Calling behaviour under climate change: geographical and seasonal variation of calling temperatures in ectotherms

Calling behaviour is strongly temperature‐dependent and critical for sexual selection and reproduction in a variety of ectothermic taxa, including anuran amphibians, which are the most globally threatened vertebrates. However, few studies have explored how species respond to distinct thermal environments at time of displaying calling behaviour, and thus it is still unknown whether ongoing climate change might compromise the performance of calling activity in ectotherms. Here, we used new audio‐trapping techniques (automated sound recording and detection systems) between 2006 and 2009 to examine annual calling temperatures of five temperate anurans and their patterns of geographical and seasonal variation at the thermal extremes of species ranges, providing insights into the thermal breadths of calling activity of species, and the mechanisms that enable ectotherms to adjust to changing thermal environments. All species showed wide thermal breadths during calling behaviour (above 15 °C) and increases in calling temperatures in extremely warm populations and seasons. Thereby, calling temperatures differed both geographically and seasonally, both in terrestrial and aquatic species, and were 8–22 °C below the specific upper critical thermal limits (CT_max) and strongly associated with the potential temperatures of each thermal environment (operative temperatures during the potential period of breeding). This suggests that calling behaviour in ectotherms may take place at population‐specific thermal ranges, diverging when species are subjected to distinct thermal environments, and might imply plasticity of thermal adjustment mechanisms (seasonal and developmental acclimation) that supply species with means of coping with climate change. Furthermore, the thermal thresholds of calling at the onset of the breeding season were dissimilar between conspecific populations, suggesting that other factors besides temperature are needed to trigger the onset of reproduction. Our findings imply that global warming would not directly inhibit calling behaviour in the study species, although might affect other temperature‐dependent features of their acoustic communication system.     

Physiological and behavioural responses to seasonal changes in environmental temperature in the Australian spiny crayfish Euastacus sulcatus

The strategies used by ectotherms to minimise the detrimental effects of suboptimal thermal environments on physiological performance are often related to whether they inhabit a terrestrial or aquatic environment. Most terrestrial ectotherms use thermoregulatory strategies to maintain body temperature within an optimal range, while many aquatic ectotherms utilise thermal acclimation to maintain performance over varying seasonal temperatures. This study aimed to elucidate the relative contributions of acclimation and behavioural thermoregulation to maintaining whole-animal performance over varying seasonal temperatures in the semi-terrestrial Lamington spiny crayfish (Euastacus sulcatus). Crayfish activity and surface temperatures were determined by means of radio tracking and behavioural observations. Field studies demonstrated that E. sulcatus is exposed to stable daily temperatures, varying only between seasons from 10°C in late winter to over 20°C in summer. Also, terrestrial behaviour corresponded to a small portion of crayfish time (1.13%), much lower than predicted, indicating that E. sulcatus has limited opportunity for behavioural thermoregulation. We also tested the effect of acclimation to either 10 or 20°C on chela strength and stamina. We found acclimation had a more marked effect on chela stamina than maximum strength measures; however, overall acclimatory capacity was limited in E. sulcatus. Thus, we found that the semi-terrestrial crayfish E. sulcatus used neither thermoregulatory behaviours nor physiological strategies to deal with seasonal changes in environmental temperature.     

Temperature-induced plasticity in membrane and storage lipid composition: thermal reaction norms across five different temperatures

Temperature is a key environmental factor inducing phenotypic plasticity in a wide range of behavioral, morphological, and life history traits in ectotherms. The strength of temperature-induced responses in fitness-related traits may be determined by plasticity of the underlying physiological or biochemical traits. Lipid composition may be an important trait underlying fitness response to temperature, because it affects membrane fluidity as well as availability of stored energy reserves. Here, we investigate the effect of temperature on lipid composition of the springtail Orchesella cincta by measuring thermal reaction norms across five different temperatures after four weeks of cold or warm acclimation. Fatty acid composition in storage and membrane lipids showed a highly plastic response to temperature, but the responses of single fatty acids revealed deviations from the expectations based on HVA theory. We found an accumulation of C_18:2n6 and C_18:3n3 at higher temperatures and the preservation of C_20:4n6 across temperatures, which is contrary to the expectation of decreased unsaturation at higher temperatures. The thermal response of these fatty acids in O. cincta differed from the findings in other species, and therefore shows there is interspecific variation in how single fatty acids contribute to HVA. Future research should determine the consequences of such variation in terms of costs and benefits for the thermal performance of species.     

