Phylogenetic and phylogeographic insights into Sri Lankan killifishes (Teleostei: Aplocheilidae).

Three nominal species of the killifish genus Aplocheilus are reported from the lowlands of Sri Lanka. Two of these, Aplocheilus dayi and Aplocheilus werneri, are considered endemic to the island, whereas Aplocheilus parvus is reported from both Sri Lanka and Peninsular India. Here, based on a collection from 28 locations in Sri Lanka, also including a dataset of Asian Aplocheilus downloaded from GenBank, we present a phylogeny constructed from the mitochondrial cytochrome b (cytb), mitochondrial cytochrome c oxidase subunit 1 (cox1), and nuclear recombination activating protein 1 (rag1), and investigate the interrelationships of the species of Aplocheilus in Sri Lanka. The endemic Sri Lankan aplocheilid clade comprising A. dayi and A. werneri is recovered as the sister group to the clade comprising A. parvus from Sri Lanka and Aplocheilus blockii from Peninsular India. The reciprocal monophyly of A. dayi and A. werneri is not supported in our molecular phylogeny. A. dayi and A. werneri display strong sexual dimorphism, but species-level differences are subtle, explained mostly by pigmentation patterns. Their phenotypes exhibit a parapatric distribution and may represent locally adapted forms of a single species. Alternatively, the present study does not rule out the possibility that A. dayi and A. werneri may represent an incipient species pair or that they have undergone introgression or hybridization in their contact zones. We provide evidence that the Nilwala-Gin region of southwestern Sri Lanka may have acted as a drought refugium for these fishes.

Sexual dichromatism in the fur of a bat: An exploration of color differences and potential signaling functions.

Sex differences in body color (i.e., sexual dichromatism) are rare in bats and, more broadly, in mammals. The eastern red bat (Lasiurus borealis) is a common tree-roosting bat that occupies much of North America and has long been described as sexually dichromatic. However, previous research on this species found that absolute body size and collection year were better predictors of fur color in preserved specimens than sex. We revisited this issue and photographed 82 live eastern red bats under standardized conditions, then used image analysis to quantify pelage hue, saturation, and value. We used an information theoretic approach to evaluate four competing hypotheses about the principal drivers of color differences in the fur of eastern red bats. Our analyses demonstrated that sex was a better predictor of pelage color than body size; males had redder, more saturated, and lighter pelages than females. Additionally, the fur color of juvenile versus adult bats differed somewhat, as juveniles were darker than adults. In general, absolute body size (i.e., forearm length in bats) was a poor predictor of color in live eastern red bats. In an exploratory post-hoc analysis, we confirm that fur color is related to body mass (i.e., a proxy for body condition in bats), suggesting color might serve as a sexually selected signal of mate quality in this partially diurnal species. Future work should investigate the functional role of sexual dichromatism in this species, which could be related to signaling or possibly thermoregulation.

Colouration matters in dull toads: ultraviolet adornment for ladies and agrochemicals fading effects.

Integument colouration can influence many aspects of fitness, and is under strong sexual selection. Amphibians often express sexual dichromatism, and ultra-violet (UV) colouration is usually biased toward males as a sexual signal. As an honest signal, colouration is related to several individual traits, but can also be related to environmental factors such as anthropogenic pollutants, to which amphibians are highly sensitive. In this study, we investigated sexual dichromatism and UV reflectance covering a large visual spectrum (wavelength ranging from 300 to 700 nm) on different body areas (throat, ventral and dorsal areas), in a widespread amphibian species, the spiny toad (Bufo spinosus). Then, we tested the impact of chronic exposure to two widespread herbicides (glyphosate's primary metabolite [AMPA] and Nicosulfuron) on their colouration. We found a strong but unexpected sexual dichromatism with females reflecting more in the UV spectrum (throat and ventral area) than males, suggesting these body parts might be critical in intra-specific signalling. Females with higher ventral UV reflectance were in better body condition, suggesting an honest signal role of UV reflectance which could influence male choice. Throat colouration was further differentially influenced by agrochemicals according to sexes. In AMPA-exposed males, throat was more saturated in yellow-orange than in control males, and Nicosulfuron exposure decreased the throat's reflectance hue in females, which can bear consequences on mate attractiveness. Future studies need to investigate the underlying mechanisms that are altered by agrochemical exposure.

Influence of sexual dimorphism and dichromatism on reproductive success in a rare native cactus.

