Effect of light quality and larval diet on morph induction in the polymorphic caterpillar Nemoria arizonaria (Lepidoptera: Geometridae)

The emerald moth Nemoria arizonaria (Geometridae) is bivoltine, with distinct broods of caterpillars hatching in the spring and summer. Caterpillars of the spring brood develop into mimics of oak catkins, while caterpillars of the summer brood develop into mimics of oak twigs. Previous rearing experiments showed that all caterpillars reared on oak catkins developed into catkin morphs, while all caterpillars reared on oak leaves developed into twig morphs, regardless of temperature or photoperiod. However, those previous rearing experiments did not control the colour of light perceived by the caterpillars independently of their dict. Since wavelengths of light perceived by some species of polymorphic caterpillars can influence their colour, it is possible that morph induction in Nemoria arizonaria is due to the characteristics of light reflected from yellow catkins or green leaves, rather than larval diet itself. The experiments reported here independently varied larval diet and light characteristics to determine if light quality is involved in morph induction. Only larval diet influenced morph induction, since all caterpillars reared on catkins developed into the catkin morph, and all caterpillars reared on oak leaves developed into the twig morph, regardless of whether they perceived yellow light, green light, or were raised in the dark.     

A taste receptor neurone dedicated to the perception of pyrrolizidine alkaloids in the medial galeal sensillum of two polyphagous arctiid caterpillars

It is shown that pyrrolizidine alkaloids are phagostimulants for the caterpillars of two polyphagous arctiid caterpillars, Estigmene acrea and Grammia geneura. The caterpillars will also eat dry glass‐fibre discs containing only pyrrolizidine alkaloid ? an example of pharmacophagy. The tip‐recording technique is used to demonstrate that each species has a neurone in the medial galeal styloconic taste sensillum responding to pyrrolizidine alkaloids, although the species differ in their sensitivities. This neurone responds to at least four different pyrrolizidine alkaloids and their N‐oxides, and experiments indicate that it is dedicated to perception of these compounds. The sensory response is phasic–tonic and during the tonic phase remains unchanged for at least 500 ms, resembling the type of response often seen in neurones that are sensitive to plant secondary compounds producing deterrent effects.     

Facilitation of tiger moths by outbreaking tussock moths that share the same host plants

1.?Ecologists have argued about the commonness and strength of interspecific competition between insect herbivores, but facilitation between herbivores has received much less consideration. We previously found that when two species of folivorous caterpillars co-occurred on a shared host plant, feeding by early season tiger moth caterpillars reduced the growth and reproduction of later season tussock caterpillars. However, densities of tussock caterpillars in summer were positively correlated with densities of tiger moth caterpillars the following spring. 2.?In this study, we experimentally manipulated numbers of feeding tussock caterpillars and found that they facilitated tiger moth caterpillars. 3.?The depth of the litter layer beneath host lupine bushes was positively correlated with the number of tussock caterpillars feeding on each bush. Experimental additions of litter beneath lupine canopies during summer resulted in increased numbers of tiger moth caterpillars in the following spring, indicating a causal role of litter. Litter potentially provides food, habitat and protection from desiccation and predation. We failed to find evidence that tussock caterpillars facilitated tiger moth caterpillars by mechanisms independent of litter. 4.?Our study demonstrates that facilitation may operate between insect herbivores, across life-stages through indirect interactions that are non-trophic. Facilitation operated by a novel mechanism, the accumulation of litter which was a by-product of feeding by one species was valuable to a second species. Facilitation persisted in time and space far beyond the creation of litter by tussock caterpillars which should be considered important ecosystem engineers from the point of view of tiger moths. Facilitations that involve habitat modification may generally connect species that do not interact directly or trophically, and have not previously been considered to affect one another.     

Nutrient utilization by caterpillars of the generalist beet armyworm, Spodoptera exigua

