Broadscale Specificity in a Bark Beetle–Fungal Symbiosis: a Spatio-temporal Analysis of the Mycangial Fungi of the Western Pine Beetle

Whether and how mutualisms are maintained through ecological and evolutionary time is a seldom studied aspect of bark beetle–fungal symbioses. All bark beetles are associated with fungi and some species have evolved structures for transporting their symbiotic partners. However, the fungal assemblages and specificity in these symbioses are not well known. To determine the distribution of fungi associated with the mycangia of the western pine beetle (Dendroctonus brevicomis), we collected beetles from across the insect’s geographic range including multiple genetically distinct populations. Two fungi, Entomocorticium sp. B and Ceratocystiopsis brevicomi, were isolated from the mycangia of beetles from all locations. Repeated sampling at two sites in Montana found that Entomocorticium sp. B was the most prevalent fungus throughout the beetle’s flight season, and that females carrying that fungus were on average larger than females carrying C. brevicomi. We present evidence that throughout the flight season, over broad geographic distances, and among genetically distinct populations of beetle, the western pine beetle is associated with the same two species of fungi. In addition, we provide evidence that one fungal species is associated with larger adult beetles and therefore might provide greater benefit during beetle development. The importance and maintenance of this bark beetle–fungus interaction is discussed.     

Effect of the Nematode Contortylenchus brevicomi on Gallery Construction and Fecundity of the Southern Pine Beetle

Field-collected Dendroctonus frontalis were reared in a controlled environment. Male-female beetle pairs retrieved from galleries 1, 2, or 5 wk after introduction into pine bolts were examined for nematode parasites. Data were obtained for each pair on gallery length, egg niche construction, egg viability, and progeny survival. In a separate study, beetle pairs were reared under laboratory conditions for 10 wk. The number of emerged adult progeny of each pair was recorded. Contortylenchus brevicomi, a nematode parasite, was found in 25% of all beetles that established galleries. After 2 and 3 wk, female beetles infected with the nematode had produced fewer eggs and shorter galleries than did uninfected females. Uninfected females mated with nematode-infected males showed similar trends, although the differences in the 2- and 3-wk tests were not significant. Progeny survival or egg viability was not affected by nematode parasitism of either parent beetle. Unikaryon minutum, a microsporidian parasite found in 65% of all colonizing beetles, had no effect on measured variables. The lower fecundity of beetles parasitized by C. brevicomi continued throughout the insect''s reproductive cycle. After 10 wk, nematode-infected beetle pairs produced fewer emerged adult progeny than did uninfected pairs.     

Temperature effects on growth of fungal symbionts of the western pine beetle,Dendroctonus brevicomis

Differences in temperature ranges and optima among poikilothermic partners in symbioses can have profound effects on their interactions and stability. In this study, we investigated how the two mutualist mycangial fungi (Ceratocystiopsis brevicomi and Entomocorticium sp. B) associated with the western pine beetle, Dendroctonus brevicomis, respond to temperature in vitro. Little variability in growth rate at the various temperatures tested occurred among isolates of C. brevicomi either within or among sites. In contrast, E. sp. B exhibited highly variable responses to mid-range temperatures among sites and within some sites, and, unlike C. brevicomi, grew not at all or only very poorly at the highest and lowest temperatures tested. This variability affected both optimal temperature and maximum growth rate. The high variability in response to some temperatures among isolates of E. sp. B in some populations indicates that the ability to capture spatial and nutritional resources can vary greatly within this species which may have considerable impact on the outcome of both inter- and intra-specific competition among the fungi within trees and the short- and long-term dynamics of the fungi with the host beetle.     

Influence of Dietary Experience on the Induction of Preference of Adult Moths and Larvae for a New Olfactory Cue

In Lepidoptera, host plant selection is first conditioned by oviposition site preference of adult females followed by feeding site preference of larvae. Dietary experience to plant volatile cues can induce larval and adult host plant preference. We investigated how the parent’s and self-experience induce host preference in adult females and larvae of three lepidopteran stem borer species with different host plant ranges, namely the polyphagous Sesamia nonagrioides, the oligophagous Busseola fusca and the monophagous Busseola nairobica, and whether this induction can be linked to a neurophysiological phenotypic plasticity. The three species were conditioned to artificial diet enriched with vanillin from the neonate larvae to the adult stage during two generations. Thereafter, two-choice tests on both larvae and adults using a Y-tube olfactometer and electrophysiological (electroantennography [EAG] recordings) experiments on adults were carried out. In the polyphagous species, the induction of preference for a new olfactory cue (vanillin) by females and 3^rd instar larvae was determined by parents’ and self-experiences, without any modification of the sensitivity of the females antennae. No preference induction was found in the oligophagous and monophagous species. Our results suggest that lepidopteran stem borers may acquire preferences for new olfactory cues from the larval to the adult stage as described by Hopkins’ host selection principle (HHSP), neo-Hopkins’ principle, and the concept of ‘chemical legacy.’     

