Wing morphology,winter ecology,and fecundity selection: evidence for sex-dependence in barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>)

Variation in wing morphology results from the combination of diverse selection pressures. Wing feather morphology within species varies with sex and ontogenetic effects, and also with ecological factors. Yet, the direction of causation for the wing morphology–ecology association remains to be elucidated. Under the ‘ecology-dependence’ hypothesis, wing morphology covaries with ecological conditions, because the latter affect feather molt. Alternatively, the ‘habitat choice’ hypothesis posits that individuals with different wing morphology choose different habitats because of the habitat-dependent advantages of a specific wing morphology. We tested these competing hypotheses in the migratory, aerially insectivorous barn swallow (Hirundo rustica). We quantified wing morphology (isometric size, pointedness, and convexity) on the same individuals during consecutive breeding seasons (i.e., before and after molt in sub-Saharan wintering areas) and located wintering areas using light-level geolocators. Wing pointedness of females but not males during 1 year negatively correlated with vegetation vigor (gauged by the Normalized Difference Vegetation Index; NDVI) in the African area where individuals spent the next winter. Partial least-squares path modelling showed that the association between wing morphology and NDVI was sex-dependent. Conversely, NDVI during wintering did not predict wing morphology in the next breeding season. Because wing morphology can have carry-over effects on subsequent performance, we investigated selection on wing traits and found strong positive fecundity selection on wing size of females. Our results suggest that female barn swallows choose their wintering habitat depending on their wing morphology. In addition, directional fecundity selection operates on females, suggesting sex-dependence of current selection on the flight apparatus.     

Wing shape,wing size,and sexual dimorphism in eye‐span in stalk‐eyed flies (Diopsidae)

The eyes of stalk‐eyed flies (Diopsidae) are positioned at the end of rigid peduncles (‘stalks’) protruding laterally from the head. Eye‐stalk length varies within the family and, in some species, varies between males and females. Larger eye‐stalks in males result from sexual selection for longer stalks, a trait that increases male reproductive success. In the present study, we examined whether an increase in eye‐stalk length results in an adjustment of wing size and shape to deal with the burden of bearing an exaggerated ‘ornament’. We compared wing morphology among ten species of stalk‐eyed flies that differ in eye‐span and the degree of sexual dimorphism. Mass‐specific wing length differed between males and females in seven out of the ten species. Nondimensional wing shape parameters differed between the species (P < 0.001), but mostly did not differ between males and females of the same species. Dimorphism in eye‐span closely correlated with dimorphism in wing length (r = 0.89, P < 0.001) and the correlation remained significant (r = 0.81, P = 0.006) after correcting for phylogenetic relationships. Once corrected for phylogenetic relatedness, the mass‐specific wing length of males (but not females) was weakly correlated with mass‐specific eye‐span (r = 0.66, P = 0.042). We propose that the observed proportional increase in wing length associated with increased eye‐span can facilitate aerial manoeuverability, which would otherwise be handicapped by the elevated moment of inertia imposed by the wider head. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 860–871.     

Discovery of the genus Iasvia Zalessky, 1934 in the Upper Permian of France (Lodève basin) (Orthoptera, Ensifera, Oedischiidae)

New material attributed to the species Iasvia reticulata ZALESSKY, 1934 and to a new species in this genus is described from the Salagou Formation (Saxonian Group, Lodève basin). Preamble to the taxonomic section, the wing venation pattern of Orthoptera is discussed. The numerous described specimens yield decisive information about variability of wing venation within the genus, previously based on a single specimen from the Russian Permian. I. reticulata is the first species from the Lodève basin that is already known from another site. The biostratigraphic implications are discussed.     

Body mass and locomotor habits of the smallest darter,Anhinga minuta (Aves,Anhingidae)

During the Neogene of South America, Anhingidae was represented by several species, mainly with greater sizes than the extant members. In the present contribution, body mass and locomotor habits of Anhinga minuta, the smallest known darter, were inferred. Body mass was estimated using two methods, one with measures of a tibiotarsus (the holotype) and the other, with measurements of a humerus; locomotor habits were inferred through muscular reconstructions and wing parameters (wing span, wing area and wing loading). Estimates of wing span and wing area were based on the length of humerus, assuming a condition of isometry with respect to Anhinga anhinga; wing loading was obtained through a relation formula between wing area and body mass. The results obtained indicate a body mass of about 729 g, a wing span of 0.958 m, a wing area of 0.117 m² and a corresponding wing loading of 61 N/m². These values and also the proximal insertion of the musculus pectoralis are consistent with those of a soaring bird but with more frequent flapping than extant anhingids. Furthermore, the inferred musculature for tibiotarsus indicates abilities for swimming, climbing and moving through the vegetation as in extant representatives.     