Abstract.  Beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), caterpillars are destructive crop pests responsible for considerable annual economic losses. These generalist herbivores are challenged with a diversity of dietary choices that can impact their survival, development and fecundity. In the present study, dietary choices of S. exigua caterpillars, based on the protein to digestible carbohydrate (P : C) ratio of the food, and the impact of nutritionally unbalanced foods on caterpillar performance are assessed. In choice experiments, individual third-instar caterpillars are offered simultaneously a P-biased and a C-biased food until pupation. Caterpillars feed nonrandomly and select a slightly P-biased diet (22P : 20C). In no-choice experiments, second instar caterpillars are reared until pupation on diets ranging in P : C ratio from extremely P- to extremely C-biased. High mortality and delayed development are observed on the C-rich, P-poor diets, highlighting the potential deleterious effects of excess carbohydrates and the importance of protein for growth and development. Diet-dependent differences in pupal weight or pupal lipid reserves are not observed. This contrasts with closely-related Spodoptera species where pupal mass and lipid stores increase on C-rich, P-poor diets. On the extremely P-biased diet, performance is similar to that of individuals reared on the self-selected diet, suggesting that these caterpillars may efficiently be deaminating excess amino acids to generate carbon skeletons, which are shunted into lipid biosynthesis. Spodoptera exigua caterpillars exhibit flexible and efficient pre-ingestive nutrient intake regulation and post-ingestive utilization, allowing these generalist feeders to cope with the heterogeneous diets they may encounter.     

Larval mortality and population regulation in the butterfly Danaus chrysippus in Ghana

An ecological study was made of a population of caterpillars of the African queen butterfly (Danaus chrysippus) feeding on three species of asclepiad plants at Nungua, Ghana, over a nine month period in 1972-73. The principal sources of mortality of caterpillars are probably the parasitic hymenopterans Apanteles chrysippi and Charops sp. There is an inverse correlation between the population of caterpillars and incidence of parasitization. This, and other evidence, suggests that the two parasites are important in limiting the population of Danaus chrysippus below the level imposed by the available food supply. Early instar caterpillars are probably palatable to birds but later instars may be edible or emetic depending on the toxicity of their food plant. It is suggested that the early instar caterpillars are cryptic whilst late instars are cryptic from a distance but conspicuous from nearby, and these may be aposematic or mimetic according to the nature of the food plant. Caterpillars parasitized by Apanteles are paler and hence more cryptic than normal caterpillars. There is also a green, cryptic morph present in the population at low frequency. It is suggested that there is a correlation between colour of caterpillars and the principal source of mortality: i.e., cryptic caterpillars are palatable to birds, suffer heavy predation but a low incidence of parasitization, whilst conspicuous caterpillars are unpalatable to many birds, suffer little predation but have a high incidence of parasitization.     

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.     

Diet mixing enhances the performance of a generalist caterpillar,Platyprepia virginalis

1. Platyprepia virginalis caterpillars are dietary generalists and feed on multiple host species within a single day. We conducted field experiments to evaluate their performance on diets consisting of only their primary food, Lupinus arboreus, or diets consisting of L. arboreus plus other acceptable host species. 2. We found that relative growth rates and rates of survival were higher when they fed on mixed diets compared to lupine only. These results were consistent with hypotheses that mixed diets provided balanced nutrition, diluted toxins, and/or allowed recovery from parasitoids, although our data did not allow us to separate these non‐exclusive explanations. 3. We assayed alkaloids in their host foliage, in the caterpillars themselves, in parasitoids within caterpillars, in food boluses passing through their guts, and in frass that they excreted. We consistently found positive assays for alkaloids in foliage and in frass but negative assays in caterpillars, parasitoids, and food boluses. This suggests that the alkaloids that they ingest are metabolised or rendered non‐reactive by unknown means during passage through the gut. We found no support for the hypothesis that mixed diets prevented caterpillars from exhausting food supplies or allowed them to sequester chemicals from their alkaloid‐containing hosts. 4. Behavioural observations revealed that previous experience influenced a caterpillar's likelihood of moving to a different host. Caterpillars that had previously fed on other hosts were more likely to move to lupine while caterpillars previously collected on lupine were equally likely to choose more lupine or a different host. 5. These results are unusual in providing a clear and consistent benefit of diet mixing in a natural field setting where multiple ecological factors act upon the caterpillars.     

Can plants mediate the humoral immunity and biochemical metabolism of entomopathogen‐infected caterpillars?

