Variation In Wing Morphology Research Articles

Wings of fringed fruit-eating bats (Artibeus fimbriatus) are highly integrated biological aerofoils from perspectives of secondary sexual dimorphism, allometry and modularity

AbstractPhenotypic variability is ubiquitous. This is especially true in bats, where families such as Phyllostomidae encompass as much phenotypic variability as some entire orders of mammals. Typically, phenotypic variability is characterized based on cranial morphology, with studies of other functionally important aspects of the phenotype, such as legs, feet and wings, being less frequent. We examined patterns of secondary sexual dimorphism and allometry of wing elements of the fringed fruit-eating bat (Artibeus fimbriatus) and examined, for the first time, the modularity of bat wings. Patterns were based on 13 wing measurements taken from 21 females and 15 males from eastern Paraguay. From a multivariate perspective, A. fimbriatus exhibited significant secondary sexual dimorphism. Females were larger than males for all 13 wing characteristics, with significant differences involving the last phalanx of the fourth and fifth digits. Female wings were also relatively larger than male wings from a multivariate perspective, as was the last phalanx of the fourth and fifth digits, after adjusting for wing size based on forearm length. Wing elements were highly variable regarding allometric relationships, with some exhibiting no allometric patterns and others exhibiting isometry or hyperallometry, depending on the element. Wings exhibited significant modularity, with metacarpals, proximal phalanges and distal phalanges each representing a discrete module. The wings of A. fimbriatus exhibit substantive patterns of dimorphism, allometry and modularity. Although the big mother hypothesis is a strong theoretical construct to explain wing dimorphism, there is not yet any sound theoretical basis for the patterns of allometry and modularity of the wing. Further investigation is required to understand the determinants of variation in wing morphology.

Variation in wing morphology is related to breeding environment in a high‐elevation specialist bird

The morphology of bird wings is subject to a variety of selective pressures, including migration, predation, habitat structure and sexual selection. Variation in wing morphology also occurs at the intraspecific and intrapopulation level, and can be related to sex, age, migration strategy and environmental factors. The relationship between environment and intraspecific variation in wing morphology is still poorly understood. In this work, we studied the relationship between wing morphology and breeding environment in a high‐elevation specialist bird, the water pipit Anthus spinoletta. We calculated wing isometric size, pointedness and convexity of 84 birds mist‐netted at breeding sites in year 2021 in the European Alps. We then searched for associations between these traits and potentially relevant breeding site characteristics (vegetation structure, elevation, latitude). For all wing traits, sex and one or more environmental factors best explained the variation, with environmental factors explaining between 3 and 8% of the variation. Wing size was negatively related to tree cover and wing convexity was negatively related to bush cover. Elevation contributed to explain variation in wing pointedness, but the direction of its effect was unclear. The negative relationship between wing size and tree cover could be due to intraspecific competition, i.e. to the relegation of smaller winged low‐quality individuals in marginal grassland areas. Higher wing convexity could improve predator escape ability in areas with scarce protecting vegetation, with possible effects on habitat choice. These findings represent one of the few demonstrated cases of wing morphology–environment relationships at the intraspecific level.

Relating wing morphology and immune function to patterns of partial and differential bat migration using stable isotopes.

