Mixed support for an alignment between phenotypic plasticity and genetic differentiation in damselfly wing shape.

The relationship between genetic differentiation and phenotypic plasticity can provide information on whether plasticity generally facilitates or hinders adaptation to environmental change. Here, we studied wing shape variation in a damselfly (Lestes sponsa) across a latitudinal gradient in Europe that differed in time constraints mediated by photoperiod and temperature. We reared damselflies from northern and southern populations in the laboratory using a reciprocal transplant experiment that simulated time-constrained (i.e. northern) and unconstrained (southern) photoperiods and temperatures. After emergence, adult wing shape was analysed using geometric morphometrics. Wings from individuals in the northern and southern populations differed significantly in shape when animals were reared in their respective native environment. Comparing wing shape across environments, we found evidence for phenotypic plasticity in wing shape, and this response differed across populations (i.e. G × E interactions). This interaction was driven by a stronger plastic response by individuals from the northern population and differences in the direction of plastic wing shape changes among populations. The alignment between genetic and plastic responses depended on the specific combination of population and rearing environment. For example, there was an alignment between plasticity and genetic differentiation under time-constrained, but not under non-time-constrained conditions for forewings. We thus find mixed support for the hypothesis that environmental plasticity and genetic population differentiation are aligned. Furthermore, although our laboratory treatments mimicked the natural climatic conditions at northern and southern latitudes, the effects of population differences on wing shape were two to four times stronger than plastic effects. We discuss our results in terms of time constraints and the possibility that natural and sexual selection is acting differently on fore- and hindwings.

Predation selects for smaller eye size in a vertebrate: effects of environmental conditions and sex.

Increased eye size in animals results in a larger retinal image and thus improves visual acuity. Thus, larger eyes should aid both in finding food as well as detecting predators. On the other hand, eyes are usually very conspicuous and several studies have suggested that eye size is associated with predation risk. However, experimental evidence is scant. In this study, we address how predation affects variation in eye size by performing two experiments using Eurasian perch juveniles as prey and either larger perch or pike as predators. First, we used large outdoor tanks to compare selection due to predators on relative eye size in open and artificial vegetated habitats. Second, we studied the effects of both predation risk and resource levels on phenotypic plasticity in relative eye size in indoor aquaria experiments. In the first experiment, we found that habitat altered selection due to predators, since predators selected for smaller eye size in a non-vegetated habitat, but not in a vegetated habitat. In the plasticity experiment, we found that fish predators induced smaller eye size in males, but not in females, while resource levels had no effect on eye size plasticity. Our experiments provide evidence that predation risk could be one of the driving factors behind variation in eye size within species.

Size-mediated priority and temperature effects on intra-cohort competition and cannibalism in a damselfly.

A shift in the relative arrival of offspring, for example a shift in hatching time, can affect competition at the intraspecific level through size-mediated priority effects, where the larger individuals gain more resources. These priority effects are likely to be affected by climate warming and the rate of intraspecific predation, that is cannibalism. In a laboratory experiment, we examined size-mediated priority effects in larvae of the univoltine damselfly, Lestes sponsa, at two different temperatures (21 and 23°C). We created three size groups of larvae by manipulating hatching time: early hatched with a large size (extra-advanced), intermediate hatched with an intermediate size (advanced) and late hatched with a small size (non-advanced). Thereafter, we reared the larvae from these groups in non-mixed and mixed groups of 12 larvae. We found strong priority and temperature effects. First, extra-advanced larvae most often had higher survival, growth and development rates than non-advanced larvae in mixed groups, compared to groups that consisted of only extra-advanced larvae. Second, temperature increased growth and development rates and cannibalism. However, the strength of priority effects did not differ between the two experimental temperatures, because there was no statistical interaction between temperature and treatments. That is, the mixed and non-mixed groups of non-advanced, advanced and extra-advanced larvae showed the same relative change in life-history traits across the two temperatures. Non-advanced and advanced larvae had similar or higher growth rate and mass in mixed groups compared to non-mixed groups, suggesting that predation from advanced larvae in the mixed group released resources for the non-advanced and advanced larvae that survived despite cannibalism risk. Thus, a thinning effect occurred due to cannibalism caused by priority effects. The results suggest that a shift in the relative arrival of offspring can cause temperature-dependent priority effects, mediated through cannibalism, growth and development, which may change the size distribution and abundance of emerging aquatic insects.

