Social Cues and Following Behavior in the Forest Tent Caterpillar

Colonies of the social caterpillar Malacosoma disstria Hubner (Lepidoptera: Lasiocampidae) travel in groups following silk trails marked with pher-omone. This study examined first, the cues involved in following behavior and second, the responses to these cues at different larval stadia. Both second and fourth instar larvae discriminated between fresh and older trails, and travelled faster in the presence of trails. In addition to trail following, young caterpillars exhibited leader following, which might be particularly important in exploring unmarked territory. Indeed, second instar caterpillars were more likely to travel together when trails were absent. Fourth instar larvae exhibited greater independent locomotion in the absence of trails than did younger larvae. These findings help explain patterns of social behavior observed in forest tent caterpillar colonies in the field.     

Fitness costs and benefits of shelter building and leaf trenching behaviour in a pyralid caterpillar

1. Shelter building and petiole trenching in the Lepidoptera is a behaviour that mediates ecological pressures including those exerted by both food plants and natural enemies. 2. Fitness costs and benefits of trenching and shelter‐building behaviour related to predation and larval performance were investigated in a pyralid species that inhabits and feeds on leaf shelters. 3. Assays comparing the performance of caterpillars feeding on trenched versus non‐trenched foliage and fresh versus dry leaves were conducted. Whereas pupal weight was positively affected by petiole trenching, larval development was delayed when caterpillars fed on dry leaves. 4. A field experiment comparing predation on caterpillars inside and outside shelters demonstrated that predation was significantly higher for exposed caterpillars. 5. No physiological costs associated with shelter building were found given that caterpillars performed equally regardless of the number of shelters they built. 6. The effect sizes of top‐down and bottom‐up forces on pupal weight, development time, and predation risk indicated that the major effect of shelters is through the reduction of predation risk. The integration of experiments and natural history observations showed that fitness benefits provided by shelters change across ontogeny.     

Stage-specific effects of temperature and dietary protein on growth and survival of Manduca sexta caterpillars

The effects of temperature and dietary protein concentration on growth and survival of Manduca sexta L. (Lepidoptera: Sphingidae) caterpillars during different larval stages were examined. Sets of caterpillars were raised from hatching at one of five constant temperatures (18, 22, 26, 30 or 34°C) and on one of two artificial diets (low or high protein concentration). Mass gain, duration (development time) and mean growth rate were measured for each caterpillar for the 1st to 3rd stadia, the 4th stadium, and the 5th stadium. Temperature significantly affected mass gain during each larval stage, resulting in smaller mass gains at higher temperatures at each stage. This effect was strongest at high temperatures during the 5th stadium. Temperature significantly affected durations of each larval stage, but the effect varied among stages: for example, the duration of stadia 1–3 decreased continuously with increasing temperature, whereas the duration of the 5th stadium was shortest at 26–30°C and increased at lower and higher temperatures. The effect of temperature on mean growth rate changed dramatically across larval stages: maximal growth rate occurred at 34°C during the 1st to 3rd stadia, at 30°C during the 4th stadium and at 26°C during the 5th stadium. Higher dietary protein concentration significantly decreased the duration of stadia 1–3 and of the 4th stadium, but had no significant effect on the duration of the 5th stadium. Temperature and dietary protein had little effect on mortality rates during any larval stadium, with one exception: mortality during the 5th stadium increased dramatically at temperatures of 30 and 34°C. These results demonstrate that the effects of temperature and dietary protein concentration on growth, development and survival in M. sexta vary markedly in different larval stadia during development; 5th instar caterpillars are particularly sensitive to higher temperatures.     

