Wing Plasticity Is Associated with Growth and Energy Metabolism in Two Color Morphs of the Pea Aphid.

The pea aphid, Acyrthosiphon pisum, is a major pest of legume crops, exhibiting distinct polymorphism in terms of wings and body color. We found that, under crowded conditions, the red morph A. pisum produced more winged offspring than the green morph. The signaling pathways involved in aphid wing determination, like insulin and ecdysone, also play important roles in regulating growth, development, and metabolism. Thus, here, we examined the association between the wing-producing ability and the growth rate, development time, reproductive capacity, and energy metabolism in these two color morphs. The growth rate of red morphs was significantly higher than that of green morphs, whereas green morphs produced more offspring during the first 6 days of the adult stage. Red morphs accumulated higher levels of glycogen and triglycerides and consumed more triglycerides during starvation; however, green aphids consumed more trehalose during food deprivation. Red aphids exhibited stronger starvation tolerance, possibly due to their higher triglyceride catabolic activity. Furthermore, the expression levels of genes involved in the insulin pathway, glycolysis, and lipolysis in red aphids were higher than those in green aphids. These results suggest that the wing-producing ability of the pea aphid may be associated with its growth and metabolism, which may be due to the shared regulatory signaling pathways.

New morphological and biological contributions to adults and immature forms of Pissonotusparaguayensis (Fulgoromorpha, Delphacidae) in wetlands of Argentina.

In the search for insects as biological control agents for the water primrose, the delphacid Pissonotusparaguayensis (Delphacidae) was found on Ludwigiagrandiflorasubsp.hexapetala (Onagraceae) in a wetland of Central East Argentina. The morphology of the unknown females (brachypterous and macropterous) and immature stages are described and illustrated. Adults and nymphs were collected in wetlands of Del Plata River Basin, from Buenos Aires to the northeastern part of Argentina. A rearing methodology was developed to perform biological studies. Both winged forms and structural features of the female genitalia are described for the first time at the genus level. Eggs and immature stages are described and keyed; fifth nymphal instars may be easily recognised by the yellowish colouration, blackish on dorsal of head, thorax and abdomen with conspicuous yellowish pits, ventrally only darkened on base of frons extended to lower level of eyes and dorsal surface of antennomeres I and II, and legs with distinctive black marks at femoro-tibial joint and apex. The geographical distribution is updated, expanding its range into Argentina, making Buenos Aires the southernmost limit of the genus in America. Biological information of the species is also reported here: life cycle, fecundity, oviposition behaviour, and host plant. Field observations showed that P.paraguayensis breeds, feeds, and causes damage to L.g.subsp.hexapetala. This delphacid presents a certain degree of specificity to the Ludwigia species in the Jussiaea section in host specificity tests. More studies are required to test this species as a potential biological control agent.

Aphid male wing polymorphisms are transient and have evolved repeatedly.

Polymorphic phenotypes have long been used to examine the maintenance of genetic variation within and between species. Most studies have focused on persistent polymorphisms, which are retained across species boundaries, and their positive effects on speciation rates. Far less is known about the macroevolutionary impacts of more transient polymorphisms, which are also common. Here we investigated male wing polymorphisms in aphids. We estimated the phylogenetic history of wing states across species, along with several other traits that could affect wing evolution. We found that male wing polymorphisms are transient: they are found in only ~4% of extant species, but have likely evolved repeatedly across the phylogeny. We reason that the repeated evolution of transient polymorphisms might be facilitated by the existence of the asexual female wing plasticity, which is common across aphids, and would maintain the wing development program even in species with wingless males. We also discovered that male wingedness correlates positively with host plant alternation and host plant breadth, and that winged morphs and wing polymorphisms may be associated with higher speciation rates. Our results provide new evolutionary insights into this well-studied group and suggest that even transient polymorphisms may impact species diversification rates.

Dispersal and Migration Patterns of Freshwater Semiaquatic Bugs.