The genetic variance but not the genetic covariance of life-history traits changes towards the north in a time-constrained insect.

Seasonal time constraints are usually stronger at higher than lower latitudes and can exert strong selection on life-history traits and the correlations among these traits. To predict the response of life-history traits to environmental change along a latitudinal gradient, information must be obtained about genetic variance in traits and also genetic correlation between traits, that is the genetic variance-covariance matrix, G. Here, we estimated G for key life-history traits in an obligate univoltine damselfly that faces seasonal time constraints. We exposed populations to simulated native temperatures and photoperiods and common garden environmental conditions in a laboratory set-up. Despite differences in genetic variance in these traits between populations (lower variance at northern latitudes), there was no evidence for latitude-specific covariance of the life-history traits. At simulated native conditions, all populations showed strong genetic and phenotypic correlations between traits that shaped growth and development. The variance-covariance matrix changed considerably when populations were exposed to common garden conditions compared with the simulated natural conditions, showing the importance of environmentally induced changes in multivariate genetic structure. Our results highlight the importance of estimating variance-covariance matrixes in environments that mimic selection pressures and not only trait variances or mean trait values in common garden conditions for understanding the trait evolution across populations and environments.

Cannibalism and activity rate in larval damselflies increase along a latitudinal gradient as a consequence of time constraints.

BACKGROUND: Predation is ubiquitous in nature. One form of predation is cannibalism, which is affected by many factors such as size structure and resource density. However, cannibalism may also be influenced by abiotic factors such as seasonal time constraints. Since time constraints are greater at high latitudes, cannibalism could be stronger at such latitudes, but we know next to nothing about latitudinal variation in cannibalism. In this study, we examined cannibalism and activity in larvae of the damselfly Lestes sponsa along a latitudinal gradient across Europe. We did this by raising larvae from the egg stage at different temperatures and photoperiods corresponding to different latitudes. RESULTS: We found that the more seasonally time-constrained populations in northern latitudes and individuals subjected to greater seasonal time constraints exhibited a higher level of cannibalism. We also found that activity was higher at north latitude conditions, and thus correlated with cannibalism, suggesting that this behaviour mediates higher levels of cannibalism in time-constrained animals. CONCLUSIONS: Our results go counter to the classical latitude-predation pattern which predicts higher predation at lower latitudes, since we found that predation was stronger at higher latitudes. The differences in cannibalism might have implications for population dynamics along the latitudinal gradients, but further experiments are needed to explore this.

Physiological mechanisms of adaptive developmental plasticity in Rana temporaria island populations.

BACKGROUND: Adaptive plasticity is essential for many species to cope with environmental heterogeneity. In particular, developmental plasticity allows organisms with complex life cycles to adaptively adjust the timing of ontogenetic switch points. Size at and time to metamorphosis are reliable fitness indicators in organisms with complex cycles. The physiological machinery of developmental plasticity commonly involves the activation of alternative neuroendocrine pathways, causing metabolic alterations. Nevertheless, we have still incomplete knowledge about how these mechanisms evolve under environments that select for differences in adaptive plasticity. In this study, we investigate the physiological mechanisms underlying divergent degrees of developmental plasticity across Rana temporaria island populations inhabiting different types of pools in northern Sweden. METHODS: In a laboratory experiment we estimated developmental plasticity of amphibian larvae from six populations coming from three different island habitats: islands with only permanent pools, islands with only ephemeral pools, and islands with a mixture of both types of pools. We exposed larvae of each population to either constant water level or simulated pool drying, and estimated their physiological responses in terms of corticosterone levels, oxidative stress, and telomere length. RESULTS: We found that populations from islands with only temporary pools had a higher degree of developmental plasticity than those from the other two types of habitats. All populations increased their corticosterone levels to a similar extent when subjected to simulated pool drying, and therefore variation in secretion of this hormone does not explain the observed differences among populations. However, tadpoles from islands with temporary pools showed lower constitutive activities of catalase and glutathione reductase, and also showed overall shorter telomeres. CONCLUSIONS: The observed differences are indicative of physiological costs of increased developmental plasticity, suggesting that the potential for plasticity is constrained by its costs. Thus, high levels of responsiveness in the developmental rate of tadpoles have evolved in islands with pools at high but variable risk of desiccation. Moreover, the physiological alterations observed may have important consequences for both short-term odds of survival and long term effects on lifespan.

