Morphology and ultrastructure of spiracles in phlebotomine sandfly larvae

The morphology and ultrastructure of the larval spiracle system of three phlebotomine sandfly species, Phlebotomus perniciosus, P. perfiliewi and P. papatasi, were examined by scanning (SEM) and transmission (TEM) electron microscopy and by confocal scanning laser microscopy (CSLM). During larval development, thoracic and abdominal spiracles show considerable modifications. In fourth instar larvae, the spiracles consist of a plate with a sclerotized central portion and a peripheral circle of papillae. The latter is distinctive in the larvae of P. papatasi, which are readily distinguished from the other species. Opening clefts across the papillae communicate with an internal chamber that encircles an electrondense plug. Many cylindrical projections cross the chamber, uniting the central plug with the larval body, forming an air filter. Spiracular development in successive larval instars has both a taxonomic and adaptive value.     

A new species of Longitarsus Latreille, 1829 (Coleoptera, Chrysomelidae, Galerucinae) pupating inside stem aerenchyma of the hydrophyte host from the Oriental Region

A new species of subaquatic Longitarsus pupating inside the stem aerenchyma of its hydrophyte host plant is described. Eggs are laid on tender leaves and buds and the larvae are open feeders. This is the first report of an Oriental flea beetle pupating inside the stem of its hydrophyte host. A key to the species of southern Indian Longitarsus is provided.     

Larval habits, host-plant associations, and speciation in nematine sawflies (Hymenoptera: Tenthredinidae)

Adaptive radiations consist of two intertwined processes, diversification of species and diversification of their ecological niches, but it is unclear whether there is a causal link between the processes. In phytophagous insects, ecological diversification mainly involves shifts in host-plant associations and in larval feeding habits (internal or external) on different plant parts, and several observations indicate that speciation is facilitated by host shifts. Data on host use in individual species suggest that internal feeders are less likely to colonize new hosts than external-feeding taxa and, consequently, increases in collective host ranges and species numbers should be slowed down in endophagous lineages. We tested these related hypotheses by using phylogenetic information to reconstruct the evolutionary history of larval resource use in the sawfly subfamily Nematinae, a group of 1000 plus species with a broad range of niches: the subfamily's combined host range includes over 20 plant families, and larvae may feed externally on leaves or needles, or internally, for example, in buds, fruits, leaves, or galls. The results show that: (1) Most internally feeding groups have evolved independently from external-feeding ancestors, but several distinct internal habits have appeared convergently multiple times; (2) Shifts among host taxa are clearly more common than changes in larval habits; (3) The majority of host switches have occurred among phylogenetically close plant groups, but many shifts are manifest among distantly related, ecologically proximate hosts; (4) Although external feeding characteristic of the common ancestor of Nematinae is associated with relatively high rates of host-shifting, internal feeders are very conservative in their host use; (5) In contrast, the effect of endophagy on speciation probabilities is more variable: net speciation rates are lowered in most internal-feeding groups, but a striking exception is found in species that induce galls on Salicaceae. The loose connection between collective host ranges and species diversity provides empirical support for theoretical models suggesting that speciation rates are a function of a complex interplay between "intrinsic" niche width and resource heterogeneity.     

Living in fast‐flowing water: Morphology of the gastromyzophorous tadpole of the bufonid Rhinella quechua (R. veraguensis group)

We describe the bufonid gastromyzophorous tadpoles of Rhinella quechua from montane forest streams in Bolivia. Specimens were cleared and stained, and the external morphology, buccopharyngeal structures, and the musculoskeletal system were studied. These tadpoles show a combination of some traits common in Rhinella larvae (e.g., emarginate oral disc with large ventral gap in the marginal papillae, labial tooth row formula 2/3, prenarial ridge, two infralabial papillae, quadratoorbital commissure present, larval otic process absent, mm. mandibulolabialis superior, interhyoideus posterior, and diaphragmatopraecordialis absent, m. subarcualis rectus I composed of three slips), some traits apparently exclusive for the described species of the R. veraguensis group (e.g., second anterior labial tooth row complete, lingual papillae absent, adrostral cartilages present), and some traits that are shared with other gastromyzophorous tadpoles (e.g., enlarged oral disc, short and wide articular process of the palatoquadrate, several muscles inserting on the abdominal sucker). In the context of the substantial taxonomic and nomenclatural changes that the former genus Bufo has undergone, and despite the conspicuous morphological differences related to the presence of an abdominal sucker, the larval morphology of R. quechua supports including it in the genus Rhinella and placing it close to species of the R. veraguensis assemblage. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Host sex preferences and transmission success by the water mite Unionicola foili (Acari: Unionicolidae) parasitic on the midge Chironomus tentans (Diptera: Chironomidae)

