First description of the female of Sarcophaga (Sarcorohdendorfia) gracilior (Chen, 1975) (Diptera,Sarcophagidae)

Sarcophaga (Sarcorohdendorfia) gracilior (Chen, 1975) is documented from specimens collected in Hubei Province, China, using morphological characters and wing interference patterns (WIPs). The female of S. (S.) gracilior is described for the first time, the male is redescribed, and both sexes are photographed. The distribution of the species is updated.     

Ecological and phylogenetic perspectives on wing stiffness in nine Theclini species (Lepidoptera: Lycaenidae)

We studied the relationship between wing stiffness and butterfly ecology and phylogeny. Nine species belonging to the tribe Theclini of the family Lycaenidae were selected and examined for the wing stiffness of dried specimens by a three‐point bending test. It was found in Japonica lutea that the wing stiffness was not affected by the humidity to which it had been exposed, but was strongly affected by wing size and sex. Comparisons of sexual differences in four species indicated that females of patrolling species had stiffer wings than conspecific males, but that males of territorial species had stiffer wings than conspecific females. Finally, the wing stiffness was compared among males of nine species that use different mate‐locating tactics, and the results revealed a tendency that males of territorial species have stiffer wings than males of patrolling species. These results, though including a few exceptional cases, are discussed from the perspective of ecological requirements and phylogenetic constraints on the species.     

The wing morphology of limnephilid caddisflies in relation to their habitat preferences

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Swarm behaviour and mate competition in mayflies (Ephemeroptera)

Although mayfly swarms are frequently cited as an example of lekking by insects, little is known about the behaviour of individuals within a swarm, or how mate-selection takes place. A study of five species of mayfly over a period of 10 consecutive years has revealed species-specific differences in the flight pattern of swarming males and in the ability of males to recognize swarms of their own species. Males of four of the five species jostle other males in the swarm at all times except when mating: mating pairs are not jostled. The pattern of jostling varies with the species. Measurements of the sperm content of the vesicula seminalis and of the wing length of members of individual swarms show that larger wing size is positively correlated with the presence of less sperm. The vesicula seminalis is always filled with sperm at the beginning of the imaginal stage and the testes regress before the beginning of the imaginal stage. If the volume of sperm in the vesicula seminalis is a valid index of mating success then males with larger wings have the highest success. Large wings may bestow an advantage during jostling. The males of Ephemera danica , which do not jostle, glide with outspread wings; these outspread wings may attract females, the largest wings being the most attractive. Females of all five species enter the swarm a few at a time, although many females may be resting beneath the swarm. This phased entry may decrease the attraction of the swarm for predators. The number of females in a swarm is not correlated with swarm size, and the factors which enable females to regulate their entry into a swarm remain obscure.     

The Role of Wing Pigmentation, UV and Fluorescence as Signals in a Neotropical Damselfly

Pigmentation patterns, ultraviolet reflection and fluorescent emission are often involved in mate recognition and mate quality functions in many animal taxa. We investigated the role of wing ultra-violet reflection, fluorescence emission, and pigmentation on age and sexual signals in the damselfly Mnesarete pudica. In this species, wings are sexually dimorphic in colour and exhibit age dependency: males and females show a smoky black colouration when young, turning red in mature males while it turns brown in females. First, we investigated wing UV patterns through reflectance and emission spectra. Second, behavioural experiments were undertaken to show male and female responses to manipulated wing pigmentation and experimentally reduced UV (UV-). Reflectance spectra of the wings of juvenile and mature males and females were used to show the differences between controls and individuals with manipulated colouration used in the behavioural experiment. UV-reduced, females with wings painted red, and control males and females were tethered and presented to conspecific males and females, and their behavioral responses were recorded. The male red wing pigmentation and females with red wings elicited an aggressive response in territorial males and a sexual response in females. Both males and females showed neutral responses towards individuals with reduced UV. Wing signals of juvenile individuals also provoked neutral responses. These results suggest that UV, together with pigmentation, plays a role during mate recognition in males and females. Other than butterflies and spiders, it seems that fluorescence signals and UV reflectance can also be part of communication in odonates.     