Latitudinal comparison of the thermal biology in the endemic lizard Liolaemus multimaculatus

Thermoregulation in ectotherms may be modulated by climatic variability across geographic gradients. Environmental temperature varies along latitudinal clines resulting in heterogeneous thermal resource availability, which generally induces ectotherms to use compensatory mechanisms to thermoregulate. Lizards can accommodate to ambient temperature changes through a combination of adaptive evolution and behavioral and physiological plasticity. We studied the thermal ecology of the endangered endemic lizard Liolaemus multimaculatus at six different sites distributed from the northern to southern areas of the distribution (700 km) in the Atlantic dune barriers of Argentina, and even including the borders areas of the distribution range. Environmental temperatures and relative humidity showed a strong contrast between northern and southern limits of the distribution range. The northern localities had operative temperatures (Te) above the range of preferred temperatures (Tset), instead, the southern localities had large proportion of Tes within the Tset. Although these different climatic conditions may constrain the thermal biology of L. multimaculatus, individuals from all localities maintained relatively similar field body temperatures (XTb = 34.07 ± 3.02 °C), suggesting that this parameter is conservative. Thermal preference partially reflected latitudinal temperature gradient, since lizards from the two southernmost localities showed the lowest Tsel and Tset. Thermoregulatory efficiency differed among localities, since E values in the northern localities (E = 0.53–0.69) showed less variability than those of southern localities (E = 0.14–0.67). Although L. multimaculatus employed a strategy of having a conservative Tb and being able to acclimatize the thermal preference to copes with latitudinal changes in the thermal environment, other local factors, such as ecological interactions, may also impose limitations to thermoregulation and this may interfered in the interpretation of results at wider spatial scale.     

Developmental plasticity of thermal tolerances in temperate and subtropical populations of Drosophila melanogaster

Variation in temperature imposes selection pressures on organisms. In variable environments, organisms must adopt fixed or plastic strategies that enable persistence over a broad range of temperatures. In coarse-grained environments, where the thermal variation among generations exceeds that within generations, selection should favor developmental plasticity. Here, we compare the degree of developmental plasticity of thermal tolerances between populations of Drosophila melanogaster from environments with relatively high (Marlton, NJ, USA) and relatively low (Miami, FL, USA) variance in temperature among generations. We predicted that flies from Marlton would exhibit a greater plasticity of thermal tolerances than would flies from Miami. Flies from both populations were reared in three ecologically relevant treatments, after which we assessed knockdown and chill-coma recovery times. Flies from both populations responded plastically to temperature, but flies from New Jersey did not exhibit greater plasticity. Our results complement previous comparative studies and indicate that selection favors plasticity of thermal tolerances equally in these populations.     

Individual variation in thermal plasticity and its impact on mass-scaling

Physiological processes vary widely across individuals and can influence how individuals respond to environmental change. Repeatability in how metabolic rate changes across temperatures (i.e. metabolic thermal plasticity) can influence mass-scaling exponents in different thermal environments. Moreover, repeatable plastic responses are necessary for reaction norms to respond to selective forces which is important for populations living in fluctuating environments. Nonetheless, only a small number of studies have explicitly quantified repeatability in metabolic plasticity, and fewer have explored how it can impact mass-scaling. We repeatedly measured standard metabolic rate of n = 42 delicate skinks Lampropholis delicata at six temperatures over the course of four months (N_{[observations]} = 4952). Using hierarchical statistical techniques, we accounted for multi-level variation and measurement error in our data in order to obtain more precise estimates of reaction norm repeatability and mass-scaling exponents at different acute temperatures. Our results show that individual differences in metabolic thermal plasticity were somewhat consistent over time (R_slope = 0.25, 95% CI = 2.48 × 10⁻⁸ – 0.67), however estimates were associated with a large degree of error. After accounting for measurement error, which decreased steadily with temperature, we show that among individual variance remained consistent across all temperatures. Congruently, temperature specific repeatability of average metabolic rate was stable across temperatures. Cross-temperature correlations were positive but were not uniform across the reaction norm. After taking into account multiple sources of variation, our estimates for mass-scaling did not change with temperature and were in line with published values for snakes and lizards. This implies that repeatable plastic responses may promote thermal stability of scaling exponents. Our work contributes to understanding how energy expenditure scales with abiotic and biotic factors and the capacity for reaction norms to respond to selection.     

