Developmental noise and phenotypic plasticity are correlated in Drosophila simulans.

Non-genetic variation is the phenotypic variation induced by the differential expression of a genotype in response to varying environmental cues and is broadly categorized into two types: phenotypic plasticity and developmental noise. These aspects of variation have been suggested to play an important role in adaptive evolution. However, the mechanisms by which these two types of non-genetic variations influence the evolutionary process are currently poorly understood. Using a machine-learning-based phenotyping tool, we independently quantified phenotypic plasticity and developmental noise in the wing morphological traits of the fruit fly Drosophila simulans. Utilizing a rearing experiment, we demonstrated plastic responses in both wing size and shape as well as non-zero heritability of both phenotypic plasticity and developmental noise, which suggests that adaptive phenotypic plasticity can evolve via genetic accommodation in the wing morphology of D. simulans. We found a positive correlation between phenotypic plasticity and developmental noise, while the correlation between the plastic response to three kinds of environmental factors that were examined (nutrient condition, temperature, and light-dark cycle) was poor. These results suggest that phenotypic plasticity and developmental noise contribute to evolvability in a similar manner, however, the mechanisms that underlie the correspondence between these two types of variation remain to be elucidated.

Phylogeography and phenotypic wing shape variation in a damselfly across populations in Europe.

BACKGROUND: Describing geographical variation in morphology of organisms in combination with data on genetic differentiation and biogeography can provide important information on how natural selection shapes such variation. Here we study genetic structure using ddRAD seq and wing shape variation using geometric morphometrics in 14 populations of the damselfly Lestes sponsa along its latitudinal range in Europe. RESULTS: The genetic analysis showed a significant, yet relatively weak population structure with high genetic heterozygosity and low inbreeding coefficients, indicating that neutral processes contributed very little to the observed wing shape differences. The genetic analysis also showed that some regions of the genome (about 10%) are putatively shaped by selection. The phylogenetic analysis showed that the Spanish and French populations were the ancestral ones with northern Swedish and Finnish populations being the most derived ones. We found that wing shape differed significantly among populations and showed a significant quadratic (but weak) relationship with latitude. This latitudinal relationship was largely attributed to allometric effects of wing size, but non-allometric variation also explained a portion of this relationship. However, wing shape showed no phylogenetic signal suggesting that lineage-specific variation did not contribute to the variation along the latitudinal gradient. In contrast, wing size, which is correlated with body size in L. sponsa, had a strong negative correlation with latitude. CONCLUSION: Our results suggest a relatively weak population structure among the sampled populations across Europe, but a clear differentiation between south and north populations. The observed geographic phenotypic variation in wing shape may have been affected by different local selection pressures or environmental effects.

Unraveling biotypes of the northern house mosquito, Culex pipiens s.l. (Diptera: Culicidae): molecular differentiation and morphometric analysis.

Geometric morphometrics was used to determine whether geographic isolation could explain differences in wing size and shape between and within continental (27°S to 41°S) and insular (Rapa Nui) populations of Culex pipiens s.s. Linnaeus and their biotypes (f. pipiens and f. molestus). Molecular protocols based on polymorphisms in the second intron of nuclear locus ace-2 (acetylcholinesterase-2) were used to differentiate Cx. pipiens s.s. from Cx. quinquefasciatus Say, and an assay based on polymorphisms in the flanking region of a microsatellite locus (CQ11) was used to identify biotypes. Culex pipiens f. molestus and hybrids shared larval habitats in all continental sites, while Cx. pipiens f. pipiens was found in 5 of the 10 sites. Only biotype molestus was found in Rapa Nui (Easter Island) Pipiens and molestus biotypes occur sympatrically in aboveground locations, and only molestus was found in the underground site (ME). Biotype molestus was dominant in rural locations and preferably anthropophilic. These results agree with the ecological descriptions previously reported for the biotypes of Cx. pipiens s.s. Procrustes ANOVA only showed differences in centroid size between biotypes in females and males and did not show significant differences in wing shape. However, we found significant differences among the geographic areas in the centroid size and wing shape of both females and males. Particularly, the population of Rapa Nui Island had shorter wings than the continental populations. The results highlight the effects of geographic and environmental processes on morphotypes in vector mosquitoes.

The Extraordinary Diversity of Merodon avidus Complex (Diptera: Syrphidae)-Adding New Areas, New Species and a New Molecular Marker.