Morphological and histological examination of short‐wing formation in the winter moth Protalcis concinnata (Insecta: Lepidoptera,Geometridae)

Winter geometrid moths exhibit sexual dimorphism in wing length and female‐specific flightlessness. Female‐specific flightlessness in insects is an interesting phenomenon in terms of sexual dimorphism and reproductive biology. In the winter geometrid moth, Protalcis concinnata (Wileman), adult females have short wings and adult males have fully developed wings. Although the developmental process for wing reduction in Lepidoptera is well studied, little is known about the morphology and the developmental pattern of short‐winged flightless morphs in Lepidoptera. To clarify the precise mechanisms and developmental processes that produce short‐winged morphs, we performed morphological and histological investigations of adult and pupal wing development in the winter geometrid moth P. concinnata. Our findings showed that (a) wing development in both sexes is similar until larval‐pupal metamorphosis, (b) the shape of the sexually dimorphic wings is determined by the position of the bordering lacuna (BL), (c) the BL is positioned farther inward in females than in males, and (d) after the short pupal diapause period, the female pupal wing epithelium degenerates to approximately two‐thirds its original size due to cell death. We propose that this developmental pattern is a previously unrecognized process among flightless Lepidoptera.     

The Sharpshooter Genus Geitogonalia Young: Descriptions of a New Brazilian Species and of the Female of Geitogonalia quatuordecimmaculata (Taschenberg) (Hemiptera: Cicadellidae)

The sharpshooter genus Geitogonalia Young was so far known from a single species from Southeastern Brasil—Geitogonalia quatuordecimmaculata (Taschenberg). A new species, Geitogonalia viridis, is described and illustrated in this paper based on males and females from the state of Rio de Janeiro. In addition, the female of G. quatuordecimmaculata is for the first time described and illustrated in detail. Geitogonalia quatuordecimmaculata is a dark species marked with bright orange on the fore wings, whereas the new species has the corium and clavus of the fore wings mostly green. The paraphyses are well developed in G. quatuordecimmaculata, with elongate rami, whereas they are inconspicuous in G. viridis, with the rami modified as a pair of very small sclerites.     

Can the narrow red bands of dragonflies be used to perceive wing interference patterns?

Despite numerous studies of selection on position and number of spectral vision bands, explanations to the function of narrow spectral bands are lacking. We investigate dragonflies (Odonata), which have the narrowest spectral bands reported, in order to investigate what features these narrow spectral bands may be used to perceive. We address whether it is likely that narrow red bands can be used to identify conspecifics by the optical signature from wing interference patterns (WIPs). We investigate the optical signatures of Odonata wings using hyperspectral imaging, laser profiling, ellipsometry, polarimetric modulation spectroscopy, and laser radar experiments. Based on results, we estimate the prospects for Odonata perception of WIPs to identify conspecifics in the spectral, spatial, intensity, polarization, angular, and temporal domains. We find six lines of evidence consistent with an ability to perceive WIPs. First, the wing membrane thickness of the studied Odonata is 2.3 μm, coinciding with the maximal thickness perceivable by the reported bandwidth. Second, flat wings imply that WIPs persist from whole wings, which can be seen at a distance. Third, WIPs constitute a major brightness in the visual environment only second after the solar disk. Fourth, WIPs exhibit high degree of polarization and polarization vision coincides with frontal narrow red bands in Odonata. Fifth, the angular light incidence on the Odonata composite eye provides all prerequisites for direct assessment of the refractive index which is associated with age. Sixth, WIPs from conspecifics in flight make a significant contribution even to the fundamental wingbeat frequency within the flicker fusion bandwidth of Odonata vision. We conclude that it is likely that WIPs can be perceived by the narrow red bands found in some Odonata species and propose future behavioral and electrophysiological tests of this hypothesis.     

Phenotypic variability of eye-spots in natural populations of <Emphasis Type="Italic">Coenonympha pamphilus</Emphasis> L. (Lepidoptera,Satyridae)

Variation of eye-spots in the wing pattern of Coenonympha pamphilus (Linnaeus, 1758) was studied in the Urals with adjacent territories and in Dzhelal-Abad Province of Kyrgyzstan. Discriminant analysis of the wing length and eye-spot diameter and the phenetic distances calculated by the Hartman’s method revealed clinal variation of the characters. The butterflies in the northern parts of the range (Sverdlovsk and Kurgan Provinces) are smaller than those in the south (Orenburg and Chelyabinsk Provinces) and have fewer eye-spots which themselves are smaller. The Kyrgyzstan specimens differ significantly from the Ural ones (the nominotypical subspecies) in both quantitative and qualitative characteristics of the wing pattern, and should probably be assigned to the subspecies C. pamphilus marginata Heyne, 1894. As in most Satyridae, females of C. pamphilus are larger than males and on the average possess more eye-spots on the wings. The degree of phenotypic variation in natural populations was studied using the method of variation spectra (Kovalenko, 1996a, 1996b, 2007, etc.). The arrangement of phenotypic combinations within the theoretical spectrum (St) matrix allows one to describe the actual (Sr) and potential (Sp) spectra for either sex and for the species as a whole. The actual spectrum was found to be considerably broader in males than in females. With minor variations, Sr and Sp showed the same general trends in different samples of both sexes. In all the samples, phenetic combinations with one spot on the fore wing (in cell M ₁-M ₂) were predominant. The hind wing had either no spots (which was typical of males, especially in the northern parts of the range) or the maximum possible number, six (mostly in females, more often in the southern parts of the range). The phenetic combinations with the maximum (4) number of spots on the fore wing and the minimum number (0) on the hind wing are prohibited for C. pamphilus.     

