Maintenance Of Colour Polymorphism Research Articles

Territorial status is explained by covariation between boldness, exploration, and thermal preference in a colour polymorphic lizard.

Colour polymorphic species often exhibit variation in morphology, physiology, and behaviour among morphs. In particular, dominance status may be signalled by the interaction between behaviour and colour morph. Behavioural traits associated with dominance include boldness, exploration, and aggression, which influence access to preferred habitat, territorial defence, and mate acquisition. In ectotherms, the social structure associated with morphs may result in the exploitation of structural niches differing in thermal quality. Hence, social interactions among morphs may generate concordant variation in thermal preference and environmental temperature. However, few studies have assessed thermal preference variation in colour polymorphic species and its covariation with behaviour. Doing so can provide insight into niche specialization and the maintenance of colour polymorphism in populations. Here, we investigated the patterns of covariation in boldness behaviour, exploratory behaviour, and thermal preference in the tree lizard, Urosaurus ornatus. We assessed trait variation between territorial and non-territorial male morphs and between orange and yellow female morphs. Boldness and exploratory behaviour were repeatable in male U. ornatus and bolder individuals were significantly more likely to incur tail loss, a potential consequence of bold behaviour. Territorial male morphs were significantly bolder and more exploratory and preferred higher body temperatures with a narrower T set than non-territorial morphs. Female morphs did not vary in behavioural or thermal traits. This study highlights behavioural mechanisms that underly ecological niche segregation and variable habitat use between morphs in a colour polymorphic species.

Pleiotropic effects of melanin pigmentation: haemoparasite infection intensity but not telomere length is associated with plumage morph in black sparrowhawks.

There is increasing recognition of the potential pleiotropic effects of melanin pigmentation, particularly on immunity, with reports of variation in haemoparasite infection intensity and immune responses between the morphs of colour-polymorphic bird species. In a population of the black sparrowhawk (Accipiter melanoleucus) in western South Africa, light morphs have a higher haemoparasite infection intensity, but no physiological effects of this are apparent. Here, we investigate the possible effects of haemoparasite infection on telomere length in this species and explore whether relative telomere length is associated with either plumage morph or sex. Using quantitative polymerase chain reaction analysis, we confirmed that dark morphs had a lower haemoparasite infection intensity than light morphs. However, we found no differences in telomere length associated with either the haemoparasite infection status or morph in adults, although males have longer telomeres than females. While differences in haemoparasite intensity between morphs are consistent with pleiotropic effects of melanin pigmentation in the black sparrowhawk, we found no evidence that telomere length was associated with haemoparasite infection. Further work is needed to investigate the implications of possible pleiotropic effects of plumage morph and their potential role in the maintenance of colour polymorphism in this species.

Colour-specific diet specialization is associated with differences in owlet weight in a polymorphic owl: influence of the trophic quality variation.

The niche divergence hypothesis proposes that the evolution and maintenance of colour polymorphism is based on a mechanism of disruptive selection. In a trophic context, the hypothesis predicts that individuals differing in colour vary in their trophic niche, either because they differ in foraging efficiency or feed in different habitats. A major evolutionary conundrum is how these expectations are affected by variation in trophic quality. Using an owl species with colour plumage polymorphism, the Eurasian scops owl Otus scops, we examined diet and habitat segregation during reproduction in relation to plumage colouration and trophic quality. Intensive sampling revealed that trophic quality for scops owls (i.e. abundance of grasshoppers and locusts) varied more among territories than between years, but scops owls did not segregate among territories of different quality by their colouration. However, we found that sex, plumage colouration and territory differences in trophic quality explained differences in the degree of dietary specialization. Brownish males delivered a higher diversity of prey to the nest than greyish ones in high trophic quality territories. We also found that the more diverse the diet provided by males, the heavier the owlets at fledging. Our study provides evidence for a different sensitivity to trophic quality of the colour morphs with potential fitness consequences in scops owls. We highlight the importance of studying the mechanisms leading to the persistence of colour polymorphism in patchy environments, since segregation may pass otherwise unnoticed if only habitats or years with similar conditions are considered.