Semiaquatic bugs (Hemiptera: Heteroptera: Gerromorpha) are mostly wing-polymorphic species with flight dispersal as an important life history trait, but the specific flight ability and dispersal pattern remain unexplored in most species. This report presents the results of a long-term survey based on the individual marking of more than 23,000 specimens of eight water striders (Gerridae) and a water cricket Velia caprai (Veliidae). Three distinct lentic habitats were sampled (solitary fishponds, systems of nearby fishponds and systems of small, often temporary pools) and one lotic habitat-a small forest stream. Recaptures revealed that three gerrid species tend to stay at the breeding site, but can differ in dispersal via the water surface. Reproductive flightless females disperse most actively via the water surface, possibly bypassing the trade-off between dispersal and reproduction. One species has a sex-dependent dispersal pattern, with females being rather philopatric, whereas males often disperse. Three other gerrid species are highly dispersive and tend to change breeding site. V. caprai, the only lotic species included in this survey, tend to move upstream and possibly compensate for the downstream drift.

An InR/mir-9a/NlUbx regulatory cascade regulates wing diphenism in brown planthoppers.

Wing polymorphism significantly contributes to the ecological success of some insect species. For example, the brown planthopper (BPH) Nilaparvata lugens, which is one of the most destructive rice pests in Asia, can develop into either highly mobile long-winged or highly fecund short-winged adult morphs. A recent study reported a highly provocative result that the Hox gene Ultrabithorax (Ubx) is expressed in BPH forewings and showed that this wing development gene is differentially expressed in nymphs that develop into long-winged versus short-winged morphs. Here, we found that Ubx may be a mir-9a target, and used dual luciferase reporter assays and injected micro RNA (miRNA) mimics and inhibitors to confirm the interactions between mir-9a and NlUbx. We measured the mir-9a and NlUbx expression profiles in nymphs and found that the expression of these two biomolecules was negatively correlated. By rearing BPH nymphs on host rice plants with different nutritional status, we were able to characterize a regulatory cascade between insulin receptor genes, mir-9a, and NlUbx that regulate wing length in BPHs. When host quality was low, NlInR1 expression in the nymph terga increased and NlInR2 expression decreased; this led to a higher mir-9a level, which in turn reduced the NlUbx transcript level and ultimately resulted in longer wing lengths. Beyond extending our understanding of the interplay between host plant status and genetic events that modulate polymorphism, we demonstrated both the upstream signal and miRNA-based regulatory mechanism that control Ubx expression in BPH forewings.

Discovery of cockroach specimens of the genus Squamoptera (Ectobiidae: Pseudophyllodromiinae) from Okinawa, Japan, showing wing polymorphism

@# Cockroach specimens of the genus, Squamoptera were collected from the Iriomote island of Okinawa prefecture, Japan. The morphological features of the specimens were characterized as having a white band on the dorsal surface of its thorax, its tegmen reduced into a tiny scale-like structure and the hindwing was absent. Ocelli was also absent and the small compound eyes not extending to apex of the head nor to the frontal face but extend further lower than the base of the antennae. When the specimens were reared in the laboratory, besides the short wing form, the long wing form began to appear in the rearing colony. In our reproductive biological study, we observed that hatching of the ootheca from the short wing female takes about 30 days, with an average of 6.6 nymphs being hatched from one ootheca. The male to female ratio of the offspring was 36:30. However, the frequency appearance of the offspring from the ootheca of the short wing female was 98.5% short wing and 1.5% long wing form. Our specimens occasionally show body polymorphism in the form of individuals having long wings instead of the usual short one. The long wing form does not show the white band on the dorsal surface of its thorax.

Association between host wing morphology polymorphism and Wolbachia infection in Vollenhovia emeryi (Hymenoptera: Myrmicinae).

Many eusocial insects, including ants, show complex colony structures, distributions, and reproductive strategies. In the ant Vollenhovia emeryi Wheeler (Hymenoptera: Myrmicinae), queens and males are produced clonally, while sterile workers arise sexually, unlike other ant species and Hymenopteran insects in general. Furthermore, there is a wing length polymorphism in the queen caste. Despite its evolutionary remarkable traits, little is known about the population structure of this ant species, which may provide insight into its unique reproductive mode and polymorphic traits. We performed in-depth analyses of ant populations from Korea, Japan, and North America using three mitochondrial genes (COI, COII, and Cytb). The long-winged (L) morph is predominant in Korean populations, and the short-winged (S) morph is very rare. Interestingly, all L morphs were infected with Wolbachia, while all Korean S morphs lacked Wolbachia, demonstrating a association between a symbiont and a phenotypic trait. A phylogenetic analysis revealed that the S morph is derived from the L morph. We propose that the S morph is associated with potential resistance to Wolbachia infection and that Wolbachia infection does not influence clonal reproduction (as is the case in other ant species).