Conditioning as a factor in parasitoid host plant preference

The relative effects of larval and adult conditioning on Diglyphus isaea host plant preference and subsequent parasitism were investigated. Parasitoid larvae were reared on Chromatomyia syngenesiae on lettuce, chrysanthemum or in isolation, and female parasitoids were exposed to hosts on either lettuce or chrysanthemum. Hosts on lettuce and chrysanthemum were then presented to assess parasitoid preference. Larval and adult D. isaea conditioning only slightly affected the females selection of host plant type on which the host larvae were located. The major effect of conditioning was a reduction in the number of eggs laid after larvae and adults were conditioned on chrysanthemum and particularly when both stages were conditioned on this host plant.     

Oviposition,larval preference,and larval performance in two polyphagous species: does the larva know best?

It is expected that females preferentially oviposit on plant hosts that allow for optimal larval performance. However, this expectation contradicts empirical evidence where adults do not always choose the best host for their descendants. Recent evidence suggests that females’ host selection depends on the number of potential hosts. Females from oligophagous species seem to be able to choose an appropriate host in terms of larval performance, whereas in polyphagous species, adult oviposition preference is not related with larval performance. This suggests that larvae in polyphagous species could be taking a more active role in host selection than their mothers. Here, we evaluated the oviposition preference and the larval preference and performance of two polyphagous species of economic importance, Copitarsia decolora (Guenée) (Lepidoptera: Noctuidae: Cuculliinae) and Peridroma saucia (Hübner) (Lepidoptera: Noctuidae: Noctuinae), on eight species of cultivated plants. In laboratory and greenhouse choice assays, we tested adult preference for oviposition and larval preference at 1 and 24 h. Larval performance was measured in terms of survival to adulthood, length of larval period, and pupal weight. We found that both adult females and larvae actively choose their hosts and that the larval preference toward the hosts is related to the females’ preference in both herbivore species. However, the females and larvae did not preferentially select the host with the best larval performance, indicating that larval performance is not related to female or larval preference and that other selective pressures are influencing the choice of the host plant in these two species.     

The life cycle of Pteronarcys scotti (Plecoptera: Pteronarcyidae) from the Southern appalachians,U.S.A.

Larval and adult Pteronarcys scotti Ricker were collected from a mountain stream in northwestern South Carolina, USA. The pattern of larval growth indicated a semivoltine life cycle. Emergence occurred in April and early May, and small larvae first appeared in May. Sexual dimorphism in adult size was apparent; males were 0.8 as large as females in head capsule width.

Female larvae possess a midventral notch on the eighth abdominal segment that can be used to sex larvae as small as 2.5 mm in head width. Female larvae as small as 1 mm in head width have a midventral break in the posterior setal row of segment 8. These characters are illustrated.     

Larval and adult morphology ofCamallanus anabantis Pearse, 1933 andC. kulasirii (Yeh, 1960) (Nematoda: Camallanidae) from freshwater fishes,with notes on the validity of some related forms

Larvae and adults ofCamallanus anabantis andC. kulasirii, recovered from the West Bengali freshwater fishes,Anabas testudineus andOphicephalus punctatus, respectively, are described on the basis of detailed morphological studies under the light microscope. Larval forms collected fromA. testudineus are deemed to be of the third and fourth stages when compared with those from experimental studies of the life cycle ofC. anabantis. Moreover, the present fourth stage female larvae are similar to the females ofC. pearsei, both morphologically and metrically.C. pearsei is, therefore, believed to represent the fourth stage female larvae ofC. anabantis. Similarly, adult males and females recovered fromO. punctatus closely resembleC. kulasirii andC. fernandoi, respectively. The larval forms from this host are fourth stage and can be distinguished as males and females, but both possess a buccal capsule bearing beaded longitudinal ridges similar to that of adult males. The late fourth stage (just prior to the final moult) female larva is, however, found to possess a buccal capsule transitional between that of the adult male and female and also betweenC. kulasirii andC. fernandoi. C. fernandoi is, therefore, presumed to represent the females ofC. kulasirii. However,C. pearsei andC. fernandoi are regarded, for the present, asspecies inquirendae.     