Thermal plasticity and microevolution enhance establishment success and persistence of a water hyacinth biological control agent

Aspects of the thermal physiology of the water hyacinth biological control agent Eccritotarsus catarinensis Carvalho (Hemiptera: Miridae) have been extensively investigated over the past 20 years to understand and improve post‐release establishment in the field. Thermal physiology studies predicted that the agent would not establish at a number of cold sites in South Africa, where it has nonetheless subsequently established and thrived. Recently, studies have begun to incorporate the plastic nature of insect thermal physiology into models of agent establishment. This study determined whether season and locality influenced the thermal physiology of two field populations of E. catarinensis, one collected from the hottest site where the agent has established in South Africa, and one from the coldest site. The thermal physiology of E. catarinensis was significantly influenced by season and site, demonstrating a degree of phenotypic plasticity, and that some post‐release local adaptation to climatic conditions has occurred through microevolution. We then determined whether cold acclimation under laboratory conditions was possible. Successfully cold‐acclimated E. catarinensis had a significantly lower critical thermal minimum (CT_min) compared to the field cold‐acclimated population. This suggests that cold acclimation of agents could be conducted in the laboratory before future releases to improve their cold tolerance, thereby increasing their chance of establishment at cold sites and allowing further adaptation to colder climates to occur in the field. Although the thermal tolerance of E. catarinensis is limited by local adaptations to climatic conditions in the native range, the plastic nature of the insect's thermal physiology has allowed it to survive in the very different climatic conditions of the introduced range, and there has been some adaptive change to the insect's thermal tolerance since establishment. This study highlights the importance of plasticity and microevolutionary processes in the success of biological control agents under the novel climatic conditions in the introduced range.     

Genetic correlation between temperature-induced plasticity of life-history traits in a soil arthropod

Temperature is considered one of the most important mediators of phenotypic plasticity in ectotherms. However, the costs and benefits shaping the evolution of different thermal responses are poorly elucidated. One of the possible constraints to phenotypic plasticity is its intrinsic genetic cost, such as genetic linkage or pleiotropy. Genetic coupling of the thermal response curves for different life history traits may significantly affect the evolution of thermal sensitivity in thermally fluctuating environments. We used the collembolan Orchesella cincta to study if there is genetic variation in temperature-induced phenotypic plasticity in life history traits, and if the degree of temperature-induced plasticity is correlated across traits. Egg development rate, juvenile growth rate and egg size of 19 inbred isofemale lines were measured at two temperatures. Our results show that temperature was a highly significant factor for all three traits. Egg development rate and juvenile growth rate increased with increasing temperature, while egg size decreased. Line by temperature interaction was significant for all traits tested; indicating that genetic variation for temperature-induced plasticity existed. The degree of plasticity was significantly positively correlated between egg development rate and growth rate, but plasticity in egg size was not correlated to the other two plasticity traits. The findings suggest that the thermal plasticities of egg development rate and growth rate are partly under the control of the same genes or genetic regions. Hence, evolution of the thermal plasticity of traits cannot be understood in isolation of the response of other traits. If traits have similar and additive effects on fitness, genetic coupling between these traits may well facilitate the evolution of optimal phenotypes. However, for this we need to know the selective forces under field conditions.     

Coping with temperature at the warm edge--patterns of thermal adaptation in the microbial eukaryote Paramecium caudatum

Background

Ectothermic organisms are thought to be severely affected by global warming since their physiological performance is directly dependent on temperature. Latitudinal and temporal variations in mean temperatures force ectotherms to adapt to these complex environmental conditions. Studies investigating current patterns of thermal adaptation among populations of different latitudes allow a prediction of the potential impact of prospective increases in environmental temperatures on their fitness.

Methodology/Principal Findings

In this study, temperature reaction norms were ascertained among 18 genetically defined, natural clones of the microbial eukaryote Paramecium caudatum. These different clones have been isolated from 12 freshwater habitats along a latitudinal transect in Europe and from 3 tropical habitats (Indonesia). The sensitivity to increasing temperatures was estimated through the analysis of clone specific thermal tolerances and by relating those to current and predicted temperature data of their natural habitats.All investigated European clones seem to be thermal generalists with a broad thermal tolerance and similar optimum temperatures. The weak or missing co-variation of thermal tolerance with latitude does not imply local adaptation to thermal gradients; it rather suggests adaptive phenotypic plasticity among the whole European subpopulation. The tested Indonesian clones appear to be locally adapted to the less variable, tropical temperature regime and show higher tolerance limits, but lower tolerance breadths.

