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Temperature has a profound effect on the growth and development of ectothermic animals. However, the extent to which ecologically driven selection pressures can adjust thermal plastic responses in growth schedules is not well understood. Comparing temperature-induced plastic responses between sexes provides a promising but underexploited approach to evaluating the evolvability of thermal reaction norms: males and females share largely the same genes and immature environments but typically experience different ecological selection pressures. We proceed from the idea that substantial sex differences in plastic responses could be interpreted as resulting from sex-specific life-history optimization, whereas similarity among the sexes should rather be seen as evidence of an essential role of physiological constraints. In this study, we performed a meta-analysis of sex-specific thermal responses in insect development times, using data on 161 species with comprehensive phylogenetic and ecological coverage. As a reference for judging the magnitude of sex specificity in thermal plasticity, we compared the magnitude of sex differences in plastic responses to temperature with those in response to diet. We show that sex-specific responses of development times to temperature variation are broadly similar. We also found no strong evidence for sex specificity in thermal responses to depend on the magnitude or direction of sex differences in development time. Sex differences in temperature-induced plastic responses were systematically less pronounced than sex differences in responses induced by variations in larval diet. Our results point to the existence of substantial constraints on the evolvability of thermal reaction norms in insects as the most likely explanation. If confirmed, the low evolvability of thermal response is an essential aspect to consider in predicting evolutionary responses to climate warming.

Assemblages of insects need to be quantitatively sampled in the context of various research questions. Light trapping is the most widely used method for sampling nocturnal Lepidoptera. Attracting moths to sugar baits offers a viable alternative. However, this method is rarely used in professional research despite its popularity among amateur lepidopterists. As the activity of insects is strongly dependent on ambient conditions, the sensitivity of any trapping method to weather parameters needs to be known for the quantitative interpretation of trapping results. In the present paper, we report data on the weather dependence of moth catches obtained by automatic bait traps. The study was performed in Estonia, representing the European hemiboreal forest zone. Portable weather stations set up next to each of the traps were used for collecting weather data. Both abundance and diversity of the moths in the catches depended strongly positively on temperature and negatively on air humidity. Diversity was also negatively correlated with air pressure and positively with the change in pressure during the night. The results show that in situ recording of weather parameters in connection to insect trapping provides useful insights for the study of insect behaviour and the interpretation of the results of monitoring projects.

Recent advances in molecular systematics have led to an emerging understanding of the phylogenetic history of the family Geometridae. These studies have basically confirmed the traditional subdivision of the subfamily Larentiinae into tribes but unsolved problems remain. Here we test the monophyly of the tribe Perizomini, and evaluate the division of this tribe into genera using Bayesian and maximum likelihood analyses of one mitochondrial and eight nuclear gene fragments. We show that the Eurasian members of Perizoma Hübner, 1825, Mesotype Hübner, 1825 and Gagitodes Warren, 1893 together form a monophyetic tribe Perizomini. However, Martania Mironov, 2000 is not closely related to these genera, but is considered to belong to Melanthiini according to the results of the phylogenetic analyses. Morphological evidence supporting this rearrangement is discussed. The Nearctic Larentia basaliata Walker, 1862 was shown to belong in the genus Martania as M. basaliata (Walker, 1862) comb. nov. and being specifically distinct from the morphologically similar Palaearctic M. taeniata. Three other studied ‘Perizoma’ species from the New World were similarly placed far from Perizomini in the phylogenetic tree, and were not related to each other. We conclude that both the tribe Perizomini and the genus Perizoma are polyphyletic which indicates that the group needs a global revision. It remains an open question whether Perizomini have a worldwide distribution as previously assumed, or is this tribe confined to the Palaearctic region.

Nola estonica Õunap sp. nov. (Lepidoptera, Nolidae, Nolinae) is described based on type material from Estonia. The lectotype is designated for Glaphyra atomosa Bremer, 1861, which is reinstated from a subspecies of Nola aerugula (Hübner, [1793]) to a full species: Nola atomosa (Bremer, 1861) stat. rev. The status of these three taxa as separate species is supported by the results of phylogenetic analysis of DNA barcodes, as well as external and genital morphology of adult specimens. Two new synonyms are established as follows: Nola atomosa (Bremer, 1861) = Nola candidalis Staudinger, 1892 syn. nov. and Nola shin Inoue, 1982 syn. nov. N. estonica occurs sympatrically with N. aerugula in Estonia, and with N. atomosa in South Korea and easternmost Russia. While the available data suggest a disjunct distribution of N. estonica (eastern Europe and the temperate Far East), it appears highly possible that the species has a wide transpalaearctic distribution.

Ecological and life-history data on the Northern European macromoth (Lepidoptera: Macroheterocera) fauna is widely available and ideal for use in answering phylogeny-based research questions: for example, in comparative biology. However, phylogenetic information for such studies lags behind. Here, as a synthesis of all currently available phylogenetic information on the group, we produce a supertree of 114 Northern European macromoth genera (in four superfamilies, with Geometroidea considered separately), providing the most complete phylogenetic picture of this fauna available to date. In doing so, we assess those parts of the phylogeny that are well resolved and those that are uncertain. Furthermore, we identify those genera for which phylogenetic information is currently too poor to include in such a supertree, or entirely absent, as targets for future work. As an aid to studies involving these genera, we provide information on their likely positions within the macromoth tree. With phylogenies playing an ever more important role in the field, this supertree should be useful in informing future ecological and evolutionary studies.

If the evolutionary determinants of longevity were mainly extrinsic, related species from different habitats should systematically differ in individual lifespans. Comparative studies of insects can significantly contribute to understanding the evolution of lifespan, as the trait can feasibly be measured in a high number of species. We recorded adult longevities for 110 species of geometrid moths from a tropical community. Comparative analyses based on an original phylogenetic reconstruction were applied to reveal the correlates of species-specific values of lifespan. Larger moth species were found to live longer, and females tended to have shorter lifespans than males. Average adult lifespans in tropical geometrids, and the relationships of lifespan with other variables, were found to be highly similar to those reported for their temperate region relatives. The among-region similarity leads to the conclusion of the dominance of intrinsic (physiological) determinants of longevity over the extrinsic (ecological) ones: the contrasting environments of tropical and temperate forests have not produced detectable differences in moth longevities.