The Rate of Seasonal Changes in Temperature Alters Acclimation of Performance under Climate Change.

How the ability to acclimate will impact individual performance and ecological interactions under climate change remains poorly understood. Theory predicts that the benefit an organism can gain from acclimating depends on the rate at which temperatures change relative to the time it takes to induce beneficial acclimation. Here, we present a conceptual model showing how slower seasonal changes under climate change can alter species' relative performance when they differ in acclimation rate and magnitude. To test predictions from theory, we performed a microcosm experiment where we reared a mid- and a high-latitude damselfly species alone or together under the rapid seasonality currently experienced at 62°N and the slower seasonality predicted for this latitude under climate change and measured larval growth and survival. To separate acclimation effects from fixed thermal responses, we simulated growth trajectories based on species' growth rates at constant temperatures and quantified how much and how fast species needed to acclimate to match the observed growth trajectories. Consistent with our predictions, the results showed that the midlatitude species had a greater capacity for acclimation than the high-latitude species. Furthermore, since acclimation occurred at a slower rate than seasonal temperature changes, the midlatitude species had a small growth advantage over the high-latitude species under the current seasonality but a greater growth advantage under the slower seasonality predicted for this latitude under climate change. In addition, the two species did not differ in survival under the current seasonality, but the midlatitude species had higher survival under the predicted climate change scenario, possibly because rates of cannibalism were lower when smaller heterospecifics were present. These findings highlight the need to incorporate acclimation rates in ecological models.

A potential pitfall in studies of biological shape: Does size matter?

The number of published studies using geometric morphometrics (GM) for analysing biological shape has increased steadily since the beginning of the 1990s, covering multiple research areas such as ecology, evolution, development, taxonomy and palaeontology. Unfortunately, we have observed that many published studies using GM do not evaluate the potential allometric effects of size on shape, which normally require consideration or assessment. This might lead to misinterpretations and flawed conclusions in certain cases, especially when size effects explain a large part of the shape variation. We assessed, for the first time and in a systematic manner, how often published studies that have applied GM consider the potential effects of allometry on shape. We reviewed the 300 most recent published papers that used GM for studying biological shape. We also estimated how much of the shape variation was explained by allometric effects in the reviewed papers. More than one-third (38%) of the reviewed studies did not consider the allometric component of shape variation. In studies where the allometric component was taken into account, it was significant in 88% of the cases, explaining up to 87.3% of total shape variation. We believe that one reason that may cause the observed results is a misunderstanding of the process that superimposes landmark configurations, i.e. the Generalized Procrustes Analysis, which removes isometric effects of size on shape, but not allometric effects. Allometry can be a crucial component of shape variation. We urge authors to address, and report, size effects in studies of biological shape. However, we do not propose to always remove size effects, but rather to evaluate the research question with and without the allometric component of shape variation. This approach can certainly provide a thorough understanding of how much size contributes to the observed shaped variation.

Repeated pulse exposures to lambda-cyhalothrin affect the behavior, physiology, and survival of the damselfly larvae Ischnura graellsii (Insecta; Odonata).