Conclusions/Significance

Due to the lack of local temperature adaptation within the European subpopulation, P. caudatum genotypes at the most southern edge of their geographic range seem to suffer from the predicted increase in magnitude and frequency of summer heat waves caused by climate change.     

Temperature, growth rate, and body size in ectotherms: fitting pieces of a life-history puzzle

The majority of ectotherms grow slower but mature at a larger body size in colder environments. This phenomenon has puzzled biologists because classic theories of life-history evolution predict smaller sizes at maturity in environments that retard growth. During the last decade, intensive theoretical and empirical research has generated some plausible explanations based on nonadaptive or adaptive plasticity. Nonadaptive plasticity of body size is hypothesized to result from thermal constraints on cellular growth that cause smaller cells at higher temperatures, but the generality of this theory is poorly supported. Adaptive plasticity is hypothesized to result from greater benefits or lesser costs of delayed maturation in colder environments. These theories seem to apply well to some species but not others. Thus, no single theory has been able to explain the generality of temperature-size relationships in ectotherms. We recommend a multivariate theory that focuses on the coevolution of thermal reaction norms for growth rate and size at maturity. Such a theory should incorporate functional constraints on thermal reaction norms, as well as the natural covariation between temperature and other environmental variables.     

Plasticity in thermal tolerance has limited potential to buffer ectotherms from global warming

Global warming is increasing the overheating risk for many organisms, though the potential for plasticity in thermal tolerance to mitigate this risk is largely unknown. In part, this shortcoming stems from a lack of knowledge about global and taxonomic patterns of variation in tolerance plasticity. To address this critical issue, we test leading hypotheses for broad-scale variation in ectotherm tolerance plasticity using a dataset that includes vertebrate and invertebrate taxa from terrestrial, freshwater and marine habitats. Contrary to expectation, plasticity in heat tolerance was unrelated to latitude or thermal seasonality. However, plasticity in cold tolerance is associated with thermal seasonality in some habitat types. In addition, aquatic taxa have approximately twice the plasticity of terrestrial taxa. Based on the observed patterns of variation in tolerance plasticity, we propose that limited potential for behavioural plasticity (i.e. behavioural thermoregulation) favours the evolution of greater plasticity in physiological traits, consistent with the ‘Bogert effect’. Finally, we find that all ectotherms have relatively low acclimation in thermal tolerance and demonstrate that overheating risk will be minimally reduced by acclimation in even the most plastic groups. Our analysis indicates that behavioural and evolutionary mechanisms will be critical in allowing ectotherms to buffer themselves from extreme temperatures.     

Preferred temperature of intertidal ectotherms: Broad patterns and methodological approaches

Preferred temperature (Tpref) has been measured in over 100 species of aquatic and 300 species of terrestrial ectotherms as a metric for assessing behavioural thermoregulation in variable environments and, as such, has been linked to ecological processes ranging from individual behaviour to population and community dynamics. Due to the asymmetric shape of performance curves, Tpref is typically lower than the optimal temperature (Topt, where physiological performance is at its peak), and the degree of this mismatch increases with variability in Tb. Intertidal ectotherms experience huge variability in Tb on a daily basis and therefore provide a good system to test whether the relationship between Tpref and variation in Tb holds in more extreme environments. A review of the literature, however, only revealed comparisons between Tpref and Topt for five intertidal species and measurements of Tpref for 23 species. An analysis of this limited literature for intertidal ectotherms showed a positive relationship between acclimation temperature and Tpref. There was, however, great variation in the methodologies employed to make these assessments. Factors contributing to behavioural thermoregulation in intertidal ectotherms including small body size; low mobility; interactions among individuals; endogenous clocks; metabolic effects; thermal sensitivity; sampling of the thermal environment and recent acclimation history were considered to varying degrees when measuring Tpref, confounding comparisons between species. The methodologies used to measure Tpref in intertidal ectotherms were reviewed in light of each of these factors, and methodologies proposed to standardize approaches. Given the theoretical predictions about the relationships between Tpref and variability in Tb, the spatial and temporal thermal variability experienced by intertidal ectotherms provides numerous opportunities to test these expectations if assessed in a standardized manner, and can potentially provide insights into the value of behavioural thermoregulation in the more thermally variable environments predicted to occur in the near future.     