Redescription of Ploiaria mellea (Heteroptera: Reduviidae: Emesinae) and its first report from India.

A macropterous species, Ploiaria mellea McAtee Malloch, 1926 (Hemiptera: Heteroptera: Reduviidae: Emesinae: Leistarchini) is reported for the first time from India; the previously unknown male is illustrated and described. The species was originally described from the Philippines. Since the species was found in three climatically different localities near Pune, it must be a well-established emesine bug that has so far remained undetected or unidentified in India.

Ultrabithorax is a key regulator for the dimorphism of wings, a main cause for the outbreak of planthoppers in rice.

Rice planthoppers, the most devastating rice pests, occur in two wing forms: the short-wing form for rapid population growth and long-wing form for long-distance migration, which together create the mechanism for outbreak. Here we show that Ultrabithorax (Ubx) is a key regulator for switching between the long- and short-wing forms of rice planthoppers. Ubx is expressed in both forewing and hindwing pads, which is different from the canonical model of Ubx expression. In brown planthoppers, expression of Ubx (NlUbx) is regulated by nutritional status of the rice host. High-quality young plants induce NlUbx expression leading to the short-wing form; low-quality ripe plants reduce NlUbx expression resulting in long-wing form. We also showed that NlUbx is regulated by the insulin receptors NlInR1 and NlInR2. The default expression of NlInR1 inhibits NlUbx resulting in long-wings, while high-quality hosts induce NlInR2 expression, which represses NlInR1 thus promoting NlUbx expression to produce short-wings.

Carabid beetles of the environs of Lake Elton: fauna, population dynamics, demography.

Background: The present paper includes the results of a year-round pitfall trapping survey of ground beetles in the region of Lake Elton, Volgograd Area, Russia. The main objectives of the project lie in studying the local fauna of Carabidae in the Lake Elton region, as well as their demographic structure and dispersal potential of the local populations of particular carabid species. A total of ten model habitats: six zonal (characteristic of that particular biogeographic area) and four azonal (present in a variety of biogeographical areas) were studied. In each model habitat, ten pitfall traps were set from 10 May 2006 until 10 May 2007 and were checked at 10-day intervals except for the period with negative temperatures (from 1 November 2006 until 31 March 2007). During the period of observation, 51,314 specimens of Carabidae, belonging to 149 species, were trapped. The resulting sampling-event dataset includes 24,291 plot-based observations (= sampling events), some of them containing zero records of particular species in a certain habitat and time. New information: This is the first sampling-event dataset of a year-round pitfall trapping survey (from May 2006 until May 2007) of ground-beetle communities and the demographic structure of local populations of particular species in the Lake Elton region, Volgograd Area, Russia.

Molecular Mechanisms of Wing Polymorphism in Insects.

Many insects are capable of developing into either long-winged or short-winged (or wingless) morphs, which enables them to rapidly match heterogeneous environments. Thus, the wing polymorphism is an adaptation at the root of their ecological success. Wing polymorphism is orchestrated at various levels, starting with the insect's perception of environmental cues, then signal transduction and signal execution, and ultimately the transmitting of signals into physiological adaption in accordance with the particular morph produced. Juvenile hormone and ecdysteroid pathways have long been proposed to regulate wing polymorphism in insects, but rigorous experimental evidence is lacking. The breakthrough findings of ecdysone receptor regulation on transgenerational wing dimorphism in the aphid Acyrthosiphon pisum and of insulin signaling in the planthopper Nilaparvata lugens greatly broaden our understanding of wing polymorphism at the molecular level. Recently, the advent of high-throughput sequencing coupled with functional genomics provides powerful genetic tools for future insights into the molecular bases underlying wing polymorphism in insects.

Diurnal and developmental differences in gene expression between adult dispersing and flightless morphs of the wing polymorphic cricket, Gryllus firmus: Implications for life-history evolution.

