Klaus Fischer

Although the temperature‐size rule, that is, an increase in egg (and body) size at lower temperatures, applies almost universally to ectotherms, the developmental mechanisms underlying this consistent pattern of phenotypic plasticity are hitherto unknown. By investigating ovarian dynamics and reproductive output in the tropical butterfly Bicyclus anynana (Butler) (Lepidoptera: Nymphalidae: Satyrinae) in relation to oviposition temperature and mating status, we tested the relevance of several competing hypotheses for temperature‐mediated variation in egg size and number. As expected, females ovipositing at a lower temperature laid fewer but larger eggs than those ovipositing at a higher temperature. Despite pronounced differences in egg‐laying rates, oocyte numbers were equal across temperatures at any given time, while oocyte size increased at the lower temperature. In contrast, there were greatly reduced oocyte numbers in mated compared to virgin females. Our results indicated that temperature‐mediated plasticity in egg size cannot be explained by reduced costs of somatic maintenance at lower temperatures, enabling the allocation of more resources to reproduction (reproductive investment was higher at the higher temperature). Furthermore, there was no indication for delayed oviposition (no accumulation of oocytes at the lower temperature, in contrast to virgin females). Rather, low temperatures greatly reduced the oocyte production (i.e., differentiation) rate and prolonged egg‐maturation time, causing low egg‐laying rates. Our data thus suggested that oocyte growth is less sensitive to temperature than oocyte production, resulting in a lower number of larger eggs at lower temperatures.

Using lines artificially selected on egg size and being subjected to a restricted and an unrestricted feeding treatment, we examined the relationship

We use artificial selection experiments targeted on egg size, development time or pupal mass within a single butterfly population followed by a common-garden experiment to explore the interactions among these life-history traits. Relationships were predicted to be negative between egg size and development time, but to be positive between development time and body size and between egg size and body size. Correlated responses to selection were in part inconsistent with these predictions. Although there was evidence for a positive genetic correlation between egg and body size, there was no support for genetic correlations between larval development time and either egg size or pupal mass. Phenotypic correlations among the three target traits of selection gave comparable results for the relationships between egg mass and development time (no association) as well as between egg mass and pupal mass (positive association), but not for the relation between development time and pupal mass (negative phenotypic correlation). In summary, correlated responses to selection as well as phenotypic correlations were rather unpredictable. The impact of variation in acquisition and allocation of energy as well as of the benign conditions used deserve further investigation.

Life history theory predicts trade‐offs arising from the costs of reproduction: individuals investing more into reproduction should have less energy available for their own somatic demands. Despite a wealth of research on such costs for females, very few studies have looked at the male side. We investigated the costs of reproduction in males of two butterfly species, Bicyclus anynana and Pieris napi. The males of these species differ tremendously in their allocation of nutrients to reproduction: virgin male spermatophores were 18.5 times larger in the latter compared with those in the former. Based on this striking difference, we expected to find a longevity cost of mating in P. napi, but not necessarily in B. anynana. In line with our predictions, we found no evidence for a survival cost of mating in B. anynana, while there was some evidence for such a cost in P. napi. Here, virgin males lived longer than once‐ or twice‐mated ones. However, this overall trend did not hold for all treatment groups, and there was no linear relationship between the number of copulations and longevity. Adult food limitation reduced longevity throughout, but had otherwise very little impact. Spermatophore dry mass decreased in later compared with first spermatophores, while water content increased, indicating that spermatophore production is indeed physiologically costly. Additionally, adult food limitation tended to reduce spermatophore fresh and dry mass, and tended to increase spermatophore water content. Based on our results we tentatively conclude that longevity and reproduction may not always be traded off against each other, as in our study organisms’ longevity seems to depend in first place on adult carbohydrate income, while male reproduction seems to be primarily (although not entirely) fuelled by larval storage reserves.

We artificially selected on egg size in a butterfly to study the consequences for fecundity, reproductive effort and offspring fitness. Correlated responses in either pupal mass, larval or pupal development time were virtually absent. Offspring size was positively related to fitness, but only partly traded off against fecundity. Rather, total reproductive effort (measured as fresh mass), egg water content and the decline of egg size with female age increased in the large-egg selected lines compared to either small-egg or control lines. Accounting for these effects showed that reproductive investment (in dry mass) was in fact similar across lines. Such mechanisms may enable increased investment in (early) offspring without a reduction in their number, revealing a much more complex picture than a simple trade-off between offspring size and number. Substantial variation among replicates suggests that there are different underlying mechanisms for change, rather than any single, unitary pathway.

Summary: The source of nutritional resources allocated to reproduction strongly influences reproductive and foraging strategies. While we are beginning to understand the role of adult and larval resources for nectivorous Lepidoptera, essentially nothing is known for the large number of species that feed on fruit as adults. We used stable isotopes to examine allocation of larval‐ and adult‐derived resources to egg production in a tropical frugivorous butterfly, Bicyclus anynana (Butler, 1879), under both ad libitum and semistarvation conditions. The butterfly's larval and adult host plants differ from each other in both carbon and nitrogen isotopic ratios. Adult fruit feeding is required for the onset of oviposition. At peak, adult fruit feeding contributed 55% of the carbon found in eggs under both feeding conditions. This is similar to values for several nectivorous Lepidoptera with similar ovarian dynamics and egg C/N. Egg ¹⁵N declined rapidly during the first week of oviposition, suggesting that the adult diet was not contributing nitrogen to egg production. Values rose during the following 2 weeks, consistent either with adult contribution or with isotopic fractionation. Our results are consistent with the hypothesis that fruit serves as a carbon source for egg production in a similar manner as nectar. However, more work is needed to elucidate fully fruit's role as a nitrogen source.

By dividing families of the tropical butterfly, Bicyclus anynana, among different larval (including early pupal) and adult (including late pupal) temperatures, we investigate the genetic and environmental effects on egg size. Both sources of variation affected egg size to similar extents. As previously found in other arthropods, egg size tended to increase at lower temperatures. Our data suggest that the plastic response in egg size can be induced during the pupal stage. Females reared as larvae at the same high temperature tended to lay larger eggs when transferred to a lower temperature, either as prepupae or pupae, compared to those remaining at the high temperature. Additionally, females reared as larvae at different temperatures, but maintained at the same temperature from the early pupal stage onwards, laid larger eggs after larval growth at a low temperature. Heritability estimates for egg size were about 0.4 (parent–offspring regression) and 0.2 (variance component estimates using the full-sib families). Although there seemed to be some variation in the plastic response to temperature among families, genotype–environment interactions were nonsignificant.