Wing Shape Variation Research Articles

Intercontinental effect on sexual shape dimorphism and allometric relationships in the beetle pest Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae)

Hind wing shape variation was examined in 686 adult Diabrotica virgifera virgifera collected from maize plants in Europe and the USA Corn Belt, using geometric morphometric techniques. Sexual dimorphism at an Intercontinental scale was assessed using canonical variates analysis, a multivariate statistical method used to find the shape characters that best distinguish among groups of specimens. Our results showed that each of the populations of D. v. virgifera investigated in this study showed high levels of sex based hind wing shape dimorphism. In particular a stronger and more obvious pattern of hind wing shape variation was found in the USA than in Europe. These results support previous studies on D. v. virgifera wing shape that show that female D. v. virgifera have more elongated wings than males. These differences raise the question of whether sexual dimorphism may be modulated by natural selection.

Describing Variability in Wing Shapes among Three Populations of Plesispareichei Using Landmark-Based Geometric Morphometric Analysis

This study was conducted to describe variability in 3 populations of a coconut pest, Plesispareichei landmark-based geometric morphometric analysis of inner wing shapes. A total of 21 landmarks were used to represent dimensions in the left and right wings. Analysis of Variance, coordinate mapping, relative warp, Euclidean Distance Matrix and Cluster Analyses were used to analyze these landmarks. Results showed that significant variations were observed among populations. Variation in the left wing is mainly seen along the proximal landmark points but is variable in the right wing which may be an indication of asymmetry. Cluster analysis showed wing shape variations between populations indicating population differentiation in the pest. Distance was not a factor which may indicate differences in genetic structure between populations.

Wing geometry of Culex coronator (Diptera: Culicidae) from South and Southeast Brazil

BackgroundThe Coronator Group encompasses Culex coronator Dyar & Knab, Culex camposi Dyar, Culex covagarciai Forattini, Culex ousqua Dyar, Culex usquatissimus Dyar, Culex usquatus Dyar and Culex yojoae Strickman. Culex coronator has the largest geographic distribution, occurring in North, Central and South America. Moreover, it is a potential vector-borne mosquito species because females have been found naturally infected with several arboviruses, i.e., Saint Louis Encephalitis Virus, Venezuelan Equine Encephalitis Virus and West Nile Virus. Considering the epidemiological importance of Cx. coronator, we investigated the wing shape diversity of Cx. coronator from South and Southeast Brazil, a method to preliminarily estimate population diversity.MethodsField-collected immature stages of seven populations from a large geographical area in Brazil were maintained in the laboratory to obtain both females and males linked with pupal and/or larval exuviae. For each individual female, 18 landmarks of left wings were marked and digitalized. After Procrustes superimposition, discriminant analysis of shape was employed to quantify wing shape variation among populations. The isometric estimator centroid size was calculated to assess the overall wing size and allometry.ResultsWing shape was polymorphic among populations of Cx. coronator. However, dissimilarities among populations were higher than those observed within each population, suggesting populational differentiation in Cx. coronator. Morphological distances between populations were not correlated to geographical distances, indicating that other factors may act on wing shape and thus, determining microevolutionary patterns in Cx. coronator. Despite the population differentiation, intrapopulational wing shape variability was equivalent among all seven populations.ConclusionThe wing variability found in Cx. coronator populations brings to light a new biological problem to be investigated: the population genetics of Cx. coronator. Because of differences in the male genitalia, we also transferred Cx. yojoae to the Apicinus Subgroup.

Genome differentiation of Drosophila melanogaster from a microclimate contrast in Evolution Canyon, Israel.

The opposite slopes of "Evolution Canyon" in Israel have served as a natural model system of adaptation to a microclimate contrast. Long-term studies of Drosophila melanogaster populations inhabiting the canyon have exhibited significant interslope divergence in thermal and drought stress resistance, candidate genes, mobile elements, habitat choice, mating discrimination, and wing-shape variation, all despite close physical proximity of the contrasting habitats, as well as substantial interslope migration. To examine patterns of genetic differentiation at the genome-wide level, we used high coverage sequencing of the flies' genomes. A total of 572 genes were significantly different in allele frequency between the slopes, 106 out of which were associated with 74 significantly overrepresented gene ontology (GO) terms, particularly so with response to stimulus and developmental and reproductive processes, thus corroborating previous observations of interslope divergence in stress response, life history, and mating functions. There were at least 37 chromosomal "islands" of interslope divergence and low sequence polymorphism, plausible signatures of selective sweeps, more abundant in flies derived from one (north-facing) of the slopes. Positive correlation between local recombination rate and the level of nucleotide polymorphism was also found.

