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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.

1. Juvenile growth trajectories evolve via the interplay of selective pressures on age and size at maturity, and developmental constraints. In insects, the moulting cycle is a major constraint on larval growth trajectories. 2. Surface area to volume ratio of a larva decreases during growth, so renewal of certain surfaces by moulting is likely needed for the maintenance of physiological efficiency. A null hypothesis of isometry, implied by Dyar’s Rule, would mean that the relative measures of growth remain constant across moults and instars. 3. We studied ontogenetic changes and allometry in instar‐specific characteristics of larval growth in 30 lepidopteran species in a phylogenetic comparative framework. 4. Relative instar‐specific mass increments (RMI) typically, but not invariably, decreased across instars. Ontogenetic change in RMIs varied among families with little within‐family variation. End‐of‐instar growth deceleration (GD) became stronger with increasing body size across instars. Across‐instar change in GD was conserved across taxa. Ontogenetic allometry was generally non‐isometric in both RMI and GD. 5. Results indicate that detailed studies on multiple species are needed for generalizations concerning growth trajectory evolution. Developmental and physiological mechanisms affecting growth trajectory evolution show different degrees of evolutionary conservatism, which must be incorporated into models of age and size at maturation.

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.

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.

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.

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.

The theory of life history evolution generally predicts a negative across-environment correlation between development time and size at maturity in response to variations in environmental quality. Deviations from this pattern occur under specific cir- cumstances. In particular, organisms may mature both early and at a small size when (1) some ultimate change (e.g. time constraint, resource exhaustion) in the environment precludes further growth, or (2) when there are predictable among-environment differences in mortality rates. The first scenario is frequently documented in insects but evidence for the second possibility is scarce. Here we report a crowding-induced plastic response resulting in a clear positive across-environment correlation between final weight and development time in a geometrid moth. The response was apparent during the entire larval period and in the last larval instar. Crowding also led to increased growth rates. As outbreaks have not been reported for this species it is unlikely that early pupation is a response to anticipated food shortage. Instead, we suggest that crowded larvae may perceive a higher risk of predation, perhaps because they are unable to distinguish conspecifics from potential predators. A possibility for a plastic increase in growth rate implies that the uncrowded larvae grow at submaximal rates, which indicates a cost of high growth rate.

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.

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.

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.

1. The majority of general life-history models treat the environment as being invariable through time, even though temporal variation in selective agents could dramatically change the outcomes, e.g. in terms of optimal size and time at maturity. For herbivorous insects, seasonal differences in food quality are reasonably well described, but seasonal dynamics of top-down selective forces are poorly documented. 2. The present study attempted to quantify seasonal changes in predation risk of folivorous insect larvae in temperate forest habitats. In a series of field experiments, artificial larvae were exposed to predators, and the resulting bird-inflicted damage was recorded. The trials were repeated regularly throughout the course of two summers. 3. A distinct peak of larval mortality was recorded in mid-June (the nestling period for most insectivorous passerine birds), after which predation risk declined to a plateau of 20–30% below the peak value. 4. The recorded pattern is interpreted as a consequence of seasonal changes in the number and behaviour of insectivorous birds, and the abundance of alternative food resources for these predators. 5. A quantitative analysis based on field data indicated that considering temporal variation in mortality in life-history models is crucial for obtaining realistic predictions concerning central life-history traits, such as final body size in different generations.

Summary 1The allometric relationship between growth rate and body mass has received considerable attention but different taxa have not been equally studied. In particular, a limited amount of information is available on growth allometry of insect larvae. 2In life-history studies, it is often assumed that insect larvae grow exponentially. This leads one to expect that potential rewards of extending growth periods are high in terms of increased adult body masses and fecundities. Therefore, it has been a challenge to find costs of large size that counterbalance the fecundity advantage of attaining larger sizes. 3This study examines the intraspecific growth allometry of lepidopteran larvae. Original methodology is proposed to address problems arising from the complexity of the insect growth curve, and the high sensitivity of growth rates to environmental conditions. To facilitate generalizations, larvae of 11 unrelated lepidopteran species were subjected to an identical study design. 4Instantaneous absolute growth rates of larvae were related to body size by an intraspecific allometric exponent in the range between 0·41 and 0·88. There were significant differences between the species but values of the exponent as high as 1 (exponential growth), and as low as 0 (linear growth) could safely be excluded. 5Instantaneous relative growth rates of larvae were typically 35% lower in their last instar when compared to the penultimate one. Using the exponential growth curve (i.e. assuming the constancy of relative growth rates) in modelling insect life-histories may therefore lead to substantially biased conclusions.