Developmental changes in the patterns of feeding in fourth- and fifth-instar Helicoverpa armigera caterpillars

Data are presented for developmental changes in feeding behaviour within and across the fourth and fifth stadium of Helicoverpa armigera (Lepidoptera, Noctuidae) caterpillars fed nutritionally homogeneous semi‐synthetic foods. We recorded the microstructure of feeding over continuous 12‐h periods on consecutive days throughout the two stadia, and in one experiment recorded continuously for 21 h. Larvae in the two stadia showed the same general pattern of macro‐events in feeding, including a similar duration of post‐ecdysis fast, which was usually broken by consumption of the exuviae, and then a sustained period in which discrete meals on the experimental food were taken regularly. There were, however, some distinct differences in the patterns of meal‐taking both between stadia and across different one‐third time segments within stadia. Considering between‐stadium differences, the proportion of time spent feeding differed significantly only in the last segment of the feeding period of the two stadia, with the value for the fourth‐instar larvae being substantially greater than for fifth‐instar larvae. As regards within stadium changes, the proportion of time feeding increased from the first to the second segment of both stadia. However, whereas the proportion of time feeding increased from the second to the final segment of the fourth stadium, it decreased across the same period in the fifth stadium. These patterns of changes in the proportion of time feeding within and between stadia, and their behavioural mechanisms (combination of meal durations and meal frequencies), can be explained only partially with reference to increasing food requirements with development. Three areas are identified where further study might help elucidate the reasons for the observed developmental changes in the microstructure of feeding: allometric constraint, the dynamic links between ingestion and post‐ingestive processing, and ecological factors such as predation.     

Daily foraging schedule of field colonies of the eastern tent caterpillar Malacosoma americanum

Summary The daily foraging patterns of seven colonies of the eastern tent caterpillar, Malacosoma americanum, were monitored photoelectronically during the last three larval stadia to provide the first detailed record of the foraging behavior of a gregarious caterpillar under field conditions. Colonies were active an average of 49.3% of each day. Three bouts of foraging, centered about 0600 h, 1500 h and 2000 h (EST), occurred daily during the fourth and fifth stadia. Although ambient temperatures were less favorable for foraging and food processing than at other times of the day, the caterpillars were most active at dusk and dawn, and spent comparatively little time away from the tent during the daylight hours. In the last (sixth) stadium, the caterpillars foraged only under the cover of darkness. A lack of relationship between the rate at which the caterpillars processed food and the spacing of their feeding bouts, indicates that this species follows a schedule of feeding and growth shaped by factors other than those directly related to feeding efficiency and ambient temperature. Colony foraging patterns may reduce caterpillar mortality by minimizing contact between larvae and day-active predators and parasitiods.     

The Role of Tactile and Chemical Stimuli in the Formation and Maintenance of the Processions of the Social Caterpillar Hylesia lineata (Lepidoptera: Saturniidae)

Colonies of the social caterpillar Hylesia lineata (Lepidoptera: Satumiidae) form long, single-file, head-to-tail processions as they move between their shelters and distant feeding sites. Although investigations of other processionary species have implicated a silk trail in the processionary process, silk plays little or no role in initiating or maintaining processions in H. lineata. Studies we report here implicate both tactile stimuli and a trail pheromone in the establishment and maintenance of processions. Processionaries elicit locomotion in the individual preceding them in line by brushing their heads against prominent sulci that project from the tips of their abdomens. Caterpillars mark their pathways with a pheromone deposited by brushing the ventral surfaces of their last abdominal segments against the substrate. The persistent pheromone is soluble in hexanes and appears to be secreted from glandular setae found on the proximal regions of the anal prolegs and the venter. In Y-choice tests, caterpillars selected newer trails over older trails and stronger trails over weaker trails. They did not distinguish between trials deposited by newly fed caterpillars and those deposited by starved caterpillars. Despite the unidirectional nature of processions, there is no indication that caterpillars can determine from the trail alone the direction in which the procession advanced. The significance of these findings to the foraging ecology of the caterpillars is discussed.     