Identifying plant sexual dimorphic traits is critical in advancing our knowledge on plant-pollinator interactions. For example, dimorphism in floral colors, or sexual dichromatism, is a crucial mediator of pollinator choice on foraging decisions. We studied Cylindropuntia wolfii, a model system, with diverse flower colors and a functionally dioecious sexual system. However, evidence suggests that sexual reproduction is limited in this species as it has a low seed set especially in naturally pollinated fruits. Thus, it is critical to this native species' conservation to investigate its relationship with pollinators. Our goals were to: (a) investigate the sexual dimorphism including the sexual dichromatism in the flowers of the cactus, and (b) determine whether sexually dimorphic traits affect the pollinator attraction of both the sexes. We measured several quantitative and qualitative traits and compared them between male and female flowers. Then we recorded the pollinator visitation rate in nature for both sexes and tracked pollinator color preference using fluorescent dyes as pollen analogues. Our study showed that male flowers of C. wolfii are bigger and brighter, and they attract more potential pollinators than females, supporting the hypothesis that sexual dimorphism influences pollinator visitation preference. Fluorescence dichromatism, in which female flowers' anthers fluoresce more than male flower anthers suggest this could be female flowers' strategy to compensate for their dark colors and small size. The results from this study showed that C. wolfii exhibits sexual dichromatism and fluorescence dichromatism, which is a novel finding in plant research.

Early origin and diverse phenotypic implementation of iridescent UV patterns for sexual signaling in pierid butterflies.

Iridescent ultraviolet (IUV) patterns on pierid butterfly wings are phenotypic adaptations commonly used as sexual signals, generated by scales with ultrastructural modifications. Pierid IUV patterns are sexually dichromatic, with reduced size in females, where conspicuous sexual signaling balances courtship against ecological predation. There have been no phylogenetic reconstructions of IUV within Pieridae and little morphological characterization of phenotypic diversity. Our genus-wide characterization of IUV revealed the uniform similarity of stacked lamellar ridges on the dorsal surface of cover scales. We tested a hypothesis of single versus multiple origins by reconstructing a phylogeny of 534 species (~43.2% described species), with all genera represented, and a trait matrix of 734 species (~59.4%) screened for IUV. A single, early dimorphic origin of IUV followed by several losses and gains received strong support, concluding that IUV patterns and structural coloration are old traits. Collectively, these results support the homology of IUV scales and patterns that diversified within several lineages, suggesting an interplay between female-mediated sexual selection and ecological predatory selection.

The extent of rapid colour change in male agamid lizards is unrelated to overall sexual dichromatism.

Dynamic colour change is widespread in ectothermic animals, but has primarily been studied in the context of background matching. For most species, we lack quantitative data on the extent of colour change across different contexts. It is also unclear whether and how colour change varies across body regions, and how overall sexual dichromatism relates to the extent of individual colour change. In this study, we obtained reflectance measures in response to different stimuli for males and females of six species of agamid lizards (Agamidae, sister family to Chameleonidae) comprising three closely related species pairs. We computed the colour volume in a lizard-vision colour space occupied by males and females of each species and estimated overall sexual dichromatism based on the area of non-overlapping male and female colour volumes. As expected, males had larger colour volumes than females, but the extent of colour change in males differed between species and between body regions. Notably, species that were most sexually dichromatic were not necessarily those in which males showed the greatest individual colour change. Our results indicate that the extent of colour change is independent of the degree of sexual dichromatism and demonstrate that colour change on different body regions can vary substantially even between pairs of closely related species.

Species differences in hormonally mediated gene expression underlie the evolutionary loss of sexually dimorphic coloration in Sceloporus lizards.

