Ecomorphological variation and sexual dimorphism in a recent radiation of dwarf chameleons (Bradypodion)

Natural selection tends to favour optimal phenotypes either through directional or stabilizing selection; however, phenotypic variation in natural populations is common and arises from a combination of biotic and abiotic interactions. In these instances, rare phenotypes may possess a fitness advantage over the more common phenotypes in particular environments, which can lead to adaptation and ecological speciation. A recently radiated clade of dwarf chameleons (Bradypodion) restricted to southern KwaZulu‐Natal Province, South Africa, is currently comprised of two species (Bradypodion melanocephalum and Bradypodion thamnobates), yet three other phenotypic forms exist, possibly indicating the clade is far more speciose. Very little genetic differentiation exists between these five phenotypic forms; however, all are allopatric in distribution, occupy different habitats and vary in overall size and coloration, which may indicate that these forms are adapting to their local environments and possibly undergoing ecological speciation. To test this, we collected morphometric and habitat data from each form and examined whether ecological relevant morphological differences exist between them that reflect their differential habitat use. Sexual dimorphism was detected in four of the five forms. Yet, the degree and number of dimorphic characters was different between them, with size‐adjusted male‐biased dimorphism being much more pronounced in B. thamnobates. Habitat differences also existed between sexes, with males occupying higher perches in more closed canopy (forested) habitats than females. Clear morphological distinctions were detected between four of the five forms, with the head explaining the vast majority of the variation. Chameleons occupying forested habitats tended to possess proportionally larger heads and feet but shorter limbs than those in open canopy habitats (i.e. grassland). These results show that this species complex of Bradypodion is morphologically variable for traits that are ecologically relevant for chameleons, and that the variation among the five phenotypic forms is associated with habitat type, suggesting that this species complex is in the early stages of ecological speciation. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 113–130.     

Sexual Dimorphism in Bite Performance Drives Morphological Variation in Chameleons

Phenotypic performance in different environments is central to understanding the evolutionary and ecological processes that drive adaptive divergence and, ultimately, speciation. Because habitat structure can affect an animal’s foraging behaviour, anti-predator defences, and communication behaviour, it can influence both natural and sexual selection pressures. These selective pressures, in turn, act upon morphological traits to maximize an animal’s performance. For performance traits involved in both social and ecological activities, such as bite force, natural and sexual selection often interact in complex ways, providing an opportunity to understand the adaptive significance of morphological variation with respect to habitat. Dwarf chameleons within the Bradypodion melanocephalum-Bradypodion thamnobates species complex have multiple phenotypic forms, each with a specific head morphology that could reflect its use of either open- or closed-canopy habitats. To determine whether these morphological differences represent adaptations to their habitats, we tested for differences in both absolute and relative bite performance. Only absolute differences were found between forms, with the closed-canopy forms biting harder than their open-canopy counterparts. In contrast, sexual dimorphism was found for both absolute and relative bite force, but the relative differences were limited to the closed-canopy forms. These results indicate that both natural and sexual selection are acting within both habitat types, but to varying degrees. Sexual selection seems to be the predominant force within the closed-canopy habitats, which are more protected from aerial predators, enabling chameleons to invest more in ornamentation for communication. In contrast, natural selection is likely to be the predominant force in the open-canopy habitats, inhibiting the development of conspicuous secondary sexual characteristics and, ultimately, enforcing their overall diminutive body size and constraining performance.     

Patterns of morphological variation and correlates of habitat use in Chameleons

The lizard family Chameleonidae is one of the most distinctive taxa of all vertebrates. Nonetheless, despite great intrafamilial diversity, little research has been conducted on morphological variation among chameleons. As a first step in this direction, we took morphological measurements on the head, limbs, spines, and tail of 56 species. Our goals were to investigate whether morphological variation correlates with differences in ecology and to examine whether correlations exist among different aspects of morphology. Based on existing information, species were classified either as arboreal or terrestrial, the latter referring to species that are known to use the ground on a regular basis. This study confirms that considerable intrafamilial variation exists among chameleons and that these traits appear to be evolutionarily quite labile. Once the effects of size are removed, functionally related traits tend to covary; however, few correlations are observed between non-functionally related traits. Many differences in the lengths of the limbs and head elements were detected between terrestrial and arboreal species, but the functional and selective significance of these differences is not clear. Further research on chameleon behaviour and ecology is required to understand the factors contributing to chameleon morphological diversity. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 91–103.     