Group size effects on survivorship and adult development in the gregarious larvae of Euselasia chrysippe (Lepidoptera, Riodinidae)

Caterpillars living in aggregations may derive several benefits that outweigh the costs, including better survivorship and improved growth rates. I tested whether larval group size had an effect on these two vital rates in Euselasia chrysippe. These caterpillars feed gregariously during all instars and move in processionary form over the host plant and even pupate together. There was a positive relationship between group size and larval survivorship in the field, although genetic variability was not taken into account in this experiment. Under laboratory conditions, there was also a positive relationship between group size, and larval growth rate and adult weight. This supports the hypothesis that aggregations facilitate feeding and larval growth. Single sixth instar larvae in the laboratory also had a lower survivorship than larvae in groups. These results provide further evidence of the benefits of group living for gregarious caterpillars.     

Development,growth and metabolic rate of Hermetia illucens larvae

The larvae of Hermetia illucens are known to successfully bio‐convert a vast range of organic substrates into high protein and fat biomass, but little is known about the larval instars. During this research, larval head capsules and biomass growth were measured daily and the specific metabolic rate of larger instars were considered. The head capsule measurements revealed that H. illucens pass through 6 actively feeding larval stadia before entering the last nonfeeding but migrating 7th stadium. Larval growth follows a sigmoid curve with slowly accelerating growth in the earlier stadia and decelerating growth in the latest stadia. In contrast, development was fast until reaching stadium 6 and then slowed down. Accordingly, the specific metabolic rate was high in instars 3, 4 and 5 and reduced in instars 6 and 7.     

Silk drives aggregation and following in the neotropical caterpillar Mechanitis menapis (Nymphalidae: Ithomiini)

Gregarious larvae that use chemical communication to feed and move together are widespread among folivorous insects, although social behaviour has been studied almost exclusively in a few temperate zone genera. The Menapis (or variable) tigerwing butterfly Mechanitis menapis mantineus Hewitson (Lepidoptera, Nymphalidae, Danainae, Ithomiini) is a neotropical species whose larvae feed gregariously on Solanaceae host plants. In laboratory experiments conducted in the Ecuador cloud forest, M. menapis caterpillars are attracted to silk produced by conspecifics and show no evidence of pheromone production. Indeed, caterpillars consistently choose arenas with silk over bare arenas but do not show a preference for arenas marked with abdominal cuticular surface residues. Mechanitis menapis caterpillars on silk‐coated plants are both more mobile and more cohesive than those on control plants. Nonetheless, caterpillars move independently over unmarked surfaces and groups do not make rapid collective choices between two food sources. Collective behaviour in M. menapis thus appears to be based on aggregation on collectively produced silk to facilitate feeding, as well as using this silk to maintain cohesion. Silk production is common in caterpillars, although M. menapis appears to be unique among species studied so far in using silk to maintain group cohesion.     

Host plant nitrogen enrichment has both positive and negative effects on the larval growth of a specialist butterfly

1. The nitrogen limitation hypothesis posits that phytophagous insects benefit from nitrogen enrichment of their host plants through a reduction of the concentration of toxic compounds and an increase of free amino acids and proteins. However, species' response to nitrogen enrichment varies substantially and high nitrogen levels are associated with population decline, suggesting there are major costs to feeding on nitrogen‐rich host plants. 2. To test the hypothesis that larval growth performance is maximal at intermediate nitrogen enrichment, nitrogen levels were measured in 18 populations of the host plant of Lycaena helle, a specialist butterfly inhabiting nutrient‐poor wet meadows. The nitrogen content of host plants was then modified to mirror average natural nitrogen levels (C), highest field‐recorded levels (T1), and levels higher than those observed across our study populations (T2). 3. Caterpillars fed with T1 leaves had a greater maximum body mass than caterpillars of the C group because of their improved food assimilation during the early stages of their development. Caterpillars of C and T2 groups had similar growth patterns but high nitrogen content had detrimental effects, as caterpillars fed with T2 leaves had a slower ingestion rate than C and T1 groups. 4. Quantifying the fitness consequences of these changes in growth performance is necessary to fully understand the implications of nitrogen enrichment for L. helle (rapid growth may result in fitness costs). However, conservation plans for this emblematic glacial relict species should also consider the preservation of its host plant quality to ensure its persistence.     