Phenotypic sexual dimorphism often involves the hormonal regulation of sex-biased expression for underlying genes. However, it is generally unknown whether the evolution of hormonally mediated sexual dimorphism occurs through upstream changes in tissue sensitivity to hormone signals, downstream changes in responsiveness of target genes, or both. Here, we use comparative transcriptomics to explore these possibilities in 2 species of Sceloporus lizards exhibiting different patterns of sexual dichromatism. Sexually dimorphic S. undulatus develops blue and black ventral coloration in response to testosterone, while sexually monomorphic S. virgatus does not, despite exhibiting similar sex differences in circulating testosterone levels. We administered testosterone implants to juveniles of each species and used RNAseq to quantify gene expression in ventral skin. Transcriptome-wide responses to testosterone were stronger in S. undulatus than in S. virgatus, suggesting species differences in tissue sensitivity to this hormone signal. Species differences in the expression of genes for androgen metabolism and sex hormone-binding globulin were consistent with this idea, but expression of the androgen receptor gene was higher in S. virgatus, complicating this interpretation. Downstream of androgen signaling, we found clear species differences in hormonal responsiveness of genes related to melanin synthesis, which were upregulated by testosterone in S. undulatus, but not in S. virgatus. Collectively, our results indicate that hormonal regulation of melanin synthesis pathways contributes to the development of sexual dimorphism in S. undulatus, and that changes in the hormonal responsiveness of these genes in S. virgatus contribute to the evolutionary loss of ventral coloration.

Subtractive colour mixing with bile pigments creates the rich wing palette of Graphium weiskei butterflies.

The wings of the purple spotted swallowtail Graphium weiskei are marked by an unusual bright colour pattern. Spectrophotometry on G. weiskei wings demonstrated the presence of a pigment with an absorption spectrum (peak wavelength λmax=676 nm) similar to that of the bile pigment sarpedobilin in the wings of the congeneric Graphium sarpedon (λmax=672 nm). Sarpedobilin alone causes cyan-blue wing areas, but the green-coloured areas of G. sarpedon wings result from subtractive colour mixing with the carotenoid lutein. Reflectance spectra of the blue-coloured areas of G. weiskei wings indicate that sarpedobilin is mixed with the short-wavelength-absorbing papiliochrome II. An enigmatic pigment, tentatively called weiskeipigment (λmax=580 nm), enhances the saturation of the blue colour. Weiskeipigment causes a purple colour in areas where the sarpedobilin concentration is low. The wings of the related papilionid Papilio phorcas contain the bile pigment pharcobilin (λmax=604 nm), as well as another sarpedobilin (λmax=663 nm). The cyan to greenish wings of P. phorcas are due to phorcabilin and sarpedobilin mixed with papiliochrome II. A survey of known subspecies of G. weiskei as well as of congeneric Graphium species of the 'weiskei' group shows various degrees of subtractive colour mixing of bilins and short-wavelength absorbers (carotenoids and/or papiliochromes) in their wings. This study illuminates the underestimated role of bile pigments in butterfly wing colouration.

Oestradiol reduces female bill colour in a mutually ornamented bird.

Carotenoid-based colour signals can be costly to produce and maintain, and trade-offs between signalling and other fitness traits are expected. In mutually ornamented species, trade-offs with reproduction may be stronger for females than males, because females often dedicate more resources to offspring production, which may lead to plastic investment in colour signals and plastic sexual dichromatism. Oestradiol is a candidate mediator of this trade-off because it regulates reproductive physiology and may also influence the expression of coloration. We tested this hypothesis by giving female common waxbills (Estrilda astrild) either oestradiol (17ß-oestradiol) or empty implants during the early breeding season and measured spectral reflectance of carotenoid-based bill coloration weekly for two months. Using a model of avian vision, we found that bill colour in oestradiol-implanted females became less saturated, less red in hue and brighter, compared with control females and with unimplanted males. This resulted in a change in bill sexual dichromatism from imperceptible to perceptible. Results support the hypothesis that female reproductive physiology influences investment in coloration through changes in oestradiol and show a form of female-driven plastic sexual dichromatism. Greater sensitivity of female colour to physiological and/or environmental conditions helps explain why differences in sexual dichromatism among species differing in ecology often evolve owing to changes in female rather than male phenotype.

Haemosporidian parasites and incubation period influence plumage coloration in tanagers (Passeriformes: Thraupidae).

Birds are highly visually oriented and use plumage coloration as an important signalling trait in social communication. Hence, males and females may have different patterns of plumage coloration, a phenomenon known as sexual dichromatism. Because males tend to have more complex plumages, sexual dichromatism is usually attributed to female choice. However, plumage coloration is partly condition-dependent; therefore, other selective pressures affecting individuals' success may also drive the evolution of this trait. Here, we used tanagers as model organisms to study the relationships between dichromatism and plumage coloration complexity in tanagers with parasitism by haemosporidians, investment in reproduction and life-history traits. We screened blood samples from 2849 individual birds belonging to 52 tanager species to detect haemosporidian parasites. We used publicly available data for plumage coloration, bird phylogeny and life-history traits to run phylogenetic generalized least-square models of plumage dichromatism and complexity in male and female tanagers. We found that plumage dichromatism was more pronounced in bird species with a higher prevalence of haemosporidian parasites. Lastly, high plumage coloration complexity in female tanagers was associated with a longer incubation period. Our results indicate an association between haemosporidian parasites and plumage coloration suggesting that parasites impact mechanisms of sexual selection, increasing differences between the sexes, and social (non-sexual) selection, driving females to develop more complex coloration.