In this paper, the Merodon avidus (Diptera, Syrphidae) species complex was revised, whereupon we discovered and described four new species for science: Merodon atroavidus Vujic, Radenkovic et Likov sp. nov., M. magnus Vujic, Kocis Tubic et Acanski sp. nov., M. nigroscutum Vujic, Radenkovic et Likov sp. nov. and M. pseudomoenium Vujic, Kocis Tubic et Acanski sp. nov. An integrative taxonomy approach was used to delimit species boundaries. Two molecular markers (the mitochondrial COI gene and nuclear 28S rRNA gene-newly analysed marker for the complex) and geometric morphometry of the wing shape, together with morphological data and distribution, successfully separated all species from the complex. The morphological variability of the analysed species is described and discussed and an illustrated diagnostic key for typical morpho-forms of species from the M. avidus complex is presented. A distribution map of all investigated species from the complex is provided. The level of endemicity of the M. avidus complex was discussed.

Wing Shape Variation between Terrestrial and Coastal Populations of the Invasive Box Tree Moth, Cydalima perspectalis, in Croatia.

The box tree moth (Cydalima perspectalis Walker, 1859; Lepidoptera: Crambidae) is an invasive species naturally distributed in Asia. The caterpillars in all developmental stages cause damage through defoliation of plants, and ultimately the death of the plant itself may occur. It is possible to recognize this species by its silk barriers and threads, and in the case of an intense attack, the entire plant will be covered with them. In Europe, this species' presence was first recorded in 2007 in Germany and the Netherlands, and it is now widely distributed. In Croatia, its existence was first recorded in 2012, in Istria, while substantial damages were recorded in 2013. This work aimed to determine the morphological variability of C. perspectalis from Croatia and assess its invasive character, the possibility of flight, and the risk of further spread. The methods of geometric morphometrics were used as the analysis of wing shape. A total of 269 moths from different locations in Croatia were collected, the upper wings of males and females were analyzed using 14 landmarks. Significant differences in wing shapes between terrestrial and coastal populations were found, as well as subtle wing shape sexual dimorphism. The implications of this variability in species invasiveness and capacity of spread are discussed in this paper. We also extrapolate the usefulness of our results and suggest strategies for predicting and managing invasive species.

Sex Ratio, Sexual Dimorphism, and Wing Geometric Morphometrics in Species of Trypoxylon Latreille, 1796 (Hymenoptera: Crabronidae).

Studying the nesting biology of wasp and bee species provides valuable insights into the ecology and evolution of these insects, shedding light on their ecological significance and aiding in their conservation efforts. Trypoxylon Latreille, 1796 is a genus of spider-hunting wasps that provisions their brood with paralyzed preys. This study focuses on aspects of nesting biology of Trypoxylon (Trypargilum) species such as sex ratio, sexual dimorphism, and morphometric variation at both inter- and intraspecific levels. The secondary sex ratio (SR) exhibited variation among species and populations. Males predominantly emerged from the first brood cells and from trap nests with smaller diameters. In comparison, females showed significantly larger body mass and linear wing measurements than males. Procrustes ANOVA values for centroid size (CS) and wing shape (SH) indicated significant differences in both wing size and shape among the five analyzed Trypoxylon species. Sexual dimorphism indices (SDI) derived from centroid size were found to be similar to those obtained from linear measurements, while SDI values based on body mass were significantly higher. Nests containing a greater number of cells tended to produce a larger number of higher-quality males and females. This observation, along with the lower coefficient of variation for female body size and high heritability, suggests that this trait may be subject to natural selection. Further studies that estimate the size of parents and their respective offspring are necessary to confirm the fitness advantage associated with larger female sizes in Trypoxylon species.

Allometric wing growth links parental care to pterosaur giantism.

Pterosaurs evolved a broad range of body sizes, from small-bodied early forms with wingspans of mostly 1-2 m to the last-surviving giants with sizes of small airplanes. Since all pterosaurs began life as small hatchlings, giant forms must have attained large adult sizes through new growth strategies, which remain largely unknown. Here we assess wing ontogeny and performance in the giant Pteranodon and the smaller-bodied anurognathids Rhamphorhynchus, Pterodactylus and Sinopterus. We show that most smaller-bodied pterosaurs shared negative allometry or isometry in the proximal elements of the fore- and hindlimbs, which were critical elements for powering both flight and terrestrial locomotion, whereas these show positive allometry in Pteranodon. Such divergent growth allometry typically signals different strategies in the precocial-altricial spectrum, suggesting more altricial development in Pteranodon. Using a biophysical model of powered and gliding flight, we test and reject the hypothesis that an aerodynamically superior wing planform could have enabled Pteranodon to attain its larger body size. We therefore propose that a shift from a plesiomorphic precocial state towards a derived state of enhanced parental care may have relaxed the constraints of small body sizes and allowed the evolution of derived flight anatomies critical for the flying giants.

