Weak sex-specific evolution of locomotor activity of Sepsis punctum (Diptera: Sepsidae) thermal experimental evolution lines.

Elevated temperatures are expected to rise beyond what the physiology of many organisms can tolerate. Behavioural responses facilitating microhabitat shifts may mitigate some of this increased thermal selection on physiology, but behaviours are themselves mediated by physiology, and any behavioural response may trade-off against other fitness-related activities. We investigated whether experimental evolution in different thermal regimes (Cold: 15 °C; Hot: 31 °C; Intergenerational fluctuation 15/31 °C; Control: 23 °C) resulted in genetic differentiation of standard locomotor activity in the dung fly Sepsis punctum. We assessed individual locomotor performance, an integral part of most behavioral repertoires, across eight warm temperatures from 24 °C to 45 °C using an automated device. We found no evidence for generalist-specialist trade-offs (i.e. changes in the breadth of the performance curve) for this trait. Instead, at the warmest assay temperatures hot-selected flies showed somewhat higher maximal performance than all other, especially cold-selected flies, overall more so in males than females. Yet, the flies' temperature optimum was not higher than that of the cold-selected flies, as expected under the 'hotter-is-better' hypothesis. Maximal locomotor performance merely weakly increased with body size. These results suggest that thermal performance curves are unlikely to evolve as an entity according to theory, and that locomotor activity is a trait of limited use in revealing thermal adaptation.

Ontogeny of sexual size dimorphism revisited: Females grow for a longer time and also faster.

Sex-specific mechanisms of the determination of insect body sizes are insufficiently understood. Here we use the common heath moth, Ematurga atomaria (Lepidoptera: Geometridae) to examine how larval growth trajectories differ between males and females. We monitored the development of 1379 larvae in controlled laboratory conditions. Sexually dimorphic development times during the first four instars were associated with sexual size dimorphism (SSD) in the beginning of the fifth (last) instar, when females were on average 15% heavier than males. Similarly, the duration of the last instar was about 13% longer in females. Further, we specifically focussed on the estimates of differential (instantaneous) growth rates of the larvae based on 24h mass increments of the 2nd, 3rd, 4th and 5th day in the beginning of the last instar. We calculated 'allometric' differential growth rates as the per-day increase in cube-root-transformed mass of the larvae. We found that allometric growth rates were slightly but significantly larger in females than in males. As this measure of growth rate (in contrast to the relative growth rate, based on the ratio of masses recorded at consecutive measurements) did not depend on body size, it allows an unambiguous separation of the effects of sex and size. We conclude that in accordance with an emerging general pattern, larger female body size in E. atomaria is achieved primarily by means of a longer growth period. Furthermore, our study shows that the differential growth rate can also be sexually dimorphic and contribute to SSD. This contribution, however, is lower than that of the development time by an order of magnitude. In addition to development periods and growth rates, other parameters of the non-linear growth curves of insect larvae also need to be considered in the context of SSD determination. In particular, weight loss prior to pupation was shown to be considerably larger in females than in males.

Longer life span is associated with elevated immune activity in a seasonally polyphenic butterfly.

Seasonal polyphenism constitutes a specific type of phenotypic plasticity in which short-lived organisms produce different phenotypes in different times of the year. Seasonal generations of such species frequently differ in their overall lifespan and in the values of traits closely related to fitness. Seasonal polyphenisms provide thus excellent, albeit underused model systems for studying trade-offs between life-history traits. Here, we compare immunological parameters between the two generations of the European map butterfly (Araschnia levana), a well-known example of a seasonally polyphenic species. To reveal possible costs of immune defence, we also examine the concurrent differences in several life-history traits. Both in laboratory experiments and in the field, last instar larvae heading towards the diapause (overwintering) had higher levels of both phenoloxidase (PO) activity and lytic activity than directly developing individuals. These results suggest that individuals from the diapausing generation with much longer juvenile (pupal) period invest more in their immune system than those from the short-living directly developing generation. The revealed negative correlation between pupal mass and PO activity may be one of the reasons why, in this species, the diapausing generation has a smaller body size than the directly developing generation. Immunological parameters may thus well mediate trade-offs between body size-related traits.