The functional basis of life history adaptation is a key topic of research in life history evolution. Studies of wing-polymorphism in the cricket Gryllus firmus have played a prominent role in this field. However, prior in-depth investigations of morph specialization have primarily focused on a single hormone, juvenile hormone, and a single aspect of intermediary metabolism, the fatty-acid biosynthetic component of lipid metabolism. Moreover, the role of diurnal variation in life history adaptation in G. firmus has been understudied, as is the case for organisms in general. Here, we identify genes whose expression differs consistently between the morphs independent of time-of-day during early adulthood, as well as genes that exhibit a strong pattern of morph-specific diurnal expression. We find strong, consistent, morph-specific differences in the expression of genes involved in endocrine regulation, carbohydrate and lipid metabolism, and immunity - in particular, in the expression of an insulin-like-peptide precursor gene and genes involved in triglyceride production. We also find that the flight-capable morph exhibited a substantially greater number of genes exhibiting diurnal change in gene expression compared with the flightless morph, correlated with the greater circadian change in the hemolymph juvenile titer in the dispersing morph. In fact, diurnal differences in expression within the dispersing morph at different times of the day were significantly greater in magnitude than differences between dispersing and flightless morphs at the same time-of-day. These results provide important baseline information regarding the potential role of variable gene expression on life history specialization in morphs of G. firmus, and the first information on genetically-variable, diurnal change in gene expression, associated with a key life history polymorphism. These results also suggest the existence of prominent morph-specific circadian differences in gene expression in G. firmus, possibly caused by the morph-specific circadian rhythm in the juvenile hormone titer.

Wing Polymorphism and Trypanosoma cruzi Infection in Wild, Peridomestic, and Domestic Collections of Mepraia spinolai (Hemiptera: Reduviidae) From Chile.

Mepraia spinolai (Porter) is a vector of Trypanosoma cruzi that causes Chagas disease. Females are always wingless, but males may be winged or wingless. We determined by PCR the infection percentage with T. cruzi of M. spinolai adults and nymphs in domestic, peridomestic, and wild collections, in different regions of Chile. In all regions, winged males were more abundant than females and wingless males. Winged males collected inside houses were less parasitized than were those from peridomestic and wild environments. Although winged males of M. spinolai have comparatively low levels of infection, this segment may still represent the greatest vector threat in this species for transmission of T. cruzi to humans and other vertebrates in domestic, wild, and peridomestic habitats. Winged males represent the dispersive form of this species that invades human dwellings. Feeding deprivation resulting from the time required to find a food source and to search for reproductive females could explain the lower infection rates (negatives) of winged males collected from inside houses in comparison with winged males collected from peridomestic and wild habitats.

Insulin receptors and wing dimorphism in rice planthoppers.

Wing polymorphism contributes significantly to the success of a wide variety of insects. However, its underlying molecular mechanism is less well understood. The migratory planthopper (BPH), Nilaparvata lugens, is one of the most extensively studied insects for wing polymorphism, due to its natural features of short- and long-winged morphs. Using the BPH as an example, we first surveyed the environmental cues that possibly influence wing developmental plasticity. Second, we explained the molecular basis by which two insulin receptors (InR1 and InR2) act as switches to determine alternative wing morphs in the BPH. This finding provides an additional layer of regulatory mechanism underlying wing polymorphism in insects in addition to juvenile hormones. Further, based on a discrete domain structure between InR1 and InR2 across insect species, we discussed the potential roles by which they might contribute to insect polymorphism. Last, we concluded with future directions of disentangling the insulin signalling pathway in the BPH, which serves as an ideal model for studying wing developmental plasticity in insects.This article is part of the themed issue 'Evo-devo in the genomics era, and the origins of morphological diversity'.

A taxonomic study of the genus Panesthia (Blattodea, Blaberidae, Panesthiinae) from China with descriptions of one new species, one new subspecies and the male of Panesthiaantennata.

One new species Panesthiaguizhouensis sp. n. and one new subspecies Panesthiastellataconcava ssp. n. are described and illustrated. The male of Panesthiaantennata Brunner von Wattenwyl, 1893 and its brachypterous form are described and illustrated for the first time. Panesthiastrelkovi Bey-Bienko, 1969 is redescribed and illustrated. Three known species, Panesthiabirmanica Brunner von Wattenwyl, 1893, Panesthiasinuata Saussure, 1839 and Panesthiaangustipenniscognata Bey-Bienko, 1969 are illustrated. In addition, a key to all species of the genus Panesthia from China is presented.

Differential DNA methylation between two wing phenotypes adults of Sogatella furcifera.