Migration is energetically expensive and is predicted to drive similar morphological adaptations and physiological trade-offs in migratory bats and birds. Previous studies suggest that fixed traits like wing morphology vary among species and individuals according to selective pressures on flight, while immune defences can vary flexibly within individuals as energy is variably reallocated throughout the year. We assessed intraspecific variation in wing morphology and immune function in silver-haired bats Lasionycteris noctivagans, a species that follows both partial and differential migration patterns. We hypothesized that if bats experience energy constraints associated with migration, then wing morphology and immune function should vary based on migratory tendency (sedentary or migratory) and migration distance. We predicted that long-distance migrants would have reduced immune function and more migration-adapted wing shapes compared to resident or short-distance migrating bats. We estimated breeding latitude of spring migrants using stable hydrogen isotope techniques. Our sample consisted primarily of male bats, which we categorized as residents, long-distance northern migrants, short-distance northern migrants and southern migrants (apparent breeding location south of capture site). Controlling for individual condition and capture date, we related wing characteristics and immune indices among groups. Some, but not all, aspects of wing form and immune function varied between migrants and residents. Long-distance northern migrants had larger wings than short-distance northern migrants and lower wing loading than southern migrants. Compared with resident bats, short-distance northern migrants had reduced IgG while southern migrants had heightened neutrophils and neutrophil-to-lymphocyte ratios. Body fat, aspect ratio, wing tip shape and bacteria killing ability did not vary with migration status or distance. In general, male silver-haired bats do not appear to mediate migration costs by substantially downregulating immune defences or to be under stronger selection for wing forms adapted for fast, energy-efficient flight. Such phenotypic changes may be more adaptive for female silver-haired bats, which migrate farther and are more constrained by time in spring than males. Adaptations for aerial hawking and the use of heterothermy by migrating bats may also reduce the energetic cost of migration and the need for more substantial morphological and physiological trade-offs.

Sex-related interannual plasticity in wing morphological design in Heliconius charithonia enhances flight metabolic performance.

Flight morphological variations and its consequences on animal performance are common in winged insects. In the butterfly Heliconius charithonia, sex-related differences in the wing morphological design have been described resulting in differences in foraging behavior, daily flight distances and flight aerodynamics. It has been suggested that these differences should be reflected in the metabolic capacities and energetic budgets associated with flight in both sexes. In this study, we analyzed the relationship between wing morphological variation and metabolic performance, flight aerodynamics and energetic reserves in females and males of Heliconius charithonia over two years. The results confirm the presence of wing shape sexual dimorphism, but also show an unexpected sex-related annual variation in wing shape, mirrored in the metabolic condition (resting metabolic rate) of individuals. However, contrary to expectation, intersexual variations in wing shape are not related to differences between the sexes in terms of flight aerodynamics, flight metabolic rates, or energetic reserves (carbohydrates, lipids and proteins). Our results indicate a considerable plasticity in H. charithonia wing shape, which we suggest is determined by a trade-off between environmental pressures and reproductive restriction of each sex, maintaining an optimum flight design. Finally, similarities in metabolic rates between young and older males and females in both years may be a consequence of the ability of Heliconius species to feed on pollen.

Do Bee Wings Adapt for Flight in Urban Environments?

Understanding how organisms respond to urban-associated environmental changes is key to protecting vulnerable species. Bees, in particular, have gained interest due to their economic and ecological roles. We used a geometric morphometric approach to describe changes in wing shape and size in the solitary bee Andrena barbara (Barbara's Miner) collected across an urban landscape. We found that, although the wing morphology suggests a limited dispersal ability in its short and narrow frame, the urban landscape did not significantly explain how wing shape or size vary. Our findings are consistent with other studies that show little variation in wing morphology in urban solitary bees, and suggests that urban habitats may potentially serve an important role in bee conservation.

Wing morphology in migratory North American monarchs: characterizing sources of variation and understanding changes through time

Abstract Monarch butterfly wing morphology varies substantially throughout their global range, both between resident and migratory populations and also within the migratory North American population. Here, we use a dataset comprising more than 1800 North American individuals collected between 1878-2017 to characterize the factors shaping continent-wide patterns of wing morphological variation. North American overwintering butterflies have forewings that are approximately 4.4% larger than those collected in summer breeding areas. Monarchs overwintering in Mexico have forewings that are approximately 1.8% larger than monarchs overwintering in California, conducive to the idea that migration distance is positively correlated with wing area. We find evidence for a latitudinal cline within North America, such that butterflies collected at higher latitudes have significantly larger and more elongated forewings. We also find a significant increase of approximately 4.9% in forewing area between 1878-2017, but no difference through time in wing elongation. This result is corroborated by a reanalysis of a recently published dataset of more than 600 butterflies from Mexican overwintering sites. We discuss possible reasons for this increase in wing size through time, including northward shifts in the monarch’s breeding range and changes in relative abundance of milkweed hosts, and present experimental data addressing the influence of larval host plant on adult wing morphology. Our analysis suggests that (1) migration is indeed an important selective force for monarch wing morphology; (2) wing size has increased through time in North America; (3) factors such as host plant identity must be considered to fully understand monarch wing morphological variation.

