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Jeffrey Podos - One of the best experts on this subject based on the ideXlab platform.

  • persistence of song types in darwin s finches Geospiza fortis over four
    2015
    Co-Authors: Eben Goodale, Jeffrey Podos
    Abstract:

    Department of Biology, Graduate Program in Organismic andEvolutionary Biology, University of Massachusetts, Amherst,MA 01003, USA*Author and address for correspondence: Section of Ecology, Behavior andEvolution, Division of Biological Sciences, University of California, SanDiego, 9500 Gilman Drive, La Jolla, CA 92093, USA (eben.goodale@gmail.com).

  • The head of the finch: the anatomy of the feeding system in two species of finches (Geospiza fortis and Padda oryzivora)
    Journal of Anatomy, 2011
    Co-Authors: Annelies Genbrugge, Jeffrey Podos, Luc Van Hoorebeke, Joris J.j. Dirckx, Matthieu Boone, Anthony Herrel, Peter Aerts, Adriaens Dominique
    Abstract:

    Despite the large number of studies devoted to the evolution of beak shape in Darwin's finches, surprisingly little is known about the morphology of the skull and jaw musculature in these birds. Moreover, it remains currently unclear whether Darwin's finches are unusual in their cranial morphology compared with other seed-cracking birds. Here, we provide a detailed description of the morphology of the cranial system in the medium ground finch (Geospiza fortis) and compare it with that of another seed-cracking bird of similar overall size and appearance, the Java finch (Padda oryzivora). Our data show an overall similarity in beak size and cranial morphology. Yet, differences in the jaw adductor size and corresponding attachments to the cranium and mandible are prominent, with the medium ground finch having much more robust jaw-closing muscles. This is reflected in differences in bite forces, with the medium ground finch biting much harder than the Java finch. These data suggest similarities in the evolution of the feeding system in birds specializing in the cracking of hard seeds, but also show the uniqueness of the cranial morphology and bite force of the medium ground finch compared with other seed-cracking birds.

  • ontogeny of the cranial skeleton in a darwin s finch Geospiza fortis
    Journal of Anatomy, 2011
    Co-Authors: Annelies Genbrugge, Luc Van Hoorebeke, Joris J.j. Dirckx, Matthieu Boone, Peter Aerts, Annesophie Heyde, Dominique Adriaens, Jeffrey Podos
    Abstract:

    Darwin's finches are a model system in ecological and evolutionary research, but surprisingly little is known about their skull morphology and development. Indeed, only the early beak development and external variation in adult beak shape has been studied. Understanding the development of the skull from embryo up to the adult is important to gain insights into how selection acts upon, and drives, variation in beak shape. Here, we provide a detailed description of the skeletal development of the skull in the medium ground finch (Geospiza fortis). Although the ossification sequence of the cranial elements is broadly similar to that observed for other birds, some differences can be observed. Unexpectedly, our data show that large changes in skull shape take place between the nestling and the juvenile phases. The reorientation of the beak, the orbit and the formation of well-developed processes and cristae suggest that these changes are likely related to the use of the beak after leaving the nest. This suggests that the active use of the jaw muscles during seed cracking plays an important role in shaping the adult skull morphology and may be driving some of the intra-specific variation observed in species such as G. fortis. Investigating the development of the jaw muscles and their interaction with the observed ossification and formation of the skull and lower jaw would allow further insights into the ecology and evolution of beak morphology in Darwin's finches.

  • persistence of song types in darwin s finches Geospiza fortis over four decades
    Biology Letters, 2010
    Co-Authors: Eben Goodale, Jeffrey Podos
    Abstract:

    Learned bird songs evolve via cultural evolution, with song patterns transmitted across generations by imitative learning. In Darwin9s finches of the Galapagos Islands, males learn songs from their fathers, and song types can be maintained across multiple generations. However, little is known about the time frame over which specific song types are preserved, in the face of copy errors and corresponding modifications to song structure. Here we investigate cultural evolution in songs of male Geospiza fortis , at Academy Bay, Santa Cruz Island, comparing songs recorded in 1961 by R. Bowman (20 individuals) to those recorded in 1999 by J. Podos (16 individuals). For each individual, we characterized four timing and six frequency parameters, and assessed inter-individual variation in song structure using multivariate analysis. Several 1961 song types persisted into 1999, some with remarkable fidelity. Variation among song types was extensive during both years, and we detected no changes in 10 vocal parameters across the sampling period. These results illustrate temporal continuity in a culturally acquired trait, and raise questions about mechanisms that promote stability in song structure.

