Modelling structural colour from helicoidal multi-layer thin films with natural disorder.

A coupled mode theory based on Takagi-Taupin equations describing electromagnetic scattering from distorted periodic arrays is applied to the problem of light scattering from beetles. We extend the method to include perturbations in the permittivity tensor to helicoidal arrays seen in many species of scarab beetle and optically anisotropic layered materials more generally. This extension permits analysis of typical dislocations arising from the biological assembly process and the presence of other structures in the elytra. We show that by extracting structural information from transmission electron microscopy data, including characteristic disorder parameters, good agreement with spectral specular and non-specular reflectance measurements is obtained.

Single and multiple mating reduces longevity of female dumpling squid (Euprymna tasmanica).

For many species, mating is a necessary yet costly activity. The costs involved can have an important influence on the evolution of life histories and senescence. Females of many species mate multiply, and this behaviour can inflict a longevity cost. Most studies investigating the effects of multiple mating on female survival have been conducted on insects, and the effects in other taxa are largely unknown. We investigate the effects of both a single mating and a second mating on longevity in female dumpling squid (Euprymna tasmanica), a species in which both sexes mate multiply. Through comparing the longevity of virgin, once-mated and twice-mated females, we found that a single mating reduced female life span by 15 days on average. A second mating resulted in an additional 8 day (on average) longevity cost, despite no difference in total clutch mass, number of clutches, single egg mass or number of eggs per clutch between once-mated and twice-mated females. This demonstrates a cost to multiple mating which may be independent of the cost of egg production. Furthermore, total clutch mass and female life span were positively correlated, whereas female life span decreased with increasing average water temperature. The presence of an additive effect of reproduction on longevity suggests that multiple mating in cephalopods may have benefits that outweigh these costs, or that there is a conflict in optimal mating frequency between males and females.

Ornament size and colour as alternative strategies for effective communication in gliding lizards.

Sexual ornamentation needs to be conspicuous to be effective in attracting potential mates and defending territories and indeed, a multitude of ways exists to achieve this. Two principal mechanisms for increasing conspicuousness are to increase the ornament's colour or brightness contrast against the background and to increase the size of the ornament. We assessed the relationship between the colour and size of the dewlap, a large extendible throat-fan, across a range of species of gliding lizards (Agamidae; genus Draco) from Malaysia and the Philippines. We found a negative relationship across species between colour contrast against the background and dewlap size in males, but not in females, suggesting that males of different species use increasing colour contrast and dewlap size as alternative strategies for effective communication. Male dewlap size also increases with increasing sexual size dimorphism, and dewlap colour and brightness contrast increase with increasing sexual dichromatism in colour and brightness, respectively, suggesting that sexual selection may act on both dewlap size and colour. We further found evidence that relative predation intensity, as measured from predator attacks on models placed in the field, may play a role in the choice of strategy (high chromatic contrast or large dewlap area) a species employs. More broadly, these results highlight that each component in a signal (such as colour or size) may be influenced by different selection pressures and that by assessing components individually, we can gain a greater understanding of the evolution of signal diversity.

Geographic variation in hybridization and ecological differentiation between three syntopic, morphologically similar species of montane lizards.

To understand factors shaping species boundaries in closely related taxa, a powerful approach is to compare levels of genetic admixture at multiple points of contact and determine how this relates to intrinsic and extrinsic factors, such as genetic, morphological and ecological differentiation. In the Australian Alps, the threatened alpine bog skink Pseudemoia cryodroma co-occurs with two morphologically and ecologically similar congeners, P. entrecasteauxii and P. pagenstecheri, and all three species are suspected to hybridize. We predicted that the frequency of hybridization should be negatively correlated with genetic divergence, morphological differentiation and microhabitat separation. We tested this hypothesis using a mitochondrial locus, 13 microsatellite loci, morphological and microhabitat data and compared results across three geographically isolated sites. Despite strong genetic structure between species, we detected hybridization between all species pairs, including evidence of backcrossed individuals at the two sites where all three species are syntopic. Hybridization frequencies were not consistently associated with genetic, morphological or ecological differentiation. Furthermore, P. entrecasteauxii and P. pagenstecheri only hybridized at the two sites where they are syntopic with P. cryodroma, but not at the largest site where P. cryodroma was not recorded, suggesting that P. cryodroma may serve as a bridging species. This study reveals the complex dynamics within a three species hybrid zone and provides a baseline for assessing the impact of climate change and anthropogenic habitat modification on future hybridization frequencies.

Marked colour divergence in the gliding membranes of a tropical lizard mirrors population differences in the colour of falling leaves.