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.

While the study of colour patterns is a traditional subject of evolutionary ecology, there are various hypotheses which suffer from a lack of experimental evidence. One intriguing possibility is a trade-off between warning efficiency and detectability. After a certain size threshold, the detrimental effect of increased detectability can outweigh the benefits of warning colouration. One may thus expect corresponding patterns at the level of ontogenetic development: as juveniles grow, they should first acquire warning colouration, and then lose it again. We analysed this possibility in Orgyia antiqua, a moth species with hairy larvae which are poly- phenic with respect to the intensity of warning colouration. We detected a regular change in colour patterns through larval life. Indeed, the larvae tend to display warning colouration at intermediate sizes while dull colours dominate in fully grown larvae. In aviary experiments, we confirmed that the colourful phenotype is the one that causes the strongest aversion in birds. Nevertheless, the effect was rather weak and most of the larvae were still eventually consumed when found. Unexpectedly, for human subjects, the warningly coloured larvae were harder, and not easier to find among natural vegetation, most likely due to the disruptive effect of the aposematic colour pattern. Importantly, the trend was reversed in the largest size class, suggesting that the disruptive colouration loses its advantage as the larva grows. This is consistent with the actual patterns of size-dependence of colouration. We present evi- dence against an alternative explanation which relates size-related change in colouration to behavioural changes prior to pupation. We conclude that even if the efficiency of the warning effect plays a role in determining the size-dependence of colouration, the pat- tern may be largely explained by the effects of size-dependent detectability alone.

Carotenoid-based integumental coloration is often associated with individual performance in various animals. This is because the limited amount of the pigment has to be allocated to different vital functions. However, most of the evidence for the carotenoid-based trade-off comes from vertebrate studies, and it is unclear if this principle can be applied to insects. This possibility was investigated in Orgyia antiqua L. (Lepidoptera: Lymantriidae). The larvae of this species are polyphenic in their coloration, varying from a highly conspicuous combination of yellow hair tufts on black background to cryptic appearance with brown hair tufts. The conspicuous larvae are aposematic, advertising their aversive hairiness. The maintenance of different colour morphs in O. antiqua requires explanation, as an aposematic signal is expected to evolve towards monomorphism. Chromatographic analysis showed that the yellow coloration of the hair is based on the carotenoid pigment lutein ( α -carotene3,3’-diol). The colour of hair tufts was dependent on their carotenoid content. This justifies an expectation of carotenoid-based physiological trade-offs between aposematic coloration and individual performance. To test this hypothesis, we monitored life histories of differently coloured larvae reared on various host plants, recording their body sizes, growth rates, and mortalities in each instar. There was a significant but relatively low heritability of tuft coloration, which allowed us to expect environmental effects. We found no phenotypic associations between hair tuft colour and performance indices in O. antiqua larvae, neither did the quality of host plant affect the frequency of colour morphs. However, the frequency of colour morphs differed between larval instars. Our results suggest that carotenoidmediated physiological trade-offs are not involved in the maintenance of colour morphs in O. antiqua larvae, and factors other than individual condition should be responsible for the observed variability.

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.

In arthropods, the evolution of sexual size dimorphism (SSD) may be constrained by a physiological limit on growth within each particular larval instar. A high SSD could, however, be attained if the larvae of the larger sex pass through a higher number of larval instars. 2. Based on a survey of published case studies, the present review shows that sex- related difference in the number of instars is a widespread phenomenon among insects. In the great majority of species with a sexually dimorphic instar number, females develop through a higher number of instars than males. 3. Female-biased sexual dimorphism in final sizes in species with sexually dimorphic instar number was found to considerably exceed a previously estimated median value of SSD for insects in general. This suggests a causal connection between high female- biased SSD, and additional instars in females. Adding an extra instar to larval development allows an insect to increase its adult size at the expense of prolonged larval development. 4. As in the case of additional instars, SSD is fully formed late in ontogeny , larval growth schedules and imaginal sizes can be optimised independently. No conflict between selective pressures operating in juvenile and adult stages is therefore expected. 5. In most species considered, the number of instars also varied within the sexes. Phenotypic plasticity in instar number may thus be a precondition for a sexual difference in instar number to evolve.