  • Divergence with gene flow as facilitated by ecological differences: within-island variation in Darwin's finches
    Philosophical Transactions of the Royal Society B, 2010
    Co-Authors: Luis F. De León, Jeffrey Podos, Eldredge Bermingham, Andrew P. Hendry
    Abstract:

    Divergence and speciation can sometimes proceed in the face of, and even be enhanced by, ongoing gene flow. We here study divergence with gene flow in Darwin's finches, focusing on the role of eco- logical/adaptive differences in maintaining/promoting divergence and reproductive isolation. To this end, we survey allelic variation at 10 microsatellite loci for 989 medium ground finches (Geospiza fortis) on Santa Cruz Island, Galapagos. We find only small genetic differences among G. fortis from different sites. We instead find noteworthy genetic differences associated with beak. Moreover, G. fortis at the site with the greatest divergence in beak size also showed the greatest divergence at neutral markers; i.e. the lowest gene flow. Finally, morphological and genetic differentiation between the G. fortis beak-size morphs was intermediate to that between G. fortis and its smaller (Geospiza fuliginosa) and larger (Geospiza magnirostris) congeners. We conclude that ecological differences associated with beak size (i.e. foraging) influence patterns of gene flow within G. fortis on a single island, providing additional support for ecological speciation in the face of gene flow. Patterns of genetic similarity within and between species also suggest that interspecific hybridization might contribute to the formation of beak-size morphs within G. fortis.

Peter R Grant - One of the best experts on this subject based on the ideXlab platform.

  • Triad hybridization via a conduit species
    Proceedings of the National Academy of Sciences of the United States of America, 2020
    Co-Authors: Peter R Grant, B. Rosemary Grant
    Abstract:

    Introgressive hybridization can affect the evolution of populations in several important ways. It may retard or reverse divergence of species, enable the development of novel traits, enhance the potential for future evolution by elevating levels of standing variation, create new species, and alleviate inbreeding depression in small populations. Most of what is known of contemporary hybridization in nature comes from the study of pairs of species, either coexisting in the same habitat or distributed parapatrically and separated by a hybrid zone. More rarely, three species form an interbreeding complex (triad), reported in vertebrates, insects, and plants. Often, one species acts as a genetic link or conduit for the passage of genes (alleles) between two others that rarely, if ever, hybridize. Demographic and genetic consequences are unknown. Here we report results of a long-term study of interbreeding Darwin’s finches on Daphne Major island, Galapagos. Geospiza fortis acted as a conduit for the passage of genes between two others that have never been observed to interbreed on Daphne: Geospiza fuliginosa, a rare immigrant, and Geospiza scandens, a resident. Microsatellite gene flow from G. fortis into G. scandens increased in frequency during 30 y of favorable ecological conditions, resulting in genetic and morphological convergence. G. fortis, G. scandens, and the derived dihybrids and trihybrids experienced approximately equal fitness. Especially relevant to young adaptive radiations, where species differ principally in ecology and behavior, these findings illustrate how new combinations of genes created by hybridization among three species can enhance the potential for evolutionary change.