Some like it hotter: Differential thermal preferences among lizard color morphs

Temperature rules the lives of ectotherms. To perform basic biological functions, ectotherms must make behavioral adjustments to keep their body temperatures near a preferred temperature (Tpref). Many color polymorphic lizards are active thermoregulators and exhibit morph differences in traits related to thermoregulation, such as color, body size, and microhabitat use. The Aegean wall lizard, Podarcis erhardii, is a heliothermic lizard with orange, white, and yellow color morphs that differ in size, behavior, and microhabitat use. Here, we tested whether P. erhardii color morphs from the same population from Naxos island, Greece, differ in Tpref. We hypothesized that orange morphs would prefer lower temperatures than white and yellow morphs because orange morphs are often found on cooler substrates and in microhabitats with more vegetation cover. We obtained Tpref for 95 individuals using laboratory thermal gradient experiments of wild-caught lizards and found that orange morphs do, indeed, prefer cooler temperatures. Average orange morph Tpref was 2.85 °C lower than average white and yellow morph Tpref. Our results add support to the idea that P. erhardii color morphs have multivariate alternative phenotypes and present the possibility that thermally heterogeneous environments play a role in the maintenance of color polymorphism in this species.

Color polymorphism and variation in microhabitat selection: a defensive strategy in males of Boana pulchella (Anura: Hylidae)?

Anurans vary in body coloration and frequently exhibit color polymorphism, with selection by predation pressure favoring more cryptic animals. Spatial differentiation may also favor color polymorphism in some species, since cryptic morphotypes can use substrates that better match their coloration. Boana pulchella is a polymorphic species with green and brown morphotypes, but its polymorphism remains poorly understood. The aim of the present study was to determine the functionality of color polymorphism in B. pulchella as a defense strategy. Artificial green and brown clay models were placed in environments with green and brown background to determine if the two morphotypes of B. pulchella experience different frequencies of attacks. We expected that more conspicuous morphs are attacked more frequently. The attack rate, which suggests predation intent, on the models was 9.57%. The marks observed in the artificial models were left by both mammals and birds. The number of damaged models did not significantly differ between the different background environments. The results indicate that B. pulchella morphotypes are equally cryptic to predators in the studied environments. In conclusion, probably color polymorphism in B. pulchella is not operating as a defensive strategy, and other selective forces may be acting in the maintenance of color polymorphism in this species.

Morph-specific differences in escape behavior in a color polymorphic lizard

Variation in color morph behavior is an important factor in the maintenance of color polymorphism. Alternative anti-predator behaviors are often associated with morphological traits such as coloration, possibly because predator-mediated viability selection favors certain combinations of anti-predator behavior and color. The Aegean wall lizard, Podarcis erhardii, is color polymorphic and populations can have up to three monochromatic morphs: orange, yellow, and white. We investigated whether escape behaviors differ among coexisting color morphs, and if morph behaviors are repeatable across different populations with the same predator species. Specifically, we assessed color morph flight initiation distance (FID), distance to the nearest refuge (DNR), and distance to chosen refuge (DR) in two populations of Aegean wall lizards from Naxos island. We also analyzed the type of refugia color morphs selected and their re-emergence behavior following a standardized approach. We found that orange morphs have different escape behaviors from white and yellow morphs, and these differences are consistent in both populations we sampled. Orange morphs have shorter FIDs, DNRs, and DRs; select different refuge types; and re-emerge less often after being approached compared to white and yellow morphs. Observed differences in color morph escape behaviors support the idea that morphs have evolved alternative behavioral strategies that may play a role in population-level morph maintenance and loss.Significance statementColor polymorphic species often differ in behaviors related to reproduction, but differences in other behaviors are relatively underexplored. In this study, we use an experimental approach in two natural populations of color populations of color polymorphic lizards to determine that color morphs have diverged in their escape behaviors. By conducting our experiments in two different populations with similar predator regimes, we show for the first time that behavioral differences among intra-specific color morphs are repeatable across populations, suggesting that alternative behavioral strategies have evolved in this species. Using this experimental approach, we demonstrate that the brightest orange morph stays closer to refuge than other morphs, uses a different refuge type (vegetation) more often than other morphs (wall crevices), and take much longer to emerge from refuge after a simulated predation event than other morphs. Thus, selective pressures from visual predators may differ between morphs and play a role in the evolution and maintenance of color polymorphisms in these types of systems. Our study species, Podarcis erhardii, belongs to a highly color polymorphic genus (19/23 spp. are color polymorphic) that contains the same three color morphs, thus we believe our results may be relevant to more than just P. erhardii.