Plant‐insect herbivore‐entomopathogen interactions are one of the hot topics in biological control and humoral immunity, and biochemical metabolism are important responses of herbivores to pathogen infection. Entomopathogens are key biocontrol agents of caterpillars, but how plants affect the responses of caterpillars to these organisms is not well understood. We studied hormonal immunity (lysozyme and phenoloxidase activities) and biochemical metabolism (total protein and lipid contents) of Beauveria bassiana‐infected beet armyworm (Spodoptera exigua) larvae that feed on five different host plants (soya bean, Chinese cabbage, edible amaranth, water convolvulus and pepper). Results indicated that plant species differentially affected lysozyme and phenoloxidase activity and lipid content, but had no effect on protein content of pathogen‐infected caterpillars. Both lysozyme and phenoloxidase activities were generally higher in entomopathogen‐infected larvae that feed on edible amaranth or water convolvulus compared with the other three plants from days 1 to 5 after treatment. Plant species did not affect in regular changes during the 5 days in the lipid content of infected or non‐infected caterpillars. Our study reveals that plants fail to affect the biochemical metabolism but plants can mediate the humoral immunity of caterpillars to defend against pathogens. This study provides insight into plant‐mediated effects on the response of herbivores to pathogens.     

Larvae of Maculinea rebeli, a large-blue butterfly and their Myrmica host ants: patterns of caterpillar growth and survival

Maculinea rebeli caterpillars spend their final instar as specific parasites of colonies of the ant Myrmica schencki . This paper describes 14 experiments in which 604 caterpillars were reared with six species of Myrmica . About 40% of caterpillars died during an initial period of integration: survival was significantly greater among the larger caterpillars, in M. schencki nests, and in nests that had ant-brood present. The growth of the survivors was extremely variable. After three weeks they had gained 2–18 times their initial body weight and appeared to belong to two distinct types: fast and slow developers. Only 11% of caterpillars died during the next 37 days of growth, before a fall in temperature to below c. 14 C caused them to overwinter. By then, the caterpillars weighed from 5–90 mg (mean 22 mg).
Caterpillars of all weights lost 33% of their mass during winter, and 8% died, before rising temperatures caused growth to resume in spring. Ofthe 363 that survived winter, 44% died during a period of spring growth and 12% pupated after about seven weeks. The latter came mainly from the largest caterpillars before winter and pupal size (mean weight 81 mg) was also related to prewinter caterpillar size. The remaining 44% of post-winter caterpillars continued to grow slowly and then entered another quiescent phase and were still alive after one year in the nest. A small number of these pupated in the second year, an average of 404 days after adoption.
It is calculated that an average-sized Myrmica schencki colony of 350 workers can produce 4–5 adult butterflies; this laboratory result agrees well with field observations. The results are discussed in the light of a previous behavioural study.     

Spatial and temporal patterns of caterpillar performance and the suitability of two host plant species

Abstract. 1. The butterfly Melitaea cinxia uses two host plant species in the Åland Islands of south-west Finland. Survey data show that host plant use is spatially variable and that the two species are not used (fed on by M . cinxia larvae) in proportion to their abundances. The pattern of host plant use by M . cinxia has been attributed in part to plant distribution and spatial variation of butterfly oviposition preference.
2. The additional roles that may be played by spatial variation in host plant quality and larval physiology (performance ability) were investigated. Seven years of field survey data and a series of laboratory experiments were used to demonstrate that neither of these variables contributed measurably to the observed pattern of host plant use.
3. Specifically, while there was great variation among individuals in both the performance ability of caterpillars and host plant suitability, the two plant species appeared equally suitable, there was no within-species geographic variation in plant suitability, and there was no evidence for adaptation of caterpillars to the locally used host plants.     

Egg cannibalism in Ascia monuste in the field; opportunistic, preferential and very frequent

Newly hatched caterpillars of the specialist herbivorous species Ascia monuste (Lepidoptera, Pieridae) feed on egg chorion before ingesting plant material. They also ingest their exoskeletons and perform egg cannibalism in both the field and the laboratory. Females lay eggs on plants which already have conspecifics (eggs and/or caterpillars) leading to the observation of cannibalism in the field. Two favorable conditions are required for such an occurrence: (1) eggs laid close together on the same leaf, and (2) the finding by older caterpillars of eggs on the same plant. Oviposition preference experiments in cages showed that females do not avoid laying on plants with eggs. However, they do avoid laying eggs on plants predated by caterpillars. All instars perform cannibalism in the field and, in addition to this, the ingested eggs are healthy and cannibalism occurs in the presence of abundant food. The ingestion of eggs from their own clutch was more intense when the clutch was large. Contact with eggs is also important for cannibalism. Caterpillars ingested healthy eggs during three successive days. According to the results, it is possible to state that egg cannibalism in A. monuste is opportunistic, preferential and highly frequent and that there is a tolerance to a diet based on animal protein in this species. Since A. monuste obtains nutrients from both animal and vegetable resources, it is probably better to consider it as opportunistic omnivorous.     