Damselflies form an essential part of the aquatic and terrestrial food web. Pesticides may, however, negatively affect their behavior, physiology, and survival. To assess this, a 42-day-lasting bioassay was conducted, during which damselfly larvae (Ischnura graellsii; n = 20) were repeatedly exposed to lambda-cyhalothrin (3 days at; 0, 10, 50, 250, 1250, and 6250ng LCH L-1), followed by recovery phases (4 days) in pesticide-free medium for six weeks. This exposure design was used to simulate frequent runoff events in the field. Variables related to the behavior (strikes against prey and capture success), growth, physiology (lipid content and fatty acid composition), as well as mortality were assessed throughout the experiment. The two highest LCH concentrations induced 100% mortality within the first 48h, whereas 85% of the test organisms survived 28 days under control conditions. The number of strikes against prey was not affected by LCH. In contrast, prey capture success decreased significantly (up to ~50% at 250ng LCH L-1, for instance, after the third pulse exposure) following LCH-exposures compared to the control. This difference was not observed after recovery phases, however, which did not counteract the enhanced energy demand for detoxification and defense mechanisms indicated by a lower growth rate (up to ~20%) and lipid content (up to ~30%) of damselflies at 50 and 250ng LCH L-1. In addition, two essential fatty acids (eicosapentaenoic acid and arachidonic acid) and two precursors (linolenic acid and α-linolenic acid) decreased in their concentrations upon exposure towards 250ng LCH L-1. Thus the results of this study indicate that long-term exposure towards LCH pulses can affect damselfly behavior, physiology and survival. Given the essential role of damselflies in food web dynamics, these effects may potentially translate into local population impairments with subsequent bottom-up directed effects within and across ecosystem boundaries.

Seasonal time constraints reduce genetic variation in life-history traits along a latitudinal gradient.

Time constraints cause strong selection on life-history traits, because populations need to complete their life cycles within a shorter time. We therefore expect lower genetic variation in these traits in high- than in low-latitude populations, since the former are more time-constrained. The aim was to estimate life-history traits and their genetic variation in an obligately univoltine damselfly along a latitudinal gradient of 2730 km. Populations were grown in the laboratory at temperatures and photoperiods simulating those at their place of origin. In a complementary experiment, individuals from the same families were grown in constant temperature and photoperiod that mimicked average conditions across the latitude. Development time and size was faster and smaller, respectively, and growth rate was higher at northern latitudes. Additive genetic variance was very low for life-history traits, and estimates for egg development time and larval growth rate showed significant decreases towards northern latitudes. The expression of genetic effects in life-history traits differed considerably when individuals were grown in constant rather than simulated and naturally variable conditions. Our results support strong selection by time constraints. They also highlight the importance of growing organisms in their native environment for correct estimates of genetic variance at their place of origin. Our results also suggest that the evolutionary potential of life-history traits is very low at northern compared to southern latitudes, but that changes in climate could alter this pattern.

Combination effects of pyrethroids and neonicotinoids on development and survival of Chironomus riparius.

Standard ecotoxicological risk assessments are conducted on individual substances, however monitoring of streams in agricultural areas has shown that pesticides are rarely present alone. In fact, brief but intense pulse events such as storm water runoff and spray drift during application subject freshwater environments to complex mixtures of pesticides at high concentrations. This study investigates the potential risks to non-target aquatic organisms exposed to a brief but intense mixture of the neonicotinoid pesticides imidacloprid and thiacloprid and the pyrethroid pesticides deltamethrin and esfenvalerate, compared to single substance exposure. All four of these pesticides have been detected in surface waters at concentrations higher than benchmark values and both classes of pesticides are known to exert adverse effects on non-target aquatic organisms under single substance exposure scenarios. First instar midge larvae of the non-target aquatic organism, Chironomus riparius, were exposed to combinations of these four pesticides at 50% of their LC50 (96 h) values in a 1h pulse. They were then reared to adulthood in uncontaminated conditions and assessed for survival, development time and fecundity. Our results show that the risk of disruption to survival and development of non-target aquatic organisms under this scenario is not negligible on account of the significant increases in mortality of C. riparius found in the majority of the pesticide exposures and the delays in development after pyrethroid exposure. While none of the deleterious effects appear to be amplified by combination of the pesticides, there is some evidence for antagonism. No effects on fecundity by any of the pesticide treatments were observed.