Avian wings can lengthen rather than shorten in response to increased migratory predation danger.

Increasing predation danger can select for safety-enhancing modifications to prey morphology. Here, we document the multi-decade wing lengthening of a Pacific flyway migrant, the western sandpiper (Calidris mauri), and contrast this with contemporaneous wing shortening of the closely related semipalmated sandpiper (C. pusilla) on the Atlantic flyway. We measured >12,000 southbound western sandpipers captured from 1978 to 2020 at a major stopover site in British Columbia. Wing length increased at 0.074 mm year-1 (SE = 0.017; p < .0003) for adults, and 0.087 mm year-1 (SE = 0.029; p < .007) for juveniles. These rates are of similarly large magnitude (4%-5% overall), but opposite in direction, to the rate we previously reported for semipalmated sandpiper adults (-0.103 mm year-1). In both species, the change is specific to wings rather than being part of a general body size change. We interpret both trends as responses to the ongoing strong increase of peregrine falcon (Falco peregrinus) populations since the mid-1970s, an important predator encountered by these species in contrasting ways during migration. Western sandpipers and peregrine migrations have temporal and spatial overlap. Longer wings enhance migratory speed and efficiency, enabling western sandpipers to decrease overlap by advancing to safer zones ahead of falcon passage. In contrast, semipalmated sandpipers primarily encounter peregrines as residents at migratory staging sites. Shorter wings improve acceleration and agility, helping migrants to escape attacks. Juvenile western sandpiper wing length also shows a component additive to the lengthening trend, shifting between years at 0.055 mm day-1 with the highly variable snowmelt date, with wings shorter following early springs. On the Pacific flyway, the timing of peregrine southward passage advances with snowmelt, increasing the relative exposure of juveniles to post-migratory resident peregrines. We interpret this annual wing length adjustment as an induced defense, made possible because snowmelt timing is a reliable cue to danger in the upcoming migration.

Wing morphology covaries with migration distance in a highly aerial insectivorous songbird.

According to classical prediction of aerodynamic theory, birds and other powered fliers that migrate over long distances should have longer and more pointed wings than those that migrate less. However, the association between wing morphology and migratory behavior can be masked by contrasting selective pressures related to foraging behavior, habitat selection and predator avoidance, possibly at the cost of lower flight energetic efficiency. We studied the handwing morphology of Eurasian barn swallows Hirundo rustica from four populations representing a migration distance gradient. This species is an aerial insectivore, so it flies extensively while foraging, and may migrate during the day using a 'fly-and-forage' migration strategy. Prolonged foraging flights may reinforce the effects of migration distance on flight morphology. We found that two wings' aerodynamic properties-isometric handwing length and pointedness, both favoring energetically efficient flight, were more pronounced in barn swallows from populations undertaking longer seasonal migrations compared to less migratory populations. Our result contrast with two recent interspecific comparative studies that either reported no relationship or reported a negative relationship between pointedness and the degree of migratory behavior in hirundines. Our results may thus contribute to confirming the universality of the rule that longer migrations are associated with more pointed wings.

Altered wing phenotypes of captive-bred migratory birds lower post-release fitness.

Captive breeding and release to the wild is a globally important conservation tool. However, captivity can result in phenotypic changes that incur post-release fitness costs, especially if they affect strenuous or risky behaviours. Bird wing shape is critical for migration success and suboptimal phenotypes are strongly selected against. In this study, I demonstrate surprising plasticity of bird wing phenotypes in captivity for 4/16 studied species. In a model species, captive-born juveniles with wild wing phenotypes (a 1-mm longer distal primary flight feather) survived post-release at 2.7 times the rate of those with captive phenotypes (i.e. a shorter distal feather). Subtle phenotypic changes and their fitness impacts are more common than widely realised because they are easily overlooked. To improve captive breeding for conservation, practitioners must surveil phenotypic changes and find ways to mitigate them.