Host stage selection of the mealybug parasitoid Anagyrus spec. nov near sinope

The mealybug parasitoid Anagyrus spec. nov near sinope (Hymenoptera: Encyrtidae) is an undescribed parasitoid of the Madeira mealybug, Phenacoccus madeirensis Green (Homoptera: Pseudococcidae). We investigated the preference of Anagyrus spec. nov near sinope for six developmental stadia (first‐ and second‐instar nymphs, third‐instar immature females, third‐ or fourth‐instar immature males, pre‐reproductive adult females, and ovipositing adult females) of P. madeirensis and the fitness consequences of the host stage selection behavior. In the no‐choice test, Anagyrus spec. nov near sinope parasitized and completed development in all host stadia except third‐instar immature males. When all host stadia were offered simultaneously, the parasitoids preferred third‐instar immature and pre‐reproductive adult females. Dissection of the stung mealybugs revealed that the clutch size (number of eggs per host) was approximately four and three in the third‐instar and pre‐reproductive females, respectively, and one egg per first‐instar nymph. Parasitoids emerged from P. madeirensis parasitized at third‐instar or pre‐reproductive adult female completed development in the shortest duration, achieved a higher progeny survival rate, larger brood and body size, and the lowest proportion of males. We showed that the continued development of mealybugs had significant influence on the fitness of the parasitoids. Although deposited as eggs in first‐ or second‐instar nymphs, parasitoids emerged from mummies that had attained third‐instar or adult development achieved similar progeny survival rate, brood size, body size, and sex ratio as those parasitoids deposited and developed in third‐instar or adult mealybugs. By delaying larval development in young mealybugs, Anagyrus spec. nov near sinope achieved higher fitness by allowing the parasitized mealybugs to grow and accumulate body size and resources. We suggest that the fitness consequence of host stage selection of a koinobiont parasitoid should be evaluated on both the time of parasitism and the time of mummification.     

Local Competition Between Foraging Relatives: Growth and Survival of Bruchid Beetle Larvae

Kin selection theory states that when resources are limited and all else is equal, individuals will direct competition away from kin. However, when competition between relatives is completely local, as is the case in granivorous insects whose larval stages spend their lives within a single seed, this can reduce or even negate the kin-selected benefits. Instead, an increase in competition may have the same detrimental effects on individuals that forage with kin as those that forage with non-kin. In a factorial experiment we assessed the effects of relatedness and competition over food on the survival and on fitness-related traits of the bruchid beetle Callosobruchus maculatus. Relatedness of competitors did not affect the survival of larvae. Larval survival substantially decreased with increasing larval density, and we found evidence that beetles maturing at a larger size were more adversely affected by competition, resulting in lower survival rates. Furthermore, females showed a reduction in their growth rate with increasing larval density, emerging smaller after the same development time. Males increased their growth rate, emerging earlier but at a similar size when food was more limited. Our results add to the growing number of studies that fail to show a relationship between relatedness and a reduction in competition between relatives in closed systems, and emphasize the importance of the scale at which competition between relatives occurs.     