  • Hybridization increases population variation during adaptive radiation
    Proceedings of the National Academy of Sciences of the United States of America, 2019
    Co-Authors: Peter R Grant, B. Rosemary Grant
    Abstract:

    Adaptive radiations are prominent components of the world’s biodiversity. They comprise many species derived from one or a small number of ancestral species in a geologically short time that have diversified into a variety of ecological niches. Several authors have proposed that introgressive hybridization has been important in the generation of new morphologies and even new species, but how that happens throughout evolutionary history is not known. Interspecific gene exchange is expected to have greatest impact on variation if it occurs after species have diverged genetically and phenotypically but before genetic incompatibilities arise. We use a dated phylogeny to infer that populations of Darwin’s finches in the Galapagos became more variable in morphological traits through time, consistent with the hybridization hypothesis, and then declined in variation after reaching a peak. Some species vary substantially more than others. Phylogenetic inferences of hybridization are supported by field observations of contemporary hybridization. Morphological effects of hybridization have been investigated on the small island of Daphne Major by documenting changes in hybridizing populations of Geospiza fortis and Geospiza scandens over a 30-y period. G. scandens showed more evidence of admixture than G. fortis. Beaks of G. scandens became progressively blunter, and while variation in length increased, variation in depth decreased. These changes imply independent effects of introgression on 2, genetically correlated, beak dimensions. Our study shows how introgressive hybridization can alter ecologically important traits, increase morphological variation as a radiation proceeds, and enhance the potential for future evolution in changing environments.

  • Adult sex ratio influences mate choice in Darwin’s finches
    Proceedings of the National Academy of Sciences of the United States of America, 2019
    Co-Authors: Peter R Grant, B. Rosemary Grant
    Abstract:

    The adult sex ratio (ASR) is an important property of populations. Comparative phylogenetic analyses have shown that unequal sex ratios are associated with the frequency of changing mates, extrapair mating (EPM), mating system and parental care, sex-specific survival, and population dynamics. Comparative demographic analyses are needed to validate the inferences, and to identify the causes and consequences of sex ratio inequalities in changing environments. We tested expected consequences of biased sex ratios in two species of Darwin’s finches in the Galapagos, where annual variation in rainfall, food supply, and survival is pronounced. Environmental perturbations cause sex ratios to become strongly male-biased, and when this happens, females have increased opportunities to choose high-quality males. The choice of a mate is influenced by early experience of parental morphology (sexual imprinting), and since morphological traits are highly heritable, mate choice is expressed as a positive correlation between mates. The expected assortative mating was demonstrated when the Geospiza scandens population was strongly male-biased, and not present in the contemporary Geospiza fortis population with an equal sex ratio. Initial effects of parental imprinting were subsequently overridden by other factors when females changed mates, some repeatedly. Females of both species were more frequently polyandrous in male-biased populations, and fledged more offspring by changing mates. The ASR ratio indirectly affected the frequency of EPM (and hybridization), but this did not lead to social mate choice. The study provides a strong demonstration of how mating patterns change when environmental fluctuations lead to altered sex ratios through differential mortality.

  • Rapid hybrid speciation in Darwin's finches.
    Science, 2017
    Co-Authors: Sangeet Lamichhaney, B. Rosemary Grant, Matthew T. Webster, Leif Andersson, Peter R Grant
    Abstract:

    Homoploid hybrid speciation in animals has been inferred frequently from patterns of variation, but few examples have withstood critical scrutiny. Here we report a directly documented example, from its origin to reproductive isolation. An immigrant Darwin’s finch to Daphne Major in the Galapagos archipelago initiated a new genetic lineage by breeding with a resident finch ( Geospiza fortis ). Genome sequencing of the immigrant identified it as a G. conirostris male that originated on Espanola >100 kilometers from Daphne Major. From the second generation onward, the lineage bred endogamously and, despite intense inbreeding, was ecologically successful and showed transgressive segregation of bill morphology. This example shows that reproductive isolation, which typically develops over hundreds of generations, can be established in only three.

