Achieving high sexual size dimorphism in insects: females add instars

Abstract.  1. 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.     

Developmental variation in the forest tent caterpillar: life history consequences of a threshold size for pupation

In insects, size and age at adult emergence depend on larval growth that occurs in discrete steps or instars. Understanding the mechanisms controlling stepwise larval growth and the onset of metamorphosis is essential to the study of insect life history. We examined the patterns of growth of forest tent caterpillars Malacosoma disstria to quantify variation in the number of instars that larvae undergo before pupation, to identify the mechanisms underlying variation in larval development, and to evaluate the life history consequences of this variation. All caterpillars were reared under the same conditions; at each molt, the date, the head capsule width and the mass of the freshly molted insect were recorded. Logistic regression analysis showed that a threshold size (measured either as mass or head capsule width) must be reached at the beginning of a stadium for pupation to occur at the next molt. This threshold size was higher for females than for males, and as a result, females attained a higher pupal mass than males. To achieve this larger size, females often required more instars than males, despite a higher growth ratio (size increase within an instar). Within each sex, slow growing individuals exhibited more larval instars and longer larval development time, but attained the same pupal mass as faster growing individuals. The combination of a threshold size for pupation, discrete growth steps and variation in the number of these steps can thus complicate relationships between growth rate, pupal mass and larval development time. In our study, growth ratio and number of instars were correlated with development time but not with pupal mass, and no relationship was observed between development time and pupal mass. These findings imply that, in species with variable instar number, one cannot extrapolate overall larval growth from growth during a single instar. Given the constraints of discrete larval growth, variation in instar number provides greater flexibility for insects to compensate for poor growing conditions. In this case, inferior larval growth conditions don't necessarily lead to smaller adult size.     

Inter- and intra-specific differences in the number of larval instars in British populations of 24 species of stoneflies (Plecoptera)

1. Ontogenetic changes during the life cycle of aquatic insects are important not only in life‐history studies but also in evaluating food‐web structure. They require information on the growth and number of larval instars but such information is lacking for many species, including Plecoptera. Therefore, the chief objectives of the present study were to determine inter‐ and intra‐specific differences in the number of larval instars in British populations of 24 species of stoneflies, to test Dyar’s hypothesis that growth followed a geometric progression, and to synthesise this information with previously published values for four British species. 2. Larvae were reared at constant temperatures in the laboratory from eggs from 63 populations (one to six populations per species). First instars from each population were divided into three batches and each batch was reared at one of three constant temperatures. For each species, the rearing temperature and source population had no significant effect on the mean size of each larval instar. 3. The relationship between the geometric mean length of each instar and instar number was well described by an exponential equation (P < 0.001, r² > 0.9 for all species), thus supporting Dyar’s hypothesis. Only one species, Brachyptera risi, had the same number of instars for males and females (12–13). For the other 15 herbivorous species and the four smaller carnivorous ones, the number of instars was higher for females than males (range 11–16 for males, 12–17 for females). The larger size of the females was due to their additional instars, not a sex difference in growth rates. In contrast, there was a clear growth separation of the sexes after the 9th or 10th instar for the four largest carnivores. The number of larval instars was highest for these four species (range 16–19 for males, 18–23 for females), and females were much larger than males. 4. A multiple regression equation with data from the present and previous studies (n = 27) showed that variability in the mean length of the first instar and the maximum number of larval instars for each species accounted for 88% and 91% of the variability in the mean length of the final instar for males and females, respectively. 5. Values for Plecoptera in other countries were in general agreement with those in the present study, especially in the same families. Two old, but widely quoted, high values are doubtful. The present study and four previous ones provide a sound basis for ontogenetic studies on 28 species of Plecoptera and their role in aquatic ecosystems.     