1. Comparative studies on insect life histories are facilitated by the increasing availability of reliable phylogenies but are hampered by the scarcity of comparable data. Fortunately, morphological proxies of some life‐history traits can be measured on preserved specimens. 2. This study compared values of size‐related life‐history traits among a tropical (Ugandan) and a temperate (Estonian) assemblage of geometrid moths. 3. A comparative analysis based on an originally derived phylogeny revealed that tropical moths were, on average, larger than temperate ones. Tropical moths also had somewhat lower relative abdomen masses than temperate ones. This indicates that the tropical rather than the temperate moths tend to use an income (rather than capital) breeding strategy. Nevertheless, no difference was found in a related index of pro‐ovigeny. When body size was accounted for, tropical moths were found to lay smaller eggs than temperate ones. 4. The differences between the two compared areas are consistent with selection on higher mobility of the moths imposed by the more diverse tropical vegetation. Relatively larger eggs of temperate moths may constitute an adaptation to overcome the presumably stronger quantitative defences of their host plants. 5. Overall, however, we conclude that the differences in ecologically relevant size‐related traits are relatively low among moth assemblages of a tropical and a temperate forest region, indicating that these environments may not impose radically different selective pressures on insect life histories.

Conspecific females and males often follow different development trajectories which leads to sex differences in age at maturity (sexual bimaturism, SBM). Whether SBM is typically selected for per se (direct selection hypothesis) or merely represents a side-effect of other sex-related adaptations (indirect selection hypothesis) is, however, still an open question. Substantial interspecific variation in the direction and degree of SBM, both in invertebrates and vertebrates, calls for multi-species studies to understand the relative importance of its evolutionary drivers. Here we use two complementary approaches to evaluate the evolutionary basis of SBM in insects. For this purpose, we assembled an extensive literature-derived data set of sex-specific development times and body sizes for a taxonomically and ecologically wide range of species. We use these data in a meta-analytic framework to evaluate support for the direct and indirect selection hypotheses. Our results confirm that protandry - males emerging as adults before females - is the prevailing form of SBM in insects. Nevertheless, protandry is not as ubiquitous as often presumed: females emerged before males (= protogyny) in about 36% of the 192 species for which we had data. Moreover, in a considerable proportion of species, the sex difference in the timing of adult emergence was negligible. In search for the evolutionary basis of SBM, we found stronger support for the hypothesis that explains SBM by indirect selection. First, across species, the direction and degree of SBM appeared to be positively associated with the direction and degree of sexual size dimorphism (SSD). This is consistent with the view that SBM is a correlative by-product of evolution towards sexually dimorphic body sizes. Second, within protandrous species, the degree of protandry typically increased with plastic increase in development time, with females prolonging their development more than males in unfavourable conditions. This pattern is in conflict with the direct selection hypothesis, which predicts the degree of protandry to be insensitive to the quality of the juvenile environment. These converging lines of evidence support the idea that, in insects, SBM is generally a by-product of SSD rather than a result of selection on the two sexes to mature at different times. It appears plausible that selective pressures on maturation time per se generally cannot compete with viability- and fecundity-mediated selection on insect body sizes. Nevertheless, exceptions certainly exist: there are undeniable cases of SBM where this trait has evolved in response to direct selection. In such cases, either the advantage of sex difference in maturation time must have been particularly large, or fitness effects of body size have been unusually weak.

Seasonal polyphenisms are cases in which individuals representing generations occurring in different times of the year systematically differ in their morphological, physiological, and/or behavioral traits. Such differences are often assumed to constitute adaptive responses to seasonally varying environments, but the evidence for this is still scarce. The adaptive character of the response would be corroborated by the pattern in which the decision about choosing a particular seasonal phenotype is made before the onset of respective environmental conditions (anticipatory plasticity). Alternatively, the between-generation differences can be caused by immediate effects of seasonally varying environments (responsive plasticity). Here we reared the larvae of the seasonally polymorphic map butterfly Araschnia levana under two different photoperiodic regimes, which provided different seasonal cues. These two treatments induced direct development and diapause pathways, respectively. Replicating the experiment at different temperatures and levels of host plant quality allowed us to evaluate both the anticipatory and the responsive components of the associated plastic changes in life-history traits. Larvae representing the direct development pathway invariably had higher growth rates and shorter development periods, although the difference between the developmental pathways was smaller at inferior host quality. Body size differences between the developmental pathways turned out to be less consistent, as the natural pattern of higher pupal mass of the directly developing individuals could only be reproduced at lower rearing temperature. Though being considerably modified by immediate environmental effects, the between-generation differences in size, growth rates, and larval are largely based on anticipatory plasticity (= responses to photoperiodic cues) and should be treated as seasonal adaptations in A. levana. In a more general context, we show how investigating the proximate basis of size differences can serve the purpose of identifying the limits of phenotypic plasticity in juvenile growth schedules.

Research on evolutionary forces determining optimal body sizes has primarily relied on experimental evaluation of respective selective pressures. Accounting for among-species variation through application of phylogenetic comparative methods is a complementary although little used approach. It enables the direct association of body size values with particular environments. Using phylogenetically explicit comparative analyses, we show that small body size is associated with diurnal (rather than nocturnal) activity of adults among temperate species of the moth family Geometridae. The association of an exclusively adult trait with species-specific body size suggests that optimal body sizes are at least partly determined by the costs being a large adult, as opposed to the more frequently considered costs of attaining large size. It appears likely that size-selective predation by insectivorous birds is the primary factor responsible for selection against large body size in day-flying moths.

