Climate change is increasing the frequency of heat waves and other extreme weather events experienced by organisms. How does the number and developmental timing of heat waves affect survival, growth and development of insects? Do heat waves early in development alter performance later in development? We addressed these questions using experimental heat waves with larvae of the tobacco hornworm, Manduca sexta. The experiments used diurnally fluctuating temperature treatments differing in the number (0-3) and developmental timing (early, middle and/or late in larval development) of heat waves, in which a single heat wave involved three consecutive days with a daily maximum temperature of 42°C. Survival to pupation declined with increasing number of heat waves. Multiple (but not single) heat waves significantly reduced development time and pupal mass; the best models for the data indicated that both the number and developmental timing of heat waves affected performance. In addition, heat waves earlier in development significantly reduced growth and development rates later in larval development. Our results illustrate how the frequency and developmental timing of sublethal heat waves can have important consequences for life history traits in insects.

Abstract Diurnal fluctuations in temperature are ubiquitous in terrestrial environments, and insects and other ectotherms have evolved to tolerate or acclimate to such fluctuations. Few studies have examined whether ectotherms acclimate to diurnal temperature fluctuations, or how natural and domesticated populations differ in their responses to diurnal fluctuations. We examine how diurnally fluctuating temperatures during development affect growth, acclimation, and stress responses for two populations of Manduca sexta: a field population that typically experiences wide variation in mean and fluctuations in temperature, and a laboratory population that has been domesticated in nearly constant temperatures for more than 300 generations. Laboratory experiments showed that diurnal fluctuations throughout larval development reduced pupal mass for the laboratory but not the field population. The differing effects of diurnal fluctuations were greatest at higher mean temperature (30°C): Here diurnal fluctuations reduced pupal mass and increased pupal development time for the laboratory population, but had little effect for the field population. We also evaluated how mean and fluctuations in temperature during early larval development affected growth rate during the final larval instar as a function of test temperature. At an intermediate (25°C) mean temperature, both the laboratory and field population showed a positive acclimation response to diurnal fluctuations, in which subsequent growth rate was significantly higher at most test temperatures. In contrast at higher mean temperature (30°C), diurnal fluctuations significantly reduced subsequent growth rate at most test temperatures for the laboratory population, but not for the field population. These results suggest that during domestication in constant temperatures, the laboratory population has lost the capacity to tolerate or acclimate to high and fluctuating temperatures. Population differences in acclimation capacity in response to temperature fluctuations have not been previously demonstrated, but they may be important for understanding the evolution of reaction norms and performance curves.

In invaded environments, formerly reliable cues might no longer be associated with adaptive outcomes and organisms can become trapped by their evolved responses. The invasion of Alliaria petiolata (garlic mustard) into the native habitat of Pieris virginiensis (West Virginia White) is one such example. Female butterflies oviposit on the invasive plant because it is related to their preferred native host plant Cardamine diphylla (toothwort), but larvae are unable to complete development. We have studied the impact of the A. petiolata invasion on P. virginiensis butterflies in the Southeastern USA by comparing oviposition preference and larval survival on both plants in North Carolina (NC) populations without A. petiolata and West Virginia (WV) populations where A. petiolata is present. Larval survival to the 3rd instar was equally low in both populations when raised on A. petiolata. Mean oviposition preference on the two plants also did not differ between populations. However, we found a seasonal effect on preference between early and late season flights within WV populations. Late season females laid 99% of total eggs on A. petiolata while early season females utilized both host plants. Late season females were also less likely to lay eggs than early season females. This change in preference toward A. petiolata could be driven by the early senescence of C. diphylla and suggests a seasonal component to the impact of A. petiolata. Therefore, the already short flight season of P. virginiensis could become further constrained in invaded populations.

Siepielski et al. (Reports, 3 March 2017, p. 959) claim that "precipitation drives global variation in natural selection." This conclusion is based on a meta-analysis of the relationship between climate variables and natural selection measured in wild populations of invertebrates, plants, and vertebrates. Three aspects of this analysis cause concern: (i) lack of within-year climate variables, (ii) low and variable estimates of covariance relationships across taxa, and (iii) a lack of mechanistic explanations for the patterns observed; association is not causation.