  • Introgressive hybridization and natural selection in Darwin's finches
    Biological Journal of The Linnean Society, 2015
    Co-Authors: Peter R Grant, B. Rosemary Grant
    Abstract:

    Introgressive hybridization, i.e. hybridization with backcrossing, can lead to the fusion of two species, but it can also lead to evolution of a new trajectory through an enhancement of genetic variation in a new or changed ecological environment. On Daphne Major Island in the Galapagos archipelago, ~1–2% of Geospiza fortis finches breed with the resident G. scandens and with the rare immigrant species G. fuliginosa in each breeding season. Previous research has demonstrated morphological convergence of G. fortis and G. scandens over a 30-year period as a result of bidirectional introgression. Here we examine the role of hybridization with G. fuliginosa in the evolutionary trajectory of G. fortis. Geospiza fuliginosa (~12 g) is smaller and has a more pointed beak than G. fortis (~17 g). Genetic variation of the G. fortis population was increased by receiving genes more frequently from G. fuliginosa than from G. scandens (~21 g). A severe drought in 2003–2005 resulted in heavy and selective mortality of G. fortis with large beaks, and they became almost indistinguishable morphologically from G. fuliginosa. This was followed by continuing hybridization, a further decrease in beak size and a potential morphological fusion of the two species under entirely natural conditions.

Bieke Vanhooydonck - One of the best experts on this subject based on the ideXlab platform.

  • wing shape variation in the medium ground finch Geospiza fortis an ecomorphological approach
    Biological Journal of The Linnean Society, 2009
    Co-Authors: Bieke Vanhooydonck, Ana Gabela, Anthony Herrel, Jeffrey Podos
    Abstract:

    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.

  • Wing shape variation in the medium ground finch (Geospiza fortis): An ecomorphological approach
    Biological Journal of the Linnean Society, 2009
    Co-Authors: Bieke Vanhooydonck, Ana Gabela, Anthony Herrel, Jeffrey Podos
    Abstract:

    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 Galápagos. 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.

Anthony Herrel - One of the best experts on this subject based on the ideXlab platform.

  • Comparative cranial osteology of Darwin's finches based on micro‐CT scanning: preliminary results
    2020
    Co-Authors: Annelies Genbrugge, Luc Van Hoorebeke, Joris J.j. Dirckx, Matthieu Boone, Anthony Herrel, Peter Aerts, Dominique Adriaens
    Abstract:

    Darwin’s finches are the textbook example of adaptive evolution. All fourteen species are considered having evolved a different beak morphology corresponding to their food preferences. Unfortunately, little is known about the details on the relation between the beak design and its performance. In continuation of the work of Bowman [1], we aim to unravel this causal link by comparing the osteological variation in beak design of the different Darwin’s finches with that of a model organism, the Java sparrow (Padda oryzivora, a large‐beaked species not closely related to Darwin’s finches), and link this to mechanical capacities of the upper beak for dealing with mechanical stress. It is hypothesized that the beak morphology reflects the performance in seed cracking. Based on micro‐CT scanning of 12 different species, including species with long, pointed beaks (e.g., Certhidea olivacea and Camarhynchus pallidus) to species with deep, broad beaks (e.g., Geospiza magnirostris and Padda oryzivora), three‐dimensional models of the beak have been made. With these models it was possible to study and compare the cranial skeletal elements to evaluate the adaptive nature of the observed skeletal variation with respect to dealing with hard seeds.

  • The head of the finch: the anatomy of the feeding system in two species of finches (Geospiza fortis and Padda oryzivora)
    Journal of Anatomy, 2011
    Co-Authors: Annelies Genbrugge, Jeffrey Podos, Luc Van Hoorebeke, Joris J.j. Dirckx, Matthieu Boone, Anthony Herrel, Peter Aerts, Adriaens Dominique
    Abstract:

    Despite the large number of studies devoted to the evolution of beak shape in Darwin's finches, surprisingly little is known about the morphology of the skull and jaw musculature in these birds. Moreover, it remains currently unclear whether Darwin's finches are unusual in their cranial morphology compared with other seed-cracking birds. Here, we provide a detailed description of the morphology of the cranial system in the medium ground finch (Geospiza fortis) and compare it with that of another seed-cracking bird of similar overall size and appearance, the Java finch (Padda oryzivora). Our data show an overall similarity in beak size and cranial morphology. Yet, differences in the jaw adductor size and corresponding attachments to the cranium and mandible are prominent, with the medium ground finch having much more robust jaw-closing muscles. This is reflected in differences in bite forces, with the medium ground finch biting much harder than the Java finch. These data suggest similarities in the evolution of the feeding system in birds specializing in the cracking of hard seeds, but also show the uniqueness of the cranial morphology and bite force of the medium ground finch compared with other seed-cracking birds.