Larval development of Diploptera punctata reared alone and in groups

Larvae of the cockroach Diploptera punctata were reared in isolation, in pairs, or in groups of 8–10. Duration of larval development, age at each ecdysis, weights at birth and ecdyses, and adult head-capsule width were measured. Duration of larval development was longer and adult size was larger in isolated animals than in animals reared in pairs and groups. The effect of isolation on development was more pronounced in males. All females had 4 larval instars, whereas males had 3 or 4 instars. The proportion of males with 4 larval instars was higher among animals reared in isolation. There was no difference in the duration of larval development or adult size between pair- and group-reared animals. The sex of animals in the group did not affect adult size or the duration of larval development. Males which underwent 3 or 4 larval instars had different schedules of moulting. Rates of growth of males of both instar types reared in isolation and pairs were similar. Greater adult weight of isolated animals and 4-instar-type males was a result of their longer duration of larval development. Both a higher rate of growth and longer duration of larval development contribute to the larger adult size of females than males.     

Determination of adult size in a folivorous moth: constraints at instar level?

1. Reaction norms for size and age at maturity were studied in Epirrita autumnata (Lepidoptera, Geometridae). Growth rates were manipulated by rearing larvae on different levels of food quality and quantity, and instar-specific final weights and development times were recorded.
2. Food level and initial weight of an instar accounted for most of the variance in final weights. Sexual dimorphism in pupal weights could be entirely ascribed to sex differences in initial weights of the last instar.
3. There were problems with considering the reaction norms optimal within the conventional demographic explanatory framework. Because fecundity increases linearly with body size and no costs of large adult size are known, one should expect female larvae to grow larger to increase their individual fitness. It is therefore likely that constraints play a major role in the determination, and evolution, of size in this species.
4. A focus on individual instars may be the best way to reveal the constraints on, and space for, adaptive evolution of insect growth. Some limit to the initial:final weight ratio of an instar, and the fixed number of instars, may represent important constraints.
5. Reaction norms like the ones described in this study lead to strong environmental determination rather than canalization of body size. Food quality throughout larval development may thus be very important for individual fitness and population dynamics in E. autumnata.     

Life history and population biology of the giant ostracod Leuroleberis zealandica (Baird, 1850) (Myodocopida)

A population of this large myodocopid ostracod was studied over 2 yr by random core-sampling of the medium sand bottom at Kaikoura, New Zealand. Leuroleberis zealandica (Baird) passes through seven instars, it is sexually mature only in the final instar and sexes were distinguishable from instar IV. Males and females were equally abundant except in the final instar when the morphologically distinct males were rarely found. The population consists of three cohorts at any one time and each cohort appears to split into fast- and slow-growing individuals during the sixth instar resulting in life times of 1.8–2.0 and 2.7–3.1 yr, respectively. Females produce only one brood of 37 eggs on average per life time that are carried throughout the 5–6 month development period during which there is no loss of embryos. Recruitment is discrete with most broods released in midsummer when the population density may exceed 350·0.1 m^?2. A second lesser recruitment may occur in early spring in some years. Hatched juveniles released from the female grow rapidly to instar IV within 6 months and, although size increments at each moult are proportionally similar, intermoult periods tend to increase with size with some variation according to seasonal growth rates. Instar life tables constructed from instar density data showed a large difference in the frequency of embryos initiating each cohort, very different mortalities at recruitment between cohorts, and that the mortality rates between instars I and VI of different cohorts appear to be independent of density. The biology of Leuroleberis is compared with the few published accounts of myodocopid biology. In addition, several aspects of the biology of myodocopids are reviewed. These include numbers of instars in different taxa, within-instar sexual size disparities, numbers of broods per female life time, egg and brood sizes in relation to adult female size in various taxa, and the question of post-adult moulting.     