Sexual differences in weight loss upon eclosion are related to life history strategy in Lepidoptera.

Given that immature and adult insects have different life styles, different target body compositions can be expected. For adults, such targets will also differ depending on life history strategy, and thus vary among the sexes, and in females depend on the degree of capital versus income breeding and ovigeny. Since these targets may in part be approximated by loss of substances upon eclosion, comparing sexual differences in such losses upon eclosion among species that differ in life history would provide insights into insect functional ecology. We studied weight loss in eclosing insects using original data on pupal and adult live weights of 38 species of Lepidoptera (mainly Geometridae) and further literature data on 15 species of Lepidoptera and six representatives of other insect orders, and applied the phylogenetic independent contrasts approach. In addition, data on live and dry weights of pupae of four species of Lepidoptera are presented. We documented that Lepidoptera typically lose a large proportion (20-80%) of their pupal weight upon adult eclosion. Sexual differences in weight loss varied between absent and strongly male biased. Most of the weight loss was water loss, and sexual differences in adult water content correlate strongly with differences in weight loss. Using feeding habits (feeds or does not feed as an adult) and female biased sexual size dimorphism as measures of degree of capital breeding, we found that the difference among the sexes in weight loss tends to be more pronounced in capital breeding species. Additionally, females of more pro-ovigenic species (large proportion of eggs mature upon emergence) tend to have higher water contents. Our results suggests that metamorphosis is generally facilitated by a high water content, while adults excrete water upon eclosion to benefit flight unless water has been allocated to eggs, or is treated as a capital resource for adult survival or future allocation to eggs.

Counterintuitive size patterns in bivoltine moths: late-season larvae grow larger despite lower food quality.

Within a season, successive generations of short-lived organisms experience different combinations of environmental parameters, such as temperature, food quality and mortality risk. Adult body size of e.g. insects is therefore expected to vary both as a consequence of proximate environmental effects as well as adaptive responses to seasonal cues. In this study, we examined intraspecific differences in body size between successive generations in 12 temperate bivoltine moths (Lepidoptera), with the ultimate goal to critically compare the role of proximate and adaptive mechanisms in determining seasonal size differences. In nearly all species, individuals developing late in the season (diapausing generation) attained a larger adult size than their conspecifics with the larval period early in the season (directly developing generation) despite the typically lower food quality in late summer. Rearing experiments conducted on one of the studied species, Selenia tetralunaria also largely exclude the possibility that the proximate effects of food quality and temperature are decisive in determining size differences between successive generations. Adaptive explanations appear likely instead: the larger body size in the diapausing generation may be adaptively associated with the lower bird predation pressure late in the season, and/or the likely advantage of large pupal size during overwintering.

Dependence of phenotypic variance in body size on environmental quality.

The recent "overhead threshold" model for optimal age and body size at maturity (Day and Rowe 2002 ) predicts that phenotypic variability in adult body size will be low under inferior environmental quality and will increase with improving conditions. The model is, however, based on a potentially restrictive assumption of a monotone increase of fecundity with increasing body size. On the basis of a numerical model, we show that introducing the concept of maximum adult body size changes the predictions of the model. The dependence of variability in adult body size on environmental quality becomes a concave function with a maximum at intermediate values. Depending on the range of environmental conditions considered, one may therefore expect to observe both increasing and decreasing functions. We test the predictions of our model on a literature-based database of 131 insect species covering all major orders. We demonstrate that, in most species, relative phenotypic variation in body size decreases when environment-specific average of adult body size increases. In the majority of cases at least, such a relationship can be interpreted as a decreased relative variation in better growing conditions. With some potentially meaningful exceptions (e.g., females of capital-breeding insects), the general pattern was largely invariable across different taxa, ecological subdivisions, and sexes.