Morphological variation in the Cape Dwarf Chameleon (Bradypodion pumilum) as a consequence of spatially explicit habitat structure differences

An organism's phenotype is to some extent influenced by costs and benefits in terms of natural and sexual selection. The intensity of natural selection can in part be driven by habitat structure, which may result in varying levels of crypsis and/or selection on traits related to maximizing performance in that habitat. This may be countered by sexual selection, which can lead to sexual dimorphism in body size and/or the expression of conspicuous ornamentation relating to maximizing reproductive success. The intensity of these forces can also be different between the sexes, resulting in complex patterns of phenotypic variation. With this in mind, we examined morphological variation within the Cape Dwarf Chameleon, Bradypodion pumilum. The species inhabits two geographically disjunct habitat types and, in the present study, we demonstrate that chameleons from the two habitats show morphological differences. Large, conspicuous individuals inhabit closed vegetation, whereas small, drab individuals inhabit open vegetation. However, when morphological traits are size‐adjusted, the open vegetation morph displays many traits that are larger for its body size than the closed vegetation morph, especially for characters related to locomotion (limbs) and bite force (head width). Sexual dimorphism is also present, although the degree and number of dimorphic characters was very different between the two morphs, with size‐adjusted male‐biased dimorphism much more pronounced in the closed morph. Overall, our findings suggest that natural selection in open habitats limits both body size and conspicuous characters, although sexual selection in closed habitats favours the development of ornamentation related to display. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 878–888.     

Functional divergence between morphs of a dwarf chameleon: differential locomotor kinematics in relation to habitat structure

Arboreal lizards are extremely effective at moving in structurally complex habitats, including surfaces of varying diameter and incline. Chameleons exemplify this by exhibiting a number of morphological specializations for moving in these habitats, including the use of prehensile feet and tail to grasp branches. Despite their unique morphology and behaviour, little is known about how locomotor movements vary between species. In addition, some species, such as the Cape Dwarf Chameleon, Bradypodion pumilum, consist of two morphs that differ in ecology, morphology, and behaviour. The two morphs can be found in either closed canopy woodland habitat or relatively open fynbos habitat. The morph that occupies the woodland habitat tends to be larger and utilizes larger diameter perches. Although their ecological and morphological divergence is established, whether this translates into differences in three‐dimensional kinematics of locomotion is not known. Given the potentially strong selective pressures from structurally different habitats, kinematic differences might reveal the functional basis of incipient speciation. We determined that the two morphs diverge significantly in multidimensional kinematic space, and that this occurs for the forelimb and hindlimb independently. These differences outweigh the effects of substrate within each morph, although the differences between morphs were more pronounced on the vertical treatments.     

Functional morphology of the tetra fish Astyanax lacustris differs between divergent habitats in the Pantanal wetlands

This study investigated whether the body morphology of the tetra fish Astyanax lacustris (previously Astyanax asuncionensis) varied between populations inhabiting one lagoon (a lentic, shallow environment, with great habitat complexity created by aquatic macrophytes) and an adjacent river (a deeper, lotic environment where aquatic macrophytes are scarce) in a seasonally flooded wetland, despite population mixing during the wet season. Morphological differences matched a priori predictions of the theory relating functional body morphology and swimming performance in fishes between lagoon and river habitats. Observed morphological variation could have resulted from adaptive habitat choice by tetras, predation by piscivores and adaptive phenotypic plasticity during development.     