This study examined whether ecoparasitic larval Unionicola foili exhibited a sex bias when infecting laboratory populations of the host insect Chironomus tentans and whether an association with male or female midges increased the likelihood of larval mites returning to the aquatic habitat. When laboratory populations of C. tentans were exposed to larval U. foili, there was a higher prevalence of mites among female hosts at emergence (17 of 30 males vs. 25 of 30 females infected by mites). However, there was no significant difference in the distribution or abundance of larvae among infected male (mean = 2.3 larvae per host) and female (mean = 2.6 larvae per host) midges. Larval mites parasitizing both male and female chironomids were more likely to return to water than could be expected by chance. Mite larvae infesting female C. tentans were more likely to return to water when female hosts deposited egg masses in water, suggesting that oviposition plays an important role in cueing larvae parasitizing female midges to detach. The mechanism responsible for increasing the likelihood that mites parasitizing male hosts return to water remains unclear. Future studies will address the possibility of parasite-mediated changes in host behavior.     

The interplay between toxin-releasing β-glucosidase and plant iridoid glycosides impairs larval development in a generalist caterpillar, Grammia incorrupta (Arctiidae)

Herbivores with polyphagous feeding habits must cope with a diet that varies in quality. One of the most important sources of this variation in host plant suitability is plant secondary chemistry. We examined how feeding on plants containing one such group of compounds, the iridoid glycosides, might affect the growth and enzymatic activity in a polyphagous caterpillar that feeds on over 80 plant species in 50 different families. Larvae of the polyphagous arctiid, Grammia incorrupta, were reared exclusively on one of two plant species, one of which contains iridoid glycosides (Plantago lanceolata, Plantaginaceae) while the other does not (Taraxacum officinale, Asteraceae). Larval weight was measured on the two host plants, and midgut homogenates of last instar larvae were then assayed for activity and kinetic properties of β-glucosidases, using both a standard substrate, 4-nitrophenyl-β-D-glucose (NPβGlc), and the iridoid glycoside aucubin, one of the two main iridoid glycosides in P. lanceolata. Larvae feeding on P. lanceolata weighed significantly less and developed more slowly compared to larvae on T. officinale. While the larval midgut β-glucosidase activity determined with NPβGlc was significantly decreased when fed on P. lanceolata, aucubin was substantially hydrolyzed and the larval β-glucosidase activity towards both substrates correlated negatively with larval weight. Our results demonstrate that host plants containing high concentrations of iridoid glycosides have a negative impact on larval development of this generalist insect herbivore. This is most likely due to the hydrolysis of plant glycosides in the larval midgut which results in the release of toxic aglycones. Linking the reduced larval weight to the toxin-releasing action of an iridoid glycoside cleaving β-glucosidase, our results thus support the detoxification limitation hypothesis, suggesting fitness costs for the larvae feeding solely on P. lanceolata. Thus, in addition to the adaptive regulation of midgut β-glucosidase activity, host plant switching as a behavioral adaptation might be a prerequisite for generalist herbivores that allows them to circumvent the negative effects of plant secondary compounds.     

Ecdysteroid titres in a tenebrionid beetle,Zophobas atratus: effects of grouping and isolation

Metamorphosis in Zophobas atratus is dependent on isolation: when kept in grouped conditions, larvae undergo numerous supernumerary moults, growing in size, without pupating. This beetle thus represents an interesting model for the analysis of possible differences in the endocrine regulation of normal vs. supernumerary larval moults. In this study, the ecdysteroid titres have been analysed in this species, using enzyme immunoassay. The hormonal variations of larvae undergoing normal or supernumerary larval cycles were particularly examined, in either grouped or isolated conditions. Normal larval cycles presented very similar ecdysteroid variations in grouped as well as isolated conditions, showing a single hormonal peak (at about 1000pg/&mgr;l). Supplementary larval cycles, occurring in grouped conditions, also showed a similar single ecdysteroid peak, but after a longer period of basal levels. Isolation of such larvae triggered their larval-pupal transformation, which was characterized by more complex hormonal fluctuations, including a small ecdysteroid peak before the main one. Interestingly, the isolation of big larvae during a large part of their cycle induced this peculiar hormonal pattern synchronously, confirming the involvement of a complex neuroendocrine control between external conditions and ecdysteroid titres.     