Allometric and non-allometric patterns in sexual dimorphism discrimination of wing shape in Ophion intricatus: might two male morphotypes coexist?

Bees and wasps could exhibit shape and size sexual dimorphism, and most of their morphological variation could depend on phenotypic responses due to environmental pressure during ontogenetic development. More complex measurement techniques related to size and shape rather than simply to mass and length should be required to analyze such a complex sexual dimorphism. In this study, differences related to wing shape and size of males and females of Ophion intricatus Brullé (Hymenoptera: Ichneumonidae) were evaluated using geometric morphometrics. Right and left wings of specimens were used, and a photographic matrix was constructed in which 18 morphological landmarks based on shape and vein patterns of the wings were digitalized. A multivariate analysis of wing shape showed significant differences between sexes and sites. The geometric variation demonstrated that the points at the intersection of radial and cubital-anal veins might be key characters to differentiate between sexes. This study also showed the presence of two clearly different male morphotypes coexisting in the same study site. However, it should be noted that the results of this study showed that the variation in wing shape is an analytical character in the determination of sexual differences in the family Ichneumonidae. These differences raise the question of whether sexual dimorphism of wing shape may be modulated by natural selection.

Latitudinal gradient effect on the wing geometry of Auca coctei (Guérin)(Lepidoptera, Nymphalidae)

Latitudinal gradient effect on the wing geometry of Auca coctei (Guérin) (Lepidoptera, Nymphalidae). When the environmental conditions change locally, the organisms and populations may also change in response to the selection pressure, so that the development of individuals may become affected in different degrees. There have been only a few studies in which the patterns of wing morphology variation have been looked into along a latitudinal gradient by means of geometric morphometrics. The aim of this work was to assess the morphologic differentiation of wing among butterfly populations of the species Auca coctei. For this purpose, 9 sampling locations were used which are representative of the distribution range of the butterfly and cover a wide latitudinal range in Chile. The wing morphology was studied in a total of 202 specimens of A. coctei (150 males and 52 females), based on digitization of 17 morphologic landmarks. The results show variation of wing shape in both sexes; however, for the centroid size there was significant variation only in females. Females show smaller centroid size at higher latitudes, therefore in this study the Bergmann reverse rule is confirmed for females of A. coctei. Our study extends morphologic projections with latitude, suggesting that wing variation is an environmental response from diverse origins and may influence different characteristics of the life history of a butterfly.

Sexual dimorphism and intraspecific variation in wing size and shape ofTongeia fischeri(Lepidoptera: Lycaenidae)

AbstractWing morphological variations are described here for the lycaenid butterfly Tongeia fischeri. A landmark‐based geometric morphometric approach based on wing venation of 197 male and 187 female butterflies collected in Japan was used to quantify wing size and shape variations between sexes and among populations. Sexual dimorphism in wing size and shape was detected. Females had significantly larger wings than males, while males showed a relatively elongated forewing with a longer apex and narrower wing tornus in comparison to females. Intraspecific variations in wing morphology among populations were revealed for the wing shape, but not wing size. Distinct wing shape differences were found in the vein intersections area around the distal part of the discal cell where median veins originated in the forewing and around the origin of the CU1 vein in the hindwing. In addition, phenotypic relationships inferred from wing shape variations grouped T. fischeri populations into three groups, reflecting the subspecies classification of the species. The spatial variability and phenotypic relationships between conspecific populations of T. fischeri detected here are generally in agreement with the previous molecular study based on mitochondrial and nuclear sequences, suggesting the presence of a phylogenetic signal in the wing shape of T. fischeri, and thus having taxonomic implications.