Parallels,nonparallels, and plasticity in population differentiation of threespine stickleback within a lake

The frequent occurrence of parallel phenotypic divergence in similar habitats is often evoked when emphasizing the role of ecology in adaptive radiation and speciation. However, because phenotypic plasticity can contribute to the observed pattern of divergence, confirmation of divergence at loci underlying phenotypic traits is important for confirming adaptive divergence. In the present study, we examine parallel morphological, neutral, and potentially adaptive genetic divergence of threespine stickleback inhabiting different habitats within a lake. Three genetic clusters best explained the neutral genetic structure within the lake; however, morphological differences were only weakly connected to genetic clusters and there was considerable phenotypic variation within clusters. Among the factors that could contribute to the observed pattern of morphological and genetic divergence are phenotypic plasticity, selective mortality of hybrids, and habitat choice based on morphology. Several loci are identified as outliers indicating divergent selection between the morphs and some parallels in morphological and adaptive genetic divergence are found in stickleback spawning at two lava sites. However, neutral genetic structure indicates considerable genetic connectivity among the two lava sites, and the parallels in morphology may therefore represent selective distribution of phenotypes rather than parallel divergence. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 803–813.     

The evolution of locomotor morphology, performance, and anti-predator behaviour among populations of Leiocephalus lizards from the Dominican Republic

Most ecomorphological studies use a comparative approach to examine adaptation by studying variation among species. A question of considerable interest is whether ecomorphological patterns observed among species also exist at the population level. We studied variation in morphology, performance, and behaviour in four populations of Leiocephalus personatus and two populations of Leiocephalus barahonensis in the Dominican Republic. We combined these data with measurements of predation intensity and habitat structure to test for convergence at the population level. We predicted that predation intensity would be higher in open habitats and that lizards in these habitats would have traits conferring higher predator evasion capacity (increased wariness, faster sprint speeds, and longer limbs). Principal components analysis suggests that sites tend to differ with respect to the abundance and spacing of low-lying vegetation (i.e. percentage of shrub cover and distance to nearest vegetation), but we did not detect any striking differences among sites in tail-break frequencies or attacks on clay lizard models. Consistent with predictions we find that in open habitats, lizards tend to have longer limbs, faster sprint speeds (relative to body size), and longer approach distances. These patterns corroborate findings in other ground-dwelling lizard species and indicate that they have evolved at least twice among populations of Leiocephalus lizards. The results of this study also suggest that these traits have evolved rapidly despite recent or ongoing gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 445–456.     

Secondary invasion of <Emphasis Type="Italic">Acer negundo</Emphasis>: the role of phenotypic responses versus local adaptation

During plant species invasions, the role of adaptive processes is particularly of interest in later stages of range expansion when populations start invading habitats that initially have not been disposed to invasions. The dioecious tree Acer negundo, primarily invasive in Europe in wet habitats along riversides and in floodplains, has increased its abundance in dry habitats of industrial wasteland and ruderal sites during the last decades in Eastern Germany. We chose 21 invasive populations from wet and from dry habitats in the region of Halle, Saxony-Anhalt, Germany, to test whether Acer negundo exhibits a shift in life-history strategy during expansion into more stressful habitats. We analyzed variables of habitat quality (pH, soil moisture, exchangeable cations, total C and N content) and determined density, sex ratio and regeneration of the populations. In addition, we conducted germination experiments and greenhouse studies with seedlings in four different soil moisture environments. Local adaptation was studied in a reciprocal transplant experiment. We found habitat type differentiation with lower nutrient and water supply at the dry sites than at the moist sites and significant differences in the number of seedlings in the field. In accordance, seeds from moist habitats responded significantly faster to germination treatments. In the transplant experiment, leaf life span was significantly larger for populations originating from dry habitat types than from moist habitats. This observed shift in life history strategy during secondary invasion of A. negundo from traits of establishment and rapid growth towards traits connected with persistence might be counteracted by high gene flow among populations of the different habitat types. However, prolonged leaf life span at dry sites contributed remarkably to the invasion of less favourable habitats, and, thus, is a first indication of ongoing adaptation.     