Density-dependent selection and the maintenance of colour polymorphism in barn owls.

The capacity of natural selection to generate adaptive changes is (according to the fundamental theorem of natural selection) proportional to the additive genetic variance in fitness. In spite of its importance for development of new adaptations to a changing environment, processes affecting the magnitude of the genetic variance in fitness-related traits are poorly understood. Here, we show that the red-white colour polymorphism in female barn owls is subject to density-dependent selection at the phenotypic and genotypic level. The diallelic melanocortin-1 receptor gene explained a large amount of the phenotypic variance in reddish coloration in the females ([Formula: see text]). Red individuals (RR genotype) were selected for at low densities, while white individuals (WW genotype) were favoured at high densities and were less sensitive to changes in density. We show that this density-dependent selection favours white individuals and predicts fixation of the white allele in this population at longer time scales without immigration or other selective forces. Still, fluctuating population density will cause selection to fluctuate and periodically favour red individuals. These results suggest how balancing selection caused by fluctuations in population density can be a general mechanism affecting the level of additive genetic variance in natural populations.

Conspicuous colours in a polymorphic orb-web spider: evidence of predator avoidance but not prey attraction

Conspicuous colours in animals are usually explained by mate attraction, predator avoidance and prey attraction. Predators may lure prey by mimicking food or other resources. Alternatively, some animals avoid predation by visually signalling unpalatability to predators. The evolution of a signal is affected by receiver processing mechanisms. Therefore, variation in sensory systems may explain the evolution of colour polymorphism of conspicuous species. We aimed to investigate the role of receivers in the evolution and maintenance of colour conspicuousness and polymorphism in Gasteracantha cancriformis (Araneidae). We tested two hypotheses: (1) conspicuous coloration is a prey lure strategy; (2) conspicuous coloration is a warning signal to predators. Furthermore, we identified the taxonomic order of prey captured and the type of predator (avian versus arthropod). To assess the role of prey, we conducted a field experiment in which webs were assigned to one of four treatments: yellow, red or black spider models, or no model. For each treatment, we counted the number of prey trapped on the webs and calculated the damaged area of the webs. To assess the role of predators, we conducted a second field experiment in which we attached yellow, red and black models to trees using nylon thread and observed the number of attack markings on each model. Our results did not corroborate the prey attraction hypothesis. In the predation experiment, black models had more attack markings, indicating that yellow and red models were less preferred. This suggests that spider coloration may play a role in predator avoidance. Colour polymorphism in this species is possibly a multifunctional attribute, where some morphs benefit from aposematism, whereas others may have alternative fitness advantages. Nonadaptive explanations should also be considered in future experiments of the evolution and maintenance of colour polymorphisms.

No evidence for differential sociosexual behavior and space use in the color morphs of the European common wall lizard (Podarcis muralis).

Explaining the evolutionary origin and maintenance of color polymorphisms is a major challenge in evolutionary biology. Such polymorphisms are commonly thought to reflect the existence of alternative behavioral or life‐history strategies under negative frequency‐dependent selection. The European common wall lizard Podarcis muralis exhibits a striking ventral color polymorphism that has been intensely studied and is often assumed to reflect alternative reproductive strategies, similar to the iconic “rock–paper–scissors” system described in the North American lizard Uta stansburiana. However, available studies so far have ignored central aspects in the behavioral ecology of this species that are crucial to assess the existence of alternative reproductive strategies. Here, we try to fill this gap by studying the social behavior, space use, and reproductive performance of lizards showing different color morphs, both in a free‐ranging population from the eastern Pyrenees and in ten experimental mesocosm enclosures. In the natural population, we found no differences between morphs in site fidelity, space use, or male–female spatial overlap. Likewise, color morph was irrelevant to sociosexual behavior, space use, and reproductive success within experimental enclosures. Our results contradict the commonly held hypothesis that P. muralis morphs reflect alternative behavioral strategies, and suggest that we should instead turn our attention to alternative functional explanations.