Evolutionary directional asymmetry and shape variation inDiabrotica virgifera virgifera(Coleoptera: Chrysomelidae): an example using hind wings

The western corn rootworm Diabrotica virgifera virgifera LeConte is a pest of maize in the USA and Europe and especially a problem in particular regions of Croatia. In this study patterns of variation in hind wing shape were examined through two objectives. The first objective examined the influence of soil type on 10 populations of D. v. virgifera sampled from three regions in Croatia that differed according to edaphic factors and climate. The second objective investigated the potential evolutionary presence of directional asymmetry on hind wings. Geometric morphometrics was used to examine these objectives by quantifying the morphological variation within and among individuals and populations. Overall, D. v. virgifera hind wing shape changed according to major soil type classifications in Croatia. The three hind wing morphotypes found varied because of basal radial vein differences, related to landmarks 1, 3, 7 and 14. This study shows that hind wing shape in D. v. virgifera can be used to differentiate populations based on edaphic factors and may have application as a monitoring tool in the integrated management of D. v. virgifera . In an evolutionary context, the presence of directional asymmetry in the hind wings of D. v. virgifera adds to the ever growing data on the evolution of insect wings.

Habitat variation and wing coloration affect wing shape evolution in dragonflies

Habitats are spatially and temporally variable, and organisms must be able to track these changes. One potential mechanism for this is dispersal by flight. Therefore, we would expect flying animals to show adaptations in wing shape related to habitat variation. In this work, we explored variation in wing shape in relation to preferred water body (flowing water or standing water with tolerance for temporary conditions) and landscape (forested to open) using 32 species of dragonflies of the genus Trithemis (80% of the known species). We included a potential source of variation linked to sexual selection: the extent of wing coloration on hindwings. We used geometric morphometric methods for studying wing shape. We also explored the phenotypic correlation of wing shape between the sexes. We found that wing shape showed a phylogenetic structure and therefore also ran phylogenetic independent contrasts. After correcting for the phylogenetic effects, we found (i) no significant effect of water body on wing shape; (ii) male forewings and female hindwings differed with regard to landscape, being progressively broader from forested to open habitats; (iii) hindwings showed a wider base in wings with more coloration, especially in males; and (iv) evidence for phenotypic correlation of wing shape between the sexes across species. Hence, our results suggest that natural and sexual selection are acting partially independently on fore- and hindwings and with differences between the sexes, despite evidence for phenotypic correlation of wing shape between males and females.

Rhodnius prolixus and Rhodnius robustus–like (Hemiptera, Reduviidae) wing asymmetry under controlled conditions of population density and feeding frequency

Habitat change in Rhodnius spp may represent an environmental challenge for the development of the species, particularly when feeding frequency and population density vary in nature. To estimate the effect of these variables in stability on development, the degree of directional asymmetry (DA) and fluctuating asymmetry (FA) in the wing size and shape of R. prolixus and R. robustus-like were measured under laboratory controlled conditions. DA and FA in wing size and shape were significant in both species, but their variation patterns showed both inter-specific and sexual dimorphic differences in FA of wing size and shape induced by nutrition stress. These results suggest different abilities of the genotypes and sexes of two sylvatic and domestic genotypes of Rhodnius to buffer these stress conditions. However, both species showed non-significant differences in the levels of FA between treatments that simulated sylvan vs domestic conditions, indicating that the developmental noise did not explain the variation in wing size and shape found in previous studies. Thus, this result confirm that the variation in wing size and shape in response to treatments constitute a plastic response of these genotypes to population density and feeding frequency.

WING SHAPE VARIATION ASSOCIATED WITH MIMICRY IN BUTTERFLIES

Mimetic resemblance in unpalatable butterflies has been studied by evolutionary biologists for over a century, but has largely focused on the convergence in wing color patterns. In Heliconius numata, discrete color-pattern morphs closely resemble comimics in the distantly related genus Melinaea. We examine the possibility that the shape of the butterfly wing also shows adaptive convergence. First, simple measures of forewing dimensions were taken of individuals in a cross between H. numata morphs, and showed quantitative differences between two of the segregating morphs, f. elegans and f. silvana. Second, landmark-based geometric morphometric and elliptical Fourier outline analyses were used to more fully characterize these shape differences. Extension of these techniques to specimens from natural populations suggested that, although many of the coexisting morphs could not be discriminated by shape, the differences we identified between f. elegans and f. silvana hold in the wild. Interestingly, despite extensive overlap, the shape variation between these two morphs is paralleled in their respective Melinaea comimics. Our study therefore suggests that wing-shape variation is associated with mimetic resemblance, and raises the intriguing possibility that the supergene responsible for controlling the major switch in color pattern between morphs also contributes to wing shape differences in H. numata.

