Dividing sister pairs of the butterfly Bicyclus anynana (reared in a common environment) between high and low temperature shows that oviposition temperature induces a plastic response in egg size. Females at a lower temperature laid significantly larger (but fewer) eggs than their sisters kept at a higher temperature, whereas total reproductive investment increased with temperature. Cross‐transfer experiments demonstrated that this plastic response in egg size is reversible. Interestingly, this pattern parallels an almost universal temperature‐induced developmental response in ectotherm body size. In both cases, however, we do not yet understand the underlying mechanisms or the potential adaptive significance. By cross‐transferring the experimentally manipulated eggs between temperatures, we showed that the larger eggs produced at a lower temperature had a higher hatching success, and yielded larger hatchlings with a slightly higher probability of reaching maturity and shorter larval development time (at the lower temperature) compared to the smaller eggs produced at a higher temperature. Overall, the lower temperature was slightly more detrimental for smaller than for larger eggs. The slight differences in survival across temperatures, together with more substantial differences in fecundity, indicate that it could pay off to produce fewer but larger offspring (with increased fitness) at a lower temperature, but more and smaller offspring at a higher temperature where offspring survival was generally high. Our data present for the first time evidence that selection may favor larger eggs at a lower temperature, and thus that a plastic response of egg size to temperature in ectotherms could be adaptive.

Although olfaction is a primary mode of communication, its importance in sexual selection remains understudied. Here, using the butterfly Bicyclus anynana, we address all the parameters of importance to sexual selection for a male olfactory signal. We show that variation in the male sex pheromone composition indicates male identity and male age. Courting males of different ages display small absolute (c. 200 ng) but large relative (100%) change of one specific pheromone component (hexadecanal) which, unlike the other components, showed no heritability. Females prefer to mate with mid-aged over younger males and the pheromone composition is sufficient to determine this preference. Surprisingly refined information is thus present in the male olfactory signal and is used for sexual selection. Our data also reveal that there may be no 'lek paradox' to resolve once the precise signal of importance to females is identified, as hexadecanal is, as expected, depleted in additive genetic variation.

K. 2005. Effects of food stress and density in different life stages on reproduction in a butterfly. Á/ Oikos 111: 514 Á/524.Availability of adequate nutrition and (rearing) density are among the most important factors affecting growth, development and reproduction in animals. In holometabolous insects diets and energetic needs change between life stages, with storing of larval resources, adult feeding and reproduction being linked strategies. Nevertheless, studies investigating nutritional (and density) effects across metamorphic boundaries are largely lacking. We aim at disentangling the functional basis of reproductive patterns by independently manipulating larval and adult (1) density and (2) access to food, respectively, in the tropical butterfly, Bicyclus anynana . (1) A high larval rearing density had, contrary to common wisdom, very little impact on body size, but reduced larval development time through increased growth rates. The latter is thought to be an adaptation to high densities, driven by the risk of larval food resources becoming exhausted before reaching metamorphosis. Larval density and male company during oviposition (i.e. adult density) had no detectable effects on female reproduction. (2) Larval food stress prolonged larval development time and reduced larval growth rate, body size, fecundity and reproductive investment. Detrimental effects on female reproduction were mediated through a reduction in body size. Additional negative effects of adult food stress on fecundity were largely confined to females being fed as larvae ad libitum, while those being previously starved showed reduced performance regardless of adult income. Effects on egg size were inconsistent and, overall, marginal. Our results show that restricted food access in different developmental stages may set different limits to reproduction, either posed by shortage of larval-derived storage reserves (i.e. nitrogenous compounds) or adult income (i.e. carbohydrates). Thus, one should be cautious when stating that one or the other type of nutrients is ultimately limiting to reproduction. Rather, our findings highlight the importance of resource congruence and of considering both, larval-and adult-derived resources for reproduction.The pattern of resource allocation has critical consequences for individual fitness and is fundamental to numerous fields of research in behavioural, evolutionary and population ecology. In particular the availability of adequate nutrition comprises one of the most important factors affecting growth, development and reproduction in animals. Food shortage during juvenile development was repeatedly found to increase development time, and to decrease growth rates, body size and reproductive output later in live (Briegel 1990, Berrigan and Charnov 1994, Blanckenhorn 1998, Fischer and Fiedler 2001a. Likewise, limited food access in the adult stage was found to diminish performance in various ways (Boggs and Ross 1993, Braby and Jones 1995, Fischer and Fiedler 2001b. Among the most obvious effects are those on...

The temperature-size rule (TSR), which states that body size increases at lower developmental temperatures, appears to be a near-universal law for ectotherms. Although recent studies seem to suggest that the TSR might be adaptive, the underlying developmental mechanisms are thus far largely unknown. Here, we investigate temperature effects on life-history traits, behaviour and physiology in the copper butterfly Lycaena tityrus in order to disentangle the mechanistic basis for the above rule. In L. tityrus the larger body size produced at a lower temperature was proximately due to a greater increase in mass, which was caused by both behavioural and physiological mechanisms: a much-increased food intake and a higher efficiency in converting ingested food into body matter. These mechanisms, combined with temperature-induced changes at the cellular level, may provide general explanations for the TSR. Body fat and protein content increased in butterflies reared at the higher temperature, indicating favourable growth conditions. As predicted from protandry theory, males showed reduced development times, caused by higher growth rates compared to females. The latter was itself related to a higher daily food consumption, while the total food consumption (due to the females' longer developmental period) and assimilation was higher in females and may underly the sexual body size dimorphism.