Melanism in a larval Lepidoptera: repeatability and heritability of a dynamic trait

Abstract.  1. Although it is well established that the deposition of melanin pigment in the cuticle of larval Lepidoptera is influenced by both environmental and genetic factors, few studies have examined intra-individual regional variation in the degree of melanism or the ontogenetic dynamics of this trait. Here, heritable and density-dependent effects on within-individual and stage-specific variation in melanism were examined in caterpillars of the Egyptian cotton leafworm, Spodoptera littoralis (Boisduval).
2. Using quantitative spectrometric methods, it is shown that cuticular melanism changes dramatically within larval stadia, showing the highest and lowest levels of melanism early (first day) and late (final day) in each larval stadium respectively. However, solitary-reared caterpillars were significantly paler than those reared gregariously at all stages of development and maintained greater levels of variation in melanism. This variation in melanism was repeatable and exhibited a significant heritable component (narrow sense heritability based on offspring–parent regressions: h ² = 0.18–0.30).
3. The degree of melanism was correlated negatively with larval body weight in solitary caterpillars, but not gregarious ones. Melanism also varied spatially, with the lateral longitudinal band being consistently darker than the dorsal or dorso-lateral bands. Crowd-rearing increased melanism in all regions of larval cuticle, but the extent of crowding-induced melanism was more pronounced in the dorsal and dorso-lateral bands than in the lateral one.
4. These results indicate that although cuticular melanism is a highly dynamic trait, ontogenetic changes in relative cuticular melanism are both predictable and repeatable within individuals and genotypes. This has implications for our understanding of the evolution of melanism and for applying artificial selection on the basis of colour.     

Adaptive trade-offs of leaf folding in Dichomeris caterpillars on goldenrods

Abstract.

• 1 Leaf folding is a common behaviour among caterpillars that has many potential benefits such as creation of a favourable microclimate and protection from predation or dislodgment.
• 2 One cost of leaf folding which has not previously been quantified is the reduction in growth rate that might result from the energetic expenditures of producing silk and applying it to fold leaves.
• 3 On caged goldenrods in field and forest habitats, early-instar Dichomeris leuconotella (Busck) caterpillars that were repeatedly forced to spin new leaf refuges (either folds or webs of silk) actually had higher growth rates than caterpillars left undisturbed.
• 4 This surprising result apparently reflected the fact that disturbed caterpillars chose relatively young green leaves for new refuges, whereas undisturbed caterpillars tended to remain in their webs or folds on ageing leaves.
• 5 In a glasshouse experiment using plants that underwent little senescence, growth rates were unaffected by the amout of refuge-making effort, indicating that the costs of such effort were either negligible or easily compensated for by these caterpillars.
• 6 On glasshouse plants, caterpillars periodically forced to change refuges had higher disappearance rates than those left undisturbed, despite the absence of predators. Wandering or falling from plants is evidently a major hazard of frequent refuge changes.
• 7 Thus, early-instar Dichomeris caterpillars face a trade-off in which risks of predation or loss of contact with the host favour the observed low rate of refuge changes, but in which growth rates could actually be improved with more refuge changes.

     

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.

     

Grouping fails to affect the growth and energy budget of a cyprinid, Phoxinus phoxinus (L.)

The effects of grouping on the growth and some components of the energy budget of the minnow, Phoxinus phoxinus (L.), were tested in the laboratory. No effects of grouping were detected on absorption efficiency, the proportion of food energy lost in nitrogenous excretion, maximum rate of food consumption and conversion efficiency. Grouping had no significant effect on either specific growth rate or the dry matter content of individual fish.     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Effects of temperature on growth and efficiency of food utilization in fifth-instar caterpillars of the tobacco hornworm, Manduca sexta