Similarity and specialization of the larval versus adult diet of European butterflies and moths

Many herbivorous insects feed on plant tissues as larvae but use other resources as adults. Adult nectar feeding is an important component of the diet of many adult herbivores, but few studies have compared adult and larval feeding for broad groups of insects. We compiled a data set of larval host use and adult nectar sources for 995 butterfly and moth species (Lepidoptera) in central Europe. Using a phylogenetic generalized least squares approach, we found that those Lepidoptera that fed on a wide range of plant species as larvae were also nectar feeding on a wide range of plant species as adults. Lepidoptera that lack functional mouthparts as adults used more plant species as larval hosts, on average, than did Lepidoptera with adult mouthparts. We found that 54% of Lepidoptera include their larval host as a nectar source. By creating null models that described the similarity between larval and adult nectar sources, we furthermore showed that Lepidoptera nectar feed on their larval host more than would be expected if they fed at random on available nectar sources. Despite nutritional differences between plant tissue and nectar, we show that there are similarities between adult and larval feeding in Lepidoptera. This suggests that either behavioral or digestive constraints are retained throughout the life cycle of holometabolous herbivores, which affects host breadth and identity.     

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.     

Comparison of lepidopteran larval communities among tree species in a temperate deciduous forest, Japan

Abstract.  1. The enormous diversity of phytophagous insects in forest canopies is hypothesised to be supported by the number of herbivorous species per host tree species or host specificity. It is therefore necessary to examine the effect of host plant species on compositional changes in the herbivore communities.
2. The lepidopteran larval communities were examined in the canopies of 10 tree species in a temperate deciduous forest of Japan. The phylogeny and leaf flush phenology of host plant species were taken into account as factors affecting the herbivore community assembly.
3. Examination of seasonal changes in the larval community structures on each tree species showed that larval species richness, abundance, and evenness decreased significantly from spring to summer. Larval species richness and abundance were characterised by family-level phylogenetic differences among tree species, whereas evenness was determined at a higher taxonomic level.
4. Compositional changes in the larval communities among tree species showed a remarkable pattern, with a phylogenetic effect at a high taxonomic level in spring, similar to evenness, but a phenological effect in summer. This suggests that host specificity could support the lepidopteran larval diversity in spring.
5. These results suggest that the differences in host utilisation of the herbivore, which reflects the phylogenetic effect of the host plants, can be important as a factor affecting the diversity of lepidopteran larval communities in temperate forests.     

Crowding-triggered phenotypic responses alleviate consequences of crowding inEpirrita autumnata (Lep., Geometridae)

Summary CrowdedEpirrita larvae had shorter larval periods than, and similar pupal masses to, their solitary siblings when reared on low quality diets. When fed on high quality diets, pupal masses of crowded larvae were lower than in singletons, and there was no difference in larval period. Because changes in food availability (absolute shortage, induced resistance in foliage) are caused by high larval densities in the field, crowding-triggered phenotypic changes may helpEpirrita to overcome detrimental consequences of high larval density. Pupal period was longer in crowded larvae than in singletons and crowded adults emerged later than their solitary siblings. Eggs of late emerging moths eclosed late in the ensuing spring, which coincides with delayed leaf flush in the year after defoliation. The reason for the faster growth of crowded individuals on poor diets was higher intake albeit less thorough processing of food in crowded, but not in solitary, larvae. On good diets solitary individuals tended to consume more than crowded larvae but there was no difference in processing. Predicted differences of host plant use between stealthy and opportunistic types of herbivores (sensu Rhoades 1985) were generally found between solitary and aggregated larvae on poor but not on good diets. The group response could not be explained by benefits to the group although the assumptions of Wilson's model of group selection were satisfied.     