Phenotypic flexibility in the metabolic response of the limpet Cellana tramoserica to thermally different microhabitats

Temperature determines all physiological responses by limiting cellular reaction rates. Daily temperature variation differs between microhabitats, which means that subpopulations of the same species may respond differently to temperature. The aim of this study is to determine how physiological responses to temperature of the limpet Cellana tramoserica differ between limpets from variable and from stable thermal environments. Oxygen consumption and anaerobic and aerobic metabolic capacities were measured over a range of temperatures in limpets from thermally stable and variable field sites in summer and winter, and in laboratory acclimation treatments. Limpets from both variable and stable sites, showed acclimatisation of anaerobic and aerobic potentials. Limpets from stable environments, but not from variable environments, showed increased oxygen consumption in winter. Comparison of field and laboratory data showed that temperature was the signal for acclimatisation. The physiological response of C. tramoserica to temperature depends on season and microhabitat. Care must therefore be taken when conducting interspecies comparisons of response to temperature to address the confounding effects of phenotypic plasticity. Differences in physiological response to temperature in phenotypically flexible species like C. tramoserica may simply reflect individual reactions to immediate environmental conditions.     

Ecophysiological variation across a forest‐ecotone gradient produces divergent climate change vulnerability within species

Climate change related risks and impacts on ectotherms will be mediated by habitats and their influence on local thermal environments. While many studies have documented morphological and genetic aspects of niche divergence across habitats, few have examined thermal performance across such gradients and directly linked this variation to contemporary climate change impacts. In this study, we quantified variation in thermal performance across a gradient from forest to gallery forest‐savanna mosaic in Cameroon for a skink species (Trachylepis affinis) known to be diverging genetically and morphologically across that habitat gradient. Based on these results, we then applied a mechanistic modelling approach (NicheMapR) to project changes in potential activity, as constrained by thermal performance, in response to climate change. As a complimentary approach, we also compared mechanistic projections with climate‐driven changes in habitat suitability based on species distribution models of forest and ecotone skinks. We found that ecotone skinks may benefit from warming and experience increased activity while forest skinks will likely face a drastic decrease in thermal suitability across the forest zone. Species distribution models projected that thermal suitability for forest skinks in coastal forests would decline but in other parts of the forest zone skinks are projected to experience increased thermal suitability. The results here highlight the utility of mechanistic approaches in revealing and understanding patterns of climate change vulnerability which may not be detected with species distribution models alone. This study also emphasizes the importance of intra‐specific physiological variation, and habitat‐specific thermal performance relationships in particular, in determining warming responses.     

Living on the edge: Lower thermal quality but greater survival probability at a high altitude mountain for the mesquite lizard (Sceloporus grammicus)

A 20-month recapture analysis of 1001 individually marked mesquite lizards (Sceloporus grammicus) suggests that variation in thermal quality across three altitudes influences survival probability. Each additional unit of deviation from the temperature selected by these lizards in previous laboratory experiments (i.e. decreased thermal quality) meant an increase of roughly 1.01% in survival probability. Survival probabilities ranged from 0.80 to 0.90 at the lowest elevation site (2600 m), from 0.76 to 0.87 at the middle elevation site (3100 m) and from 0.90 to 0.94 at the highest elevation site (4150 m). These results suggest that in poor thermal quality environments mesquite lizards may employ thermoregulatory strategies (behavioral, physiological and/or morphological) to decrease their metabolic expenditure and their exposure to predators, maximizing survival. These findings highlight the relevance of thermal quality of the habitat in determining survival probability of ectotherms.     

Seasonal variation in basal and plastic cold tolerance: Adaptation is influenced by both long‐ and short‐term phenotypic plasticity

Understanding how thermal selection affects phenotypic distributions across different time scales will allow us to predict the effect of climate change on the fitness of ectotherms. We tested how seasonal temperature variation affects basal levels of cold tolerance and two types of phenotypic plasticity in Drosophila melanogaster. Developmental acclimation occurs as developmental stages of an organism are exposed to seasonal changes in temperature and its effect is irreversible, while reversible short‐term acclimation occurs daily in response to diurnal changes in temperature. We collected wild flies from a temperate population across seasons and measured two cold tolerance metrics (chill‐coma recovery and cold stress survival) and their responses to developmental and short‐term acclimation. Chill‐coma recovery responded to seasonal shifts in temperature, and phenotypic plasticity following both short‐term and developmental acclimation improved cold tolerance. This improvement indicated that both types of plasticity are adaptive, and that plasticity can compensate for genetic variation in basal cold tolerance during warmer parts of the season when flies tend to be less cold tolerant. We also observed a significantly stronger trade‐off between basal cold tolerance and short‐term acclimation during warmer months. For the longer‐term developmental acclimation, a trade‐off persisted regardless of season. A relationship between the two types of plasticity may provide additional insight into why some measures of thermal tolerance are more sensitive to seasonal variation than others.     

