, with the magnitude of the effect varying greatly among closely related species and genera. Specifically, five species spanning three genera (Dascyllus, Neopomacentrus and Pomacentrus) showed severe reductions in swimming performance with U_crit reduced in these species by 21.3–27.9% and U_p?c by 32.6–51.3%. Furthermore, five species spanning all four genera showed significant reductions in metabolic performance with aerobic scope reduced by 24.3–64.9%. Comparisons of remaining performance capacities with field conditions indicate that 32 °C water temperatures will leave multiple species with less swimming capacity than required to overcome the water flows commonly found in their respective coral reef habitats. Consequently, unless adaptation is possible, significant loss of species may occur if ocean warming of ≥3 °C arises.     

Resistance to thermal stress in preadult Drosophila buzzatii: variation among populations and changes in relative resistance across life stages

Resistance to a short term exposure to a high temperature stress was examined in eggs, larvae and pupae of Drosophila buzzfltii from seven localities. Across development, pupae were most resistant, followed by eggs, and then first and third-instar larvae. Variation among populations for resistance to heat stress was significant in all life stages. However, there was much less variation among populations where measured as eggs and pupae than for both first and third instar larvae. Older larvae showed large changes both in viability and developmental time, while exposure of young larvae to heat stress led to a decline in viability without delayed development. Populations that had the shortest developmental time at 25^oC were relatively the most resistant to heat stress as larvae. High relative resistance at one preadult life stage was not necessarily associated with relatively high resistance at another, or with previous measurements of resistance for adults from these populations. Comparison of populations that were more similar in their pattern of change in resistance across development suggested a relationship with the climate of origin. The possibility that developmental variation in the expression of heat shock proteins may cause variation in resistance to thermal stress for different life stages is discussed.     

Adaptive plasticity in wing melanisation of a montane butterfly across a Himalayan elevational gradient

1. Traits that are significant to the thermal ecology of temperate or montane species are expected to prominently co-vary with the thermal environment experienced by an organism. The Himalayan Pieris canidia butterfly exhibits considerable variation in wing melanisation. We investigated: (i) whether variation in wing melanisation and (ii) activity period of this montane butterfly was influenced by the seasonally and elevationally changing thermal landscape.
2. We discovered that wing melanisation varied across elevation, seasons, sex, and wing surfaces, with the variation strongly structured in space and time: colder seasons and higher elevations produced more melanic individuals. Notably, melanisation did not vary uniformly across all wing surfaces: (i) melanisation of the ventral hindwing co-varied much more prominently with elevation, but (ii) melanisation on all other surfaces varied with seasonal changes in the thermal environment.
3. Observed wing surface-specific patterns indicated thermoregulatory function for this variation in melanisation. Such wing surface-specific responses to seasonal and elevational variation in temperature have rarely been reported in montane insects.
4. Moreover, daily and seasonal thermal cycles were found to strongly influence activity periods of this species, suggesting the potential limits to wing melanisation plasticity.
5. Overall, these results showed that the seasonal and elevational gradients in temperature influence the thermal phenotype as well as activity periods of this Himalayan butterfly. It will be critical to study the phenotypic evolution of such montane insects in response to the ongoing climate change, which is already showing significant signs in this iconic mountain range.

     

Local adaptation and cogradient selection in the alpine plant, Poa hiemata, along a narrow altitudinal gradient

Alpine environments are particularly susceptible to environmental changes associated with global warming but there is potential for alpine plants to adapt to warming if local adaptation occurs and gene flow allows genotypes adapted to low altitudes to colonize higher altitude sites. Here we examine the adaptive potential of a common alpine grass, Poa hiemata, within the restricted alpine habitat of Australian mountains, across a narrow altitudinal gradient replicated in three areas. Grasses at high altitude sites had shorter leaf lengths and larger circumferences than those at lower sites. Transplant experiments with clonal material and plants grown from seed indicated that these differences were partly genetic, with environmental and genetic factors both contributing to the differences between altitudes. Differences in altitudinal forms were also evident in a common garden experiment. Plants showed a home-site advantage in terms of survival. A fitness analysis indicated that at high altitude sites, selection favored plants with short leaves and larger circumferences, whereas these traits were selected in the opposite direction at the low altitude sites. These findings indicate cogradient selection and potential for both plastic and genotypic shifts in response to climate change in P. hiemata.     

Dimorphic growth patterns and sex‐specific reaction norms in the butterfly Lycaena hippothoe sumadiensis

Directly developing larvae of the butterfly Lycaena hippothoe sumadiensis exhibited two growth strategies with one cohort passing four larval instars at high growth rates, and the other five instars at lower growth rates. The 4‐instar‐cohort displayed decreased development times, in combination with slightly reduced pupal and adult weights. In addition to adjustment of growth rate, omitting a larval instar may comprise a further mechanism to decrease development time when needed. Using the 4‐instar‐cohort, sex‐related differences in reaction norms were investigated over a temperature gradient. At high temperatures, protandrous males showed early emergence at a reduced size, whereas weight of females remained similar throughout. These differences suggest that large size is more important for female than for male fitness. The pattern is similar to that previously reported for alpine L. tityrus, indicating that sex‐specific reaction norms might be widespread in species living under severe time constraints.     

Acclimation to predicted ocean warming through developmental plasticity in a tropical reef fish

Determining the capacity of organisms to acclimate and adapt to increased temperatures is key to understand how populations and communities will respond to global warming. Although there is evidence that elevated water temperature affects metabolism, growth and condition of tropical marine fish, it is unknown whether they have the potential to acclimate, given adequate time. We reared the tropical reef fish Acanthochromis polyacanthus through its entire life cycle at present day and elevated (+1.5 and+3.0 °C) water temperatures to test its ability to thermally acclimate to ocean temperatures predicted to occur over the next 50–100 years. Fish reared at 3.0 °C greater than the present day average reduced their resting oxygen consumption (RMR) during summer compared with fish reared at present day temperatures and tested at the elevated temperature. The reduction in RMR of up to 69 mg O₂ kg^?1 h^?1 in acclimated fish could represent a significant benefit to daily energy expenditure. In contrast, there was no acclimation to summer temperatures exhibited by fish reared at 1.5 °C above present day temperatures. Fish acclimated to +3.0 °C were smaller and in poorer condition than fish reared at present day temperatures, suggesting that even with acclimation there will be significant consequences for future populations of tropical fishes caused by global warming.     

Variation in adult stress resistance does not explain vulnerability to climate change in copper butterflies

Ongoing climate change is a major threat to biodiversity. However, although many species clearly suffer from ongoing climate change, others benefit from it, for example, by showing range expansions. However, which specific features determine a species’ vulnerability to climate change? Phenotypic plasticity, which has been described as the first line of defence against environmental change, may be of utmost importance here. Against this background, we here compare plasticity in stress tolerance in 3 copper butterfly species, which differ arguably in their vulnerability to climate change. Specifically, we investigated heat, cold and desiccation resistance after acclimatization to different temperatures in the adult stage. We demonstrate that acclimation at a higher temperature increased heat but decreased cold tolerance and desiccation resistance. Contrary to our predictions, species did not show pronounced variation in stress resistance, though plastic capacities in temperature stress resistance did vary across species. Overall, our results seemed to reflect population—rather than species‐specific patterns. We conclude that the geographical origin of the populations used should be considered even in comparative studies. However, our results suggest that, in the 3 species studied here, vulnerability to climate change is not in the first place determined by stress resistance in the adult stage. As entomological studies focus all too often on adults only, we argue that more research effort should be dedicated to other developmental stages when trying to understand insect responses to environmental change.