Variation in the morphology of the wings of the endangered grass-feeding butterfly Coenonympha oedippus (Lepidoptera: Nymphalidae) in response to contrasting habitats

At the southern limit of its range the endangered butterfly Coenonympha oedippus inhabits grasslands (wet, dry) that differ significantly in the abundance of its larval hostplants (wet > dry) and mean annual air temperature (wet dry) or in degree, in the two sexes the relative area of eyespots on hindwing had opposite patterns (males: dry > wet; females: wet > dry) and was more pronounced in males than in females. The differences in wing shape between ecotypes were detected only in the hindwings of males, with more rounded apex in the dry than in the wet ecotype. We discuss the life-history traits, behavioural strategies and selection mechanisms, which largely account for the sex- and ecotype-specific variation in wing morphology.

Climate, isolation and intraspecific competition affect morphological traits in an endangered steppe bird, the Dupont’s Lark Chersophilus duponti

ABSTRACTCapsule: We assess biometric variation in the threatened Dupont’s Lark along a wide fraction of its distribution area analysing the largest data set for this bird species available to date, comprising a 28-year period and including birds captured in Spain, Morocco and Tunisia.Aims: To analyse Dupont’s Lark morphology evaluating five potential sources of divergence: (I) sexual dimorphism, (II) macro-regional differences, (III) climate, (IV) isolation and (V) intraspecific competition.Methods: Multivariate analysis was used to summarize biometric data. Sexual dimorphism and macro-regional divergence were assessed by generalized linear mixed models. Climate, isolation and intraspecific competition effects on phenotype were explored by means of model averaging.Results: Sexes differed in wing shape suggesting a sexual selection pressure on males for aerial displays. Males showed longer bills after controlling for body size. We found an increasing Spain < Morocco < Tunisia trend in body size and bill length and volume. Tunisian populations showed more pointed wings than Moroccan and Iberian ones. Maximum temperature increased bill size as predicted by Allen’s rule. We found support for Bergmann’s rule in relation to temperature in the breeding season and water availability. Intraspecific competition was associated with more rounded wing shapes suggesting a pressure related to breeding display performance. Isolation reduced wingtip pointedness.Conclusion: We found convincing results for wing morphology variation in relation to intrasexual competition, as males seem to be subjected to a sexual selection pressure for aerial display and this adaptation strengthens when intraspecific competition increases. We also found solid support for Allen’s rule, with results suggesting that bill size plays an important role in the bird’s thermoregulation, which does not occur in the case of tarsus. Although Bergmann’s rule is not supported in relation to minimum annual temperature, we found a relationship with thermal conditions in the breeding season, as well as water availability.

Wing morphology, winter ecology, and fecundity selection: evidence for sex-dependence in barn swallows (Hirundo rustica).

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 1year 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.