PROPERTIES OF SPONTANEOUS MUTATIONAL VARIANCE AND COVARIANCE FOR WING SIZE AND SHAPE INDROSOPHILA MELANOGASTER

We estimated mutational variance-covariance matrices, M, for wing shape and size in two genotypes of Drosophila melanogaster after 192 generations of mutation accumulation. We characterized 21 potentially independent aspects of wing shape and size using geometric morphometrics, and analyzed the data using a likelihood-based factor-analytic approach. We implement a previously unused analysis that describes those directions with the greatest difference in evolvability between pairs of matrices. There are significant mutational effects on 19 of 21 possible aspects of wing form, consistent with the high dimensionality of standing genetic variation for wing shape previously identified in D. melanogaster. Mutations have partially recessive effects, consistent with average dominance around 0.25. Sex-specific matrices are relatively similar, although male-specific matrices are slightly larger, as expected due to dosage compensation on the X chromosome. Genotype-specific matrices are quite different. Matrices may differ both because of sampling error based on small samples of mutations with large phenotypic effects, and because of the mutational properties of the genotypes. Genotypic differences are likely to be involved, as the two genotypes have different molecular mutation rates and properties.

JOINT ALLELIC EFFECTS ON FITNESS AND METRIC TRAITS

Theoretical explanations of empirically observed standing genetic variation, mutation, and selection suggest that many alleles must jointly affect fitness and metric traits. However, there are few direct demonstrations of the nature and extent of these pleiotropic associations. We implemented a mutation accumulation (MA) divergence experimental design in Drosophila serrata to segregate genetic variants for fitness and metric traits. By exploiting naturally occurring MA line extinctions as a measure of line-level total fitness, manipulating sexual selection, and measuring productivity we were able to demonstrate genetic covariance between fitness and standard metric traits, wing size, and shape. Larger size was associated with lower total fitness and male sexual fitness, but higher productivity. Multivariate wing shape traits, capturing major axes of wing shape variation among MA lines, evolved only in the absence of sexual selection, and to the greatest extent in lines that went extinct, indicating that mutations contributing wing shape variation also typically had deleterious effects on both total fitness and male sexual fitness. This pleiotropic covariance of metric traits with fitness will drive their evolution, and generate the appearance of selection on the metric traits even in the absence of a direct contribution to fitness.

The phenetic structure of Aedes albopictus

The large and recent migrations of the main vector of the Chikungunya virus, Aedes albopictus, raise questions about the epidemiological impact of possible microevolutionary changes in new territories. Quantitative traits are suitable to detect such changes as induced by environmental adaptations, local competition and founder effects. Using landmark-based geometric morphometrics, we compared the size and shape of 22 populations (1572 females) of Ae. albopictus. The collection sites covered five countries around the world, with special emphasis on Asia, which is assumed to be the origin of the vector. Some collections came from places where an important epidemic outbreak of Chikungunya has recently occurred.Quantitative comparisons were based on 18 anatomical landmarks on the wing. To contrast geographic and possible interspecific shape variation, close species were introduced in the sample, namely five Aedes riversi and nine Ae. flavopictus from Japan. The three species had similar size, but they were clearly separated by shape.Within Ae. albopictus, there was general agreement on size variation with the available climatic data. Shape variation was less related to climatic data than to either geography or a known itinerary of past territorial expansion. Thus, two main clusters were distinguished by wing shape variation: the first one composed of the Southeast Asian sample, the second one grouping Japan, Florida, Hawaii and the Reunion Island samples.The Southeast Asian countries, assumed to be at the origin of the geographic expansion of the mosquito, had similar wings and constituted a distinct group where localities clustered into northern and southern localities. Contrasting with this homogeneous group, very distant localities such as United States (US) and Japan shared a common shape pattern. The US Ae. albopictus samples (Hawaii and Florida) were indeed very similar to the Japanese samples, with Florida behaving exactly like a northern Japanese locality, close to Tokyo and Yokohama. Shape proximity among these distant areas could be explained by a common and relatively recent ancestor, as generally suggested in the literature. The three Reunion Island samples conformed a group external to the Southeast Asian countries, loosely connected to the remaining localities.Thus, areas where recent epidemics took place, the Reunion Island and the southernmost localities of Thailand, did not cluster together, but represented separate groups.In sum, the size of the wing was apparently under the influence of climatic factors, while its shape could contribute information on species and geographic differences. Apparent departures from expectations based on geography were explained by known past and present migratory routes. These patterns of shape variation were compatible with genetic drift, suggesting microevolutionary changes probably induced by the expansion of the Ae. albopictus mosquito.