Shelter-building caterpillars in the cerrado: seasonal variation in relative abundance, parasitism, and the influence of extra-floral nectaries

Caterpillar shelters provide protection against desiccation and natural enemies, whereas extra-floral nectaries (EFNs) may be an anti-herbivore adaptation that reduces herbivore abundance by attracting predators and parasites. We used a large, long-term dataset for caterpillars found in the Brazilian cerrado to examine temporal variations in the relative abundance of shelter-building caterpillars and exposed caterpillars, and to determine how much variation depends on the season and the presence of EFNs on host plants. We also compared the patterns of parasitism between sheltered and exposed caterpillars, between seasons, and between different host plants. The cerrado has a marked dry season, and its vegetation is a mixture of mostly deciduous shrubs and trees. Leaf production occurs mainly during the rainy season, and many plant species bear EFNs. Our results show that 60?% of cerrado caterpillars build shelters. These caterpillars were found to be proportionally more abundant during the dry season and less parasitized than exposed ones. The proportion of caterpillars building shelters was highest on plants with new leaves (functional EFNs), and parasitism of caterpillars on these plants was higher. Even though our study includes a taxonomically diverse suite of caterpillars that build many different types of shelter and a diverse set of plants and EFN types, our results suggest that EFNs play an important role in structuring caterpillar assemblages in the cerrado, and that the prolific use of shelters by caterpillars may be a result of their effectiveness in protecting caterpillars from natural enemies and desiccation.     

Does anti‐parasitoid defense explain host‐plant selection by a polyphagous caterpillar?

While studies of tri-trophic interactions have uncovered a variety of mechanisms influencing the dietary specialization of insect herbivores, such studies have neglected host-plant selection by generalists. Here, we report an initial investigation on how host-plant quality and a tachinid parasitoid interact to affect the survival and host-plant selection by a polyphagous herbivore. This herbivore, Grammia geneura (Strecker) (Lepidoptera: Arctiidae), is a food-mixing caterpillar that feeds preferentially on forbs. A previous study suggested that G. geneura might eat certain host species for reasons other than benefits of physiological utilization. We hypothesized that host-plant mediated defenses could act against parasitoids, the major mortality agents of late instar G. geneura . Field observations indicated that caterpillars sometimes survived an attack by the parasitoid Exorista mella Walker (Diptera: Tachinidae) in nature. Laboratory experiments showed that the survival of parasitized caterpillars increased on acceptable but nutritionally inferior host-plant species, indicating that anti-parasitoid defense may explain host-plant selection in this dietary generalist. We found no indication that host-plant selection changed according to the parasitism status of individual caterpillars.     

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.     

Semiquinone and ascorbyl radicals in the gut fluids of caterpillars measured with EPR spectrometry

The biological activity of phenolic compounds ingested by caterpillars is commonly believed to result from their oxidation, although the products of oxidation have been well-characterized in only a few cases. The initial oxidation products of phenols (semiquinone or phenoxyl radicals) can be measured with electron paramagnetic resonance (EPR) spectrometry. In this study semiquinone radicals formed from tannic acid and gallic acid in the gut fluids of two species of caterpillars were measured. In Orgyia leucostigma, in which ingested phenols are not oxidized, semiquinone radicals were absent or at very low intensities. By contrast, in Malacosoma disstria, in which ingested phenols are oxidized, high semiquinone radical intensities were measured. In the absence of detectable levels of semiquinone radicals, ascorbyl radicals were detected in the EPR spectra instead. High molar ratios of ascorbate to phenols in an artificial diet produced ascorbyl radicals in the midgut fluids of both species, while diets containing low molar ratios produced semiquinone radicals. Similar results were obtained in M. disstria fed the leaves of red oak or sugar maple. The results of this study provide further evidence that ascorbate is an essential antioxidant that prevents the oxidation of phenols in the gut fluids of caterpillars, and demonstrate that EPR spectrometry is a valuable method for determining the degree of oxidative activation of phenols ingested by herbivorous insects.     

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.     