Flower heating following anthesis and the evolution of gall midge pollination in Schisandraceae

? Premise of the study: Flower heating is known from a few species in 11 of the c. 450 families of flowering plants. Flowers in these families produce heat metabolically and are adapted to beetles or flies as pollinators. Here, we focus on the Schisandraceae, an American/Asian plant family known to exhibit flower heating in some species, but not others, raising the question of the adaptive function of heat production. ? Methods: We used field observations, experiments, and ancestral trait reconstruction on a molecular phylogeny for Schisandraceae that includes the investigated species. ? Key results: At least two Chinese species of Illicium are exclusively pollinated by gall midges that use the flowers as brood sites (not for pollen feeding). Continuous monitoring of flower temperatures revealed that the highest temperatures were attained after the flowers' sexual functions were over, and experiments showed that post-anthetic warming benefited larval development, not fruit development. Midge larvae in flowers with trimmed tepals (and hence a lower temperature) died, but fruit set ratios remained unchanged. Based on the DNA phylogeny, gall midge pollination evolved from general fly/beetle pollination several times in Schisandraceae, with some species adapted to flower-breeding midges, others to pollen-feeding midges. ? Conclusions: Flower heating may be an ancestral trait in Schisandraceae that became co-opted in species pollinated by flower-breeding midges requiring long-persistent warm chambers for larval development.     

Fungal and oomycete parasites of Chironomidae,Ceratopogonidae and Simuliidae (Culicomorpha,Diptera)

Members of the families Chironomidae (chironomids or non-biting midges), Ceratopogonidae (ceratopogonids or biting midges) and Simuliidae (simulids or blackflies) are ubiquitous dipterans of the infraorder Culicomorpha. They are extremely diversified in ecological strategies. Their larvae play major roles in aquatic food webs as detritivores or predators, whereas their adults can be general predators (Chironomidae), hemolymphagous or hematophagous predators (Ceratopogonidae and Simuliidae) or pollinators. Both larval and adult stages are commonly infected by bacteria, viruses, protists, nematodes, true fungi and oomycetes. These phylogenetically diverse assemblages of microorganisms can simultaneously infect multiple species of chironomids, ceratopogonids and simulids, and each host may become trophically interrelated with other hosts by sharing their parasites. Here, we review the information on fungal and oomycete parasites of these dipteran groups with special reference to the natural regulation of host populations, the impact of parasitism in food webs, and the potential of these parasites as biocontrol agents.     

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.     

Diversity of birch sawfly responses to seasonally atypical diets

Most insect herbivores are specialised on a particular plant taxon. To have a better understanding of host shift functions and consequences for insect herbivores, it is essential to gather more information on the effects of variation in host quality on specialists across species and environments. We examined the effects of seasonally atypical food on mortality, developmental time, and final body mass of six sawfly species (Hymenoptera: Symphyta) feeding on the foliage of mountain birch (Betula pubescens ssp. czerepanovii), whose pooled larval feeding periods form a gradient and cover the growing season. Insect phenology was manipulated so that the larvae of early-season species would feed on atypically mature leaves and mid- or late-season species would feed on atypically young leaves of their major host plant. Mortality increased dramatically for all species when the larval feeding schedule was advanced or delayed. This indicates a high degree of specialisation not only on a particular host but also to its phenological phases. The main cause of mortality on novel food was a rejection of the diet by the young larvae and their subsequent starvation. An interesting observation was that late-season species showed this response on nutritious young foliage. The effects of seasonally atypical diets on larval development and growth were species-specific and milder than the effects on mortality. Interestingly, for those individuals that accepted it, atypical food seemed to be most beneficial for species appearing at both ends of the seasonal gradient, which might be related to a wider exposure to variable food quality in natural conditions compared with other species. The diversity of responses to atypical food among closely related herbivore species with overlapping feeding periods on the same host plant is the most crucial finding of this study.     