Fifth-instar larvae of Manduca sexta were reared from hatching on artificial diet at 15, 20, 25, 30 and 35°C. Total development time decreased with increasing temperature. Very few larvae (12%) survived at 15°C, so this temperature was not considered further. There was some mortality at 30°C (11%), and at 35°C (50%).The absolute rate of growth in the fifth instar was faster at 25 than at 20°C, but was similar at 25, 30 and 35°C. This was true both for caterpillars that were chronically exposed to experimental temperatures (i.e. since hatching) and for those acutely exposed (i.e. reared up to fifth instar at 25°C).There was a progressive decrease with higher rearing temperatures in both the initial and final sizes of chronically exposed fifth-instar larvae. Acutely exposed caterpillars matched for initial size showed smaller temperature related differences in final size. Because of these size differences there were differences in relative growth rate which did not reflect true differences in absolute growth rate.Total food consumed by chronically exposed caterpillars was greatest at the lowest temperature (20°C), and decreased progressively with increasing temperature. The absolute rate of food consumption increased from 20 to 25°C, but did not vary significantly between 25 and 35°C. Differences in the sizes of the insects at the different temperatures meant that there were differences among relative measures of consumption that did not reflect absolute food consumption.For chronically exposed caterpillars, none of the three usual indices of food conversion efficiency (AD, ECI and ECD) varied significantly with temperature between 20 and 35°C. This implies that the effects of temperature on metabolic costs are closely matched to food consumption.Oxygen consumption increased with temperature between 20 and 25°C but was temperature compensated between 25 and 35°C.These findings are discussed in terms of their implications for the optimal temperature for growth in Manduca.     

Consequences of constitutive and induced variation in plant nutritional quality for immune defence of a herbivore against parasitism

The mechanisms through which trophic interactions between species are indirectly mediated by distant members in a food web have received increasing attention in the field of ecology of multitrophic interactions. Scarcely studied aspects include the effects of varying plant chemistry on herbivore immune defences against parasitoids. We investigated the effects of constitutive and herbivore-induced variation in the nutritional quality of wild and cultivated populations of cabbage (Brassica oleracea) on the ability of small cabbage white Pieris rapae (Lepidoptera, Pieridae) larvae to encapsulate eggs of the parasitoid Cotesia glomerata (Hymenoptera, Braconidae). Average encapsulation rates in caterpillars parasitised as first instars were low and did not differ among plant populations, with caterpillar weight positively correlating with the rates of encapsulation. When caterpillars were parasitised as second instar larvae, encapsulation of eggs increased. Caterpillars were larger on the cultivated Brussels sprouts plants and exhibited higher levels of encapsulation compared with caterpillars on plants of either of the wild cabbage populations. Observed differences in encapsulation rates between plant populations could not be explained exclusively by differences in host growth on the different Brassica populations. Previous herbivore damage resulted in a reduction in the larval weight of subsequent herbivores with a concomitant reduction in encapsulation responses on both Brussels sprouts and wild cabbage plants. To our knowledge this is the first study demonstrating that constitutive and herbivore-induced changes in plant chemistry act in concert, affecting the immune response of herbivores to parasitism. We argue that plant-mediated immune responses of herbivores may be important in the evaluation of fitness costs and benefits of herbivore diet on the third trophic level.     

Short‐term larval food stress and associated compensatory growth reduce adult immune function in a damselfly

Abstract 1. In animals with a complex life cycle, larval stressors may carry over to the adult stage. Carry‐over effects not mediated through age and size at metamorphosis have rarely been studied. The present study focuses on the poorly documented immune costs of short‐term food stress both in the larval stage and after metamorphosis in the adult stage. 2. The present study quantified immune function [number of haemocytes, activity of prophenoloxidase (proPO) and phenoloxidase (PO)] in an experiment where larvae of the damselfly Lestes viridis were exposed to a transient starvation period. 3. Directly after starvation, immune variables were reduced in starved larvae. Levels of proPO and PO remained low after starvation, even after metamorphosis. In contrast, haemocyte numbers were fully compensated by the end of the larval stage, yet were lower in previously starved animals after metamorphosis. This can be explained as a cost of the observed compensatory growth after starvation. Focusing only on potential costs of larval stressors within the larval stage may therefore be misleading. 4. The here‐identified immunological cost in the adult stage of larval short‐term food stress and associated compensatory growth strongly indicates that physiological costs may explain hidden carry‐over effects bridging metamorphosis. This adds to the increasing awareness that the larval and adult stages in animals with a complex life cycle should be jointly studied, as trade‐offs may span metamorphosis.