Fearful symmetry? Intra-individual comparisons of asymmetry in cryptic vs. signalling colour patterns in butterflies

The signalling role of asymmetry has attracted considerable recent interest among evolutionary biologists. Although it has been studied primarily within the context of sexual selection, symmetry of signals may play a role also in inter-specific communication, such as predator–prey interactions. Both theory and experimental evidence suggest that asymmetry may impair the efficacy of visual warning signals used to deter potential predators, but increase the protective value of non-signalling, cryptic colour patterns used to decrease the risk of detection. Here we tested the prediction from this hypothesis by means of intra-individual comparisons of asymmetry in colour pattern elements in three species of moths (Arctia caja (L.), Noctua orbona (L.), Smerinthus ocellata (L.)) that possess cryptic fore wing patterns and signalling hind wing patterns. Mean asymmetries constituted 4.3% (range 2.1–7.0%) of trait size for colour pattern elements, whereas individual asymmetry levels reached as high as 26%. Asymmetry tended to be somewhat larger in cryptic patterns on fore wings than in signalling patterns on hind wings in five of six comparisons, but in only one case was the difference statistically significant. In addition, pattern elements were somewhat more asymmetric on fore wings also in Saturnia pavonia (L.), which possesses identical signalling eyespots on both fore and hind wings. The relatively low levels of asymmetry also in cryptic patterns imply either that selection does not favour increased asymmetry in cryptic patterns, or that the evolution of pronounced asymmetry is developmentally or genetically constrained.     

Flight muscle breakdown in the soldier caste of the gall-inducing thrips species, Kladothrips intermedius Bagnall

Within the gall-inducing thrips of Australia, genus Kladothrips, is a single origin of a soldier caste. A subsequent radiation has led to at least seven social species, and two species that are likely to have independently lost the soldier caste. Both losses of soldiers are connected to a shift in the insects’ host plant. A third inferred host shift is correlated with life history changes in a species with soldiers, K. intermedius, which might suggest this defensive caste may be in transition. The soldiers of this species have variable wing lengths that overlap with that of the flight-capable dispersers. Our study was designed to assess the possibility that some soldiers in this species retain the ability to disperse by wing. A morphological assessment shows that a proportion of soldiers have body dimensions and wing lengths that fall within the range of dispersers, with males more so than females. However, longer wing length in soldiers (sample of primarily females) did not correspond with a disperser like walking behaviour. Furthermore, histological sections suggest that wing muscles of soldiers are deteriorated or absent, which is similar to what was observed in their foundress mothers. The presence of long wings did not correspond with being flight capable in this caste.     

Altitude and life‐history shape the evolution of Heliconius wings

Phenotypic divergence between closely related species has long interested biologists. Taxa that inhabit a range of environments and have diverse natural histories can help understand how selection drives phenotypic divergence. In butterflies, wing color patterns have been extensively studied but diversity in wing shape and size is less well understood. Here, we assess the relative importance of phylogenetic relatedness, natural history, and habitat on shaping wing morphology in a large dataset of over 3500 individuals, representing 13 Heliconius species from across the Neotropics. We find that both larval and adult behavioral ecology correlate with patterns of wing sexual dimorphism and adult size. Species with solitary larvae have larger adult males, in contrast to gregarious Heliconius species, and indeed most Lepidoptera, where females are larger. Species in the pupal‐mating clade are smaller than those in the adult‐mating clade. Interestingly, we find that high‐altitude species tend to have rounder wings and, in one of the two major Heliconius clades, are also bigger than their lowland relatives. Furthermore, within two widespread species, we find that high‐altitude populations also have rounder wings. Thus, we reveal novel adaptive wing morphological divergence among Heliconius species beyond that imposed by natural selection on aposematic wing coloration.     