The evolution of sex similarities in social signals: Climatic seasonality is associated with lower sexual dimorphism and greater elaboration of female and male signals in antbirds (Thamnophilidae).

Selection on signals that mediate social competition varies with resource availability. Climate regulates resource availability, which may affect the strength of competition and selection on signals. Traditionally, this meant that more seasonal, colder, or dryer-overall harsher-environments should favor the elaboration of male signals under stronger male-male competition, increasing sexual dimorphism. However, females also use signals to compete; thus, harsher environments could strengthen competition and favor elaboration of signals in both sexes, decreasing sexual dimorphism. Alternatively, harsher environments could decrease sexual dimorphism due to scarcer resources to invest in signal elaboration in both sexes. We evaluated these contrasting hypotheses in antbirds, a family of Neotropical passerines that varies in female and male signals and occurs across diverse climatic regimes. We tested the association of sexual dimorphism of plumage coloration and songs with temperature, precipitation, and their seasonality. We found that greater seasonality is associated with lower sexual dimorphism in plumage coloration and greater elaboration of visual signals in both sexes, but not acoustic signals. Our results suggest that greater seasonality may be associated with convergent elaboration of female and male visual signals, highlighting the role of signals of both sexes in the evolution of sexual dimorphism.

Sexual repurposing of juvenile aposematism in locusts.

Adaptive plasticity requires an integrated suite of functional responses to environmental variation, which can include social communication across life stages. Desert locusts (Schistocerca gregaria) exhibit an extreme example of phenotypic plasticity called phase polyphenism, in which a suite of behavioral and morphological traits differ according to local population density. Male and female juveniles developing at low population densities exhibit green- or sand-colored background-matching camouflage, while at high densities they show contrasting yellow and black aposematic patterning that deters predators. The predominant background colors of these phenotypes (green/sand/yellow) all depend on expression of the carotenoid-binding "Yellow Protein" (YP). Gregarious (high-density) adults of both sexes are initially pinkish, before a YP-mediated yellowing reoccurs upon sexual maturation. Yellow color is especially prominent in gregarious males, but the reason for this difference has been unknown since phase polyphenism was first described in 1921. Here, we use RNA interference to show that gregarious male yellowing acts as an intrasexual warning signal, which forms a multimodal signal with the antiaphrodisiac pheromone phenylacetonitrile (PAN) to prevent mistaken sexual harassment from other males during scramble mating in a swarm. Socially mediated reexpression of YP thus adaptively repurposes a juvenile signal that deters predators into an adult signal that deters undesirable mates. These findings reveal a previously underappreciated sexual dimension to locust phase polyphenism, and promote locusts as a model for investigating the relative contributions of natural versus sexual selection in the evolution of phenotypic plasticity.

Sex roles in birds: Phylogenetic analyses of the influence of climate, life histories and social environment.

Sex roles describe sex differences in courtship, mate competition, social pair-bonds and parental care. A key challenge is to identify associations among the components and the drivers of sex roles. Here, we investigate sex roles using data from over 1800 bird species. We found extensive variation and lability in proxies of sex roles, indicating remarkably independent evolution among sex role components. Climate and life history showed weak associations with sex roles. However, adult sex ratio is associated with sexual dimorphism, mating system and parental care, suggesting that social environment is central to explaining variation in sex roles among birds. Our results suggest that sex differences in reproductive behaviour are the result of diverse and idiosyncratic responses to selection. Further understanding of sex roles requires studies at the population level to test how local responses to ecology, life histories and mating opportunities drive processes that shape sex role variation among higher taxa.

Secondary Sexual Trait Melanization in "Black" Scavenger Flies: Nutritional Plasticity and Its Evolution.