Understanding Color Polymorphism in the Sea Star, Pisaster ochraceus, Through Activity Levels and Dietary Preference

Color polymorphism is found in a wide array of organisms ranging from copepods to black bears. Pisaster ochraceus, an intertidal sea star on the North Pacific West Coast, shows a striking color polymorphism including a range of orange, brown, maroon, and purple. Pisaster shows extensive geographic color variation, with >95% frequency of bright purple Pisaster in the Vancouver region; while the open pacific coast has a frequency distribution of 6-28% orange, 68-90% reddish-brown to dull purple, and a small percentage of bright purple. Maintenance of color polymorphism remains unknown, but one hypothesis suggests an underlying genetic component with regional-scale variation controlled by an ecological factor. Two ecological factors suggested are diet and salinity. This study analyzes the differences in dietary preference and activity levels between the color morphs in Bamfield, British Columbia. Pisaster was binned into two colors, orange and purple, for statistical analysis. Feeding laboratory trials showed no preference between the color morphs for Mytilus edilus or Mytilus californianus. Self-righting trials were performed in a low (20 psu) and controlled (35 psu) salinity and no significant difference was found between the color morphs. Interestingly, the activity levels of orange color morphs were not significantly different between low and control salinity. The results suggest there are ecological differences between the color morphs that still need to be evaluated.

Melanin-based coloration and immunity in polymorphic population of pied flycatcher, Ficedula hypoleuca

A specific interest in the persistence of color polymorphism in some populations of birds and other vertebrates is often linked to ideas about the signaling honesty of bright coloration. The evolution of conspicuous ornamentation could be associated with physiological costs including limitations of the immune system. The study of this process is crucial for an understanding of the maintenance of polymorphic coloration. Here we summarized the results of a study of a pied flycatcher population from the Moscow region (Russia) in 2010–2013. We experimentally induced antibody production by injecting sheep red blood cells (SRBC) and inflammatory swelling by injecting phytohaemagglutinin (PHA) after which we estimated the immune response in breeding males. We used leucocytes-to-erythrocytes and heterophils-to-lymphocytes (H/L) ratios as indicators of infectious, inflammatory processes and stress. The results showed that the feeding rates of males treated with SRBC decreased and negatively related to the intensity of their immune responses. Non-molting males of different color types did not significantly differ in antibody production. Among molting breeders, the immune response to SRBC was significantly higher in pale males than in bright ones with rich melanin-based coloration. In contrast to non-molting males, molting pale males had an increased antibody titer after immunization. The lower humoral immune response was associated with the higher H/L stress index before immunization. The change in H/L after immunization positively correlated with the intensity of the humoral immune response. As opposed to humoral immunity, we did not find any significant predictors, including coloration, molt, or their two-way interaction, to explain the variation in cutaneous inflammatory response to PHA. The results suggest that the apparent advantage of a cryptic male phenotype over a conspicuous phenotype occurring in one of two types of immune response has an impact on the maintenance of color polymorphism in the pied flycatcher.

The effect of colour polymorphism on thermoregulation in an orb web spider

Spiders that build aerial webs in open areas face the risk of overheating due to incident solar radiation. These spiders can counteract overheating by either moving the web to another site or by adopting behavioural thermoregulation within the web. Since moving can be costly, studies have suggested that a passive but effective method of reducing heat load is by light reflectance through body colouration. We explored the interaction between colour and thermoregulation in a colour polymorphic species, under both field and laboratory conditions. We show that in natural conditions, the spiders maintain their body temperature below that of the ambient, but with no difference in surface temperature between colour morphs. In laboratory experiments with internal temperature measurements, white morphs bore the risk of overheating better than the yellow morphs since they heated up slower and cooled faster. We suggest that the thermoregulatory properties of colour polymorphism in Verrucosa arenata have physiological consequences and may play an important role in the maintenance of colour polymorphism in this species.

Differential foraging success across a light level spectrum explains the maintenance and spatial structure of colour morphs in a polymorphic bird.

Detectability of different colour morphs under varying light conditions has been proposed as an important driver in the maintenance of colour polymorphism via disruptive selection. To date, no studies have tested whether different morphs have selective advantages under differing light conditions. We tested this hypothesis in the black sparrowhawk, a polymorphic raptor exhibiting a discrete white and dark morph, and found that prey provisioning rates differ between the morphs depending on light condition. Dark morphs delivered more prey in lower light conditions, while white morphs provided more prey in brighter conditions. We found support for the role of breeding season light level in explaining the clinal pattern of variation in morph ratio across the species range throughout South Africa. Our results provide the first empirical evidence supporting the hypothesis that polymorphism in a species, and the spatial structuring of morphs across its distribution, may be driven by differential selective advantage via improved crypsis, under varying light conditions.