Mitigating thermal effect of behaviour and microhabitat on the intertidal snail Littorina saxatilis (Olivi) over summer

High shore intertidal ectotherms must withstand temperatures which are already close, at or beyond their upper physiological thermal tolerance. Their behaviour can provide a relief under heat stress, and increase their survival through thermoregulation. Here, we used infrared imaging to reveal the thermoregulatory behavioural strategies used by the snail Littorina saxatilis (Olivi) on different microhabitats of a high shore boulder field in Finistère (western France) in summer. On our study site, substrate temperature is frequently greater than L. saxatilis upper physiological thermal limits, especially on sun exposed microhabitats. To maintain body temperatures within their thermal tolerance window, withdrawn snails adopted a flat posture, or elevated their shells and kept appended to the rock on the outer lip of their aperture with dried mucous (standing posture). These thermal regulatory behaviours lowered snail body temperatures on average by 1–2 °C. Aggregation behaviour had no thermoregulatory effect on L. saxatilis in the present study. The occupation of biogenic microhabitats (barnacles) was associated with a 1 °C decrease in body temperatures. Barnacles and microhabitats that experienced low sun exposure, low thermal fluctuations and low thermal maxima, could buffer the heat extremes encountered at high shore level especially on sun exposed microhabitats.     

Ontogenetic Variation in the Thermal Biology of Yarrow's Spiny Lizard,Sceloporus jarrovii

Climate change is rapidly altering the way current species interact with their environment to satisfy life-history demands. In areas anticipated to experience extreme warming, rising temperatures are expected to diminish population growth, due either to environmental degradation, or the inability to tolerate novel temperature regimes. Determining how at risk ectotherms, and lizards in particular, are to changes in climate traditionally emphasizes the thermal ecology and thermal sensitivity of physiology of adult members of a population. In this study, we reveal ontogenetic differences in thermal physiological and ecological traits that have been used to anticipate how ectotherms will respond to climate change. We show that the thermal biological traits of juvenile Yarrow’s Spiny Lizards (Sceloporus jarrovii) differ from the published estimates of the same traits for adult lizards. Juvenile S. jarrovii differ in their optimal performance temperature, field field-active body temperature, and critical thermal temperatures compared to adult S. jarrovii. Within juvenile S. jarrovii, males and females exhibit differences in field-active body temperature and desiccation tolerance. Given the observed age- and sex-related variation in thermal physiology, we argue that not including physiological differences in thermal biology throughout ontogeny may lead to misinterpretation of patterns of ecological or evolutionary change due to climate warming. Further characterizing the potential for ontogenetic changes in thermal biology would be useful for a more precise and accurate estimation of the role of thermal physiology in mediating population persistence in warmer environments.     

TEMPORAL VARIATION FAVORS THE EVOLUTION OF GENERALISTS IN EXPERIMENTAL POPULATIONS OF DROSOPHILA MELANOGASTER

In variable environments, selection should favor generalists that maintain fitness across a range of conditions. However, costs of adaptation may generate fitness trade‐offs and lead to some compromise between specialization and generalization that maximizes fitness. Here, we evaluate the evolution of specialization and generalization in 20 populations of Drosophila melanogaster experimentally evolved in constant and variable thermal environments for 3 years. We developed genotypes from each population at two temperatures after which we measured fecundity across eight temperatures. We predicted that constant environments would select for thermal specialists and that variable environments would select for thermal generalists. Contrary to our predictions, specialists and generalists did not evolve in constant and spatially variable environments, respectively. However, temporal variation produced a type of generalist that has rarely been considered by theoretical models of developmental plasticity. Specifically, genotypes from the temporally variable selective environment were more fecund across all temperatures than were genotypes from other environments. These patterns suggest certain allelic effects and should inspire new directions for modeling adaptation to fluctuating environments.