While the ultimate causes and adaptive significance of sexual size dimorphism (SSD) have been extensively studied, the developmental mechanisms behind this phenomenon have received little attention. Going through an additional larval instar may form a specific way of achieving SSD in arthropods. In the present study, the mechanisms of SSD determination of two lymantriid moths, with marked SSD, were studied. In both species, females tended to go through an additional instar compared to males, and form pupae that were more than twice the weight of the males. To reveal the role of an extra instar, larval growth was monitored in the laboratory and the growth parameters were analysed as dependent on sex and developmental type (number of instars). Prolonga- tion of growth by means of adding an additional larval instar in females turned out to be the key mechanism in the determination of the highly female-biased SSD in the species studied. There is thus a developmental mechanism available that permits achieving a larger size by means of extending the growth period. This provides evidence against constraint-based evolutionary explanations for body sizes in insects. There was no considerable accumulation of SSD during earlier larval life when females went through more instars than males. In contrast, in those cases in which males and females had the same number of instars, SSD accumulated gradu- ally during the course of several larval instars. Longer growing period turned out to be a crucial mechanism leading to the female- biased SSD even when instar number did not differ between sexes, although higher instantaneous relative growth rates of females also played a complementary role in the latter case. Within sexes, an additional instar was characteristic of initially smaller larvae, as predicted by the "threshold size" hypothesis.

We studied if the population density experienced during larval development affects the reproductive schedule of Yponomeuta evonymellus L. (Lepidoptera, Yponomeutidae), a moth species characterized by outbreak population dynamics. More specifically, we predicted that reproduction would be delayed to facilitate emigration from sites with suboptimally high densities of conspecifics. We manipulated larval densities in the laboratory, as well as in those pupae collected from the extremes of natural densities. As the response, we also recorded the timing and sizes of egg clutches laid. The results did not support our initial predictions: the timing of oviposition was not dependent on larval growth conditions. This apparent lack of adaptation might be related to the loss of ‘memory’ during metamorphosis in holometabolous insects. However, oviposition schedules were also only minimally sensitive to elevated adult density. An inability to respond to high larval densities may contribute to the outbreak dynamics in this species.

It is often claimed that some organisms maximise individual growth rates (within limits set by physiological and environmental constraints) whereas others do not. Adequate experimental protocol to test for these hypotheses has not been available. We proceeded from the idea that a maximiser should be unable to further improve its growth performance, i.e. there should be no adaptive plasticity in relative growth rates. In the case of an insect larva always maximising its weight gain, we should thus expect no increase in growth rate following a short-term environmental perturbation. Another prediction is that initial and final weights of a larval instar should be highly correlated. We applied this approach to test for the growth rate maximiser status of the geometrid moth Epirrita autumnata. Based on various lines of ecological evidence it had previously been suggested that this spring-feeding species is strongly selected for high growth rates, and may be growing at its maximum physiological potential. Contrary to expectations, the larvae responded to starvation treatments by a subsequent compensatory increase in their relative growth rates. Such a plasticity may indicate that the rates of gaining weight are not maximised even in a ‘classical’ example a time-constrained organism. This observation supports the idea that endogenous regulation of growth rates is widespread, and it remains to be shown if true maximisation ever exists in nature. In contrast, possibilities to compensate for adverse conditions experienced early in larval development appeared to be limited. This was indicated by the strong correlation between initial weight in the last instar, and pupal weight. Though consistent with our expectations, such an inability to compensate can hardly be ascribed to growth rate maximisation, as indicated by quantitatively similar patterns found in an unrelated species with different ecology, the butterfly Polygonia c-album.

Refuge theory proposes that a determinant of species richness (and percent parasitism) is the presence of host refuges. Plant structural refuges are good predictors of species richness for endophytic herbivores, but not for exophytic herbivores. For exophytic herbivores other traits such as relative abundance may provide refuge from parasitism. Using unbiased data on both relative abundance and larval parasitism of species in macrolepidopteran assemblages we tested the null hypothesis that percent parasitism was independent of abundance. Numerically subdominant species do not gain refuge from parasitism by persisting at low abundance. Parasitism was not different from what would be expected based on their numbers. Among numerically dominant species there was, nevertheless, a significant positive association between abundance and parasitism rate. However, dominant herbivores displayed high levels of parasitism even in the years when their abundance was low relative to other years. This suggests that dominant species may possess other traits that enhance their susceptibility to parasitoids.