The effect of host developmental stage at parasitism on sex‐related size differentiation in a larval endoparasitoid

• 1 For their larval development, parasitoids depend on the quality and quantity of resources provided by a single host. Therefore, a close relationship is predicted between the size of the host at parasitism and the size of the emerging adult wasp. This relationship is less clear for koinobiont than for idiobiont parasitoids.
• 2 As size differentiation in host species exhibiting sexual size dimorphism (SSD) is likely to occur already during larval development, in koinobiont larval endoparasitoids the size of the emerging adult may also be constrained based on the sex of the host caterpillar.
• 3 Sex‐specific growth trajectories were compared in unparasitised Plutella xylostella caterpillars and in second and fourth instar hosts that were parasitised by the solitary larval koinobiont endoparasitoid Diadegma semiclausum. Both species exhibit SSD, where females are significantly larger than males.
• 4 Healthy female P. xylostella caterpillars developed significantly faster than their male conspecifics. Host regulation induced by D. semiclausum parasitism depended on the instar attacked. Parasitism in second‐instar caterpillars reduced growth compared to healthy unparasitised caterpillars, whereas parasitism in fourth‐instar caterpillars arrested development. The reduction in growth was most pronounced in hosts producing male D. semiclausum.
• 5 Parasitism itself had the largest impact on host growth. SSD in the parasitoid is mainly the result of differences in growth rate of the parasitoid–host complex producing male and female wasps and differences in exploitation of the host resources. Female wasps converted host biomass more efficiently into adult biomass than males.

     

Searching for constraints by cross‐species comparison: reaction norms for age and size at maturity in insects

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.     

Different paths to sexual size dimorphism in two praying mantids,Pseudomantis albofimbriata and Hierodula majuscula

Sexual size dimorphism （SSD） is widespread among diverse animal taxa and has attracted the attention of evolutionary biologists for over a century. SSD is likely to be adaptive and the result of divergent selection on different size optima for males and females, given their different roles in reproduction. The developmental trajectory leading to SSD may help us to understand how selection acts on male and female size. Here, we describe the growth and development of two Australian praying mantids, Pseudomantis albofimbriata and Hierodula majuscula including the number of moults, time to adulthood, size at each moult, and the degree of SSD. While both species exhibit the common pattern of female-biased SSD, the number of moults required for individuals to reach adulthood differed between males and females and between species. Despite their larger adult size, P. albofimbriata females require fewer moults and less time than males to reach adulthood, but are significantly larger than males from the second instar onwards. In contrast, H. majuscula males reached adulthood in fewer moults, and less time than females, however males and females did not differ in size until females went through their final moult into adulthood. H. majuscula also required more time and more moults to reach adulthood than 17. albofimbriata. We discuss these different developmental pathways in light of the existing knowledge of reproductive biology for each species. We also suggest that these differences may relate to the different phenologies that occur in strongly seasonal temperate environments compared with those in the tropics. This study provides evidence that SSD can result from two different patterns of growth and development in closely related species.     

Regulation of body size during larval development in the German cockroach, Blattella germanica

It may be advantageous for insects to attain a certain standard size when they become adult. Recent studies have demonstrated that in some species of insects, a threshold size must be reached in the last instar in order that the adult will attain the standard size. It has been revealed also, however, that there may be another type of regulation of body size during larval development in the German cockroach. Head width and other characteristics were measured in every larval instar up to the adult. Body size is distributed clearly into a small group and a large group in the 5th instar, and even in the 4th instar. This suggests that a larva has already determined the number of following moults by the late 3rd instar. Adult size is almost the same in 5- and 6-instar types. Among individuals which require the same number of instars to reach the adult stage, the coefficient of variation in body size increases until the 3rd instar then decreases to the adult stage. Smaller individuals in a particular instar after the 3rd tend to grow more than larger ones during that instar. These results suggest that a larva regulates its body size after the 3rd instar in order to attain the ‘norm’ for adult size.     