Populations of the Bornean gliding lizard, Draco cornutus, differ markedly in the colour of their gliding membranes. They also differ in local vegetation type (mangrove forest versus lowland rainforest) and consequently, the colour of falling leaves (red and brown/black in mangrove versus green, brown and black in rainforest). We show that the gliding membranes of these lizards closely match the colours of freshly fallen leaves in the local habitat as they appear to the visual system of birds (their probable predators). Furthermore, gliding membranes more closely resembled colours of local fallen leaves than standing foliage or fallen leaves in the other population's habitat. This suggests that the two populations have diverged in gliding membrane coloration to match the colours of their local falling leaves, and that mimicking falling leaves is an adaptation that functions to reduce predation by birds.

Local adaptation and divergence in colour signal conspicuousness between monomorphic and polymorphic lineages in a lizard.

Population differences in visual environment can lead to divergence in multiple components of animal coloration including signalling traits and colour patterns important for camouflage. Divergence may reflect selection imposed by different receivers (conspecifics, predators), which depends in turn on the location of the colour patch. We tested for local adaptation of two genetically and phenotypically divergent lineages of a rock-inhabiting lizard, Ctenophorus decresii, by comparing the visual contrast of colour patches to different receivers in native and non-native environments. The lineages differ most notably in male throat coloration, which is polymorphic in the northern lineage and monomorphic in the southern lineage, but also differ in dorsal and lateral coloration, which is visible to both conspecifics and potential predators. Using models of animal colour vision, we assessed whether lineage-specific throat, dorsal and lateral coloration enhanced conspicuousness to conspecifics, increased crypsis to birds or both, respectively, when viewed against the predominant backgrounds from each lineage. Throat colours were no more conspicuous against native than non-native rock but contrasted more strongly with native lichen, which occurs patchily on rocks inhabited by C. decresii. Conversely, neck coloration (lateral) more closely matched native lichen. Furthermore, although dorsal coloration of southern males was consistently more conspicuous to birds than that of northern males, both lineages had similar absolute conspicuousness against their native backgrounds. Combined, our results are consistent with local adaptation of multiple colour traits in relation to multiple receivers, suggesting that geographic variation in background colour has influenced the evolution of lineage-specific coloration in C. decresii.

Phylogeographic structure, demographic history and morph composition in a colour polymorphic lizard.

In polymorphic species, population divergence in morph composition and frequency has the potential to promote speciation. We assessed the relationship between geographic variation in male throat colour polymorphism and phylogeographic structure in the tawny dragon lizard, Ctenophorus decresii. We identified four genetically distinct lineages, corresponding to two polymorphic lineages in the Northern Flinders Ranges and Southern Flinders Ranges/Olary Ranges regions respectively, and a monomorphic lineage in the Mt Lofty Ranges/Kangaroo Island region. The degree of divergence between these three lineages was consistent with isolation to multiple refugia during Pleistocene glacial cycles, whereas a fourth, deeply divergent (at the interspecific level) and monomorphic lineage was restricted to western New South Wales. The same four morphs occurred in both polymorphic lineages, although populations exhibited considerable variation in the frequency of morphs. By contrast, male throat coloration in the monomorphic lineages differed from each other and from the polymorphic lineages. Our results suggest that colour polymorphism has evolved once in the C. decresii species complex, with subsequent loss of polymorphism in the Mt Lofty Ranges/Kangaroo Island lineage. However, an equally parsimonious scenario, that polymorphism arose independently twice within C. decresii, could not be ruled out. We also detected evidence of a narrow contact zone with limited genotypic admixture between the polymorphic Olary Ranges and monomorphic Mt Lofty Ranges regions, yet no individuals of intermediate colour phenotype. Such genetic divergence and evidence for barriers to gene flow between lineages suggest incipient speciation between populations that differ in morph composition.

Discrete colour polymorphism in the tawny dragon lizard (Ctenophorus decresii) and differences in signal conspicuousness among morphs.

Intraspecific colour variation is common in nature and can vary from the coexistence of discrete colour variants in polymorphic species to continuous variation. Whether coloration is continuous or discrete is often ambiguous and many species exhibit a combination of the two. The nature of the variation (discrete or continuous) has implications for both the genetic basis of the colour variation and the evolutionary processes generating and maintaining it. Consequently, it is important to qualify the existence of discrete morphs, particularly in relation to the animal's visual system. In this study, we quantified male throat colour variation in Ctenophorus decresii tawny dragon lizard and tested for morphological and ecological correlates of the colour variants. We confirmed that discrete throat colour morphs can be defined based on colour and pattern analyses independent of the human visual system. We also found that the colour variants differed in their conspicuousness from the background, to the lizard's visual system, which has implications for signalling. However, the morphs did not differ in morphology or microhabitat use, which suggests that these characteristics are not involved in the evolutionary maintenance of the polymorphism.