Effects of host plant and maternal feeding experience on population vital rates of a specialized leaf beetle

In herbivorous insects, the interaction between adult preference and progeny performance on specific host plants is modified by maternal feeding experience and host plant quality. Ultimately, changes in the strength of this interaction can affect insect population dynamics. In this study, we hypothesized that adult host plant preference influences progeny performance through a maternal feeding experience × host plant interaction, that is, the effect of adult feeding experience on progeny performance will depend on the host plant. Second, that decoupling of the preference–performance relationship due to host switching results in different population vital rates changing population dynamics. An increase in development time and a decrease in body size of individuals in the alternate host should decrease population growth. We tested these hypotheses using two lines of the tortoise beetle Chelymorpha varians Blanchard fed with two hosts (Convolvulus arvensis and Calystegia sepium). Maternal feeding experience treatments were crossed with host plant species, and the offspring’s developing time and adult size were measured. The host plant influence on the beetle’s population vital rates was tested using stage-structured matrix population models and life table response experiments. Host plant preference affected offspring body size through a host plant effect that contributed to adaptive life history responses only in the better quality host. C. varians’ population growth was positive when fed with either host; comparatively, however, C. sepium had a negative effect on growth by reducing all transition probabilities of the life cycle stages of the beetle. Here, we show that individuals of C. varians prefer and perform differently on distinct hosts and that these patterns influence population vital rates in different ways. When beetles prefer the host plant where their progeny performs best, life history responses and life stage transitions lead to higher population growth; otherwise, growth rate decreases.     

Asynchronous male and female life cycles in the sexually dimorphic, harem-forming isopod Paragnathia formica (Crustacea: Isopoda)

The complex life history of the sexually dimorphic, harem-forming isopod Paragnathia formica Hesse is described, combining published information with new observations. The results of a two-and-a-half year field study, carried out within the animal's saltmarsh habitat, are presented, revealing significant differences in the life cycles of males and females. Settlement to the breeding habitat of final stage male and female larvae, derived from the same annual generation, was recorded at very different times of the year. The total lifespan of males was shown to be twice that of females.
The larvae, temporary ectoparasites of estuarine fish, and the non-feeding, burrow-living adults are described and categorized into several developmental phases, whose numbers were recorded during the course of the study.
The annual production of larvae during a limited period in the autumn, and the subsequent settlement of final stage larval females and males the following year, were monitored. Larval females entered the burrows of adult males in the spring to breed, and died after releasing viviparous broods in the autumn. Larval males settled later in the summer, reaching adulthood in the autumn and overwintering before breeding some 18 months after their birth and dying at the end of the breeding season. Adult males thus bred with females from the next generation. The differences in male and female life cycles led to great seasonal variations in adult sex ratios, a huge winter bias towards males contrasting with a brief excess of females in the summer.     

Effects of crowding and temperature on Wolbachia infection density among life cycle stages of Aedes albopictus

Species of the genus Wolbachia are a group of Rickettsia-like, maternally-inherited bacteria (gram negative), which cause various reproductive alterations in their arthropod and nematode hosts including cytoplasmic incompatibility (CI), male-killing, parthenogenesis and feminization. They can be divided into supergroups such as A and B based on phylogenetic analysis of 16S rDNA sequences. In this study, we examined the relative infection densities of Wolbachia strains among life cycle stages in the mosquito, Aedes albopictus in terms of crowding effect and temperature effect. A. albopictus is known to be superinfected with both A- and B-supergroup Wolbachia which cause CI. The relative Wolbachia densities within each individual mosquito were determined and quantified by using real-time quantitative PCR assay based on the wsp gene. We found that B-supergroup Wolbachia strain densities in this host species were consistently and significantly higher than in the A-supergroup. Larval crowding also reduced adult size of mosquitoes. Our results show clearly that the higher densities of mosquito larvae cause lower densities of Wolbachia strains. Examination of the effect of temperature on Wolbachia density in each stage of the mosquito clearly revealed a significant decrease in bacterial density following exposure to elevated temperature (37 °C) in both males and females.     