Geographic variation of wing morphology of great fruit-eating bats (Artibeus lituratus): environmental, genetic and spatial correlates of phenotypic differences

Responses of species to environmental gradients are important and frequent determinants of geographic phenotypic variation that can drive adaptive processes. Nonetheless, random genetic processes such as drift can also result in geographic variation in phenotypes, and should be evaluated before implicating selection as the process driving phenotypic change. We examined geographic variation in wing morphology of Artibeus lituratus among 18 different sites distributed across interior Atlantic Forest of Paraguay and Argentina. Moreover, we contrasted geographic variation with environmental, spatial, and genetic variation to test hypotheses related to selection and drift and their impacts on wing morphology. For A. lituratus distributed across interior Atlantic Forest, significant differences among sites characterized variation in wing morphology. Geographic variation was significantly related to climatic variables but not spatial or genetic distances. Such a pattern suggests that phenotypic variation is related to selection for particular environmental regimes, and not genetic drift. Four significant dimensions of phenotypic variation were determined. Three dimensions were related to variation among individuals in terms of wing tips, whereas one was related to overall body size. Wing tips are important for manoeuverability during flight and differences among sites likely reflect differences in forest and vegetation structure that must be managed during foraging. Although climate provides good surrogates for environmental variation, it is probably only an indirect cue of selection regimes that determine variation in wing morphology. Future studies should evaluate more direct environmental measures such as vegetation structure when attempting to interpret geographical variation in wing morphology.

Intra-specific variation in wing morphology and its impact on take-off performance in blue tits (Cyanistes caeruleus) during escape flights.

ABSTRACTDiurnal and seasonal increases in body mass and seasonal reductions in wing area may compromise a bird's ability to escape, as less of the power available from the flight muscles can be used to accelerate and elevate the animal's centre of mass. Here, we investigated the effects of intra-specific variation in wing morphology on escape take-off performance in blue tits (Cyanistes caeruleus). Flights were recorded using synchronised high-speed video cameras and take-off performance was quantified as the sum of the rates of change of the kinetic and potential energies of the centre of mass. Individuals with a lower wing loading, WL (WL=body weight/wing area) had higher escape take-off performance, consistent with the increase in lift production expected from relatively larger wings. Unexpectedly, it was found that the total power available from the flight muscles (estimated using an aerodynamic analysis) was inversely related to WL. This could simply be because birds with a higher WL have relatively smaller flight muscles. Alternatively or additionally, variation in the aerodynamic load on the wing resulting from differences in wing morphology will affect the mechanical performance of the flight muscles via effects on the muscle's length trajectory. Consistent with this hypothesis is the observation that wing beat frequency and relative downstroke duration increase with decreasing WL; both are factors that are expected to increase muscle power output. Understanding how wing morphology influences take-off performance gives insight into the potential risks associated with feather loss and seasonal and diurnal fluctuations in body mass.

Making quantitative morphological variation from basic developmental processes: Where are we? The case of the Drosophila wing.

One of the aims of evolutionary developmental biology is to discover the developmental origins of morphological variation. The discipline has mainly focused on qualitative morphological differences (e.g., presence or absence of a structure) between species. Studies addressing subtle, quantitative variation are less common. The Drosophila wing is a model for the study of development and evolution, making it suitable to investigate the developmental mechanisms underlying the subtle quantitative morphological variation observed in nature. Previous reviews have focused on the processes involved in wing differentiation, patterning and growth. Here, we investigate what is known about how the wing achieves its final shape, and what variation in development is capable of generating the variation in wing shape observed in nature. Three major developmental stages need to be considered: larval development, pupariation, and pupal development. The major cellular processes involved in the determination of tissue size and shape are cell proliferation, cell death, oriented cell division and oriented cell intercalation. We review how variation in temporal and spatial distribution of growth and transcription factors affects these cellular mechanisms, which in turn affects wing shape. We then discuss which aspects of the wing morphological variation are predictable on the basis of these mechanisms. Developmental Dynamics 244:1058-1073, 2015. © 2015 Wiley Periodicals, Inc.

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.

Resource partitioning by insectivorous bats in Jamaica.