Understanding how and why some groups have become more species-rich than others, and how past biogeography may have shaped their current distribution, are questions that evolutionary biologists have long attempted to answer. We investigated diversification patterns and historical biogeography of a hyperdiverse lineage of Lepidoptera, the geometrid moths, by studying its most species-rich tribe Boarmiini, which comprises ca. 200 genera and ca. known 3000 species. We inferred the evolutionary relationships of Boarmiini based on a dataset of 346 taxa, with up to eight genetic markers under a maximum likelihood approach. The monophyly of Boarmiiniis strongly supported. However, the phylogenetic position of many taxa does not agree with current taxonomy, although the monophyly of most major genera within the tribe is supported after minor adjustments. Three genera are synonymized, one new combination is proposed, and four species are placed in incertae sedis within Boarmiini. Our results support the idea of a rapid initial diversification of Boarmiini, which also implies that no major taxonomic subdivisions of the group can currently be proposed. A time-calibrated tree and biogeographical analyses suggest that boarmiines appeared in Laurasia ca. 52 Mya, followed by dispersal events throughout the Australasian, African and Neotropical regions. Most of the transcontinental dispersal events occurred in the Eocene, a period of intense geological activity and rapid climate change. Diversification analyses showed a relatively constant diversification rate for all Boarmiini, except in one clade containing the species-rich genus Cleora. The present work represents a substantial contribution towards understanding the evolutionary origin of Boarmiini moths. Our results, inevitably biased by taxon sampling, highlight the difficulties with working on species-rich groups that have not received much attention outside of Europe. Specifically, poor knowledge of the natural history of geometrids (particularly in tropical clades) limits our ability to identify key innovations underlying the diversification of boarmiines.

Latitudinal and elevational trends in body size are found in numerous animal taxa, with various adaptive explanations proposed. It is however debatable whether geographic trends in adult body size are accompanied by corresponding differences in juvenile growth rate (= mass gain per unit time). Respective studies have been complicated by conceptual and methodological problems related to defining and measuring this variable, particularly in organisms with discontinuous growth like arthropods. Using an original method for estimating differential (instantaneous) juvenile growth rates, we compared geographically distant European populations of six insect species in a common garden experiment. We found no among population differences in differential growth rate in any of the species. This result is in concert with concurrent increase in both adult size and developmental time towards the south. While opposite examples exist, we interpret our results as challenging the view that growth rate is a trait that readily responds to environmentally based selective pressures. Our results thus advocate the more classical view of growth rate maximisation within its physiological limits. We discuss the advantages of using differential (rather than integral) measures of growth rate in evolutionary ecological studies and evaluate the reasons for the detected latitudinal trends in life history traits.

Specificity is one of the fundamental concepts in ecology. Host specificity of phytophagous insects has been of particular interest because of its crucial role in diversification and life-history evolution. However, the majority of tropical insects remain insufficiently explored with respect to their host-plant relations. A lack of respective data is also hindering the debate over whether higher levels of host-plant specificity prevail in tropical insects compared to temperate ones. We investigated host-plant specificity of forest geometrid moths (Lepidoptera: Geometridae) in equatorial Africa using host-plant acceptability trials with neonate larvae, with the addition of field observations. We compare our experimental data to the (well-known) host-specificity patterns of closely related temperate (hemiboreal) species. Similarly to the temperate region, there were broadly polyphagous tropical species in several clades of Geometridae utilizing hosts belonging to different plant families. Phylogenetic comparative analysis returned no significant differences in host specificity between the two regions. Our study contributes to the evidence that host-plant specificity of herbivores is not necessarily substantially higher in tropical than temperate regions.

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.

In litter-bearing mammals, the environment and social interactions during early life often have a substantial effect on future behaviour of the animal. Most information though derives from lab rodents, pets or farm animals while comparable data are scarce for non-domesticated species, and endangered carnivores in particular. In this study, we focused on social behaviour of juvenile European mink, with the practical aim to provide information for enhancing the ex-situ breeding programme of this critically endangered species. As the first step, we compiled a detailed ethogram of social behaviour observed among the European mink cubs. For the 13 captive born litters available, we then systematically recorded the relative duration of different types of behaviour during a two months period. The behaviour of the captive cubs was found to be diverse, containing all elements characteristic of congeneric mustelids, with no indication of litters deviating from the typical pattern. In all broods, a considerable and approximately equal share of time was allocated to social play, a suggested indicator of positive welfare. Aggressive behaviour of the mother towards her offspring was minimal when the cubs were young and, increased only with the litter dispersal period approaching. Bites between the cubs during play fighting did not increase with the age of the juveniles. We found no evidence that captive environment adversely affects the behaviour of the juveniles during the first months of their life.

Predictive classifications of life histories are essential for evolutionary ecology. While attempts to apply a single approach to all organisms may be overambitious, recent advances suggest that more narrow ordination schemes can be useful. However, these schemes mostly lack easily observable proxies of the position of a species on respective axes. It has been proposed that, in insects, the degree of capital (vs. income) breeding, reflecting the importance of adult feeding for reproduction, correlates with various ecological traits at the level of among-species comparison. We sought to prove these ideas via rigorous phylogenetic comparative analyses. We used experimentally derived life-history data for 57 species of European Geometridae (Lepidoptera), and an original phylogenetic reconstruction. The degree of capital breeding was estimated based on morphological proxies, including relative abdomen size of females. Applying Brownian-motion-based comparative analyses (with an original update to include error estimates), we demonstrated the associations between the degree of capital breeding and larval diet breadth, sexual size dimorphism, and reproductive season. Ornstein-Uhlenbeck model based phylogenetic analysis suggested a causal relationship between the degree of capital breeding and diet breadth. Our study indicates that the gradation from capital to income breeding is an informative axis to ordinate life-history strategies in flying insects which are affected by the fecundity vs. mobility trade off, with the availability of easy to record proxies contributing to its predictive power in practical contexts.

Understanding ecological requirements of endangered species is a primary precondition of successful conservation practice. Regrettably, we know surprisingly little about the life history of numerous threatened insects, and about their use of larval host plants in particular. The brown butterflies (Nymphalidae: Satyrinae) have traditionally been considered polyphagous on grasses and indiscriminatory in their oviposition behavior. However, detailed studies on several species have revealed local specialization in host plant use as well as the decisive role of microlimatic conditions as determinants of habitat quality. The present study addresses host plant relationships in the endangered brown butterfly Coenonympha hero (L.) at the northern limit of its European distribution. We combine laboratory-based host preference and performance tests with an analysis of microhabitat use by adult butterflies in the field. Both lines of evidence suggest that C. hero is polyphagous enough not to be associated with one particular host species. Oviposition choices of C. hero are not driven by host plant species but rather by structural characteristics of the substrate. The preferred rigid needle-like structures may serve as cues of ‘transparent’ vegetation which allows the larvae to benefit from sunlight reaching the lower strata of the tuft. Our results suggest that conservation efforts should prioritize microclimatic parameters, rather than the presence of any particular host plant species, as decisive determinants of habitat quality in C. hero.