Adaptation to a novel host modifies host discrimination by the seed beetle Callosobruchus maculatus

Females of the seed beetle Callosobruchus maculatus avoid adding eggs to seeds that already bear eggs. Geographical variation in this behaviour has been thought to depend on differences in host size. In populations that attack small-seeded legumes, only one or two larvae can develop within a seed, and females are especially adept at detecting and rejecting occupied (egg-laden) seeds. We performed a mass-selection experiment in which replicate lines of a population associated with a small host (mung bean, Vigna radiata) were either maintained on this host or were transferred to a larger host (cowpea, Vigna unguiculata) that can support several larvae per seed. After more than 40 generations, we estimated the strength of host discrimination by presenting females a choice of egg-free and egg-laden seeds, and by quantifying how uniformly females spread their eggs among egg-free seeds. Compared to females maintained on mung bean, females from cowpea lines were more likely to accept occupied seeds in choice tests. They also distributed their eggs less uniformly, especially when cowpea served as the test host. Cowpea lines thus evolved to resemble populations that have long been associated with the larger host. A separate study showed that weaker host discrimination in the cowpea lines was accompanied by a decline in larval competitiveness, which may have further relaxed selection for avoidance of occupied hosts. Our results demonstrate that switching to a novel resource can produce rapid and predictable changes in a fitness-related insect behaviour. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Sound and vibration are not obligatory host finding stimuli for the bark beetle parasite,Coeloides brunneri [Hymenoptera: Braconidae]

Previous researchers, have speculated or concluded thatCoeloides brunneri Viereck females detect their bark beetle hosts by perception of the vibrations or sound made by boring larvae. However, when placed on logs containing various actively mining stages ofDendroctonus pseudotsugae Hopkins,C. brunneri females actively searched for the host only on logs infested with young or maturing brood larvae, and oviposited only in logs with maturing brood larvae. Moreover, when offered larvae in logs that had been frozen at approximately ? 50°C, and then allowed to thaw at room temperature for 2 days, they found the motionless, dead larvae, and oviposited through the bark on to them. Therefore,C. brunneri is able to find ist host by perception of some stimulus (or stimuli) other than sound or vibration.     

Variable correspondence of female host preference and larval performance in a phytophagous ladybird beetle Epilachna pustulosa (Coleoptera: Coccinellidae)

Genetic and environmental factors causing intraspecific variation of the thistle Cirsium kamtschaticum Ledeb. as a host plant of the phytophagous ladybird Epilachna pustulosa Kôno were investigated through simple food-choice tests and rearing of larvae. Two thistle clones (T1U₁ and T4H₂) were used, originally growing approximately 12 km apart. A previous study showed that adult female ladybirds preferred T1U₁ to T4H₂, and that larval performance was better on T1U₁, when leaves from the clones in situ were examined. The two clones retained their characteristics with respect to beetle preference after transplantation into a common garden. However, the difference between T1U₁ and T4H₂ with respect to larval performance was reduced after the transplantation. When leaves from shoots of T1U₁ exposed to different sunlight intensities were offered, adult female ladybirds did not show obvious preferences. Larval eclosion rates increased significantly with the increase in leaf sunlight intensity exposure. These results suggest strongly that both genetic and environmental factors are involved in interclonal variation of thistle quality in beetle preference and/or performance. It is suggested that the quality of thistle leaves for larval performance is largely affected by environmental factors, while leaf quality for beetle preference may be determined strictly by genetic factors. Under certain conditions, E. pustulosa females may behave maladaptively, preferring plants not appropriate for larval growth, or not choosing plants appropriate for the larval growth.     

Gut microbiota of dung beetles correspond to dietary specializations of adults and larvae

Vertebrate dung is central to the dung beetle life cycle, constituting food for adults and a protective and nutritive refuge for their offspring. Adult dung beetles have soft mandibles and feed primarily on nutritionally rich dung particles, while larvae have sclerotized mandibles and consume coarser dung particles with a higher C/N ratio. Here, using the dung beetles Euoniticellus intermedius and E. triangulatus, we show that these morphological adaptations in mandibular structure are also correlated with differences in basic gut structure and gut bacterial communities between dung beetle life stages. Metagenome functional predictions based on 16S rDNA characterization further indicated that larval gut communities are enriched in genes involved in cellulose degradation and nitrogen fixation compared to adult guts. Larval gut communities are more similar to female gut communities than they are to those of males, and bacteria present in maternally provisioned brood balls and maternal ‘gifts’ (secretions deposited in the brood ball along with the egg) are also more similar to larval gut communities than to those of males. Maternal secretions and maternally provisioned brood balls, as well as dung, were important factors shaping the larval gut community. Differences between gut microbiota in the adults and larvae are likely to contribute to differences in nutrient assimilation from ingested dung at different life history stages.     