AbstractThe black scavenger fly Sepsis thoracica exhibits polyphenic development resulting in alternate small black and large amber male morphs. Although the behavior, ecology, and physiology of both morphs are being scrutinized, the evolutionary origins of the nutritional polyphenism remain poorly understood. I here use a comparative approach to study variation in the degree of melanization of the forefemur-a secondary sexual trait. Melanization showed nutritional plasticity in all species, and character mapping suggests polyphenic development to represent the ancestral character state that was lost repeatedly. That is, interspecific variation among the studied species is mainly caused by the loss and not the gain of polyphenic development. Coevolution between male melanization and mating system differences further implicates sexual selection in the evolution of male melanization. These findings highlight the usefulness of comparative and natural history data in shedding new light on the evolution of phenotypic variation.

Patterns of variations in dorsal colouration of the Italian wall lizard Podarcis siculus.

Research on animal colouration has grown exponentially in the last decade thanks to multidisciplinary approaches. Most studies are focused on trade-offs between communication and mimicry, which represent the two main constraints and drivers of the evolution of body colourations. Reptiles are excellent model species for investigating this field of study and lizards in particular show great variability of body colourations and their functions. We studied the lizard Podarcis siculus, analysing the variations of dorsal colour of three populations and obtained clear patterns of seasonal and ontogenetical variation of dorsal colour. According to baseline colour, males were greener and brighter than females, although no difference in saturation was recorded. According to seasonal variations, analyses showed that both sexes significantly vary in colour over the year: males reached higher peaks of hue and saturation later than females during spring, while females showed higher peaks of brightness and reached earlier similarly to hue and saturation. Ontogenetic variations were recorded only in males, which become greener, less bright and saturated with growing size. Therefore, our results suggest the occurrence of two opposing strategies in colour expression between sexes: males' dorsal colouration plays a major role in communication, while females are more crypsis-oriented.

Does the addition of a new signalling trait enhance receiver responses in diurnal geckos?

Animal signals in multiple modalities expands the opportunity for effective communication. Among diurnal geckos of the genus Cnemaspis, chemical signalling traits preceded the evolution of visual traits. Males of all species possess chemical secreting ventral glands, but only in some species, males also express yellow gular patches. This difference in the expression of unimodal or multimodal signalling traits between closely related species provided us with an opportunity to understand the use of multimodal signals for communication. We studied receiver responses in Cnemaspis indica, a sexually monochromatic species, and in C. littoralis, a species where males possess yellow gulars. We performed behavioural trials where individuals of each species were exposed to only chemical stimuli, only visual stimuli, or both chemical and visual stimuli simultaneously from male and female conspecifics. Our results show that only chemical stimuli were necessary and sufficient to elicit responses in males and females of C. indica as well as in females of C. littoralis. However, males of the dimorphic C. littoralis required the multimodal stimulus to elicit movement-based responses. Our results suggest that the evolution of colour traits in diurnal geckos is associated with a partial shift in some receiver responses toward multimodal communication, with no addition to the behavioural repertoire.

Carotenoid deprivation and beta-carotene's effects on male and female turtle color.

Carotenoid-colored integuments commonly function as sexually selected honest signals because carotenoid pigments can be costly to obtain, ingest, absorb, metabolize or transport before being deposited into the integument. As such, carotenoid pigmentation is often sexually dichromatic, with males being more colorful than females. Sexual dichromatism may also occur in ultraviolet (UV) wavelengths, which is visible to organisms who possess UV-sensitive photoreceptors. The stripes and spots of painted turtles (Chrysemys picta) are carotenoid-based and reflect UV wavelengths. This research describes UV sexual dichromatism in painted turtles and shows how carotenoid deprivation changes spot and stripe color in male and female painted turtles. Adult turtles were fed a diet that was supplemented with carotenoids (i.e., C diet) or deprived of carotenoids (C-). Stripe and spot color were measured with UV-vis spectrometry, and blood was drawn from all turtles before and after the dietary treatment. HPLC analysis revealed five carotenoids (4 xanthophylls and beta-carotene) circulating in turtle blood. C-diet reduced yellow chroma and increased brightness of yellow and red stripes or spots, relative to the C diet, but there was no sexually dimorphic effect of carotenoid deprivation on color, nor did carotenoid deprivation affect UV reflectance. Carotenoid deprivation reduced all circulating carotenoids, but beta-carotene was the only pigment with a significant effect on post-experimental carotenoids, implying that changes in color were due in part to reduction in circulating levels of beta-carotene. Color generation appears to be complex in turtles and have dietary as well as non-dietary components.