Morphological and histological examination of polyphenic wing formation in the pea aphid <Emphasis Type="Italic">Acyrthosiphon pisum</Emphasis> (Hemiptera,Hexapoda)

Aphids display divergent adult phenotypes, depending on environmental conditions experienced during their embyonic and nymphal stages in their complex life cycles. The plastic developmental mode is an extreme case of phenotypic plasticity, so-called “polyphenism”, in which discrete multiple phenotypes are produced based on a single genome. For example, winged and wingless adult females are derived from a single genotype. However, the developmental mechanisms producing these polyphenic traits according to the extrinsic stimuli, such as density conditions, still remain unknown. In this study, to analyze the developmental processes underlying the wing polyphenism, we extensively observed and compared wing development in the winged and wingless individuals in parthenogenetic generations of the aphid Acyrthosiphon pisum (Harris), using scanning electron microscopy and histological sectioning. At the first-instar stage, the wing primordia were observed both in the future winged (W) and wingless (WL) nymphs. Developmental differences can be seen from the second-instar stage, when wing primordia degenerate in the WL nymphs, while they develop and become more thickened in the W nymphs, suggesting that the developmental programs should be launched prior to this stage. Furthermore, during the third- to fifth-instar stages, wing buds and flight muscles were well developed in the W nymphs, while wing primordia completely disappeared in the WL ones. In addition, the observation on the detailed developmental process of wing primordia during the third-instar W nymphs showed that the wing buds become swollen especially at the basal part, even during the intermolt period. This was caused by the development of wing epithelia under the cuticle of this instar nymph. Actually on the surface of the cuticle of wing-bud bases, there were numerous furrows, which gradually expand during the intermolt period. The similar situation was also observed at the forth-instar nymphs, in which the wings are formed in the complicated manner inside the wing pads. Furthermore, the developmental process of flight muscles was also described in detail. These dynamic developmental differences between the wing morphs should be regulated under the gene expression cascades that switch according to environmental stimuli.     

Gap junction distribution in the Drosophila wing disc mutants vg,l(2)gd,l(3)c43hs1, and l(2)gl4

The density of gap junctions in four Drosophila melanogaster mutants with abnormal wing disc development has been determined using quantitative electron microscopy and compared with the gap junction density in wild-type wing discs. No appreciable differences relative to wild-type controls were found in the cell death mutant vestigial or in the mildly hyperplastic mutant lethal giant disc which could not be accounted for in terms of altered lateral plasma membrane surface density or as an extension of the gap junction growth which normally occurs during the third larval stage of development in wild-type wing discs. However, both the severely hyperplastic mutant l₍₃₎c43^hs1 and the neoplastic mutant lethal giant larva have significant reductions in the gap junction surface density, the number of gap junctions, and the gap junction areal fraction of the lateral plasma membrane compared with wild-type controls. These differences cannot be attributed to altered lateral plasma membrane surface densities which are not significantly different from wild-type control wing discs. The reduced gap junction density in severely hyperplastic and neoplastic wing discs suggests that alterations in the number or distribution of gap junctions may be as disruptive to normal growth and development as their complete absence.     

Ergatoid reproductives in Nasutitermes columbicus (Isoptera,Termitidae)

Numerous functional ergatoid replacement reproductives were found in one colony of Nasutitermes columbicus in Panama. Their morphology was mainly workerlike, although several imaginal characters such as the compound eyes and variable wing buds were more or less developed. The sex organs were fully mature and the fat body of the females, not of the males, was of the “royal” type. The development of the eyes was not accompanied by the differentiation of the optic lobes of the brain, nor was the presence of wing buds correlated with a development of the wing muscles.     