Climate change has the potential to affect the ecology and evolution of every species on Earth. Although the ecological consequences of climate change are increasingly well documented, the effects of climate on the key evolutionary process driving adaptation-natural selection-are largely unknown. We report that aspects of precipitation and potential evapotranspiration, along with the North Atlantic Oscillation, predicted variation in selection across plant and animal populations throughout many terrestrial biomes, whereas temperature explained little variation. By showing that selection was influenced by climate variation, our results indicate that climate change may cause widespread alterations in selection regimes, potentially shifting evolutionary trajectories at a global scale.

Although many selection estimates have been published, the environmental factors that cause selection to vary in space and time have rarely been identified. One way to identify these factors is by experimentally manipulating the environment and measuring selection in each treatment. We compiled and analyzed selection estimates from experimental studies. First, we tested whether the effect of manipulating the environment on selection gradients depends on taxon, trait type, or fitness component. We found that the effect of manipulating the environment was larger when selection was measured on life-history traits or via survival. Second, we tested two predictions about the environmental factors that cause variation in selection. We found support for the prediction that variation in selection is more likely to be caused by environmental factors that have a large effect on mean fitness but not for the prediction that variation is more likely to be caused by biotic factors. Third, we compared selection gradients from experimental and observational studies. We found that selection varied more among treatments in experimental studies than among spatial and temporal replicates in observational studies, suggesting that experimental studies can detect relationships between environmental factors and selection that would not be apparent in observational studies.

In many ectotherms, exposure to high temperatures can improve subsequent tolerance to higher temperatures. However, the differential effects of single, repeated, or continuous exposure to high temperatures are less clear. We measured the effects of single heat shocks and of diurnally fluctuating or constant rearing temperatures on the critical thermal maximum temperatures (CTmax) for final instar larvae of Manduca sexta. Brief (2h) heat shocks at temperatures of 35°C and above significantly increased CTmax relative to control temperatures (25°C). Increasing mean temperatures (from 25 to 30°C) or greater diurnal fluctuations (from constant to ±10°C) during larval development also significantly increased CTmax. Combining these data showed that repeated or continuous temperature exposure during development improved heat tolerance beyond the effects of a single exposure to the same maximum temperature. These results suggest that both acute and chronic temperature exposure can result in adaptive plasticity of upper thermal limits.

ABSTRACT Most terrestrial ectotherms experience diurnal and seasonal variation in temperature. Because thermal performance curves are non-linear, mean performance can differ in fluctuating and constant thermal environments. However, time-dependent effects – effects of the order and duration of exposure to temperature – can also influence mean performance. We quantified the non-linear and time-dependent effects of diurnally fluctuating temperatures for larval growth rates in the tobacco hornworm, Manduca sexta L., with four main results. First, the shape of the thermal performance curve for growth rate depended on the duration of exposure: for example, optimal temperature and thermal breadth were greater for growth rates measured over short (24 h during the last instar) compared with long (the entire period of larval growth) time periods. Second, larvae reared in diurnally fluctuating temperatures had significantly higher optimal temperatures and maximal growth rates than larvae reared in constant temperatures. Third, for larvae maintained at three mean temperatures (20, 25 and 30°C) and three diurnal temperature ranges (±0, ±5 and ±10°C), diurnal fluctuations had opposite effects on mean growth rates at low versus high mean temperature. Fourth, both short- and long-term thermal performance curves yielded poor predictions of the non-linear effects of fluctuating temperature on mean growth rates (compared with our experimental results) at higher mean temperatures. Our results suggest caution in using constant temperature studies to model the consequences of variable thermal environments. Summary: Growth rates measured at constant temperatures do not correctly predict mean growth under fluctuating conditions because of both non-linear and time-dependent effects. This has important implications for modeling ecological responses to climate change.