Sogatella furcifera is a major rice pest with wing dimorphism. DNA methylation is an important epigenetic modification that plays a role in gene regulation and phenotype variation in most organisms. The objective of the current research was to survey the frequencies and variation of cytosine methylation at CCGG sequences in macropterous female adults (MFA) and brachypterous female adults (BFA) of S. furcifera, and to determine the occurrence of methylation changes associated with wing phenotypes by using methylation-sensitive amplification polymorphism (MSAP). No differences were found in the average proportions of methylated CCGG sites between MFA and BFA, but there were significant differences for methylation patterns between MFA and BFA. The fully methylated ratio was 5.81% in BFA, much higher than 2.40% in MFA; while the hemi-methylated ratio was 4.35% in BFA, much lower than 8.35% in MFA. These results provide circumstantial evidence that DNA methylation might be related to wing phenotype variation in S. furcifera. We also cloned and got 14 satisfactory sequences, which displayed variable cytosine methylation patterns between MFA and BFA. All these data will facilitate the researches on the epigenetic mechanisms of insect wing polymorphism.

A new species and karyotype variation in the bordering distribution of Mepraia spinolai (Porter) and Mepraia gajardoi Frías et al (Hemiptera: Reduviidae: Triatominae) in chile and its parapatric model of speciation

In the present study, the morphology, color pattern, chromosomal complement and aspects of meiosis in natural populations at the borders of the distributions of Mepraia gajardoi Frías et al and Mepraia spinolai (Porter) are described. The males of these bordering populations are brachypterous or macropterous, while females are always micropterous. Morphological and cytogenetic data indicated that the populations that border the distributions of M. gajardoi and M. spinolai, belong to a different species of parapatric origin.

EVOLUTIONARY ENDOCRINOLOGY OF JUVENILE HORMONE ESTERASE: FUNCTIONAL RELATIONSHIP WITH WING POLYMORPHISM IN THE CRICKET, GRYLLUS FIRMUS.

The existence, nature, and physiological consequences of genetic variation for juvenile hormone esterase (JHE) activity was studied in the wing-polymorphic cricket, Gryllus firmus. Hemolymph (blood) JHE activity was sixfold lower in nascent short-winged (SW) females, relative to nascent long-winged (LW) females during the last juvenile stadium (stage). Morph-associated genetic variation for JHE activity had two causes, variation in loci: (1) regulating whole-organism enzyme activity; and (2) controlling the degree to which JHE is secreted into the blood Reduced JHE activity in nascent SW-selected individuals was associated with reduced in vivo juvenile hormone catabolism. This suggests that variation in JHE activity during juvenile development may have important physiological consequences with respect to the regulation of blood levels of juvenile hormone and consequent specification of wing morph. This is the first definitive demonstration of genetic variation for hormonal metabolism in any insect and a genetic association between hormone metabolism and the subsequent expression of morphological variation (wing morph). However, we have not yet firmly established whether these associations represent causal relationships In contrast to the clear association between JHE activity and wing morph development, we observed no evidence indicating that variation in JHE activity plays any direct or indirect role in causing the dramatic differences in ovarian growth between adult wing morphs. Variation in JHE activity also does not appear to be important in coordinating the development of wing morph with the subsequent expression of reproductive differences between adult morphs. Finally genetic variation for the developmental profiles of JHE activity during juvenile and adult stages are remarkably similar in three Gryllus species. This suggests that genetic correlations between JHE activities during different periods of development, which underlie these activity profiles, have been conserved since the divergence of the three Gryllus species.

Age and size of European saltmarshes and the population genetic consequences for ground beetles.

Enzyme and dispersal polymorphisms of the saltmarsh carabid beetles Pogonus chalceus and Dicheirotrichus gustavii were studied in European populations varying in size and in isolation in space and time. D. gustavii, a constantly fullwinged species, has a larger genetic diversity and a smaller genetic differentiation between populations than the wing-polymorphic P. chalceus. Clear relationships between population or site characteristics and genetic structure were not observed, except for the special position taken by some small populations in both species. The dispersal power of P. chalceus in small populations is larger than in large populations, suggesting that these populations are unstable and/or young. Small populations, however, do not always show a lower genetic diversity than large populations, as would be expected from genetic drift. Dispersal power in P. chalceus declines with increasing age of the saltmarsh, probably due to continuous emigration of winged individuals. Age and size of saltmarshes, although difficult to study independently, both appear to be important in determining the genetic structure of saltmarsh beetles. Maximum diversity in both parameters is therefore recommended as an optimal nature conservation strategy.