Sexual selection is positively associated with ecological generalism among agamid lizards.

Natural and sexual selection shape the evolution of species but the interplay between them is poorly understood. Two phylogenetic studies on birds have suggested that species with greater sexual dichromatism have a broader habitat use. We show that in agamid lizards, species with more elaborate secondary sexual traits are also ecologically more opportunistic. Species with greater dimorphism in head size and ornamentation have greater altitudinal range and broader habitat use, respectively, and species with greater sexual dichromatism have wider microhabitat use. Body size was positively associated with sexual and ecological generalism, but associations between ecological and sexual traits remained after accounting for body size. We suggest that sexual and natural selection may be linked either because sexual selection can promote generalism at the population level by favouring 'good genes', or because higher population densities may be associated with both stronger sexual selection and broader resource use.

The contribution of structural-, psittacofulvin- and melanin-based colouration to sexual dichromatism in Australasian parrots.

Colour ornamentation in animals is exceptionally diverse, but some colours may provide better signals of individual quality or more efficient visual stimuli and, thus, be more often used as sexual signals. This may depend on physiological costs, which depend on the mechanism of colour production (e.g. exogenously acquired colouration in passerine birds appears to be most sexually dichromatic). We studied sexual dichromatism in a sample of 27 Australasian parrot species with pigment- (melanin and psittacofulvin) and structural-based colouration, to test whether some of these types of colouration are more prominent in sexual ornamentation. Unlike passerines, in which long wavelength colouration (yellow to red) usually involves exogenous and costly carotenoid pigments, yellow to red colouration in parrots is based on endogenously synthesized psittacofulvin pigments. This allows us to assess whether costly exogenous pigments are necessary for these plumage colours to have a prominent role in sexual signalling. Structural blue colouration showed the largest and most consistent sexual dichromatism, both in area and perceptually relevant chromatic differences, indicating that it is often ornamental in parrots. By contrast, we found little evidence for consistent sexual dichromatism in melanin-based colouration. Unlike passerines, yellow to red colouration was not strongly sexually dichromatic: although the area of colouration was generally larger in males, colour differences between the sexes were on average imperceptible to parrots. This is consistent with the idea that the prominent yellow to red sexual dichromatism in passerines is related to the use of carotenoid pigments, rather than resulting from sensory bias for these colours.

Sex-specific ecomorphological variation and the evolution of sexual dimorphism in dwarf chameleons (Bradypodion spp.).

Natural selection can influence the evolution of sexual dimorphism through selection for sex-specific ecomorphological adaptations. The role of natural selection in the evolution of sexual dimorphism, however, has received much less attention than that of sexual selection. We examined the relationship between habitat structure and both male and female morphology, and sexual dimorphism in size and shape, across 21 populations of dwarf chameleon (genus Bradypodion). Morphological variation in dwarf chameleons was strongly associated with quantitative, multivariate aspects of habitat structure and, in most cases, relationships were congruent between the sexes. However, we also found consistent relationships between habitat and sexual dimorphism. These resulted from both differences in magnitude of ecomorphological relationships that were otherwise congruent between the sexes, as well as in sex-specific ecomorphological adaptations. Our study provides evidence that natural selection plays an important role in the evolution of sexual dimorphism.

Ornament evolution in dragon lizards: multiple gains and widespread losses reveal a complex history of evolutionary change.

The expression in females of ornaments thought to be the target of sexual selection in males is a long-standing puzzle. Two main hypotheses are proposed to account for the existence of conspicuous ornaments in both sexes (mutual ornamentation): genetic correlation between the sexes and sexual selection on females as well as males. We examined the pattern of ornament gains and losses in 240 species of dragon lizards (Agamidae) in order to elucidate the relative contribution of these two factors in the evolution of mutual ornamentation. In addition, we tested whether the type of shelter used by lizards to avoid predators predicts the evolutionary loss or constraint of ornament expression. We found evidence that the origin of female ornaments is broadly consistent with the predictions of the genetic correlation hypothesis. Ornaments appear congruently in both sexes with some lineages subsequently evolving male biased sexual dimorphism, apparently through the process of natural selection for reduced ornamentation in females. Nevertheless, ornaments have also frequently evolved in both sexes independently. This suggests that genetic correlations are potentially weak for several lineages and sexual selection on females is responsible for at least some evolutionary change in this group. Unexpectedly, we found that the evolutionary loss of some ornaments is concentrated more in males than females and this trend cannot be fully explained by our measures of natural selection.