Competitive interactions modify the temperature dependence of damselfly growth rates.

Individual growth rates and survival are major determinants of individual fitness, population size structure, and community dynamics. The relationships between growth rate, survival, and temperature may thus be important for predicting biological responses to climate change. Although it is well known that growth rates and survival are affected by competition and predation in addition to temperature, the combined effect of these factors on growth rates, survival, and size structure has rarely been investigated simultaneously in the same ecological system. To address this question, we conducted experiments on the larvae of two species of damselflies and determined the temperature dependence of growth rate, survival, and cohort size structure under three scenarios of increasing ecological complexity: no competition, intraspecific competition, and interspecific competition. In one species, the relationship between growth rate and temperature became steeper in the presence of competitors, whereas that of survival remained unchanged. In the other species, the relationship between growth rate and temperature was unaffected by competitive interactions, but survival was greatly reduced at high temperatures in the presence of interspecific competitors. The combined effect of competitive interactions and temperature on cohort size structure differed from the effects of these factors in isolation. Together, these findings suggest that it will be challenging to scale up information from single-species laboratory studies to the population and community level.

Effects of temperature and resource level on interspecific interactions in two species of Odonata larvae.

Identifying how temperature and food resources affect interactions between species is important for understanding how climate change will shape community structure in the future. Here, we tested how temperature and resource density affect survival and growth in the larval stage of two coexisting odonates: the damselfly Lestes sponsa and the dragonfly Sympetrum vulgatum. We performed a laboratory experiment at two temperatures (21 and 24°C) with two resource densities. We estimated the timing of egg hatching of individual egg clutches and thereafter the larval growth rate-, survival- and size-mediated priority effects under interspecific conditions. Eggs of both species hatched slightly faster at 24°C, and S. vulgatum eggs started hatching approximately 1 day earlier than L. sponsa eggs. However, this earlier hatching did not result in a size-mediated priority effect, that is, a higher predation on the later hatching L. sponsa. Nevertheless, L. sponsa larvae were significantly larger than S. vulgatum at hatching. Growth rate and survival were significantly higher: (1) at 24°C compared with 21°C, (2) at high compared with low-resource density and (3) in L. sponsa compared with S. vulgatum. Several significant interaction effects between resource density and temperature and between temperature and species were found. At high temperature, L. sponsa had a higher growth rate than S. vulgatum, but no difference in growth rate between species was found at low temperature. Additionally, a high-resource density resulted in a higher growth rate in both species, but only under high temperature. There was a negative relationship between growth rate and survival in both species, suggesting that the higher growth rate of larvae was to some degree driven by intraguild predation and/or cannibalism. Our results imply that resource levels interact with temperature to affect interactions between the species.

Wing shape allometry and aerodynamics in calopterygid damselflies: a comparative approach.

BACKGROUND: Wing size and shape have important aerodynamic implications on flight performance. We explored how wing size was related to wing shape in territorial males of 37 taxa of the damselfly family Calopterygidae. Wing coloration was also included in the analyses because it is sexually and naturally selected and has been shown to be related to wing shape. We studied wing shape using both the non-dimensional radius of the second moment of wing area (RSM) and geometric morphometrics. Lower values of the RSM result in less energetically demanding flight and wider ranges of flight speed. We also re-analyzed previously published data on other damselflies and dragonflies. RESULTS: The RSM showed a hump-shaped relationship with wing size. However, after correcting for phylogeny using independent contrast, this pattern changed to a negative linear relationship. The basal genus of the study family, Hetaerina, was mainly driving that change. The obtained patterns were specific for the study family and differed from other damselflies and dragonflies. The relationship between the RSM and wing shape measured by geometric morphometrics was linear, but relatively small changes along the RSM axis can result in large changes in wing shape. Our results also showed that wing coloration may have some effect on RSM. CONCLUSIONS: We found that RSM showed a complex relationship with size in calopterygid damselflies, probably as a result of other selection pressures besides wing size per se. Wing coloration and specific behavior (e.g. courtship) are potential candidates for explaining the complexity. Univariate measures of wing shape such as RSM are more intuitive but lack the high resolution of other multivariate techniques such as geometric morphometrics. We suggest that the relationship between wing shape and size are taxa-specific and differ among closely-related insect groups.