Intraspecific variability in number of larval instars in insects.

The number of larval instars varies widely across insect species. Although instar number is frequently considered to be invariable within species, intraspecific variability in the number of instars is not an exceptional phenomenon. However, the knowledge has remained fragmentary, and there are no recent attempts to synthesize the results of relevant studies. Based on published case studies, we show that intraspecific variability in the number of larval instars is widespread across insect taxa, occurring in most major orders, in both hemimetabolous and holometabolous insects. We give an overview of various factors that have been observed to affect the number of instars. Temperature, photoperiod, food quality and quantity, humidity, rearing density, physical condition, inheritance, and sex are the most common factors influencing the number of instars. We discuss adaptive scenarios that may provide ultimate explanations for the plasticity in instar number. The data available largely support the compensation scenario, according to which instar number increases in adverse conditions when larvae fail to reach a species-specific threshold size for metamorphosis. However, in Orthoptera and Coleoptera, there are some exceptional species in which instar number is higher in favorable conditions. In more specific cases, the adaptive value of the variability in instar number may be in reaching or maintaining the developmental stage adapted to hibernation, producing additional generations in multivoltine species, or increasing the probability of surviving in long-lasting adverse conditions.

Maintenance of larval color polymorphism in Orgyia antiqua (Lepidoptera: Lymantriidae): evaluating the role of thermal adaptation.

Intraspecific color polymorphism is widespread in insects, and various mechanisms have been proposed to explain its maintenance. Some explanations rely on the effect of body color on the organism's thermal physiology. Darker individuals accumulate solar energy more efficiently, and therefore, dark body coloration in insects is frequently presumed to be an adaptation to low temperature conditions. However, it is largely unclear what is the importance of the thermal biology in comparison to other potential selective forces on body coloration. In this study, we evaluated the role of temperature as a potential selective factor maintaining color polymorphism in aposematic larvae of the moth Orgyia antiqua L. It was found that darker, and thus less aposematic, larvae accumulated solar energy more efficiently. However, in a set of laboratory and outdoor experiments, we found no evidence of temperature-dependent performance of different color morphs or in development of different morphs induced by rearing temperature. We conclude that the effects related to thermal physiology are not likely important determinants of optimal coloration in O. antiqua. The reasons may lie in high mobility of the larvae, which allows for effective behavioral thermoregulation, which is also shown in this study. Our results caution against an uncritical extrapolation of results obtained for model organisms and indicate the need for giving more attention to the species-specific ecological background in ecophysiological studies.

No evidence for costs of being large in females of Orgyia spp. (Lepidoptera, Lymantriidae): larger is always better.

Strong correlation between female body size and potential fecundity is often observed in insects. Directional selection favouring increased body sizes is thus predicted in the absence of opposing selection pressure. The evolutionary forces capable of counterbalancing such a 'fecundity advantage' are poorly documented. This study focuses on revealing the costs of large body size in the wingless females of Orgyia antiqua and O. leucostigma, two related species of lymantriid moths. Extreme behavioural simplicity of these animals allows systematic assessment of various fitness components in conditions that are close to natural. A linear relationship between pupal weight and potential fecundity was observed. This association was found to be independent of particular rearing conditions. There was no evidence that the relationship between fecundity and body mass becomes asymptotic when body sizes increases. No component of fitness showed a negative phenotypic correlation with body size; some displayed a weakly positive one. In particular, pupal mortality, adult longevity, mating and oviposition success, as well as egg hatching rate and egg size, were established as independent of body size in a series of field and laboratory experiments. There was a very high overall efficiency of converting resources accumulated during the larval stage to egg masses. With no costs of large adult size, selective forces balancing the fecundity advantage should operate in the course of immature development. The strong dependence of realized fecundity on body size is considered characteristic of the capital breeding strategy.