A new form of <Emphasis Type="Italic">Kailasius autocrator</Emphasis> (Lepidoptera,Papilionidae) from the eastern Pamirs

A new subspecies, Kailasius autocrator pshartanus, from the eastern Pamirs (the Muzkol Ridge, Sasyk River) is described. The new subspecies is distinguished from the nominative subspecies by the wing pattern and ecology. This subspecies has the most primitive wing pattern in the genus. The range of this species is considered to be the center of Kailasius genus origin.     

Crowding leads to higher incidence of brachypterous females in the tobacco thrips,Frankliniella fusca (Hinds) (Thysanoptera: Thripidae)

The effects of larval density on the wing form determination of female tobacco thrips, Frankliniella fusca, were investigated by rearing thrips on leaf disks at 27.5 °C. The developmental period, head width, body length, and forewing length of individuals in each wing morph were determined to assess the relationships among larval density, growth, and wing form. Data showed that higher rearing densities increased the production of female F. fusca brachypters. There was no consistent difference in the mean developmental periods between the two wing morphs or among all 5 density treatments. The body length of females tended to decrease with increasing rearing density, but there was no significant difference in body size between the two wing morphs when they were reared under the same density level.     

Phylogenetic Signal in the Wing Shape in the Subfamily Dolichopodinae (Diptera,Dolichopodidae)

Interspecific and sex-related variations in the wing shape of 22 species of the fly subfamily Dolichopodinae, family Dolichopodidae were analyzed using geometric morphometrics. Mapping morphometric traits onto phylogeny revealed a clear phylogenetic signal in the interspecific variation and sexual dimorphism of wing shape. In some cases, not too closely related species occupied the same portion of the morphometric space, indicating some degree of homoplasy. Interspecific variation was associated with an increased wing area due to both elongation and widening or only elongation of the wing. An increase in wing area was accompanied by extension of the posterior crossvein to the apical part of the wing. The variation in wing shape related to sexual dimorphism involved the same structures (the posterior crossvein and the apical part of CuA₁), but variation associated with sexual dimorphism was distributed in fewer dimensions than interspecific variation. The allometric component of sexual dimorphism varied between species, and in most cases it was not the leading factor in wing shape variability.     

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.     

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.     

Male-specific flight apparatus development in Acyrthosiphon pisum (Aphididae, Hemiptera, Insecta): comparison with female wing polyphenism

Wing polymorphisms observed in many Insecta are important topics in developmental biology and ecology; these polymorphisms are a consequence of trade-offs between flight and other abilities. The pea aphid, Acyrthosiphon pisum, possesses 2 types of wing polymorphisms: One is a genetic wing polymorphism occurring in males, and the other is an environmental wing polyphenism seen in viviparous females. Although genetic and environmental cues for the 2 wing polymorphisms have been studied, differences in their developmental regulation have not been elucidated. In particular, there is little knowledge regarding the developmental processes in male wing polymorphism. Therefore, in this study, the development of flight apparatuses and external morphologies was compared among 3 male wing morphs (winged, wingless, and intermediate). These male developmental processes were subsequently compared with those of female wing morphs. Developmental differences between the male and female polymorphisms were identified in flight muscle development and degeneration but not in wing bud development. Furthermore, the nymphal periods of wingless and intermediate males were significantly shorter than that of winged males, indicating the adaptive significance of male winglessness. Overall, this study indicates that the male and female wing polymorphisms are based on different regulatory systems for flight apparatus development, which are probably the result of different adaptations under different selection pressures.     

<Emphasis Type="Italic">Kailasius autocrator</Emphasis> (Avinov, 1913) (Lepidoptera,Papilionidae) found in the alai mountain ridge (the Pamir-Alai)

A new subspecies, Kailasius autocrator murun Korb, ssp. n., of the species Kailasius autocrator (Avinov, 1913) from the eastern part of the Alai Mountain Ridge is described. This subspecies differs from the other known ones in the smaller size (the fore-wing length is less than 23 mm) and in the presence of 5 black rounded spots in the submarginal area of the hind-wing upper side (4 spots in the nominotypical subspecies and K. a. pshartanus). The new subspecies is shown to be the most archaic one among the subspecies of the genus Kailasius by the wing pattern. Possible ways for the origin of local and small populations of morphologically different subspecies of the genus are discussed.     