Experimental Evidence That Phenotypic Evolution but Not Plasticity Occurs along Genetic Lines of Least Resistance in Homogeneous Environments.

AbstractGenetic correlations concentrate genetic variation in certain directions of the multivariate phenotype. Adaptation and, under some models, plasticity is expected to occur in the direction of the phenotype containing the greatest amount of genetic variation (gmax). However, this may hinge on environmental heterogeneity, which can affect patterns of genetic variation. I use experimental evolution to test whether plasticity and phenotypic evolution follow gmax during adaptation to environments that varied in environmental heterogeneity. For >25 generations, Drosophila melanogaster populations were exposed to six homogeneous or spatially and temporally heterogeneous treatments involving hot (25°C) and cold (16°C) temperatures. Five wing traits were assayed in both temperatures. Wing morphology diverged between populations evolving in homogeneous hot and cold temperatures in a direction of the phenotype containing a large proportion of genetic variance and that aligned closely with gmax at 16°C but not at 25°C. Spatial heterogeneity produced an intermediate phenotype, which was associated with similar genetic variance across assay temperatures compared with all other treatments. Surprisingly, plasticity across assay temperatures was in a different direction to phenotypic evolution and aligned better with maternal variance than gmax. Together, these results provide experimental evidence for evolution along genetic lines of least resistance in homogeneous environments but no support for predicting plastic responses from the orientation of genetic variation. These results also suggest that spatial heterogeneity could maintain genetic variation that increases the stability of genetic variance across environments.

Complexities of recapitulating polygenic effects in natural populations: replication of genetic effects on wing shape in artificially selected and wild-caught populations of Drosophila melanogaster.

Identifying the genetic architecture of complex traits is important to many geneticists, including those interested in human disease, plant and animal breeding, and evolutionary genetics. Advances in sequencing technology and statistical methods for genome-wide association studies have allowed for the identification of more variants with smaller effect sizes, however, many of these identified polymorphisms fail to be replicated in subsequent studies. In addition to sampling variation, this failure to replicate reflects the complexities introduced by factors including environmental variation, genetic background, and differences in allele frequencies among populations. Using Drosophila melanogaster wing shape, we ask if we can replicate allelic effects of polymorphisms first identified in a genome-wide association studies in three genes: dachsous, extra-macrochaete, and neuralized, using artificial selection in the lab, and bulk segregant mapping in natural populations. We demonstrate that multivariate wing shape changes associated with these genes are aligned with major axes of phenotypic and genetic variation in natural populations. Following seven generations of artificial selection along the dachsous shape change vector, we observe genetic differentiation of variants in dachsous and genomic regions containing other genes in the hippo signaling pathway. This suggests a shared direction of effects within a developmental network. We also performed artificial selection with the extra-macrochaete shape change vector, which is not a part of the hippo signaling network, but showed a largely shared direction of effects. The response to selection along the emc vector was similar to that of dachsous, suggesting that the available genetic diversity of a population, summarized by the genetic (co)variance matrix (G), influenced alleles captured by selection. Despite the success with artificial selection, bulk segregant analysis using natural populations did not detect these same variants, likely due to the contribution of environmental variation and low minor allele frequencies, coupled with small effect sizes of the contributing variants.

Mixed support for an alignment between phenotypic plasticity and genetic differentiation in damselfly wing shape.

The relationship between genetic differentiation and phenotypic plasticity can provide information on whether plasticity generally facilitates or hinders adaptation to environmental change. Here, we studied wing shape variation in a damselfly (Lestes sponsa) across a latitudinal gradient in Europe that differed in time constraints mediated by photoperiod and temperature. We reared damselflies from northern and southern populations in the laboratory using a reciprocal transplant experiment that simulated time-constrained (i.e. northern) and unconstrained (southern) photoperiods and temperatures. After emergence, adult wing shape was analysed using geometric morphometrics. Wings from individuals in the northern and southern populations differed significantly in shape when animals were reared in their respective native environment. Comparing wing shape across environments, we found evidence for phenotypic plasticity in wing shape, and this response differed across populations (i.e. G × E interactions). This interaction was driven by a stronger plastic response by individuals from the northern population and differences in the direction of plastic wing shape changes among populations. The alignment between genetic and plastic responses depended on the specific combination of population and rearing environment. For example, there was an alignment between plasticity and genetic differentiation under time-constrained, but not under non-time-constrained conditions for forewings. We thus find mixed support for the hypothesis that environmental plasticity and genetic population differentiation are aligned. Furthermore, although our laboratory treatments mimicked the natural climatic conditions at northern and southern latitudes, the effects of population differences on wing shape were two to four times stronger than plastic effects. We discuss our results in terms of time constraints and the possibility that natural and sexual selection is acting differently on fore- and hindwings.