The conservation of forest biodiversity is increasingly relying on forest landscapes managed for timber production. A precondition for successful biodiversity conservation in intensively managed landscapes is a good understanding of habitat- and landscape-level drivers of species diversity. In this study, we investigated the effects of habitat characteristics and landscape context on the species richness and composition of butterflies in temperate forest landscapes routinely managed by clear-cutting. Our focus was on forest-dependent species. Even though about a quarter of the entire butterfly fauna in Europe are associated with forest habitats, the drivers of their diversity are poorly known. We used data from an unprecedentedly extensive butterfly survey, conducted in more than 400 sites distributed across all major forest types in Estonia (Northern Europe). Our data indicate that, generally, forests harvested for timber production remain suitable as habitats for forest butterflies: across the study sites, we recorded almost the full complement of their total regional species richness. By far the most influential driver of local butterfly species richness in such forests appears to be the proportion of forest cover in the surrounding landscape. The species richness of forest butterflies starts to show signs of decline at remarkably high levels of forest cover in the landscape. Surprisingly, landscape-scale forest heterogeneity around study sites had no discernible effect on local species richness, the pattern plausibly resulting from low mobility of forest butterflies. The most important habitat-scale predictor of butterfly species richness was soil moisture, with butterfly communities being most species-rich in forests on soils of intermediate moisture. Different forest types contribute to the regional species pool of forest butterflies in a complementary manner. No single forest type alone provides habitat for the entire species pool: peaks of abundance of different species are distributed across multiple forest types. Accordingly, a substantial part of the variation in species assemblages could be attributed to characteristic soil conditions (moisture and pH) of different forest types. Our findings imply that both the extent and heterogeneity of forest landscapes are important for maintaining the diversity of forest butterflies at the landscape scale.

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.

Diet breadth and the degree of capital breeding have been established as major determinants of species-specific ecology of herbivorous insects. Both of these variables are related to resource foraging and therefore can be expected to have effects on sensory capacity. However, such effects have remained poorly studied, let alone in phylogenetically explicit multi-species comparisons. We contribute to filling this gap in a study of 60 species of geometrid moths (Lepidoptera: Geometridae), using adult head measures (eye size, antennal length and shape, forehead width) as indices of sensory capacity. When controlled for body size, eyes of the more capital breeding species (i.e. those with low contribution of adult feeding to reproduction) were found to be smaller, and female antennae shorter, than in income breeders. Feathery (vs simple filiform) male antennae were more frequently present in the capital breeders and in larger species. Regarding diet breadth, generalist species were found to have relatively wider male foreheads than specialists. The results suggest that (a) breeding strategy rather than diet breadth predicts sensory capacity in geometrids, (b) capital breeding (vs income breeding) is related to low sensory capacity in females, and (c) in contrast, males of the capital breeding species have evolved towards elevated olfactory capacity.

Regionally oligophagous insects are often host plant specialists at smaller geographical scales, so conservation planning should preferably rely on locally derived information. Host use of the endangered butterfly Euphydryas aurinia was investigated close to the northern limit of its European distribution. We experimentally studied host preference of ovipositing females and growth performance of pre-diapause larvae on different hosts, as well as made field observations on host use by surveying larval webs. In the field, Succisa pratensis has remained the only confirmed host of E. aurinia in Estonia. Also, in the laboratory, the females consistently preferred S. pratensis as oviposition substrate. Larval performance was the highest on S. pratensis though the insects were also able to develop on Knautia arvensis. By contrast, species of Plantago, or Valeriana officinalis, were not suitable for larval development. E. aurinia appears thus to be a host specialist in Estonia, even if the regional list includes a higher number of plant species. Our results confirm a pattern repeatedly recorded in Melitaeine butterflies, i.e. geographic differences in host use and local monophagy. As a methodological contribution, we showed that different lab-based methods yield consistent information on host use, though tend to somewhat overestimate the actual host range of the species in the field. Using larval growth rate as an index of host suitability appears problematic for species not time-stressed during their larval development, and survival is a more appropriate parameter to record.

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 43.8 days (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.

An evolutionary explanation should consider the balance between environmentally-based selective pressures, and the resistance of the organism's phenotype to adaptive evolution, with the latter being captured by the concept of constraint. The limited attention to non-adaptive explanations in evolutionary ecology is at least partly caused by methodological difficulties with respect to identifying and quantifying constraints. As an example of an experimental approach evaluating a constraint-based explanation, we present a cross-species comparison of the shape of reaction norms for size and age at maturity. Instar- and sex-specific development times and final sizes were recorded for two distantly-related species of insects (Lepidoptera), with larval growth rates being manipulated by means of refined starvation treatments. We found that (1) the ‘classical’ L-shaped reaction norms for final size and development time are characteristic also of individual larval instars; (2) these responses show a high degree of quantitative similarity across the species, different larval instars, and sexes within species; and (3) the similarity among species and sexes is higher for the penultimate than for the final instar. The high degree of similarity suggests that some physiological mechanisms determining such reaction norms are evolutionarily conservative. An alternative explanation (i.e. quantitative similarity of ecologically based selective pressures) appears less likely. The results of a previous study on a third lepidopteran species not only support our general conclusions, but also provide a clear case of adaptive evolution in some aspects of such reaction norms. The present study shows one way how the data required to measure evolutionary conservatism in reaction norms for body size can be obtained empirically. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114, 296–307.