The effects of age,sex, and habitat on body size and shape of the blackstripe topminnow,Fundulus notatus (Cyprinodontiformes: Fundulidae) (Rafinesque 1820)

Lake and stream habitats pose a variety of challenges to fishes due to differences in variables such as water velocity, habitat structure, prey community, and predator community. These differences can cause divergent selection on body size and/or shape. Here, we measured sex, age, length, and eight different morphological traits of the blackstripe topminnow, Fundulus notatus, from 19 lake and stream populations across four river drainages in central Illinois. Our goal was to determine whether size and shape differed consistently between lake and stream habitats across drainages. We also considered the effects of age and sex as they may affect size and morphology. We found large differences in body size of age 1 topminnows where stream fish were generally larger than lake fish. Body shape mainly varied as a function of sex. Adult male topminnows had larger morphological traits (with the exception of body width) than females, in particular longer dorsal and anal base lengths. Subtle effects of habitat were present. Stream fish had a longer dorsal fin base than lake fish. These phenotypic patterns may be the result of genetic and/or environmental variation. As these lakes are human‐made, the observed differences, if genetic, would have had to occur relatively rapidly (within about 100 years). © 2013 The Linnean Society of London     

The relationship between cranial morphology,bite performance,diet and habitat in a radiation of dwarf chameleon (Bradypodion)

Many animals show unique morphological and behavioural adaptations to specific habitats. In particular, variation in cranial morphology is known to influence feeding performance, which in turn influences dietary habits and, ultimately, fitness. Dietary separation is an important means of partitioning ecological niches and avoiding inter‐ and intraspecific competition. Consequently, differences in dietary resources may help explain phenotypic divergence in closely‐related species occupying different habitats, as well as sexual dimorphism. We test this hypothesis on five phenotypic forms of a recent radiation of dwarf chameleons (Bradypodion) that vary extensively in habitat use and cranial morphology. By examining stomach contents, the dietary composition of each phenotypic form is compared to investigate potential differences in feeding strategies. Overall, chameleons in the present study exhibit considerable dietary overlap (at both inter‐ and intraspecific levels), indicating that diet is not a major driver of variation in cranial morphology within this radiation. However, the stomachs of closed‐canopy females were found to contain more prey items than male stomachs, possibly indicating that females require a greater caloric intake than their male counterparts.     

Personality traits and behavioral syndromes in differently urbanized populations of house sparrows (Passer domesticus)

Urbanization creates novel environments for wild animals where selection pressures may differ drastically from those in natural habitats. Adaptation to urban life involves changes in various traits, including behavior. Behavioral traits often vary consistently among individuals, and these so-called personality traits can be correlated with each other, forming behavioral syndromes. Despite their adaptive significance and potential to act as constraints, little is known about the role of animal personality and behavioral syndromes in animals' adaptation to urban habitats. In this study we tested whether differently urbanized habitats select for different personalities and behavioral syndromes by altering the population mean, inter-individual variability, and correlations of personality traits. We captured house sparrows (Passer domesticus) from four different populations along the gradient of urbanization and assessed their behavior in standardized test situations. We found individual consistency in neophobia, risk taking, and activity, constituting three personality axes. On the one hand, urbanization did not consistently affect the mean and variance of these traits, although there were significant differences between some of the populations in food neophobia and risk taking (both in means and variances). On the other hand, both urban and rural birds exhibited a behavioral syndrome including object neophobia, risk taking and activity, whereas food neophobia was part of the syndrome only in rural birds. These results indicate that there are population differences in certain aspects of personality in house sparrows, some of which may be related to habitat urbanization. Our findings suggest that urbanization and/or other population-level habitat differences may not only influence the expression of personality traits but also alter their inter-individual variability and the relationships among them, changing the structure of behavioral syndromes.     