Latitudinally structured variation in the temperature dependence of damselfly growth rates.

The Metabolic Theory of Ecology predicts that the slope of the rate-temperature relationship, E, remains consistent across traits and organisms, acting as a major determinant of large-scale ecological patterns. Although E has recently been shown to vary systematically, we have a poor understanding of its ecological significance. To address this question, we conducted a common-garden experiment involving six damselfly species differing in distribution, estimating E at the level of full-sib families. Each species was sampled throughout its latitudinal range, allowing us to characterise variation in E along a latitudinal gradient spanning 3600 km. We show that E differs among populations and increases with latitude. E was right-skewness across species, but this was largely an artefact of the latitudinal trend. Increased seasonality towards higher latitude may contribute to the latitudinal trend in E. We conclude that E should be seen as a trait involved in local adaptation.

Phenotypic plasticity in the hepatic transcriptome of the European common frog (Rana temporaria): the interplay between environmental induction and geographical lineage on developmental response.

Phenotypic plasticity might facilitate adaptation to new environmental conditions through the enhancement of initial survival of organisms. Once a population is established, further adaptation and diversification may occur through adaptive trait evolution. While several studies have found evidence for this mechanism using phenotypic traits, much less is known at the level of gene expression. Here, we use an islands system of frog populations that show local adaptation and phenotypic plasticity to pool drying conditions in development time until metamorphoses. We examined gene expression differences in Rana temporaria tadpole livers with respect to pool drying at the source population and in response to simulated pool drying in the laboratory. Using a MAGEX cDNA microarray and quantitative real-time polymerase chain reaction (qPCR), we identified an increase in several gene transcripts in response to artificial pool drying including thyroid hormone receptor alpha and beta, carbamoyl phosphate synthetase 1, ornithine transcarbamylase and catalase. In addition, these gene transcripts also showed greater abundance in island populations that developed faster. Hence, the gene transcripts were related to both constitutive response (higher levels in island populations that developed faster) and plastic response (increased abundance under decreasing water levels). This pattern is in accordance with genetic accommodation, which predicts similarities between plastic gene expression and constitutive expression in locally adapted populations.

Generalists and specialists along a latitudinal transect: patterns of thermal adaptation in six species of damselflies.

Tropical organisms colonizing temperate environments face reduced average temperatures and dramatic thermal fluctuations. Theoretical models postulate that thermal specialization should be favored either when little environmental variation is experienced within generations or when among-generation variation is small relative to within-generation variation. To test these predictions, we studied six temperate species of damselflies differing in latitudinal distribution. We developed a computer model simulating how organisms experience environmental variation (accounting for diapause and voltinism) and performed a laboratory experiment assaying thermal sensitivities of growth rates. The computer model showed opposing latitudinal trends in among- and within-generation thermal variability: within-generation thermal variability decreased toward higher latitudes, whereas relative levels of among-generation thermal variability peaked at midlatitudes (where a shift in voltinism occurred). The growth experiment showed that low-latitude species were more thermally generalized than mid- and high-latitude species, supporting the prediction that generalists are favored under high levels of within-generation variation. Northern species had steeper, near-exponential reaction norms suggestive of thermal specialization. However, they had strikingly high thermal optima and grew very slowly over most of the thermal range they are expected to experience in the field. This observation is at present difficult to explain. These results highlight the importance of considering interactions between life history and environmental variation when deriving expectations of thermal adaptation.

Pushed Northward by Climate Change: Range Shifts With a Chance of Co-occurrence Reshuffling in the Forecast for Northern European Odonates.