宁波滑蜥两性异形和雌性繁殖

蜥蜴的雌性繁殖特征对理解两性异形的进化原因起着重要作用。于2011年4月在安徽滁州采集宁波滑蜥(Scincella modesta),定量研究该种形态特征的两性异形和雌性繁殖特征,检验成体形态特征两性异形与雌性繁殖的相关性。研究共采集43条(17♀♀,26♂♂)宁波滑蜥,雄性和雌性个体的最大体长分别为47.4 mm和46.6 mm。雌雄两性在体长和头宽上没有差异,而在腹长和头长上差异显著,雄性有较大的头长,雌性有较大的腹长。宁波滑蜥年产单窝卵。窝卵数和窝卵重与雌体体长及腹长呈正相关,卵重与雌体体长无相关性。窝卵数及卵重的变异系数分别为0.20和0.12。卵长径与窝卵数呈负相关,而卵短径与窝卵数无关。雌体主要通过增加窝卵数来增加繁殖输出。这些结果表明,宁波滑蜥是雌雄个体大小同形的两性异形模式,性选择使得雄性有着较大的头长,以具有较高的交配成功率,生育力选择使得雌性有着较大的腹长,以具有较大的生育力和繁殖输出。     

Catch‐up growth in the rhinoceros beetle Trypoxylus dichotomus (Coleoptera: Scarabaeidae): Smaller neonates gain relatively more body mass during larval development

Body size often varies among conspecific neonates. As larger adults generally have higher fitness than smaller conspecifics, it is adaptive for smaller neonates to subsequently gain relatively more size increments during larval development (catch‐up growth). Although catch‐up growth has been suggested in insects, inappropriate methods have been used to examine the size dependence of growth increments. Therefore, it remains unclear to what extent catch‐up growth is common among insects. The present study examined the size dependence of growth increments among larvae of Trypoxylus dichotomus using reduced major axis regression of final to initial body masses. Catch‐up growth was found consistently for larval instars. Furthermore, simulations of the size increments revealed that not only sexual divergence of the mean size, but also catch‐up growth within sexes plays a role in the development of sexual divergence in the body size distribution of T. dichotomus. The significance of catch‐up growth in body size evolution was discussed.     

A Quantitative Analysis of Growth and Size Regulation in Manduca sexta: The Physiological Basis of Variation in Size and Age at Metamorphosis

Body size and development time are important life history traits because they are often highly correlated with fitness. Although the developmental mechanisms that control growth have been well studied, the mechanisms that control how a species-characteristic body size is achieved remain poorly understood. In insects adult body size is determined by the number of larval molts, the size increment at each molt, and the mechanism that determines during which instar larval growth will stop. Adult insects do not grow, so the size at which a larva stops growing determines adult body size. Here we develop a quantitative understanding of the kinetics of growth throughout larval life of Manduca sexta, under different conditions of nutrition and temperature, and for genetic strains with different adult body sizes. We show that the generally accepted view that the size increment at each molt is constant (Dyar’s Rule) is systematically violated: there is actually a progressive increase in the size increment from instar to instar that is independent of temperature. In addition, the mass-specific growth rate declines throughout the growth phase in a temperature-dependent manner. We show that growth within an instar follows a truncated Gompertz trajectory. The critical weight, which determines when in an instar a molt will occur, and the threshold size, which determines which instar is the last, are different in genetic strains with different adult body sizes. Under nutrient and temperature stress Manduca has a variable number of larval instars and we show that this is due to the fact that more molts at smaller increments are taken before threshold size is reached. We test whether the new insight into the kinetics of growth and size determination are sufficient to explain body size and development time through a mathematical model that incorporates our quantitative findings.     

Dimorphic growth patterns and sex‐specific reaction norms in the butterfly Lycaena hippothoe sumadiensis

Directly developing larvae of the butterfly Lycaena hippothoe sumadiensis exhibited two growth strategies with one cohort passing four larval instars at high growth rates, and the other five instars at lower growth rates. The 4‐instar‐cohort displayed decreased development times, in combination with slightly reduced pupal and adult weights. In addition to adjustment of growth rate, omitting a larval instar may comprise a further mechanism to decrease development time when needed. Using the 4‐instar‐cohort, sex‐related differences in reaction norms were investigated over a temperature gradient. At high temperatures, protandrous males showed early emergence at a reduced size, whereas weight of females remained similar throughout. These differences suggest that large size is more important for female than for male fitness. The pattern is similar to that previously reported for alpine L. tityrus, indicating that sex‐specific reaction norms might be widespread in species living under severe time constraints.     