Size does matter: intraspecific geometric morphometric analysis of wings of the blowfly Chrysomya albiceps (Diptera: Calliphoridae).

Blowflies have forensic, sanitary and veterinary importance, as well as being pollinators, parasitoids and ecological bioindicators. There is still little work with real data and from experiments assessing the relationship between blowflies' morphologic features and environmental and demographic factors. The present work tests whether the variation, in the shape and size, of Chrysomya albiceps (Wiedemann, 1819) wings is influenced by the following factors: 1) time; 2) temperature; 3) sex and; 4) different types of carcasses (pig, dog/cat and whale). Male and female wings from four different sites collected in six different years were used to obtain wing size and shape of C. albiceps. Analyses between wing shape and the variables tested had low explanatory power, even though they had statistical support. However, it was possible to identify differences in wing shape between males and females, with good returns in sex identification. The comparison between wing size and the variables tested showed that wing size has a negative relationship with temperature, significant differences between sexes, slight variation over time and no influence by carcass types. Furthermore, wing size influenced wing shape. Understanding population-specific characteristics of C. albiceps provide important insights about how the species reacts under specific conditions.

Morphological Stasis in Time? A Triatoma brasiliensis brasiliensis Study Using Geometric Morphometrics in the Long Run.

Triatoma brasiliensis brasiliensis Neiva, 1911 is one of the most important vectors of Chagas disease in the Brazilian semiarid regions in the north-east. The risk imposed by T. b. brasiliensis to the human populations, due to frequent invasions and/or colonization of the domiciles, demands constant monitoring and control actions as well as an understanding of its evolutionary process. In this context, the following research studies the pattern of shape adaptation over time using a large dataset from 102 years of specimen collections in order to identify the morphological plasticity of this vector in Brazil. This dataset was analyzed using geometric morphometrics tools and the timescale was divided into eight different groups, containing specimens from 1912 to 2014. Geometric morphometrics analysis showed an interesting morphological stasis in the wing shape of T. b. brasiliensis, which allowed us to understand the high capacity of adaptation to changes in climate condition through time, and the invasive status which Triatoma species have around the world. Moreover, these results showed novel findings as an interesting phenotypic pattern, with no modifications in more than 100 years, leading us to understand the shape evolution in Triatominae as a vector species of diseases.

Sex and age-specific differences in wing pointedness and wing length in blackcaps Sylvia atricapilla migrating through the southern Baltic coast.

The blackcap Sylvia atricapilla shows a complex migratory pattern and is a suitable species for the studies of morphological migratory syndrome, including adaptations of wing shape to different migratory performance. Obligate migrants of this species that breed in northern, central, and Eastern Europe differ by migration distance and some cover shorter distance to the wintering grounds in the southern part of Europe/North Africa or the British Isles, although others migrate to sub-Saharan Africa. Based on ˃40 years of ringing data on blackcaps captured during autumn migration in the Southern Baltic region, we studied age- and sex-related correlations in wing pointedness and wing length of obligate blackcap migrants to understand the differences in migratory behavior of this species. Even though the recoveries of blackcaps were scarce, we reported some evidence that individuals which differ in migration distance differed also in wing length. We found that wing pointedness significantly increased with an increasing wing length of migrating birds, and adults had longer and more pointed wings than juvenile birds. This indicates stronger antipredator adaptation in juvenile blackcaps than selection on flight efficiency, which is particularly important during migration. Moreover, we documented more pronounced differences in wing length between adult and juvenile males and females. Such differences in wing length may enhance a faster speed of adult male blackcaps along the spring migration route and may be adaptive when taking into account climatic effects, which favor earlier arrival from migration to the breeding grounds.

Adaptive shifts underlie the divergence in wing morphology in bombycoid moths.