In this investigation, we use variation in wing morphology, echolocation behaviour, patterns of habitat use and molecular diet analysis to demonstrate that six species of sympatric insectivorous bats in Jamaica show significant differences that could explain resource partitioning among the species. High-intensity echolocating species that used shorter, broadband signals and had shorter, broader wings (Pteronotus macleayii, Pteronotus quadridens, Mormoops blainvillii) foraged most in edge habitats, but differed in timing of peak activity. P. macleayii and M. blainvillii differed in diet, but low sample size precluded diet analysis for P. quadridens. High-intensity echolocating species that used longer, more narrowband signals and had longer, narrower wings (Molossus molossus, Tadarida brasiliensis) foraged most in open areas and differed in diet from the other species. Two disparate species were most active in clutter (dense vegetation). Pteronotus parnellii used high-duty-cycle echolocation apparently specialized for detecting fluttering targets in clutter. Macrotus waterhousii used low-intensity, broadband echolocation calls and presumably uses prey-generated sounds when foraging. These two species also differed in diet. Our data show that differences in morphology and echolocation behaviour coincide with differences in habitat use and diet, resulting in minimal overlap in resource use among species.

Morphological and Physiological Idiosyncrasies Lead to Interindividual Variation in Flight Metabolic Rate in Worker Bumblebees (Bombus impatiens)

Although intraspecific variation in metabolic rate is associated with variation in body size, similarly sized individuals nonetheless vary greatly. At similar masses, hovering bumblebee workers (Bombus impatiens) can differ in metabolic rate up to twofold. We examined how such interindividual variation arises by studying covariation of flight metabolic rate with morphological and other physiological parameters. Body size alone explained roughly half the variation in flight metabolic rate. The remaining variation could be explained as the outcome of variation in wing morphology and possibly an association with variation in flight muscle metabolic enzyme activities. As shown using statistical models, for a given mass, higher metabolic rate was correlated with both higher thoracic temperature and higher wing stroke frequency, in turn resulting from smaller wing surface area. When organismal and cellular metabolism for a given mass were linked, variation in metabolic rate was positively correlated with the activities of trehalase and hexokinase. Altogether, covariation with morphology and other physiological parameters explains up to 75% of the variation in metabolic rate. We also investigated the role of flight experience and show that neither flight restriction nor the number of lifetime flights affected flight energetics or flight muscle phenotype. Additionally, manipulating the level of wing asymmetry increased flight wing stroke frequency, metabolic rate, and thoracic temperature, but it did not alter enzyme activity. We conclude that idiosyncrasies in body morphology largely explained interindividual variation in flight metabolic rate but flight muscle metabolic phenotype shows little variation associated with differences in flight experience.

Population Structure of Migrating and Wintering Reed BuntingsEmberiza Schoeniclus Schoeniclusin Northern Spain

Iberia is a very important wintering area for several species of European seed-eating passerines, including the reed runting Emberiza schoeniclus. The dynamics and structure of its wintering population remains unknown over large areas, especially in northern Iberia. This study aimed to test (1) whether northern Iberia is a stop-over region rather than a wintering area for non-breeding nominate race reed buntings; (2) whether sex and age ratios indicate a higher proportion of immature males and whether such ratios differ from those found in regions further south; (3) whether wing morphology varies at different stages of the non-breeding season, thus possibly revealing differential migration by different populations. Reed buntings were mist-netted weekly from October to April in a reedbed in the Ebro Valley, northern Iberia, between 2003 and 2009. The zone was used mainly by reed buntings stoppingover in this area during the autumn migration period but also, to a lesser extent, by an apparently stable population that overwintered in this zone. The high proportion of females supports the hypothesis that Iberia plays a major role as a destination area for female reed buntings during the non-breeding period. Seasonal variation in wing morphology suggests that populations of very distinct origin occur but the temporal patterns of their occurrence still remain unresolved. Our results suggest that longwinged reed buntings, which presumably have travelled from more distant regions, are more abundant in mid-winter.