Size and scaling of sexually-selected traits in the lizard, Uta palmeri

Differences between the sexes in overall body size and in the size of other morphological traits, relative to overall body size, are common in many animals. In this study, patterns of growth and scaling of sexually dimorphic tratis are assessedin a lilzard and then used to sugest general developmental mechanisms responsible for sexual size dimorphism (SSD). Adult make Uta palmeri lizards are larger than adult females inoverall body size (snout-vent length, SVL), body mass, jaw length head width, and head depth. Two general growth processes produce this adult SSD. First, juvenile males have greater annual SVL growth rates than do juvenile females, contributing to adult SSD because males will be larger than females in any trait positively correlated with SVL. Secondly, males and females differ in age-related changes in growth of the three head size traits, relative to growth in SVL. Comparing slopes from reduced major axis regressions of each trait on SVL reveals that the sexes do not differ in the scaling of these traits as juveniles, but as adults males have greater slopes than adult females, indicating ontogenetic differences in scaling of these traits in males. Two other topics in SSD are addressed with these data. First, comparing these data on scaling to those of an earlier analysis that used ordinary least squares regression reveals that conclusions about underlying mechanisms in an analysis of scaling can be altered by the choice of a regression model. Secondly, these data indicate that postmaturational differences in scaling contribute to adult sexual size differences, contrary to an earlier study. Shine (1990) found that for many ectotherms, which continue to grow after sexual maturation, post-maturational events contribute little to sexual differences in overall body size. Results for U. palmeri suggest that these findings may only hold for measures of overall body size (e.g. SVL) and may not generalize to traits that exhibit sex difference in scaling.     

Variation in growth and instar number in field and laboratory Manduca sexta

The tobacco hornworm Manduca sexta has been an important model system for understanding physiological control of growth, development and metamorphosis of insects for more than half a century. Like all Manduca, M. sexta typically has five larval instars, with developmental commitment to metamorphosis occurring early in the 5th (final) instar. Here we show that M. sexta from a field population in North Carolina (USA) shows substantial intraspecific variation in the number of larval instars when feeding on a modified artificial diet. Individuals with six instars consistently exhibited slower growth rates during early larval development than individuals with five instars. The frequency of individuals with six instars decreased with increased rearing temperature. In contrast, M. sexta from a laboratory colony consistently had five instars, and had more rapid larval growth rates than M. sexta from the field. We identify a threshold body size at the start of the 5th instar that predicts whether an individual will have five (greater than 600mg) or six instars (less than 600mg). Variation in field populations in Manduca provides an important resource for understanding physiological control, developmental plasticity and evolution of growth rate, body size and instar number.     

Sexual differences in the absolute growth pattern of the Indo-Pacific sandy shore crab Matuta lunaris

The absolute growth pattern of Matuta lunaris (Forskål) is assessed, based on moult increment data and analyses of size frequency distributions. Both sexes show determinate growth with maturity occurring at the terminal puberty moult. However, Matuta lunaris is unusual as the sexes differ markedly in their growth patterns. Males mature at a larger size than females and grow to a larger maximum size. Male growth is characterized by relatively constant moult increments and an unknown and possibly variable number of immature instars with widely overlapping size ranges. In contrast, female growth is characterized by relatively large moult increments and a fixed number of instars, each with restricted size ranges.     