Leaf beetle grazing does not induce willow trichome defence in the coppicing willow Salix viminalis

Abstract 1 Willows are frequently attacked and defoliated by adult leaf beetles (Phratora vulgatissima L.) early in the season and the plants are then attacked again when new larvae emerge. The native willow Salix cinerea has previously been shown to respond to adult grazing by producing new leaves with an increased trichome density. Subsequent larval feeding was reduced on new leaves. This type of induced plant response may reduce insect damage and could potentially be utilized for plant protection in agricultural systems. 2 Here, we investigated if the willow species most commonly used for biomass production in short rotation coppice, Salix viminalis, also responds to adult beetle grazing by increasing trichome density. Larval performance and feeding behaviour on plants previously exposed to adult beetles was compared with that on undefoliated control plants in a greenhouse. 3 We found an overall decrease in trichome density within all the plants (i.e. trichome density was lower on new leaves compared to that for older basal leaves on S. viminalis). However, leaves of beetle defoliated plants had a higher trichome density compared to control plants. Larval growth and feeding was not affected by this difference between treatments. Larvae appeared to remove trichomes when feeding on S. viminalis, a behaviour that might explain the lack of difference between treatments.     

Win,Lose, or Draw: Effects of Residency,Size, Sex,and Kinship on High‐Stakes Larval Contests in a Moth

Many factors can affect the outcome of a competitive interaction. One such factor is the relatedness between competitors as competitive intensity may decrease between kin. Because adult females lay eggs in clusters, larvae of the moth Utetheisa ornatrix are likely to be found in high densities of their siblings. Larval U. ornatrix actively compete for access into seedpods of their host plant (Crotalaria spp.), and successful competitors will reap numerous reproductive benefits during adulthood. The objective of this study was to determine whether residency, size, sex, and relatedness affect competition over seedpods in U. ornatrix larvae. In one‐on‐one trials for access into artificial seedpods, we monitored occupancy and weight change of larvae varying in residency, size, sex, and relatedness. We found that larger larvae have a competitive advantage over smaller larvae. This finding has consequences for the mating system of U. ornatrix in that females, by selecting males based on pheromone levels that are correlated with body size, can rear larger offspring that will have an advantage in competition over seedpods. These data did not support our hypothesis that males would outcompete females, perhaps because the rewards of acquiring a seedpod are substantial for both sexes. Finally, our data show that resident larvae are more likely to maintain control of seedpods over sibling than non‐siblings intruders, which suggests that relatedness affects competitive interactions.     

Attraction of the predator,Harmonia axyridis (Coleoptera: Coccinellidae), to its prey,Myzus persicae (Hemiptera: Aphididae), feeding on Chinese cabbage

The predatory multicolored Asian lady beetle, Harmonia axyridis, was attracted to volatiles released from Chinese cabbage infested by the green peach aphid, Myzus persicae, in T-tube olfactometer choice tests. However, lady beetle adults and larvae did not respond to clean air, Chinese cabbage alone or green peach aphid alone. Of different prey densities, H. axyridis adults were most attracted to Chinese cabbage infested by 60 M. persicae adults after 24 h. However, H. axyridis larvae were not attracted to Chinese cabbage infested by M. persicae. Mechanically damaged Chinese cabbage attracted neither lady beetle adults nor larvae. Predatory adults were attracted to 60 M. persicae adults after 24 and 48 h, and to 90 M. persicae adults after 12 h, suggesting that the predatory response depends on the prey density. Lady adult beetles did not prefer the volatiles induced by Diamondback moth, Plutella xylostella, indicating that specific host insect specificity attracts respective natural enemies. It can be explained that the volatile compounds emitted from the host plant as a result of herbivore attack preferred by the specific insect species.     

Dietary complementation across life stages in the polyphagous lady beetle Coleomegilla maculata

We investigated the life history consequences of changes in diet between larval and adult life stages in the polyphagous lady beetle Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae). Beetles were reared on three larval diets: greenbug, Schizaphis graminum Rondani (Homoptera: Aphididae), eggs of the flour moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), and bee pollen. The reproductive performance of females was then evaluated on an adult diet of either greenbug or moth eggs. Moth eggs appeared to be the most suitable diet for larvae, yielding the largest adults, and pollen the least suitable, resulting in the smallest adults and greatly extended developmental time. Pollen‐reared beetles tended to have lower fecundity and fertility than those reared on animal protein, regardless of adult diet. Female fitness was generally increased by a change in diet upon emergence to the alternative source of animal protein, suggesting that dietary complementation occurred across life stages. Among females reared on greenbug, a change of diet to moth eggs reduced the period required for production of 12 clutches and increased egg fertility compared to continued feeding on greenbug. Among females reared on moth eggs, a change of diet to greenbug increased fecundity compared to continued feeding on moth eggs. Among females fed an adult diet of greenbug, those fed moth eggs as larvae had faster production of 12 clutches and higher fecundity. We discuss these novel results in the context of coccinellid life history and ecology and their potential implications for other insects that are predatory as both larvae and adults.