Getting real with real-time qPCR: a case study of reference gene selection for morphological variation in Drosophila melanogaster wings

Accurate estimation of gene expression differences during development requires sensitive techniques combined with gold-standard normalization procedures. This is particularly true in the case of quantitative traits, where expression changes might be small. Nevertheless, systematic selection and validation of reference genes has been overlooked, even in Drosophila studies. Here, we tested the stability of six traditional reference genes across samples of imaginal wing disks from morphologically divergent strains of Drosophila melanogaster, in a two-class comparison: quantitative or qualitative variation in wing morphology. Overall, we identified and validated a pair of genes (RpL32 and Tbp) as being stably expressed in both experimental comparisons. These genes might be considered as a bona fide pair of reference genes for gene expression analyses of morphological divergence in D. melanogaster wings. They might also be taken as good candidates for experimental identification of stable reference genes in other morphological comparisons using Drosophila or other insect species. Besides, we found that some genes traditionally used as reference in qPCR experiments were not stably expressed in wing disks from the different fly strains. In fact, a significant bias was observed when the expression of three genes of interest, which are involved in the regulation of growth and patterning during imaginal wing development, was normalized with such putative reference genes. Our results demonstrate how inaccurate findings and opposite conclusions might be drawn if traditional reference genes are arbitrarily used for internal normalization without proper validation in the given experimental condition, a practice still common in qPCR experiments.

Wing shape variations in an invasive moth are related to sexual dimorphism and altitude

Wing morphology has great importance in a wide variety of aspects of an insect's life. Here, we use a geometric morphometric approach to test the hypothesis that variation, in insect wing morphology patterns, occurs between sexes and along altitudinal gradients for invasive species, despite their recent association to this environment. We explored the variation in wing morphology between 12 invasive populations of the invasive potato pest, Tecia solanivora, at low and high altitude in the central highlands of Ecuador. After characterizing sexual dimorphism in wing shape, we investigated if moths at higher elevations differ in wing morphology from populations at lower altitudes. Results indicate wing shape and size differences between sexes and between altitudinal ranges. Females showed larger, wider wings than males, while high altitude moths showed larger, narrow-shaped wings by comparison to low-altitude moths. GLM analyses confirmed altitude was the only significant determinant of this gradient. Our study confirms a sexual dimorphism in size and wing shape for the potato moth. It also confirms and extends predictions of morphological changes with altitude to an invasive species, suggesting that wing morphology variation is an adapted response contributing to invasion success of the potato moth in mountainous landscapes. Ours is one of the first studies on the morphology of invasive insects and represents a valuable contribution to the study of insect invasions because it both offers empirical support to previous genetic studies on T. solanivora as well as proving broader insight into the mechanisms behind morphological evolution of a recently introduced pest.

Wing shape variation in the medium ground finch (Geospiza fortis): an ecomorphological approach

Wing design in birds is subject to a suite of interacting selective pressures. As different performance traits are favoured in different ecological settings, a tight link is generally expected between variation in wing morphology and variation in ecological parameters. In the present study, we document aspects of variation in wing morphology in the medium ground finch (Geospiza fortis) on Isla Santa Cruz in the Galapagos. We compare variation in body size, simple morphometric traits (body mass, last primary length, wing length, wing chord, and wing area) and functional traits (wing loading, aspect ratio and wing pointedness) across years, among populations, and between sexes. Functional traits are found to covary across years with differences in climatic conditions, and to covary among populations with differences in habitat structure. In dry years and arid locations, wing aspect ratios are highest and wings are more pointed, consistent with a need for a low cost of transport. In wet years and cluttered habitats, wing loading is lowest and wings are more rounded, suggesting enhanced capabilities for manoeuvrability. Sexes differ in wing loading, with males having lower wing loadings than females. Superior manoeverability might be favoured in males for efficient territory maintenance. Lastly, in contrast to functional traits, we found little consistent inter-annual or inter-site variation in simple morphometric traits. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98, 129–138.

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

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.