Summary Spatiotemporal variation in the degree of melanism is often considered in the context of thermal adaptation, melanism being advantageous under suboptimal thermal conditions. Yet, other mutually nonexclusive explanations exist. Analysis of geographical patterns combined with laboratory experiments on the mechanisms of morph induction helps to unveil the adaptive value of particular cases of polyphenism. In the context of the thermal melanism hypothesis and seasonal adaptations, we explored an array of environmental factors that may affect the expression and performance of nonmelanic vs. melanic larval morphs in different latitudinal populations of the facultatively bivoltine moth Chiasmia clathrata (Lepidoptera: Geometridae). Geographical variation in larval coloration was independent of average temperatures experienced by the populations in the wild. The melanic morph was, however, more abundant in dry than in mesic habitats. In the laboratory, the melanic morph was induced especially under a high level of incident radiation but also at relatively high temperatures, but independently of photoperiod. Melanic larvae had higher growth rates and shorter development times than the nonmelanic ones when both temperature and the level of incident radiation were high. Our results that melanism is induced and advantageous in warm desiccating conditions contradict the thermal melanism hypothesis for this species. Neither has melanism evolved to compensate time constraints due to forthcoming autumn. Instead, larvae solve seasonal variation in the time available for growth by an elevated growth rate and a shortened larval period in the face of autumnal photoperiods. The phenotypic response to the level of incident radiation and a lack of adaptive adjustment of larval growth trajectories in univoltine populations underpin the role of deterministic environmental variation in the evolution of irreversible adaptive plasticity and seasonal polyphenism.

The effect of larval body size of Epirrita autumnata (Lepidoptera, Geometridae) on the risk of parasitism was studied in a field experiment. The experiment involved three pairwise exposures of different larval instars to parasitoids. Three hymenopteran species were responsible for most of the parasitism. Parasitism risk was found to be host-instar independent. This result was consistent across parasitoid species and experiments. The results suggest that host use by larval parasitoids cannot constrain selection for larger body size in E. autumnata. However, high mortality due to parasitism may select for a short developmental period (the slow-growth/high-mortality hypothesis), and smaller body sizes as a by-product. A strong selective effect of parasitism on the timing of larval development in E. autumnata is also unlikely. The larger was the host, the larger was the adult size of the parasitoid and the shorter its development time (for one species). We suggest that the lack of a preference-performance linkage in the system studied may be related to the time stress associated with the short phenological window of host vulnerability.

For three species of geometrid moths with noneruptive population dynamics, physiological and behavioral responses of ovipositing females to larval host deprivation were recorded. Single substrate trials were used. Availability of adult food was also varied to manipulate physiological condition of the moths. Egg production rate decreased only weakly in the absence of host plant, whereas it was strongly influenced by adult feeding. The time from the beginning of the experiment to initiation of oviposition was largely determined by presence of suitable host. This variable, oviposition latency, was only weakly affected by nutritional status of the females. Oviposition rates also remained lower on unsuitable hosts after oviposition was initiated. This indicates that the effect of inferior oviposition substrate was not limited to postponing oviposition. Host deprivation and adult feeding both lead to accumulation of chorionated eggs in abdomens. However, high egg loads of the fed females did not alter their behavior in the direction predicted by optimality models. We conclude that oviposition latency was the only variable in which female responses to unsuitable hosts were sufficiently strong to have potential significance for population dynamics. Latency presumably corresponds to dispersing behavior in nature. Such a response, not having been observed in an outbreaking geometrid studied earlier, may reflect a behavioral mechanism capable of stabilizing local population dynamics. The results are discussed in the light of the gradation from capital to income breeding. In herbivorous insects, oviposition latency may serve as an index of overall sensitivity of ovipositing females to adverse environmental conditions.

We sought to distinguish between two potential effects of larval crowding on developmental schedules in Epirrita autumnata, an outbreaking folivorous moth. We asked if consequences of crowding can be entirely attributed to deterioration of environment, caused by mutual disturbance of larvae, or if there are specific modifications of reaction norms involved. In a laboratory rearing experiment, we found that most of the consequences of larval crowding could be explained as unspecific environmental effects. However, there was some evidence of crowding-related modifications of reaction norms. In particular, crowding during early development more strongly affected further growth of larvae than expected from the direct influence of inferior environmental quality: larvae crowded early in development had shorter growth periods in the final instar and attained lower final weights. It is not clear if these modifications of reaction norms are adaptive. Nevertheless, these observations document a type of plasticity which is different from unidimensional responses to variation in environmental quality, and may thus contribute to understanding evolution of body size in insects. However, recorded modifications of reaction norms are apparently weak enough to have only a minor role in population dynamics of E. autumnata.