Convergent body flattening in a clade of tropical rock-using lizards (Scincidae: Lygosominae)

The repeated occurrence of similar morphologies in organisms from similar habitats provides good evidence of convergent selection, and convergent patterns of evolutionary change. In lizards, a flattened morphology has often been noted; however, whether this trait is convergent in specific habitats has never been tested using phylogenetic methods. The present study examined patterns of morphological convergence in 18 species of tropical Lygosomine skinks from three broad habitat categories (generalist, leaf litter-dwelling, and rock-using species). In general, although there where relatively few morphological differences of species from different habitats, phylogenetic analyses revealed that rock-using species have consistently and repeatedly evolved a dorsoventrally flattened head and body. The adaptive basis of this flattened morphology is consistent with both biomechanical predictions of performance (e.g. climbing locomotion) and ecology (e.g. use of rock crevices, camouflage) of species that occupy rocky habitats.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 399–411.     

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 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.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 129–138.     

ASYNCHRONOUS EVOLUTION OF PHYSIOLOGY AND MORPHOLOGY IN ANOLIS LIZARDS

Species‐rich adaptive radiations typically diversify along several distinct ecological axes, each characterized by morphological, physiological, and behavioral adaptations. We test here whether different types of adaptive traits share similar patterns of evolution within a radiation by investigating patterns of evolution of morphological traits associated with microhabitat specialization and of physiological traits associated with thermal biology in Anolis lizards. Previous studies of anoles suggest that close relatives share the same “structural niche” (i.e., use the same types of perches) and are similar in body size and shape, but live in different “climatic niches” (i.e., use habitats with different insolation and temperature profiles). Because morphology is closely tied to structural niche and field active body temperatures are tied to climatic niches in Anolis, we expected phylogenetic analyses to show that morphology is more evolutionarily conservative than thermal physiology. In support of this hypothesis, we find (1) that thermal biology exhibits more divergence among recently diverged Anolis taxa than does morphology; and (2) diversification of thermal biology among all species often follows diversification in morphology. These conclusions are remarkably consistent with predictions made by anole biologists in the 1960s and 1970s.     

Complex trait differentiation between host-populations of the pea aphid Acyrthosiphon pisum (Harris): implications for the evolution of ecological specialisation

Variation in traits affecting preference for, and performance on, new habitats is a key factor in the initiation of ecological specialisation and adaptive speciation. However, habitat and resource use also involves other traits whose influence on ecological and genetic divergence remains poorly understood. In the present study, we investigated the extent of variation of life-history traits among sympatric populations of the pea aphid Acyrthosiphon pisum , which shows several host races that are specialised on various plants of the family Fabaceae plants and is an established model for ecological speciation. First, we assessed the community structure of microbial partners within host populations of the pea aphid. The effect of these microbes on host fitness is uncertain, although there is growing evidence that they may modulate various important adaptive traits of their host such as plant utilisation and resistance against natural enemies. Second, we performed a multivariate analysis on several ecologically relevant features of host populations recorded in the present and previous studies (including microbial composition, colour morph, reproductive mode, and male dispersal phenotype), enabling the identification of correlations between phenotypic traits. We discuss the ecological significance of these associations of traits in relation to the habitat characteristics of pea aphid populations, and their consequences for the evolution of ecological specialisation and sympatric speciation.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 718–727.     

Is variation in tail vertebral morphology linked to habitat use in chameleons?

Chameleons (Chamaeleonidae) are known for their arboreal lifestyle, in which they make use of their prehensile tail. Yet, some species have a more terrestrial lifestyle, such as Brookesia and Rieppeleon species, as well as some chameleons of the genera Chamaeleo and Bradypodion. The main goal of this study was to identify the key anatomical features of the tail vertebral morphology associated with prehensile capacity. Both interspecific and intra-individual variation in skeletal tail morphology was investigated. For this, a 3D-shape analysis was performed on vertebral morphology using μCT-images of different species of prehensile and nonprehensile tailed chameleons. A difference in overall tail size and caudal vertebral morphology does exist between prehensile and nonprehensile taxa. Nonprehensile tailed species have a shorter tail with fewer vertebrae, a generally shorter neural spine and shorter transverse processes that are positioned more anteriorly (with respect to the vertebral center). The longer tails of prehensile species have more vertebrae as well as an increased length of the processes, likely providing a greater area for muscle attachment. At the intra-individual level, regional variation is observed with more robust proximal tail vertebrae having longer processes. The distal part has relatively longer vertebrae with shorter processes. Although longer, the small size and high number of the distal vertebrae allows the tail to coil around perches.     