Biodiversity is heavily influenced by ongoing climate change, which often results in species undergoing range shifts, either poleward or uphill. Range shifts can occur provided suitable habitats exist within reach. However, poleward latitudinal shifts might be limited by additional abiotic or biotic constraints, such as increased seasonality, photoperiod patterns, and species interactions. To gain insight into the dynamics of insect range shifts at high latitudes, we constructed ecological niche models (ENMs) for 57 Odonata species occurring in northern Europe. We used citizen science data from Sweden and present-day climatic variables covering a latitudinal range of 1,575 km. Then, to measure changes in range and interactions among Odonata species, we projected the ENMs up to the year 2080. We also estimated potential changes in species interactions using niche overlap and co-occurrence patterns. We found that most Odonata species are predicted to expand their range northward. The average latitudinal shift is expected to reach 1.83 and 3.25 km y-1 under RCP4.5 and RCP8.5 scenarios, respectively, by 2061-2080. While the most warm-dwelling species may increase their range, our results indicate that cold-dwelling species will experience range contractions. The present-day niche overlap patterns among species will remain largely the same in the future. However, our results predict changes in co-occurrence patterns, with many species pairs showing increased co-occurrence, while others will no longer co-occur because of the range contractions. In sum, our ENM results suggest that species assemblages of Odonata-and perhaps insects in general-in northern latitudes will experience great compositional changes.

Weak effects on growth and cannibalism under fluctuating temperatures in damselfly larvae.

The Earth's climate is changing with a trend towards higher mean temperatures and increased temperature fluctuations. Little attention has been paid to the effects of thermal variation on competition within species. Understanding the temperature-dependence of competition is important since it might affect dynamics within and between populations. In a laboratory experiment we investigated the effects of thermal variation on growth and cannibalism in larvae of a damselfly. The temperature treatments included three amplitudes between 20 and 26 °C with an average of 23 °C, and a constant control at 23 °C. Larvae were also raised at five constant temperatures for an estimation of the thermal performance curve, which showed that the thermal optimum for growth was 26.9 °C. Cannibalism was significantly positively correlated with initial body size variance. There was neither a difference among the temperature variation treatments, nor between the constant and the variation treatments in growth and cannibalism. Hence, positive and negative effects of temperature variation within the linear range of a species thermal performance curve might cancel each other out. Since our study mimicked natural temperature conditions, we suggest that the increase in temperature variation predicted by climate models will not necessarily differ from the effects without an increase in variation.

Phenological Shifts in a Warming World Affect Physiology and Life History in a Damselfly.

Under climate warming, temperate ectotherms are expected to hatch earlier and grow faster, increase the number of generations per season, i.e., voltinism. Here, we studied, under laboratory conditions, the impact of artificial warming and manipulated hatching dates on life history (voltinism, age and mass at emergence and growth rate) and physiological traits (phenoloxidase (PO) activity at emergence, as an indicator of investment in immune function) and larval survival rate in high-latitude populations of the damselfly Ischnura elegans. Larvae were divided into four groups based on crossing two treatments: early versus late hatching dates and warmer versus control rearing temperature. Damselflies were reared in groups over the course of one (univoltine) or two (semivoltine) growth seasons, depending on the voltinism. Warming temperature did not affect survival rate. However, warming increased the number of univoltine larvae compared to semivoltine larvae. There was no effect of hatching phenology on voltinism. Early hatched larvae reared under warming had elevated PO activity, regardless of their voltinism, indicating increased investment in immune function against pathogens. Increased PO activity was not associated with effects on age or mass at emergence or growth rate. Instead, life history traits were mainly affected by temperature and voltinism. Warming decreased development time and increased growth rate in univoltine females, yet decreased growth rate in univoltine males. This indicates a stronger direct impact of warming and voltinism compared to impacts of hatching phenology on life history traits. The results strengthen the evidence that phenological shifts in a warming world may affect physiology and life history in freshwater insects.