Interference by the mantispid Mantispa uhleri with the development of the spider Lycosa rabida

Abstract. 1. The larvae of Mantispa uhleri Banks (Neuroptera: Mantispidae) board spiders to await the production of an egg sac containing their obligate developmental food. While aboard the spider, larvae maintain themselves by feeding on spider blood. This parasitic behaviour was investigated by allowing larvae to board sixth instar Lycosa rabida Walckenaer (Araneae: Lycosidae). Larval parasitism has a direct and indirect effect on the developmental physiology of the spider.
2. The direct effect, equal in both spider sexes, is an increase in development time and a decrease in adult size.
3. The indirect effect on development time and adult size is brought about by the loss of an instar in female spiders only. Parasitized females were mature at nine or ten instars; control females at ten or eleven. Male instar number was not affected; both control and parasitized males were mature at nine or ten instars.
4. The net result is that parasitized female spiders are even smaller than would be predicted from the direct effect alone, but actually mature faster than control females. In males there is only the direct effect. The adaptive significance of this sexually dimorphic response is discussed.     

Larvae of lycaenid butterflies that parasitize ant colonies provide exceptions to normal insect growth rules

Fourth instar larvae of Maculinea species of lycaenid butterfly live as social parasites inside Myrmica ant nests. They show highly unusual growth patterns, with small but regular growth in early phytophagous instars, followed by >10 times the growth predicted by extrapolating the early growth rate (following Dyar's rule) during the final carnivorous instar. This produces striking allometry between head and body size in full-grown larvae (ratios of 4–5% compared with 8–10%). Larvae of the Myrmica ant hosts have a similar growth. Data for c. 150 other lycaenid species showed that species with similar life-histories exhibit the same unusual growth pattern (Phengaris spp., Lepidochrysops spp., Niphanda fused); all others have regular growth throughout their larval life, including the carnivorous species that are parasitic on ants from the first instar. It is suggested that Maculinea-type growth pattern has arisen convergently in at least three unrelated lineages of lycaenids. Selection pressures might include the need for reduced early growth to produce late instars that are small enough to be integrated as brood mimics into ant social systems, combined with the need to achieve at least the same adult size as the ancestral species. Trophic pressures that operate on both sedentary ant and butterfly larvae, which must survive long periods of starvation and grow rapidly when food is abundant, may also be involved.     

Venturia canescens parasitizing Galleria mellonella and Anagasta kuehniella: differing suitability of two hosts with highly variable growth potential

Venturia canescens (Grav.) (Hymenoptera: Ichneumonidae) is a solitary larval koinobiont endoparasitoid, ovipositing in several larval instars of different pyralid moth species that are pests of stored food products. After oviposition, the host larva continues to feed and grow for at least several days, the precise time doing so depending on the stage attacked. We examined the relationship between host stage and body mass on parasitoid development in late second to fifth instars of two hosts with highly variable growth potential: the wax moth, Galleria mellonella (L) and the flour moth, Anagasta kuehniella (Zeller)(Lepidoptera: Pyralidae). G. mellonella is the largest known host of V. canescens, with healthy larvae occasionally exceeding 400mg at pupation, whereas those of A. kuehniella rarely exceed 40 mg at the same stage. Parasitoid survival was generally higher in early instars of G. mellonella than in later instars. By contrast, percentage adult emergence in A. kuehniella was highest in late fifth instar and lowest in late second instar. A. kuehniella was the more suitable host species, with over 45% adult emergence in all instars, whereas in G. mellonella we found less than 35% adult emergence in all instars. Adult parasitoid size increased and egg-to-adult development time decreased in a host size- and instar-specific manner from A. kuehniella. The relationship between host size and stage and these fitness correlates was less clear in G. mellonella. Although both host species were parasitized over a similar range of fresh weights, the suitability weight-range of A. kuehniella was considerably wider than G. mellonella for the successful development of V. canescens. However, in hosts of similar weight under 5 mg when parasitized, larger wasps emerged from G. mellonella than from A. kuehniella. Parasitoid growth and development is clearly affected by host species, and we argue that patterns of host utilization and resource acquisition by parasitoids have evolved in accordance with host growth potential and the nutritional requirements of the parasitoid.