Wing pigmentation in territorial male damselflies, Calopteryx haemorrhoidalis: a possible relation to sexual selection

One striking characteristic in adult males of some odonate species is the presence of wing pigmentation. In Calopteryx species, males show a series of pre- and postcopulatory behavioural displays during which they face females while showing their pigmented wings. One hypothesis to explain the precopulatory flying displays and the associated wing pigmentation is that they may serve a sexual selection function. I investigated this in the territorial damselfly Calopteryx haemorrhoidalis. Males of this species defend aquatic substrates that females use for oviposition. Observational evidence indicated that males with a higher proportion of wing pigmentation were more likely to defend a territory, obtained more matings, had fewer gut parasites, survived in the study site and stayed in territories for longer. Experimental evidence suggested that the relationship mating success and wing pigmentation still held when controlling for the size of the substrate defended by territorial males. Similar to other studies in the Calopterygidae, these results suggest that wing pigmentation may be favoured by sexual selection. I discuss, however, whether an alternative function for male copulatory courtship displays and wing pigmentation, as sexual and/or species recognition, may also explain the evolution of these traits. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Seasonal variability of wing length and eyespots in populations of <Emphasis Type="Italic">Erebia ligea</Emphasis> (L.) (Lepidoptera,Satyridae) in the Middle Urals

The flight period of Erebia ligea (Linnaeus, 1758) in the Sysert District of Sverdlovsk Province lasts from early July until early August only in the odd-numbered years. The species is characterized by protandry, its males emerging 5–7 days before females. Erebia ligea manifests sexual and seasonal variation of the wing length and diameter of the eyespots in the Middle Urals. According to the literature and newly obtained data, the relative male/female wing length varies within the species range. The vectors of the seasonal variation of the body size are opposite in the males and females: in early summer the largest males and the smallest females occur, then mediumsized individuals appear, while the smallest males and the largest females fly at the end of the warm period. Although the males of Erebia ligea are larger than females, the latter have larger eyespots on the wings. Seasonal variation of the diameters of the forewing eyespots is largely similar in the two sexes: individuals with larger spots appear earlier in the season than those with small spots. The eyespots treated as phenes in this study develop on the wings only if the remaining spots of the wing pattern attain a threshold size. A threshold nature was shown for P4 spot in M ₃-Cu ₁ cell of the fore wing and G2 spot in cell M ₁–M ₂ of the hind wing. Individuals with and without phene G2 have opposite vectors of seasonal variation of the sizes of the rest spots of the hind wing pattern. It is apparently the threshold character of variation of these elements of the wing pattern that defines the variation of the proportion of phenetic combinations of the eyespots in the individuals emerging in different periods of the season. Describing phenetic variation at the population level, one should remember that the percentage of the “morphs” or phenetic combinations may vary during a season, especially in the protandric species.     

Development of a wingless morph in the ladybird beetle, Adalia bipunctata

SUMMARY Many taxa of winged insects have independently lost the ability to fly and often possess reduced wings. Species exhibiting natural variation in wing morphology provide opportunities to investigate the genetics and developmental processes underlying the evolution of alternative wing morphs. Although many wing dimorphic species of beetles are known, the underlying mechanisms of variation are not well understood in this insect order. Here, we examine wing development of wild type and natural wingless morphs of the two-spot ladybird beetle, Adalia bipunctata . We show that both pairs of wings are distally truncated in the wingless adults. A laboratory population of the wingless morph displays heritable variation in the degree of wing truncation, reflecting reduced growth of the larval wing discs. The coexistence of variable wingless morphs supports the idea that typical monomorphic wingless insects may be the result of a gradual evolution of wing loss. Gene expression patterns in wing discs suggest that the conserved gene network controlling wing development in wild-type Adalia is disrupted in the dorsoventral patterning pathway in the wingless morphs. Previous research on several species of ant has revealed that the anteroposterior wing patterning pathway is disrupted in wingless workers. Future investigations should confirm whether interruptions in both taxa are limited to the patterning pathways found thus far, or whether there are also shared interruption points. Nevertheless, our results highlight that diverse mechanisms of development are likely to underlie the evolution of wingless insects.     

Age-associated male mating success in three swarming caddis fly species (Trichoptera: Leptoceridae)

Abstract. 1. In the three caddis fly species, Athripsodes albifrons (L.), A. cinereus (Curtis) and Mystacides azurea (L.) (Leptoceridae; Trichoptera), males swarm above the water surface of lakes or rivers. Receptive females fly to swarms and are chased and/or courted by males. After one of the swarming males has grasped an approaching female, the pair flies in tandem to the shore where they copulate.
2. In males, wing wear indices were negatively correlated with the ratio of fat/dry weight. In the only species in which comparisons were possible between newly emerged and swarming males (M. azurea), the former had significantly lower indices. Unmated females on average had lower wing wear indices than spent females. These facts suggest that wing wear reflects relative age.
3. The tandem males had significantly less wing wear than those in swarms, and are probably therefore younger. Age is therefore likely to be significant in relation to mating success.
4. Among males of the same relative age, tandem males had higher fat ratio than swarming ones, indicating that male mating success was also influenced by traits other than age. It is suggested that the shortest possible duration of the period of adult prematurity is adaptive, especially in insects with marginal adult food intake.     