Tomasson, K., Tammru, T. & Kurina, O. 2014: Harvestmen (Arachnida: Opiliones) in Estonia: results of the Estonian Malaise Trap Project. — Entomol. Fennica 25: 142–156. The species composition of Estonian harvestmen (Opiliones) was revised based on a critical review of published data and examination of the material from the Estonian Malaise Trap Project (EMTP), which is briefly introduced. Four years ofcollecting(from2008to2011)with“Czechtype”Malaisetrapsfrom15localities throughout Estonia resulted in 4,102 specimens of 9 species. Two of them, Lophopilio palpinalis (Herbst, 1799) and Leiobunum tisciae (Avram, 1968), were found from Estonia for the first time. Combined with reliable published data, the number of harvestmen species known from Estonia is now set at 12. Distribution and habitat preferences of harvestmen in Estonia were examined subjectingtheMalaisetrappingdatatoaquantitativeanalysisusingenvironmentalvariablesrecordedforeachtraplocation.Onlytreelayerdensity(i.e.thepresence of forest as such) was identified as a significant predictor of harvestmen abundance and diversity which indicates a low degree of habitat specificity in boreo-nemoral harvestmen. K.Tomasson&O.Kurina,InstituteofAgriculturalandEnvironmentalSciences

The post-glacial recolonisation of northern Europe has left distinct signatures in the genomes of many organisms, both due to random demographic processes and divergent natural selection. However, ...

Volatile organic compounds (VOCs) released by host plants attract gravid European corn borer (ECB) female moths for oviposition. Despite extensive studies, little is known about VOCs emitted by maize under natural conditions or the odorscape of a maize field, particularly at the time of ECB oviposition. Here, we characterized VOCs released by undamaged maize plants and VOCs in the maize field odorscape. VOCs were collected throughout the diel cycle with solid-phase microextraction fibres. VOCs were identified by GC-MS and quantified with calibration curves. Four replicates per time period were collected; i.e., dusk, night, dawn, and day. VOC patterns differed between the maize plants and the maize field odorscape throughout the diel cycle. At night, the period of ECB oviposition, the VOC pattern was characterized by an increase in monoterpenes, a decrease in sesquiterpenes, and the presence of methyl salicylate, α-copaene, and Z-3-hexenyl acetate. An apparent discrepancy between maize plant and field odorscape VOC compositions was observed. Key compounds were identified as putative host-cues, including methyl salicylate, α-pinene, 3-carene, p-cymene, limonene, and dimethyl nonatriene. This study showed that VOCs were released by maize in a diel pattern, and host-characteristic cues were present for nocturnal ECB oviposition.

The distribution of butterflies was mapped throughout entire Estonia in 2016 and 2017. Butterflies were surveyed during three phenologically targeted visits at more than 1200 pre-selected sites. In total, over 180 000 individuals belonging to 97 species were recorded. We compare the resulting distribution maps with historical records, and discuss changes in the species' distribution. We conclude that, within the time frame of century, the changes in the Estonian butterfly fauna were moderate. For about 75% of species, there was no conclusive evidence of a change in distribution or abundance. In case of the remaining 25%, there were more butterfly species with an increasing rather than a decreasing trend in abundance and/or distribution. Some but not all of those changes could be associated with ongoing climate change. The decline of several dry-meadow specialists can be regarded as the most urgent conservation concern.

Our study aims to investigate the relationships of the major lineages within the moth family Geometridae, with a focus on the poorly studied Oenochrominae-Desmobathrinae complex, and to translate some of the results into a coherent subfamilial and tribal level classification for the family. We analyzed a molecular dataset of 1,206 Geometroidea terminal taxa from all biogeographical regions comprising up to 11 molecular markers that includes one mitochondrial (COI) and 10 protein-coding nuclear gene regions (wingless, ArgK, MDH, RpS5, GAPDH, IDH, Ca-ATPase, Nex9, EF-1alpha, CAD). The molecular data set was analyzed using maximum likelihood as implemented in IQ-TREE and RAxML. We found high support for the subfamilies Larentiinae, Geometrinae and Ennominae in their traditional scopes. Sterrhinae becomes monophyletic only ifErgaviaWalker,AmetrisHübner andMacrotesWestwood, which are currently placed in Oenochrominae, are formally transferred to Sterrhinae. Desmobathrinae and Oenochrominae are found to be polyphyletic. The concepts of Oenochrominae and Desmobathrinae required major revision and, after appropriate rearrangements, these groups also form monophyletic subfamily-level entities. Oenochrominaes.str. as originally conceived by Guenée is phylogenetically distant fromEpidesmiaand its close relatives. The latter is hereby described as the subfamily Epidesmiinae Murillo-Ramos, Sihvonen & Brehm,subfam. nov.Epidesmiinae are a lineage of “slender-bodied Oenochrominae” that include the generaEcphyasTurner,SystaticaTurner,AdeixisWarren,DichromodesGuenée,PhrixocomesTurner,AbraxaphantesWarren,EpidesmiaDuncan & Westwood andPhratariaWalker. Archiearinae are monophyletic whenDirceandAcalyphesare formally transferred to Ennominae. We also found that many tribes were para- or polyphyletic and therefore propose tens of taxonomic changes at the tribe and subfamily levels. Archaeobalbinistat. rev.Viidalepp (Geometrinae) is raised from synonymy with Pseudoterpnini Warren to tribal rank. Chlorodontoperini Murillo-Ramos, Sihvonen & Brehm,trib. nov.and Drepanogynini Murillo-Ramos, Sihvonen & Brehm,trib. nov.are described as new tribes in Geometrinae and Ennominae, respectively.

In North-East Estonia, considerable amounts of toxicants (e.g. polycyclic aromatic hydrocarbons (PAHs), phenols, heavy metals) leach into water bodies through discharges from the oil-shale industry. In addition, natural and anthropogenic hypoxic events in water bodies affect the health of aquatic organisms. Here we report a study on the combined effects of contaminated sediment and hypoxia on the physiology of gibel carp (Carssius auratus gibelio). We conducted a laboratory exposure study that involved exposure to polluted sediments from oil-shale industries (River Purtse) and sediments from a relatively clean environment (River Selja), together with sediments spiked with PAHs. The oxygen content (saturation vs. hypoxia (< 2 mg/L)) was changed to reflect hypoxia. A multi-biomarker approach was chosen to enable the combined effects to be assessed comprehensively and integratively. We used HPLC to measure the PAH concentration in sediment and fish muscle, fixed wavelength fluorescence (FF) analyses to indicate the presence of PAH metabolites in fish bile, and nuclear abnormalities in erythrocytes as markers of geno- and cyto-toxicity; and we monitored the change in body condition and measured EROD activity to indicate CYP1A induction. High levels of PAH conjugates in fish bile were found in the group exposed to the Purtse River sediment under hypoxia. The results suggested that induction of the CYP1A gene was modulated by hypoxia as well as by heavy metals. We found a correlation between several erythrocyte abnormalities (8-shaped nuclei and blebbed nuclei) and PAH metabolite content in fish. In conclusion, a measurable effect of pollution from the oil-shale industry on fish health parameters was clear under different oxygen levels.