BackgroundThe ability to withstand thermal stress is considered to be of crucial importance for individual fitness and species' survival. Thus, organisms need to employ effective mechanisms to ensure survival under stressful thermal conditions, among which phenotypic plasticity is considered a particularly quick and effective one.Methodology/Principal FindingsIn a series of experiments we here investigate phenotypic adjustment in temperature stress resistance following environmental manipulations in the butterfly Bicyclus anynana. Cooler compared to warmer acclimation temperatures generally increased cold but decreased heat stress resistance and vice versa. In contrast, short-time hardening responses revealed more complex patterns, with, e.g., cold stress resistance being highest at intermediate hardening temperatures. Adult food stress had a negative effect on heat but not on cold stress resistance. Additionally, larval feeding treatment showed interactive effects with adult feeding for heat but not for cold stress resistance, indicating that nitrogenous larval resources may set an upper limit to performance under heat stress. In contrast to expectations, cold resistance slightly increased during the first eight days of adult life. Light cycle had marginal effects on temperature stress resistance only, with cold resistance tending to be higher during daytime and thus active periods.Conclusions/SignificanceOur results highlight that temperature-induced plasticity provides an effective tool to quickly and strongly modulate temperature stress resistance, and that such responses are readily reversible. However, resistance traits are not only affected by ambient temperature, but also by, e.g., food availability and age, making their measurement challenging. The latter effects are largely underexplored and deserve more future attention. Owing to their magnitude, plastic responses in thermal tolerance should be incorporated into models trying to forecast effects of global change on extant biodiversity.

K. 2000. Sex-related differences in reaction norms in the butterfly Lycaena tityrus (Lepidoptera: Lycaenidae). -Oikos 90: 372 -380.We investigate sexual differences in reaction norms in directly developing individuals of the copper butterfly Lycaena tityrus predicted from sexual selection theory. As recent studies on butterflies revealed a high degree of adaptive plasticity in growth and development, which may undermine the basic trade-offs assumed in life-history theory, we focus on effects of temperature, trying to drive growth rates to their physiological upper limit and thus disclosing otherwise potentially concealed responses. Development time strongly depended on temperature, leading in accordance with a central assumption in life-history theory to a larger size at low temperatures, and vice versa. At all temperatures larval development time of males was significantly shorter compared to females, as was predicted by protandry theory. This was partially due to an invariably higher growth rate of males. However, sexes responded in different ways to developmental time constraints caused by increasing temperatures. Despite the shorter larval time of males, both sexes achieved similar body sizes at lower temperatures, because males avoided a reduction in weight due to plastic growth. At high temperatures, in contrast, males were forced to make a trade-off in which they favoured early emergence over large size, leading to a dramatic weight loss. Weight of females, however, remained similar throughout showing no trade-off. These different reaction norms reflect divergent selective pressures acting on males and females, which can be explained in relation to the reproductive system. The strong selection for early emergence in males is likely to be due to monandry, discrete non-overlapping generations (as was already predicted by theory), and territoriality, because prior ownership of a territory seems to be a major advantage for successful reproduction. On the other hand, the preference of females for large body size was expected due to the close relationship between this trait and fecundity. Thus, our results highlight the extraordinary importance of the specific reproductive system, which can influence central life-history traits in manifold ways.

Temperature is an important selective agent in nature. Consequently, temperature-induced plasticity which may help buffering detrimental effects of temperature variation has received considerable attention over recent decades. Laboratory studies have almost exclusively used constant temperatures, while in nature, temperature typically shows pronounced daily fluctuations. Using a factorial design with constant versus fluctuating temperatures and a higher versus a lower mean temperature, we here investigate in the butterfly Lycaena tityrus whether the use of constant temperatures is justified. Fluctuating compared to constant temperatures caused shorter development times, increased heat but decreased cold stress resistance, decreased heat-shock protein expression, and increased immunocompetence. Thus, overall, fluctuating temperatures were more beneficial to the butterflies compared to constant ones. However, despite substantial variation across temperature regimes, the ranking of trait values among treatments remained largely unaffected (e.g. lower constant as well as fluctuating temperatures caused increased pupal mass). Thus, we tentatively conclude that there is no general reason for concern about using constant temperatures in studies investigating phenotypic plasticity, which seem to comprise a fair proxy. However, substantial differences in mean values as well as interactive effects suggest that one needs to be cautious. We further demonstrate negative effects of high temperatures on butterfly immune function, which seem to result from a trade-off between the latter and the heat shock response.

Global warming and its associated increase in temperature extremes pose a substantial challenge on natural systems. Tropical ectotherms, living close to their (upper) critical thermal limits, may be particularly vulnerable to global warming, yet they are as a group understudied. Most studies assessing fitness effects under global warming focused on life-history correlates such as body size and largely neglected immune function. Furthermore they did not consider to what extent temperature effects may be modulated under resource-based trade-offs. Against this background we here investigate effects of temperature extremes on fitness-related adult traits (viz. body mass, fat content, and two key parameters of arthropod immune function: phenoloxidase (PO) activity and haemocyte numbers) at different levels of larval and adult food stress in the tropical butterfly Bicyclus anynana. Body mass and PO activity decreased after short-term larval food stress, but not fat content and haemocyte numbers (probably owing to compensatory mechanisms during further development). Longer-term food deprivation in the adult stage, in contrast, diminished performance throughout, confirming that the feeding treatments chosen imposed stress. Temperature manipulations yielded contrary responses between life-history correlates and immune function: while body mass and fat content increased by increasing temperatures, PO activity and haemocyte numbers decreased. The latter was particularly pronounced under adult food stress, suggesting a resource-allocation trade-off. Our data suggest that global warming will not only reduce performance through direct effects of thermal stress, but also through secondary effects on adult immune function, which may be missed when exclusively focussing on other life-history correlates.