Aerial Contests, Sexual Selection and Flight Morphology in Solitary Pompilid Wasps

Aerial contest competition has proven to be a challenging phenomenon to interpret in many territorial insects. Because the duels often consist of elaborate and/or high speed ascending maneuvers, the hypothesis that they are settled due to asymmetries in flight performance is intuitively appealing. We evaluated this hypothesis by contrasting differences in known morphological determinants of flight performance between (1) residents vs. non-residents of the territorial wasp, Hemipepsis ustulata and between (2) H. ustulata vs. a non-territorial relative, Pepsis thisbe . In the first contrast, resident male H. ustulata were seen to be larger, and had a tendency for reduced wing loading, but they did not possess greater flight musculature or wing aspect ratios (i.e., more elongated wings) than their non-resident counterparts. In the second contrast, male H. ustulata exhibited clearly greater flight musculature and greater sexual dimorphism in this parameter (males more muscular), and also exhibited a slight tendency for greater wing loading and smaller aspect ratios than males of the patrolling species P. thisbe . Interestingly, although size is linked with territorial success in H. ustulata , males of this species were not larger than male P. thisbe , nor did the former species exhibit greater sexual size dimorphism. These results do not support the hypothesis that the repeated ascending contests of H. ustulata require, and select for, a high acceleration design. However, the observed intraspecific patterns of flight musculature suggest that high acceleration is favored in males of the perching species, perhaps for the ability to intercept passing receptive females.     

Sexual dimorphism of auditory function and structure in praying mantises (Mantodea; Dictyoptera)

Sexual dimorphism of tympanate auditory systems in insects has bees described in only a few taxonomically isolated cases. However, widespread sexual dimorphism occurs in the ultrasound-sensitive, midline ear of the praying mantis.
In dimorphic species, it is always the female mantis that shows a reduction in ultrasonic hearing. The dimorphism may be mild—a difference in tuning and small reduction in sensitivity—or extreme with no evidence of audition in the female. In all but the mildest cases, the reduction in hearing is accompanied by significant anatomical divergence from the male ear structure. Two distinct metathoracic groove ('ear') types are linked to hearing reduction in the females.
Anatomical evidence of auditory sexual dimorphism appears in 34% of the 183 mantis genera examined. The dimorphic genera are widely but non-uniformly distributed within three of the four largest mantis families.
Auditory sexual dimorphism is closely correlated with dimorphism in wing length. In general, mantises with functional wings have sensitive ultrasonic hearing while those with short wings do not. These findings support the hypothesis that ultrasonic hearing in mantises is part of a defensive system against attack by echolocating bats.     

Insights into the molecular mechanisms underlying diversified wing venation among insects

Insect wings are great resources for studying morphological diversities in nature as well as in fossil records. Among them, variation in wing venation is one of the most characteristic features of insect species. Venation is therefore, undeniably a key factor of species-specific functional traits of the wings; however, the mechanism underlying wing vein formation among insects largely remains unexplored. Our knowledge of the genetic basis of wing development is solely restricted to Drosophila melanogaster. A critical step in wing vein development in Drosophila is the activation of the decapentaplegic (Dpp)/bone morphogenetic protein (BMP) signalling pathway during pupal stages. A key mechanism is the directional transport of Dpp from the longitudinal veins into the posterior crossvein by BMP-binding proteins, resulting in redistribution of Dpp that reflects wing vein patterns. Recent works on the sawfly Athalia rosae, of the order Hymenoptera, also suggested that the Dpp transport system is required to specify fore- and hindwing vein patterns. Given that Dpp redistribution via transport is likely to be a key mechanism for establishing wing vein patterns, this raises the interesting possibility that distinct wing vein patterns are generated, based on where Dpp is transported. Experimental evidence in Drosophila suggests that the direction of Dpp transport is regulated by prepatterned positional information. These observations lead to the postulation that Dpp generates diversified insect wing vein patterns through species-specific positional information of its directional transport. Extension of these observations in some winged insects will provide further insights into the mechanisms underlying diversified wing venation among insects.