ALTITUDINAL CLINAL VARIATION IN WING SIZE AND SHAPE IN AFRICANDROSOPHILA MELANOGASTER: ONE CLINE OR MANY?

Geographical patterns of morphological variation have been useful in addressing hypotheses about environmental adaptation. In particular, latitudinal clines in phenotypes have been studied in a number of Drosophila species. Some environmental conditions along latitudinal clines-for example, temperature-also vary along altitudinal clines, but these have been studied infrequently and it remains unclear whether these environmental factors are similar enough for convergence or parallel evolution. Most clinal studies in Drosophila have dealt exclusively with univariate phenotypes, allowing for the detection of clinal relationships, but not for estimating the directions of covariation between them. We measured variation in wing shape and size in D. melanogaster derived from populations at varying altitudes and latitudes across sub-Saharan Africa. Geometric morphometrics allows us to compare shape changes associated with latitude and altitude, and manipulating rearing temperature allows us to quantify the extent to which thermal plasticity recapitulates clinal effects. Comparing effect vectors demonstrates that altitude, latitude, and temperature are only partly associated, and that the altitudinal shape effect may differ between Eastern and Western Africa. Our results suggest that selection responsible for these phenotypic clines may be more complex than just thermal adaptation.

Lead-Induced Variation in Wing Size and Shape in Populations ofDrosophila subobscura

The aim of the current study was to analyze the direction and range of changes in wing size and shape in both sexes of Drosophila subobscura (Collin, 1936) flies that originated from two natural populations with different evolutionary history (sampled from ecologically distinct habitats) maintained during seven generations in laboratory conditions on different lead (Pb) concentrations. The results showed significant wing size variability differences across seven generations of rearing on lead for both populations. Wing size is negatively correlated with lead level, in contrast with wing shape in which significant variation was observed in just one of the populations. According to our results, wing size seems to be more affected by lead pollution in both tested populations compared with wing shape. Our data suggest that presence of lead in higher concentration over extended period of time may reduce the stability of wing morphology and consequently reduce the fitness of exposed individuals. Therefore, specific stress that persists over multiple generations could increase the probabilities for extinction of populations composed of sensitive individuals.

Microgeographic spatial structuring of Triatoma infestans (Hemiptera: Reduviidae) populations using wing geometric morphometry in the Argentine Chaco.

We investigated the occurrence of spatial structuring in Triatoma infestans (KLug) (Hemiptera: Reduviidae) populations 12 yr after the last community-wide insecticide spraying campaign in rural Pampa del Indio, in the Gran Chaco of northeastern Argentina. In total, 172 male and 149 female right wings collected at 16 georeferenced sites with at least 10 individuals of the same sex were analyzed using geometric morphometry. Mean female body length and wing centroid size (CS) were significantly larger than for males. Log-transformed CS and length were significantly and positively correlated both for males and females. Males collected in domiciles had significantly smaller CS than those collected in peridomestic structures both closed (kitchens or storerooms) or open (chicken coops), in agreement with our previous results elsewhere in the dry Argentine Chaco. Female wing CS was not significantly different between ecotopes. Wing shape analyses showed the occurrence of significant geographic structuring in males and females combined and in males only. Male wings showed a strong association between Mahalanobis distance and geographic distance. In general, Mahalanobis distances were significantly different between collection sites located > 4 km apart. For collection sites located < 4 km apart, the greater the geographic distance the larger the difference in wing shape variables. Among females, only a partial correspondence between geographic groups and Mahalanobis distances was recorded. The strong spatial structuring found in T. infestans populations may be useful for the identification of putative reinfestation sources after vector control interventions.