High among-individual variation in mating success often causes problems in conservation breeding programs. This is also the case for critically endangered European mink and may jeopardize the long-term maintenance of the species' genetic diversity under the European mink EEP Program. In this study, breeding success of wild and captive born European minks at Tallinn Zoological Garden are compared, and the mating behavior of the males is analyzed. Results show that wild born males successfully mate significantly more often than captive born males (89% and 35%, respectively). On the basis of an extensive record of mating attempts, both male aggressiveness and passivity are identified as primary causes of the observed mating failures. All other potential determinants have only a minor role. Mating success as well as a male's aggressiveness and passivity are shown to depend more strongly on the male than the female partner. We did not find any evidence that the behavior of an individual is dependent on the identity of its partner. We suggest that aggressiveness and passivity are two expressions of abnormal behavior brought about by growing up in captivity: the same individuals are likely to display both aggressive and passive behavior. The results point to the need to study and modify maintenance conditions and management procedures of mink to reduce the negative impact of the captive environment on the long-term goals of the program.

Most insect populations are exploited by a complex of different parasitoid species, providing ample opportunities for competitive interactions among the latter. Despite this, resource-mediated competition (i.e., exploitative competition) among insect parasitoids remains poorly documented in natural systems. Here we propose a novel way to infer the presence of competitive interactions from covariance patterns in parasitism levels, and illustrate the use of this approach on a relatively well-defined and simple host–parasitoid system. The parasitism levels caused by three parasitoid species on a shared host showed a highly consistent negative covariance among samples. With the levels of parasitism by one species increasing, the levels of parasitism attributable to the two others decreased. Importantly, negative covariance between parasitism levels by different species appeared at high abundance, but not at low abundance of the phenologically earlier parasitoid species. This as well as several other lines of evidence indicates the importance of competitive interactions in this system. Feeding biology and phenology of the parasitoids suggest that competition in this parasitoid assemblage is primarily resource-mediated rather than occurring through direct interference. The species attacking earlier stages of the host are competitively superior to those attacking their host later in the season. Better dispersal ability and use of alternative host species by the inferior species could contribute to the coexistence of these competing parasitoids.

Numerous studies have suggested a general relationship between the degree of host specialization and body size in herbivorous animals. In insects, smaller species are usually shown to be more specialized than larger-bodied ones. Various hypotheses have attempted to explain this pattern but rigorous proof of the body size-diet breadth relationship has been lacking, primarily because the scarceness of reliable phylogenetic information has precluded formal comparative analyses. Explicitly using phylogenetic information for a group of herbivores (geometrid moths) and their host plant range, we perform a comparative analysis to study the body size-diet breadth relationship. Considering several alternative measures of body size and diet breadth, our results convincingly demonstrate without previous methodological issues-a first for any taxon-a positive association between these traits, which has implications for evaluating various central aspects of the evolutionary ecology of herbivorous insects. We additionally demonstrate how the methods used in this study can be applied in assessing hypotheses to explain the body size-diet breadth relationship. By analyzing the relationship in tree-feeders alone and finding that the positive relationship remains, the result suggests that the body size-diet breadth relationship is not solely driven by the type of host plant that species feed on.

The aim of the present study was to evaluate—in a geographic perspective—the role of host plant as a determinant of habitat quality for Lopinga achine, a satyrine butterfly endangered over much of its European range. Laboratory trials were performed to record host choices made by the ovipositing females as well as by neonate larvae. In rearing experiments, growth performance and mortality on different host plants was determined. Oviposition was found to be indiscriminate but larvae were shown to be able to choose between host plants, with the choices made broadly consistent with growth performance of the larvae on particular hosts. Nevertheless, most grasses and sedges offered were found to support larval development reasonably well. No clear superiority of the previously suggested primary host plant Carex montana could be shown. Importantly, no differences in host plant relationships were found between the populations of Sweden, western Estonia and eastern Estonia. In particular, the larvae originating from eastern Estonian populations developed on C. montana equally well even if the plant is absent from their native habitat. In the context of species conservation, one should conclude that L. achine is polyphagous enough on various grasses and sedges so that the presence of any particular host species cannot be a critical component of habitat quality. Nevertheless, some preference to broad- and soft-leaved hosts, as well as sensitivity to host wilting, may partly explain the butterfly’s preference to moist forest habitats, further emphasizing the central role of habitat management in the conservation practice of this species. In turn, the absence of ecological differences between geographic populations should enable conservationists to successful transfer their experience across national boundaries.

Seasonal generations of short-lived organisms often differ in their morphological, behavioural and life history traits, including body size. These differences may be either due to immediate effects of seasonally variable environment on organisms (responsive plasticity) or rely on presumably adaptive responses of organisms to cues signalizing forthcoming seasonal changes (anticipatory plasticity). When directly developing individuals of insects are larger than their overwintering conspecifics, the between-generation differences are typically ascribed to responsive plasticity in larval growth. We tested this hypothesis using the papilionid butterly Iphiclides podalirius as a model species. In laboratory experiments, we demonstrated that seasonal differences in food quality could not explain the observed size difference. Similarly, the size differences are not likely to be explained by the immediate effects of ambient temperature and photoperiod on larval growth. The qualitative pattern of natural size differences between the directly developing and diapausing butterflies could be reproduced in the laboratory as a response to photoperiod, indicating anticipatory character of the response. Directly developing and diapausing individuals followed an identical growth trajectory until the end of the last larval instar, with size differences appearing just a few days before pupation. Taken together, various lines of evidence suggest that between-generation size differences in I. podalirius are not caused by immediate effects of environmental factors on larval growth. Instead, these differences rather represent anticipatory plasticity and are thus likely to have an adaptive explanation. It remains currently unclear, whether the seasonal differences in adult size per se are adaptive, or if they constitute co-product of processes related to the diapause. Our study shows that it may be feasible to distinguish between different types of plasticity on the basis of empirical data even if fitness cannot be directly measured, and contributes to the emerging view about the predominantly adaptive nature of seasonal polyphenisms in insects.