Foraging behavior and growth of isolated larvae of a social caterpillar, Malacosoma americanum

In laboratory experiments, isolated eastern tent caterpillars, Malacosoma americanum (Fabricius) (Lepidoptera: Lasiocampidae), grew more slowly than their grouped siblings even though factors previously reported to give grouped caterpillars an advantage were eliminated from our experimental design. Analysis of time-lapse videorecordings of daily foraging bouts showed that, despite their slower growth, isolated individuals fed significantly more often than their grouped siblings. This finding is consistent with previous observations showing that the rate at which tent caterpillars assimilate food is largely independent of foraging frequency and suggests that increased metabolic costs associated with superfluous activity may cause isolated caterpillars to grow slower. More rapid growth of grouped caterpillars also appears attributable to the significantly longer periods of time solitary caterpillars spent inactive during episodes of molting. Our study shows that the distinctive temporal pattern of foraging characteristic of intact colonies of eastern tent caterpillars is a emergent property of the group.     

Camouflage and selective predation in caterpillars of the poplar and eyed hawkmoths (Laothoe populi and Smerinthus ocellata)

Final instar caterpillars of the poplar hawkmoth ( Laothoe populi ) are usually yellow-green when they have been feeding on Salix fragilis but white when on Populus alba. Similarly final instar caterpillars of the eyed hawkmoth (Smerinthus ocellata) vary in colour from yellow-green when fed on Salix cinerea or S. fragilis to grey-green when fed on Salix ripens or S. alba. Most caterpillars of these two moths are thus very cryptic to the human eye. It has already been shown that colour matching is brought about by young caterpillars changing colour depending on the colour of the substrate perceived by their eyes. Evidence is presented here for selective predation, probably by birds. Yellow-green poplar hawk caterpillars disappeared from P. alba more rapidly than did white caterpillars, but there was no significant difference in the survival of the two colour forms on S. fragilis. Yellow-green caterpillars survived better on S. fragilis than on P. alba , but there was no significant difference in the survival of white caterpillars on the two plants. Reasons why selective predation did not occur in all of the experiments are discussed.     

Evolution and losses of spines in slug caterpillars (Lepidoptera: Limacodidae)

Larvae of the cosmopolitan family Limacodidae, commonly known as “slug” caterpillars, are well known because of the widespread occurrence of spines with urticating properties, a morpho‐chemical adaptive trait that has been demonstrated to protect the larvae from natural enemies. However, while most species are armed with rows of spines (“nettle” caterpillars), slug caterpillars are morphologically diverse with some species lacking spines and thus are nonstinging. It has been demonstrated that the evolution of spines in slug caterpillars may have a single origin and that this trait is possibly derived from nonstinging slug caterpillars, but these conclusions were based on limited sampling of mainly New World taxa; thus, the evolution of spines and other traits within the family remains unresolved. Here, we analyze morphological variation in slug caterpillars within an evolutionary framework to determine character evolution of spines with samples from Asia, Australia, North America, and South America. The phylogeny of the Limacodidae was reconstructed based on a multigene dataset comprising five molecular markers (5.6 Kbp: COI, 28S, 18S, EF‐1α, and wingless) representing 45 species from 40 genera and eight outgroups. Based on this phylogeny, we infer that limacodids evolved from a common ancestor in which the larval type possessed spines, and then slug caterpillars without spines evolved independently multiple times in different continents. While larvae with spines are well adapted to avoiding generalist predators, our results imply that larvae without spines may be suited to different ecological niches. Systematic relationships of our dataset indicate six major lineages, several of which have not previously been identified.     

Chemical disguise as particular caste of host ants in the ant inquiline parasite Niphanda fusca (Lepidoptera: Lycaenidae)

The exploitation of parental care is common in avian and insect 'cuckoos' and these species engage in a coevolutionary arms race. Caterpillars of the lycaenid butterfly Niphanda fusca develop as parasites inside the nests of host ants (Camponotus japonicus) where they grow by feeding on the worker trophallaxis. We hypothesized that N. fusca caterpillars chemically mimic host larvae, or some particular castes of the host ant, so that the caterpillars are accepted and cared for by the host workers. Behaviourally, it was observed that the host workers enthusiastically tended glass dummies coated with the cuticular chemicals of larvae or males and those of N. fusca caterpillars living together. Cuticular chemical analyses revealed that N. fusca caterpillars grown in a host ant nest acquired a colony-specific blend of cuticular hydrocarbons (CHCs). Furthermore, the CHC profiles of the N. fusca caterpillars were particularly close to those of the males rather than those of the host larvae and the others. We suggest that N. fusca caterpillars exploit worker care by matching their cuticular profile to that of the host males, since the males are fed by trophallaxis with workers in their natal nests for approximately ten months.