Females show greater changes in wing colour with latitude than males in the green‐veined white butterfly,Pieris napi (Lepidoptera: Pieridae)

In butterflies, wing colour may simultaneously be under sexual selection in the context of mating selection and natural selection in the context of thermoregulation. In the present study, we collected mated females of the green‐veined white butterfly (Pieris napi) from locations spanning 960 km of latitude across Fennoscandia, and investigated sex‐specific latitudinal wing colour variation in their offspring raised under identical conditions. We measured wing colour characteristics, including reflectance at wavelengths 300–700 nm and the degree of wing melanization. At all latitudes, females reflected more light in the short wavelengths (< 400 nm) and less in the long wavelengths (> 450 nm), and they were more melanized than males. However, female wing colour varied more with latitude than that of males. Among females, long wavelength reflectance decreased, whereas short wavelength reflectance and melanization increased, towards the north. By contrast, among males, latitudinal variation was found only in the ventral hindwing melanization. These results are consistent with the idea that the balance between natural and sexual selection acting on wing colour changes with latitude differently in males than females. The dark wing colour of females in the north may be a thermoregulatory adaptation, although males may be constrained from evolving the dark dorsal wing colour favoured by natural selection because of constant sexual selection across latitudes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Phenotypic variation of the scarce heath <Emphasis Type="Italic">Coenonympha hero</Emphasis> (L.) (Lepidoptera,Satyridae) from the central part of its range

Variation of the fore and hind wing lengths and the diameters of wing eyespots in two subspecies of Coenonympha hero (L.) from the central part of the species range was analyzed. Clinal variation was revealed, manifested in a gradual increase of the wing size and eyespot diameters from west to east in the Urals. Both in C. h. hero in the Urals and in C. h. perseis in Siberia, the northern peripheral populations had a specific wing pattern with eyespots smaller than in more southern parts of the corresponding ranges. In most cases, such climatic factors as the mean annual air temperature, precipitation, duration of the frost-free period, and humidity were shown to affect significantly the variation of morphological traits of C. hero. Chrono-geographical analysis of variation showed that in some cases the differences between the samples collected in the same locality in different years were comparable to or greater than geographic variation within the subspecies range. Thus, the population as a whole is influenced by climatic factors and weather conditions in a particular part of the range during a particular season; C. hero may therefore be a suitable object for monitoring the impact of environmental changes on insects.     

Thermoregulatory significance of wing melanization in Pieris butterflies (Lepidoptera: Pieridae): physics,posture, and pattern

Summary Pieris butterflies use a novel behavioral posture for thermoregulation called reflectance basking, in which the wings are used as solar reflectors to reflect radiation to the body. As a means of exploring the thermoregulatory significance of wing melanization patterns, I examine the relation of basking posture and wing color pattern to body temperature. A mathematical model of the reflectance process predicts certain combinations of dorsal wing melanization pattern and basking posture that maximize body temperature. Laboratory experiments and field observations show that this model correctly predicts qualitative differences in the relation of body temperature to basking posture based on differences in the extent of dorsal melanization on the wing margins, both between species and between sexes within species of Pieris. This is the first demonstration in insects that coloration of the entire wing surface can affect thermoregulation. Model and experimental results suggest that, in certain wing regions, increased melanization can reduce body temperature in Pieris; this effect of melanization is exactly the opposite of that found in other Pierid butterflies that use their wings as solar absorbers. These results are discussed in terms of the evolution of wing melanization pattern and thermoregulatory behavior in butterflies.     

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.     

New species of the genus <Emphasis Type="Italic">Pyrausta</Emphasis> (Lepidoptera,Crambidae) from Southern Siberia and Transcaucasia

Two new species of the pyralid genus Pyrausta Schrank, 1802, P. anastasia Shodotova, sp. n. from southern Siberia and P. alexandra Shodotova, sp. n. from the Transcaucasia, are described. P. anastasia sp. n. is very similar to P. chrysitis (Butler, 1881) in coloration and the wing pattern, but differs in the darker brown fore wings, a wider pale yellow band on the hind wing, and the structure of the aedeagus. P. alexandra sp. n. is very similar to P. falcatalis (Guenée, 1854) in coloration and the wing pattern, but differs in the darker brown wings, in the number of pale yellow spots on the fore and hind wings, and in the structure of the male and female genitalia. The holotypes and paratypes of both species are deposited in the Zoological Institute, Russian Academy of Sciences (St. Petersburg).