The relative roles of genetic differentiation and developmental plasticity in generating latitudinal gradients in life histories remain insufficiently understood. In particular, this applies to determination of voltinism (annual number of generations) in short-lived ectotherms, and the associated trait values. We studied different components of variation in development of Chiasmia clathrata (Lepidoptera: Geometridae) larvae that originated from populations expressing univoltine, partially bivoltine or bivoltine phenology along a latitudinal gradient of season length. Indicative of population-level genetic differentiation, larval period became longer while growth rate decreased with increasing season length within a particular phenology, but saw-tooth clines emerged across the phenologies. Indicative of phenotypic plasticity, individuals that developed directly into reproductive adults had shorter development times and higher growth rates than those entering diapause. The most marked differences between the alternative developmental pathways were found in the bivoltine region suggesting that the adaptive correlates of the direct development evolve if exposed to selection. Pupal mass followed a complex cline without clear reference to the shift in voltinism or developmental pathway probably due to varying interplay between the responses in development time and growth rate. The results highlight the multidimensionality of evolutionary trajectories of life-history traits, which either facilitate or constrain the evolution of integrated traits in alternative phenotypes.

The role of pathogens in insect ecology is widely appreciated but remains insufficiently explored. Specifically, there is little understanding about the sources of the variation in the outcome of insect-pathogen interactions. This study addresses the extent to which immune traits of larvae and pupae of the mothOrgyia antiquaL. (Lepidoptera: Lymantriidae) depend on the host plant species and individual condition of the insects. The two host plants,Salix myrsinifoliaSalisb. andS. viminalisL., were chosen because they differ in the concentration of phenolic glycosides, harmful to most polyphagous insects. Individual condition was assumed to be reflected in body weight and development time, and was manipulated by rearing larvae either singly or in groups of four. The resistance traits recorded were survival and time to death after fungal infection in the larval stage and the efficiency of encapsulating a nylon implant by the pupae. The survival of the infected larvae was mainly determined by the species of the host plant. Encapsulation response was not associated with the resistance to the pathogen, suggesting that the host plant affected the pathogen rather than the immune system of the insect. Interestingly, the host plant supporting better larval growth led to inferior resistance to the pathogen, indicating a trade-off between different aspects of host plant 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.

It is not clear which selective pressures balance the strong fecundity advantage associated with large female body size in insects. A positively size-dependent mortality risk could provide a solution. In aviary experiments with artificial larvae, we studied if larger larvae of folivorous insects are more readily found (= detectability) and/or attacked (= acceptability) by birds. As size and colouration are likely to interact in determining birds’ responses, both cryptic and conspicuous prey items were used. As detectability is likely to be context-dependent, both simple (smooth) and complex (plants) backgrounds were used in respective experiments. In the conspicuous larvae, acceptability correlated negatively with prey size. However, their detectability was context dependent, being positively correlated with size on the simple background, whereas no significant effect was found on the complex background. Surprisingly, cryptic larvae showed no correlation between detectability and size, and there was only a weak tendency for birds to attack large larvae more readily. On the basis of a quantitative model, we conclude that the effect of positively size dependent bird predation, as a single factor, is not likely to counterbalance the fecundity advantage in cryptic species, and may thus not be crucial in determining the optimum for body sizes in these insects. In conspicuous species, there is a potential for different outcomes, because detectability and acceptability affect survival in different directions. The net outcome is, therefore, likely to be highly context-dependent. Furthermore, our results provide an explanation for the recently reported absence of systematic body-size differences between cryptic and conspicuous Lepidopteran larvae: although conspicuous larvae benefit from increasing their warning signal when growing larger, they also suffer a much sharper rise in detectability.

Potential fecundity of Epirrita autumnata females is strongly correlated with pupal weight. Adult feeding weakly increases fecundity but does not eliminate its dependence on body size. Daily numbers of eggs laid are positively correlated with body size. However, night minimum temperature is the main determinant of oviposition rate under fluctuating natural temperatures. Adult mortality was estimated to be low in a subarctic forest in autumn. Based on these results, we constructed a simulation model predicting oviposition rate and realized fecundity under given weather conditions. The model indicates that expected realized fecundity strongly depends on female size also under field conditions. This causes selection for large body (abdomen) size which is a probable reason for lack of fine-tuned adult behaviour in E. autumnata. Factors acting during the adult stage can be considered relatively unimportant in the demography of this species; populations are largely regulated by biotic interactions of the larval stage, involving delayed density-dependence. This may be an explanation for population cycles, characteristic for this species.