We have studied the phylogeography of the red-listed Palearctic butterfly Lopinga achine (Nymphalidae: Satyrinae) based on 1,450 base pairs of mitochondrial DNA sequences from 86 individuals representing 12 populations. Our results indicate a strong structuring of genetic variation, with among-population differences accounting for ca. 67% of the variation and almost all populations being significantly differentiated from each other. We surmise that the insular nature of populations as well as the low dispersal ability of the species has given rise to such a pattern. The genetic diversity within populations is low compared to that in other butterflies. Our results point to a scenario where the species originated in the Eastern Palearctic and expanded into Europe. Based on the analyses, we suggest that the Czech population merits the highest conservation priority. The two Swedish populations represent a distinct evolutionary lineage, and hence merit high conservation attention. The Estonian and Asian populations had the highest genetic diversity, and although we do not consider them to be under immediate threat, their genetic diversity should be conserved in the long term.

Patterns of variability in quantitative traits across environmental gradients have received relatively little attention in evolutionary ecology. A recent meta-analysis showed that relative phenotypic variability in body size tends to decrease with improving environmental conditions. This pattern was explained by introducing the concept of upper threshold size to a general optimality model of individual growth but alternative explanations certainly exist. In particular, it is frequently observed in insects that variability in individual growth rates decreases with improving environmental conditions. Here we explore the effect of this phenomenon on environment-specific variability in adult sizes. A quantitative model shows that relative variability in adult sizes is independent of environmental quality if absolute variability in growth rates remains constant across the gradient of environmental quality. Deviations from this borderline case are definitely realistic in both directions. Both negative and positive relationships between relative variability of body size and environmental quality can thus be predicted to arise as a consequence of environment-specific variability in growth rates. The variability itself can be both genetic or environmental in its nature. We present empirical data which support both the assumptions and conclusions of our model-based analysis, as well as emphasize the advantages of controlled experiments for understanding the proximate sources of phenotypic variance.

It is generally believed that harsh climate inhibits growth and development of ectotherms at high latitudes. However, this environmental effect may be counterbalanced by countergradient genetic variation and habitat selection. While there is laboratory evidence for genetically based increases in growth and development rates in amphibians living at high latitudes, it remains unclear how environmental and genetic effects combine in the field to produce gradients of phenotypic variation. We performed a field study on the growth and development of tadpoles and habitat selection of breeding adults of the natterjack toad (Bufo calamita) in Denmark and in Estonia where the average onset of breeding differs by 12 days. Although growth and development of Estonian tadpoles trailed Danish conspecifics during the first half of the larval period, Estonian tadpoles caught up in both body length and developmental stage to Danish tadpoles in the last third of the larval period. Breeding ponds in Denmark were significantly larger and deeper than in Estonia, however, the water temperatures in the ponds did not vary substantially between the countries. In Estonia the toads selected shallower breeding ponds with higher temperature and oxygen concentrations, suggesting, that at higher latitudes active selection of specific water bodies ensures faster growth and development of larvae. The results, together with previous laboratory evidence, indicate that amphibian distribution limits are shaped both by adaptive differences in developmental rates and behavioural plasticity. To cope with harsh climates, amphibians in the north apparently require shallow water bodies — a habitat particularly vulnerable to anthropogenic degradation.

We present a phylogenetic comparative study assessing the evolutionary determinants of egg size in the moth family Geometridae. These moths were found to show a strong negative allometric relationship between egg size and maternal body size. Using recently developed comparative methods based on an Ornstein-Uhlenbeck process, we show that maternal body size explains over half the variation in egg size. However, other determinants are less clear: ecological factors, previously hypothesized to affect egg size, were not found to have a considerable influence in the Geometridae. The limited role of such third factors suggests a direct causal link between egg size and body size rather than an indirect correlation mediated by some ecological factors. Notably, no large geometrid species lay small eggs. This pattern suggests that maternal body size poses a physical constraint on egg size, but within these limits, there appears to be a rather invariable selection for larger eggs.

Assumptions about mortality rates form a cornerstone for models of life-history evolution. When seeking adaptive explanations for body sizes, the size dependence of predation risk is of particular interest. Here, we review published studies that provide (1) estimates of the daily predation rates experienced by insect larvae feeding on tree leaves or (2) evidence concerning the relationship between predation risk and larval size. Larvae were found to experience an average avian predation rate of 3.1% per day and an average arthropod predation rate of 10.5% per day. In some studies, mortality rates were systematically dependent on parameters of the larvae (e.g. coloration) or of the environment (host plant, season), but not to the extent that would render generalizations meaningless. Nevertheless, mortality rates varied considerably more for arthropod than avian predators, making an estimate of avian predation rate more reliable for use in quantitative models. Moreover, birds tend to be a more important predator guild exploiting the larger larval stages, as indicated by the predominantly positive size dependence of avian predation risk. By contrast, predation by arthropods was generally negatively size dependent. Based on the available data, we estimate that avian predation rates increase approximately 3.6-fold, while arthropod predation rates decrease approximately 4.9-fold, in response to a 2-fold increase in the linear size of prey. A modelling exercise showed that realistic mortality rates – if assumed to be independent of size – cannot serve as a basis for adaptive explanations for observed body sizes. However, by assuming a positive size dependence of mortality risk within the limits observed for bird predation, it is possible to explain a wide range of body sizes within an optimality framework. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 1–18.

Although the general concept of capital vs. income breeding has become widely used in insect ecology, finding easy-to-measure indices for quantifying the role of larval- vs. adult-derived nutrients in egg production has remained a challenge. When searching for possible candidates for this task, we evaluated the applicability of three morphometric ratios to be measured on freshly eclosed adults: (1) relative size of female abdomen, (2) sexual size dimorphism, and (3) proportion of volume of mature eggs relative to total volume of female abdomen. We report the values of these indices in five species of geometrid moths [Hypomecis punctinalis Scopoli, Ematurga atomaria L., Selenia tetralunaria Hufnagel, Semiothisa clathrata L., and Epirrhoe alternata Muller (all Lepidoptera: Geometridae)] and compare them to the degrees of income breeding measured directly by oviposition experiments. As a comparison, we also recorded the ovigeny index and the effect of income on lifespan. All morphometric indices varied considerably between the species studied, and, as predicted, the values of all three proposed indices were higher in the more capital-breeding species. The only exception to this pattern was a low proportion of mature eggs in abdomens in one primarily capital-breeding species, S. tetralunaria. Unlike other capital breeders, this species also proved to be largely synovigenic, indicating that capital breeding and pro-ovigeny are not strictly linked. The reproductive traits measured concord with ecological niches occupied by the species studied: the oligophagous meadow dwellers were largely income breeding and synovigenic, whereas the polyphagous forest dwellers were capital breeders. The high and predictable among-species variation in egg-production strategies suggests that geometrid moths form a promising target group for comparative studies on respective traits. Simple anatomical indices, in turn, appear to be applicable as easy-to-record proxies of egg production strategies.