Body size and climate as predictors of plumage colouration and sexual dichromatism in parrots.

Psittaciformes (parrots, cockatoos and lorikeets) comprise one of the most colourful clades of birds. Their unique pigments and safe cavity nesting habits are two potential explanations for their colourful character. However, plumage colour varies substantially between parrot species and sometimes also between males and females of the same species. Here, we use comparative analyses to evaluate what factors correlate with colour elaboration, colour diversity and sexual dichromatism. Specifically, we test the association between different aspects of parrot colouration and (a) the intensity of sexual selection and social interactions, (b) variation along the slow-fast life-history continuum and (c) climatic variation. We show that larger species and species that live in warm environments display more elaborated colours, yet smaller species have higher levels of sexual dichromatism. Larger parrots tend to have darker and more blue and red colours. Parrots that live in wetter environments are darker and redder, whereas species inhabiting warm regions have more blue plumage colours. In general, each of the variables we considered explain small to moderate amounts of variation in parrot colouration (up to 15%). Our data suggest that sexual selection may be acting more strongly on males in small, short-lived parrots leading to sexual dichromatism. More elaborate colouration in both males and females of the larger, long-lived species with slow tropical life histories suggests that mutual mate choice, social selection and reduced selection for crypsis may be important in these species, as has been shown for passerines.

Macroevolutionary Patterns of Sexual Size Dimorphism Among African Tree Frogs (Family: Hyperoliidae).

Sexual size dimorphism (SSD) is shaped by multiple selective forces that drive the evolution of sex-specific body size, resulting in male or female-biased SSD. Stronger selection on one sex can result in an allometric body size scaling relationship consistent with Rensch's rule or its converse. Anurans (frogs and toads) generally display female-biased SSD, but there is variation across clades and the mechanisms driving the evolution of SSD remain poorly understood. We investigated these topics in a diverse family of African treefrogs (Hyperoliidae). Hyperoliids display traits considered rare among amphibians, including sexual dichromatism and protogynous sex change. Using phylogenetic comparative methods, we tested if adult ecology, sexual dichromatism, and sex change were predictors of body size or SSD. We also tested whether hyperoliids displayed allometric interspecific body size scaling relationships. We found a majority of hyperoliid taxa display female-biased SSD, but that adult ecology and sexual dichromatism are poor predictors of sex-specific body size and SSD. Regardless of the groupings analyzed (partitioned by clades or traits), we found support for isometric body size scaling. However, we found that sex change is a significant predictor of SSD variation. Species in the Hyperolius viridiflavus complex, which putatively display this trait, show a significant reduction in SSD and are frequently sexually monomorphic in size. Although protogynous sex change needs to be validated for several of these species, we tentatively propose this trait is a novel mechanism influencing anuran body size evolution. Beyond this association, additional factors that shape the evolution of anuran body size and SSD remain elusive.

Mite load predicts the quality of sexual color and locomotor performance in a sexually dichromatic lizard.

Since Darwin, the maintenance of bright sexual colors has recurrently been linked to mate preference. However, the mechanisms underpinning such preferences for bright colors would not be resolved for another century. Likely, the idea of selection for colors that could decrease the chances of survival (e.g., flashy colors that can inadvertently attract predators) was perceived as counterintuitive. It is now widely accepted that these extreme colors often communicate to mates the ability to survive despite a "handicap" and act as honest signals of individual quality when they are correlated with the quality of other traits that are directly linked to individual fitness. Sexual colors in males are frequently perceived as indicators of infection resistance, in particular. Still, there remains considerable discord among studies attempting to parse the relationships between the variables associating sexual color and infection resistance, such as habitat type and body size. This discord may arise from complex interactions between these variables. Here, we ask if sexual color in male Florida scrub lizards (Sceloporus woodi) is an honest signal of resistance to chigger mite infection. To this end, we use linear modeling to explore relationships between mite load, different components of sexual color, ecological performance, body size, and habitat type. Our data show that that the brightness of sexual color in scrub lizards is negatively associated with the interaction between mite load and body size, and scrub lizards suffer decreased endurance capacity with increases in mite load. Our data also indicate that mite load, performance, and sexual color in male scrub lizards can vary between habitat types. Collectively, these results suggest that sexual color in scrub lizards is an honest indicator of individual quality and further underscore the importance of considering multiple factors when testing hypotheses related to the maintenance of sexual color.