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.

Weak and inconsistent associations between melanic darkness and fitness-related traits in an insect.

The idea that the fitness value of body coloration may be affected by biochemically mediated trade-offs has received much research attention. For example, melanization is believed to interact with other fitness-related traits via competition for substrates, costs associated with the synthesis of melanin or pleiotropic effects of the involved genes. However, genetic correlations between coloration and fitness-related traits remain poorly understood. Here, we present a quantitative-genetic study of a coloration trait correlated to melanin-based cuticular darkness ('darkness', hereafter) in a geometrid moth, Ematurga atomaria. This species has considerable variation in larval appearance. We focus on correlations between larval darkness and fitness-related growth performance traits. Both a half-sib analysis and an 'animal model' approach revealed moderately high heritabilities of larval darkness and indices of growth performance. Heritability estimates of darkness derived from the animal model were, however, considerably higher than those based on the half-sib model suggesting that the determination of coloration includes genetic interactions and epigenetic effects. Importantly, on the host plant with the largest sample size, we found no evidence for either genetic or environmental correlations between darkness and growth parameters. On an alternative host plant, there was some indication of positive genetic and negative environmental correlation between these traits. This shows that respective relationships are environment-specific. Nevertheless, the overall pattern of weak and inconsistent correlations between larval coloration and growth parameters does not support universal trade-offs between these traits and suggests that physiological costs of producing colour patterns do not necessarily interfere with adaptive evolution of coloration.

Sources of variance in immunological traits: evidence of congruent latitudinal trends across species.

Among-population differences in immunological traits allow assessment of both evolutionary and plastic changes in organisms' resistance to pathogens. Such knowledge also provides information necessary to predict responses of such traits to environmental changes. Studies on latitudinal trends in insect immunity have so far yielded contradictory results, suggesting that multispecies approaches with highly standardised experimental conditions are needed. Here, we studied among-population differences of two parameters reflecting constitutive immunity-phenoloxidase (PO) and lytic activity, using common-garden design on three distantly related moth species represented by populations ranging from northern Finland to Georgia (Caucasus). The larvae were reared at different temperatures and on different host plants under a crossed factors experimental design. Haemolymph samples for measurement of immune status were taken from the larvae strictly synchronously. Clear among-population differences could be shown only for PO activity in one species (elevated activity in the northern populations). There was some indication that the cases of total absence of lytic activity were more common in southern populations. The effects of temperature, host and sex on the immunological traits studied remained highly species specific. Some evidence was found that lytic activity may be involved in mediating trade-offs between immunity and larval growth performance. In contrast, PO activity rarely covaried with fitness-related traits, and neither were the values of PO and lytic activity correlated with each other. The relatively inconsistent nature of the detected patterns suggests that studies on geographic differences in immunological traits should involve multiple species, and rely on several immunological indices if general trends are a point of interest.

Reproductive parameters of critically endangered European mink (Mustela lutreola) in captivity.

Founding captive populations is often the last chance for saving endangered species from extinction. Ensuring successful reproduction is typically most critical for the maintenance of captive populations, with purposeful selection of individuals for breeding being one of the crucial aspects. Comparable cross-species data on the determinants of reproduction success are most useful for solving problems in captive species programs. In the present study, we provide an overview of a 20-year captive breeding program of the critically endangered European mink. The mating season starts in March, reaching its peak in the middle of April. The average gestation length was 43.8days (mode 43), the mean litter size being 4.4 (mode 4). Litter size and cub survival were negatively correlated with maternal age but this effect was entirely due to the lower performance of the females over 4 years of age. Female body weight also showed a positive correlation with litter size, with the weight itself having increased by 10% during the 20- year period. We did not find any signs of a cost of reproduction: the number of litters the female had delivered earlier in her life did not have an effect on her litter size in the focal year. Beyond the effect of age and size, individual females did not differ in litter sizes. Consistently, we found the heritability of litter size to be low. We conclude that, when selecting females for breeding, there is little need to consider aspects other than genetic relatedness crucial for avoiding progressive inbreeding.

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.