The evolution of flapping flight is linked to the prolific success of insects. Across Insecta, wing morphology diversified, strongly impacting aerodynamic performance. In the presence of ecological opportunity, discrete adaptive shifts and early bursts are two processes hypothesized to give rise to exceptional morphological diversification. Here, we use the sister-families Sphingidae and Saturniidae to answer how the evolution of aerodynamically important traits is linked to clade divergence and through what process(es) these traits evolve. Many agile Sphingidae evolved hover feeding behaviours, while adult Saturniidae lack functional mouth parts and rely on a fixed energy budget as adults. We find that Sphingidae underwent an adaptive shift in wing morphology coincident with life history and behaviour divergence, evolving small high aspect ratio wings advantageous for power reduction that can be moved at high frequencies, beneficial for flight control. By contrast, Saturniidae, which do not feed as adults, evolved large wings and morphology which surprisingly does not reduce aerodynamic power, but could contribute to their erratic flight behaviour, aiding in predator avoidance. We suggest that after the evolution of flapping flight, diversification of wing morphology can be potentiated by adaptative shifts, shaping the diversity of wing morphology across insects.

Exploratory Analysis of Color Forms' Variability in the Invasive Asian Lady Beetle Harmonia axyridis (Pallas 1773).

The Asian ladybird (Harmonia axyridis Pallas), native to Asia, is one of the 100 most invasive species in the world and has spread worldwide. This study aimed to characterize color forms of H. axyridis in Croatia and to analyze the variability of wing shape between populations and indicated forms. Geometric morphometric methods were used to analyze a total of 129 left and right wings in males and 126 left and right wings in females of H. axyridis collected from four different sites in Croatia. The results show a significant difference in wing shapes between the studied forms. Each form had its own specific morphotype that likely originated under the influence of genetic changes in the species. This study demonstrates that the use of geometric morphometric analysis is effective in studying the variability in H. axyridis populations. As this study is the first of its kind, for further clarity, it is necessary to conduct additional studies on a larger number of sites and an equal number of individuals of all forms.

Punctuational ecological changes rather than global factors drive species diversification and the evolution of wing phenotypes in Morpho butterflies.

Assessing the relative importance of geographical and ecological drivers of evolution is paramount to understand the diversification of species and traits at the macroevolutionary scale. Here, we use an integrative approach, combining phylogenetics, biogeography, ecology and quantified phenotypes to investigate the drivers of both species and phenotypic diversification of the iconic Neotropical butterfly genus Morpho. We generated a time-calibrated phylogeny for all known species and inferred historical biogeography. We fitted models of time-dependent (accounting for rate heterogeneity across the phylogeny) and paleoenvironment-dependent diversification (accounting for global effect on the phylogeny). We used geometric morphometrics to assess variation of wing size and shape across the tree and investigated their dynamics of evolution. We found that the diversification of Morpho is best explained when considering variable diversification rates across the tree, possibly associated with lineages occupying different microhabitat conditions. First, a shift from understory to canopy was characterized by an increased speciation rate partially coupled with an increasing rate of wing shape evolution. Second, the occupation of dense bamboo thickets accompanying a major host-plant shift from dicotyledons towards monocotyledons was associated with a simultaneous diversification rate shift and an evolutionary 'jump' of wing size. Our study points to a diversification pattern driven by punctuational ecological changes instead of a global driver or biogeographic history.

Genomics of altitude-associated wing shape in two tropical butterflies.

Understanding how organisms adapt to their local environment is central to evolution. With new whole-genome sequencing technologies and the explosion of data, deciphering the genomic basis of complex traits that are ecologically relevant is becoming increasingly feasible. Here, we studied the genomic basis of wing shape in two Neotropical butterflies that inhabit large geographical ranges. Heliconius butterflies at high elevations have been shown to generally have rounder wings than those in the lowlands. We reared over 1,100 butterflies from 71 broods of H. erato and H. melpomene in common-garden conditions and showed that wing aspect ratio, that is, elongatedness, is highly heritable in both species and that elevation-associated wing aspect ratio differences are maintained. Genome-wide associations with a published data set of 666 whole genomes from across a hybrid zone, uncovered a highly polygenic basis to wing aspect ratio variation in the wild. We identified several genes that have roles in wing morphogenesis or wing aspect ratio variation in Drosophila flies, making them promising candidates for future studies. There was little evidence for molecular parallelism in the two species, with only one shared candidate gene, nor for a role of the four known colour pattern loci, except for optix in H. erato. Thus, we present the first insights into the heritability and genomic basis of within-species wing aspect ratio in two Heliconius species, adding to a growing body of evidence that polygenic adaptation may underlie many ecologically relevant traits.