Organisms have evolved morphological and behavioral traits that reduce their susceptibility to predation. However, few studies have considered how morphological structures work to provide informational inputs necessary for effective behavioral responses to predation risk. In this study, we demonstrate that the hairs of Orgyia leucostigma (Lymantriidae) caterpillars not only function as physical barriers that deter predators but also act to provide sensory inputs triggering behavior that reduces predation risk. In particular, the way in which caterpillars respond when their hairs are touched is predator specific. Mechanical parameters of the interaction determining the response were identified correlatively and confirmed manipulatively. Caterpillars predominantly dropped in response to high hair bending velocities and predominantly walked away in response to low hair bending velocities. These stimulus-specific responses appear to be adaptive as they led to increased survival. Our results demonstrate a functional link between morphology and behavior. The ability to respond effectively only after the initiation of a predator attack should reduce the costs associated with antipredator behaviors. Copyright 2011, Oxford University Press.

Environmental perturbations such as starvation and poor diet often prevent animals from attaining their optimal sizes. When the perturbation has a transient char- acter, compensatory responses are expected in terms of faster growth or a prolonged developmental period. In the case of insect larvae, details of such responses are insuf- ficiently known at the proximate level. Attention to responses at the level of particular larval instars should promote an understanding of insect developmental plasticity also in a more general context. To provide an instar-specific analysis of compensatory growth, larvae of the moth Orgyia antiqua (L.) are reared on inferior diet during one larval instar. Responses in growth parameters are recorded in the course of the manipulated instars, as well as at the level of the entire larval period. The negative relationship between development time and size in response to the inferior food quality, typical of the entire larval periods, is also observed within the manipulated instars taken separately. The manipulated larvae remain smaller than the larvae of the control group (significant in males only), even by the end of the subsequent instar during which all individuals are provided with superior host. In males, close to full size compensation by the time of pupation is achieved only by means of adding an extra larval instar. The inability of larvae to fully compensate during one and even two instars is considered as an indication of the presence of constraints on the within- instar growth pattern. An alternative, adaptational explanation for the incomplete compensation could be based on the cost of prolonged development period. Given the ecological context of the species' life history, such an explanation appears less likely.

Separating genetic and environmental causes of the latitudinal differences among populations is crucial when evaluating the potential for microevolutionary responses to the changing environment. We studied among-population and environmental components of variation in several life-history traits of a lichen-feeding moth Eilema depressum when offspring of replicate Swiss and Finnish females were reared in a common-garden factorial experiment. A partial second generation was produced only among Swiss larvae, more likely so at higher temperature regime and higher host quality, and more frequently among the offspring of particular females. Growth rates of larvae that chose the diapause development were higher in northern individuals. Our results thus reveal adaptive differences between latitudinal populations in studied life-history traits, allowing to expect rapid adaptation of the species to further environmental changes. In contrast, invariable responses of the growth rates of the larvae to temperature and host quality support the idea that some basic parameters of insect growth show a high degree of evolutionary conservatism.

1. Body size is positively correlated with fecundity in various animals, but the factors that counterbalance the resulting selection pressure towards large size are difficult to establish. Positively size-dependent predation risk has been proposed as a selective factor potentially capable of balancing the fecundity advantage of large size. 2. To construct optimality models of insect body size, realistic estimates of size-dependent predation rates are necessary. Moreover, prey traits such as colouration should be considered, as they may substantially alter the relationship between body size and mortality risk. 3. To quantify mortality patterns, we conducted field experiments in which we exposed cryptic and conspicuous artificial larvae of different sizes to bird predators, and recorded the incidence of bird attacks. 4. The average daily mortality rate was estimated to vary between 4% and 10%. In both cryptic and conspicuous larvae, predation risk increased with prey size, but the increase tended to be steeper in the conspicuous group. No main effect of colour type was found. All the quantitative relationships were reasonably consistent across replicates. 5. Our results suggest that the size dependence of mortality risk in insect prey is primarily determined by the probability of being detected by a predator rather than by a size-dependent warning effect associated with conspicuous colouration. Our results therefore imply that warningly coloured insects do not necessarily benefit more than the cryptic species from large body size, as has been previously suggested.

Different levels of sexual size dimorphism (SSD) have usually been explained by selective forces operating in the adult stage. Developmental mechanisms leading to SSD during the juvenile development have received less attention. In particular, it is often not clear if the individuals of the ultimately larger sex are larger already at hatching/birth, do they grow faster, or do they grow for a longer time. In the case of insects, the question about sexually dimorphic growth rates is still open because most previous studies fail to adequately consider the complexity of larval growth curve, the existence of distinct larval instars in particular. Applying an instar-specific approach, we analysed ontogenetic determination of female-biased SSD in a number of distantly related species of Lepidoptera. The species studied showed a remarkable degree of similarity: SSD appeared invariably earlier than in the final instar, and tended to accumulate during development. The higher weight of the females was shown to be primarily a consequence of longer development within several larval instars. There was some evidence of higher instantaneous growth rates of females in the penultimate instar but not in the final instar. Egg size, studied in one species, was found not to be sexually dimorphic. The high across-species similarity may be seen as an indication of constraints on the set of possible mechanisms of size divergence between the two sexes. The results are discussed from the perspective of the evolution of insect body size in general. In particular, this study confirms the idea about limited evolvability of within-instar growth increments. An evolutionary change towards larger adult size appears always to be realised via moderate changes in relative increments of several larval instars, whereas a considerable change in just one instar may not be feasible.