Wing size and shape variation of Phlebotomus papatasi (Diptera: Psychodidae) populations from the south and north slopes of the Atlas Mountains in Morocco

The wing shape and size morphology of populations of the medically important phlebotomine sand fly, Phlebotomus papatasi, were examined in two endemic (south of the Atlas Mountains) and nonendemic (north of the Atlas Mountains) foci of cutaneous leishmaniasis by using geometric morphometrics in Morocco. Although it is present in all of Morocco, P. papatasi is the main vector of Leishmania major in only southern part of the Atlas Mountains. There are four major mountain ranges that serve as geographical barriers for species distribution in the study area and at least four gaps were recognized among these barriers. We found statistically significant differences in wing shape morphology between southern and northern populations. Analysis clearly recognized two main groups of populations on both sides of the mountains. The graphical depiction of Principal Component Analysis (PCA) and Canonical Variates Analysis (CVA) confirmed our morphometric study suggesting that the difference in wing morphology between the populations indicates that the population of P. papatasi shows phenotypic plasticity in the study area. According to centroid size analyses, which were used as measures of wing size differences among different sites, the north population of P. papatasi had relatively larger wings than the south population.

The relationship between molecular variation and variation in the wing shape of three aphid parasitoid species: Aphidius uzbekistanicus Luzhetzki, Aphidius rhopalosiphi De Stefani Perez and Aphidius avenaphis (Fitch) (Hymenoptera: Braconidae: Aphidiinae)

This paper examines the relationship between molecular and phenotypic variability among three closely related cereal aphid parasitoids: Aphidius rhopalosiphi and Aphidius uzbekistanicus from Euro-Asia and Aphidius avenaphis from North America. A. rhopalosiphi has the same distribution and shares the same aphid hosts as A. uzbekistanicus. The genetic variation of these parasitoids was explored using a barcoding area of the mitochondrial COI gene. Phenotypic variability based on the wing shape and stigma shape was explored using geometric morphometrics. Molecular analyses revealed the existence of two distinct clades: the A. rhopalosiphi clade, which is characterized by a high level of within-clade genetic variation, and the A. uzbekistanicus/A. avenaphis clade, which has almost no variation at the barcoding area of the mitochondrial COI gene. Generally, a high level of variation in wing shape has been shown within populations of both clades. Our results indicate that there is no relationship between the level of the variation at the barcoding area of the mitochondrial COI gene and the level of morphological variation. The genetically uniform A. uzbekistanicus/A. avenaphis clade had even greater variation in stigma shape than A. rhopalosiphi clade, which is characterized by a high level of genetic variation.

Morphometric and genetic differentiation of two sibling gossamer-wing damselflies, Euphaea formosa and E. yayeyamana, and adaptive trait divergence in subtropical East Asian islands.

Insular species frequently demonstrate different tendencies to become smaller or larger than their continental relatives. Two sibling gossamer—wing damselflies, Euphaea formosa (Odonata: Euphaeidae) from Taiwan and E. yayeyamana from the Yaeyama Islands of Japan, have no clear structural differentiation, and can only be recognized by their geographical distribution, sizes, and subtle differences in wing shape and coloration. This study combined morphometric and genetic techniques to investigate the adaptive significance of trait divergence and species status in these two Euphaea damselflies. Phylogenetic analyses of the mitochondrial cox2 sequences demonstrated that the two damselflies are monophyletic lineages and constitute valid phylogenetic species. The landmark—based geometric morphometrics indicated that the two damselflies are different morphological species characterized by distinctive wing shapes. The larger E. formosa exhibited broader hind wings, whereas E. yayeyamana had narrower and elongated forewings. The body size and wing shape variations among populations of the two species do not follow the expected pattern of neutral evolution, suggesting that the evolutionary divergence of these two traits is likely to be subjected to natural or sexual selection. The decreased body size, elongated forewings, and narrower hind wings of E. yayeyamana may represent insular adaptation to limited resources and reduced territorial competition on smaller islands.