  • mechanical stress fracture risk and beak evolution in darwin s ground finches Geospiza
    Philosophical Transactions of the Royal Society B, 2010
    Co-Authors: Joris Soons, Annelies Genbrugge, Jeffrey Podos, Anthony Herrel, Peter Aerts, Dominique Adriaens, Yoni De Witte, Patric Jacobs
    Abstract:

    Darwin's finches have radiated from a common ancestor into 14 descendent species, each specializing on distinct food resources and evolving divergent beak forms. Beak morphology in the ground finches (Geospiza) has been shown to evolve via natural selection in response to variation in food type, food availability and interspecific competition for food. From a mechanical perspective, however, beak size and shape are only indirectly related to birds' abilities to crack seeds, and beak form is hypothesized to evolve mainly under selection for fracture avoidance. Here, we test the fracture-avoidance hypothesis using finite-element modelling. We find that across species, mechanical loading is similar and approaches reported values of bone strength, thus suggesting pervasive selection on fracture avoidance. Additionally, deep and wide beaks are better suited for dissipating stress than are more elongate beaks when scaled to common sizes and loadings. Our results illustrate that deep and wide beaks in ground finches enable reduction of areas with high stress and peak stress magnitudes, allowing birds to crack hard seeds while limiting the risk of beak failure. These results may explain strong selection on beak depth and width in natural populations of Darwin's finches.

  • Scaling and shear transformations capture beak shape variation in Darwin’s finches
    Proceedings of the National Academy of Sciences of the United States of America, 2010
    Co-Authors: Otger Campàs, Anthony Herrel, Ricardo Mallarino, Arkhat Abzhanov, Michael P. Brenner
    Abstract:

    Evolution by natural selection has resulted in a remarkable diversity of organism morphologies that has long fascinated scientists and served to establish the first relations among species. Despite the essential role of morphology as a phenotype of species, there is not yet a formal, mathematical scheme to quantify morphological phenotype and relate it to both the genotype and the underlying developmental genetics. Herein we demonstrate that the morphological diversity in the beaks of Darwin’s Finches is quantitatively accounted for by the mathematical group of affine transformations. Specifically, we show that all beak shapes of Ground Finches (genus Geospiza) are related by scaling transformations (a subgroup of the affine group), and the same relationship holds true for all the beak shapes of Tree, Cocos, and Warbler Finches (three distinct genera). This analysis shows that the beak shapes within each of these groups differ only by their scales, such as length and depth, which are genetically controlled by Bmp4 and Calmodulin. By measuring Bmp4 expression in the beak primordia of the species in the genus Geospiza, we provide a quantitative map between beak morphology and the expression levels of Bmp4. The complete morphological variation within the beaks of Darwin’s finches can be explained by extending the scaling transformations to the entire affine group, by including shear transformations. Altogether our results suggest that the mathematical theory of groups can help decode morphological variation, and points to a potentially hierarchical structure of morphological diversity and the underlying developmental processes.

  • wing shape variation in the medium ground finch Geospiza fortis an ecomorphological approach
    Biological Journal of The Linnean Society, 2009
    Co-Authors: Bieke Vanhooydonck, Ana Gabela, Anthony Herrel, Jeffrey Podos
    Abstract:

    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.

Ana Gabela - One of the best experts on this subject based on the ideXlab platform.

  • wing shape variation in the medium ground finch Geospiza fortis an ecomorphological approach
    Biological Journal of The Linnean Society, 2009
    Co-Authors: Bieke Vanhooydonck, Ana Gabela, Anthony Herrel, Jeffrey Podos
    Abstract:

    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.

  • Wing shape variation in the medium ground finch (Geospiza fortis): An ecomorphological approach
    Biological Journal of the Linnean Society, 2009
    Co-Authors: Bieke Vanhooydonck, Ana Gabela, Anthony Herrel, Jeffrey Podos
    Abstract:

    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 Galápagos. 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.