No-substrate and low-moisture conditions during pupating adversely affect Ectropis grisescens (Lepidoptera: Geometridae) adults

Ectropis grisescens Warren (Lepidoptera: Geometridae) is an important defoliator pest of the tea plant, Camellia sinensis L. Our previous studies showed that E. grisescens pupae had significantly lower body water content when they pupated in the dry soil (20%-moisture) than those that pupated in the wetter soil (50%- or 80%-moisture). However, it is still unclear whether low-moisture conditions during pupating would further influence the physiological and fitness traits of adults. In the present study, E. grisescens individuals were allowed to pupate in 20%-, 50%- or 80%-moisture soil (silt loam), or in containers without soil (no-substrate conditions) in the laboratory. No significant difference in emergence was detected among different pupating conditions. However, female moths that emerged from 20%-moisture soil had significantly lower body water content compared to 80%-moisture soil, whereas the body water content of males was similar among the four treatments. The paired moths that emerged from 20%-moisture soil laid significantly fewer eggs than those that emerged from 50%-moisture soil. In addition, the number of hatched offerings (larvae) were significantly less when the adults previously pupated in no-soil containers compared to 50%-moisture soil. The longevity of adults was shortest when they emerged from 20%-moisture soil. Our study showed that the no-substrate and low-moisture conditions during pupating adversely affected E. grisescens adults.     

Control techniques for Culicoides biting midges and their application in the U.K. and northwestern Palaearctic

The recent emergence of bluetongue virus (Reoviridae: Orbivirus) (BTV) in northern Europe, for the first time in recorded history, has led to an urgent need for methods to control the disease caused by this virus and the midges that spread it. This paper reviews various methods of vector control that have been employed elsewhere and assesses their likely efficacy for controlling vectors of BTV in northern Europe. Methods of controlling Culicoides spp. (Diptera: Ceratopogonidae) have included: (a) application of insecticides and pathogens to habitats where larvae develop; (b) environmental interventions to remove larval breeding sites; (c) controlling adult midges by treating either resting sites, such as animal housing, or host animals with insecticides; (d) housing livestock in screened buildings, and (e) using repellents or host kairomones to lure and kill adult midges. The major vectors of BTV in northern Europe are species from the Culicoides obsoletus (Meigen) and Culicoides pulicaris (L.) groups, for which there are scant data on breeding habits, resting behaviour and host-oriented responses. Consequently, there is little information on which to base a rational strategy for controlling midges or for predicting the likely impact of interventions. However, data extrapolated from the results of vector control operations conducted elsewhere, combined with some assessment of how acceptable or not different methods may be within northern Europe, indicate that the treatment of livestock and animal housing with pyrethroids, the use of midge-proofed stabling for viraemic or high-value animals and the promotion of good farm practice to at least partially eliminate local breeding sites are the best options currently available. Research to assess and improve the efficacy of these methods is required and, in the longer term, efforts should be made to develop better bait systems for monitoring and, possibly, controlling midges. All these studies will need better methods of analysing the ecology and behaviour of midges in the field than are currently in use. The paucity of control options and basic knowledge serve to warn us that we must be better prepared for the possible emergence of other midge-borne diseases, particularly African horse sickness.     

Diverse population trajectories among coexisting species of subarctic forest moths

Records of 232 moth species spanning 26 years (total catch of ca. 230,000 specimens), obtained by continuous light-trapping in Kevo, northernmost subarctic Finland, were used to examine the hypothesis that life-history traits and taxonomic position contribute to both relative abundance and temporal variability of Lepidoptera. Species with detritophagous or moss-feeding larvae, species hibernating in the larval stage, and species pupating during the first half of the growing season were over-represented among 42 species classified as abundant during the entire sampling period. The coefficients of variation in annual catches of species hibernating as eggs averaged 1.7 times higher than those of species hibernating as larvae or pupae. Time-series analysis demonstrated that periodicity in fluctuations of annual catches is generally independent of life-history traits and taxonomic affinities of the species. Moreover, closely related species with similar life-history traits often show different population dynamics, undermining the phylogenetic constraints hypothesis. Species with the shortest (1 year) time lag in the action of negative feedback processes on population growth exhibit the largest magnitude of fluctuations. Our analyses revealed that only a few consistent patterns in the population dynamics of herbivorous moths can be deduced from life-history characteristics of the species. Moreover, the diversity of population behaviour in one moth assemblage challenges any conventional wisdom suggesting predictable patterns. Our results raise several questions about perceptions and paradigms in insect population dynamics and stress the need for research on detritivorous insect population dynamics, as well as the need for more assemblage-wide studies using common trapping methods to provide comparative data on related and unrelated species with different life-history traits.