Colonization of novel White Sands habitat is associated with changes in lizard anti‐predator behaviour

Colonization of novel habitats is often associated with differences in ecological community composition. For small diurnal animals, differences in predator diversity and abundance can lead to behavioural shifts in the novel habitat. The eastern fence lizard Sceloporus undulatus (Bosc and Daudin, 1801) recently colonized the gypsum dunes of White Sands, a predator‐poor community relative to the predator‐rich community of the surrounding Chihuahuan dark‐soil habitat. We used field experiments to assess S. undulatus anti‐predator behaviour in white‐sand versus dark‐soil habitats, and used laboratory assays to determine whether behavioural differences could be mediated by hormonal regulation. Overall, we found that white‐sand lizards were less vigilant but more wary than their dark‐soil counterparts; it took them longer to detect a simulated predator, but once detected they were more likely to retreat from their perches than dark‐soil lizards. At the proximate level, differences in anti‐predator behaviour could not be explained by differences in plasma hormone levels (corticosterone and testosterone); we detected elevated corticosterone for lizards in our stress treatment relative to control treatment, but found no differences between habitats in baseline or acute corticosterone levels. At the evolutionary level, we suggest that differences in anti‐predator behaviour may be explained by differences across habitats in predation environment, habituation, and/or the cost of retreating. Our study implicates changes in predator community composition in mediating ecological divergence in behaviour. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 657–667.     

Personality‐related differences in response to habitat in Mediterranean blue tits

Habitat‐specific selection pressures have been widely recognized, but whether selection favours different personality types in different habitats has rarely been evaluated. This study aimed to test whether personality‐related differences in annual reproductive success differed between two populations of blue tits (Cyanistes caeruleus) occupying different Mediterranean habitats (oakwood and pinewood). We measured exploration and parental provisioning behaviours and used a path analysis to ask how the interplay between these two behavioural traits affected reproductive success in each of these two habitats. We found that blue tits breeding in the pinewood were slow‐exploring compared to blue tits breeding in the oakwood, suggesting the occurrence of personality‐related differences in settlement, or behavioural plasticity in response to habitat. Exploration behaviour and feeding rates were positively associated, suggesting that they affect each other or that there is an environmental factor affecting both traits simultaneously. Finally, fast explorers were favoured in the pinewood, while there was no selection acting on exploration behaviour in the oak habitat. These findings emphasize the importance of integrating habitat selection, plasticity and personality in the study of behavioural evolution.     

Colour plasticity and background matching in a threespine stickleback species pair

Examining differences in colour plasticity between closely‐related species in relation to the heterogeneity of background colours found in their respective habitats may offer important insight into how cryptic colour change evolves in natural populations. In the present study, we examined whether nonbreeding dorsal body coloration has diverged between sympatric species of stickleback along with changes in habitat‐specific background colours. The small, limnetic species primarily occupies the pelagic zone and the large, benthic species inhabits the littoral zone. We placed benthic and limnetic sticklebacks against extremes of habitat background colours and measured their degree of background matching and colour plasticity. Benthics matched the littoral background colour more closely than did the limnetics, although there was no difference between species in their resemblance to the pelagic background colour. Benthics were able to resemble both background colours by exhibiting greater directional colour plasticity in their dorsal body coloration than limnetics, which may be an adaptive response to the greater spectral heterogeneity of the littoral zone. The present study highlights how habitat‐specific spectral characteristics may shape cryptic coloration differences between stickleback species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 902–914.