Behavioural differences across contexts may indicate morph-specific strategies in the lizard Ctenophorus decresii

Discrete colour polymorphisms are often genetically correlated with other traits under natural and sexual selection, such as behaviour, life history and physiology. Elucidating such correlations is essential to understand the adoption of alternative strategies between morphs and the role they play in the maintenance of colour polymorphisms within a population. Using field experiments, we tested the hypothesis that four visually discrete morphs (orange, yellow, yellow with a central orange patch (orange-yellow) and grey) of the tawny dragon lizard, Ctenophorus decresii, display alternative behavioural strategies. Specifically, we compared the response of colour morphs to simulated conspecific territorial intruders and predators in the wild. Although the orange-yellow morph can be objectively classified, it may behaviourally resemble the orange or yellow morph; therefore we compared statistical models in which the orange-yellow morph was considered a separate morph (four-morph model) or grouped with either pure orange or pure yellow individuals (three-morph models). For aggression, a three-morph model with orange-yellow individuals grouped as yellow morphs best fitted the data. The orange morph showed consistently high aggression to all morphs, while the grey morph showed consistently low aggression. Aggression of the yellow morph was conditional on the morph of the intruder. In addition to being the least aggressive, the grey morph was the least bold. Although the orange morph was the most aggressive, it was only the boldest under a three-morph model, which was equally likely compared to a four-morph model. Overall our results support the view that tawny dragon lizard morphs adopt different behavioural strategies, the orange and grey morphs exhibiting more aggressive and cautious strategies, respectively, and the yellow morph changing its aggression depending on its competitor's colour.

Variation in male fertility in a polymorphic moth, Parasemia plantaginis

The maintenance of multiple morphs in warning signals is enigmatic because directional selection through predator avoidance should lead to the rapid loss of such variation. Opposing natural and sexual selection is a good candidate driving the maintenance of multiple male morphs but it also includes another enigma: when warning signal efficiency differs between male morphs, why would females choose a phenotype with lower survival? We tested the hypothesis that indirect responses to selection on correlated characters through sexual selection may substantially shape the evolution of male coloration. If male phenotypes differ in their fertilization ability, female choice against the best surviving phenotype can evolve. The wood tiger moth, Parasemia plantaginis , has two coexisting male morphs in Europe. Previous studies have shown that yellow males are better defended against predators, but that white males have a higher mating success. We examined differences in fertility between white and yellow males in terms of sperm production, number of sperm transferred and rate of sperm replenishment, and association between these fertility traits, female mate choice and reproductive output. If white morphs have greater fertility than yellow males, then this could explain why females prefer to mate with white males. However, we did not find any difference between male colour morphs either in mating probability, fertility (i.e. sperm availability and sperm transferred) or reproductive success (i.e. number of eggs laid and hatching success). We discuss our results in relation to context-dependent mating success and maintenance of colour polymorphism within populations.

Background matching ability and the maintenance of a colour polymorphism in the red devil cichlid

The evolution and maintenance of colour polymorphisms remains a topic of considerable research interest. One key mechanism thought to contribute to the coexistence of different colour morphs is a bias in how conspicuous they are to visual predators. Although individuals of many species camouflage themselves against their background to avoid predation, differently coloured individuals within a species may vary in their capacity to do so. However, to date, very few studies have explicitly investigated the ability of different colour morphs to plastically adjust their colouration to match their background. The red devil (Amphilophus labiatus) is a Neotropical cichlid fish with a stable colour polymorphism, with the gold morph being genetically dominant and having a myriad of documented advantages over the dark morph. However, gold individuals are much rarer, which may be related to their heightened conspicuousness to would-be predators. Here, we tested the ability of differently coloured individuals to phenotypically adjust the shade of their body colour and patterns to match their background. In particular, we filmed dark, gold and mottled (a transitioning phase from dark to gold) individuals under an identical set-up on light vs. dark-coloured substrates. We found that, in contrast to individuals of the dark morph, gold and mottled individuals were less capable of matching their body colouration to their background. As a result, gold individuals appeared to be more conspicuous. These results suggest that a difference in background matching ability could play an important role in the maintenance of colour polymorphisms.

Sensory limitations and the maintenance of colour polymorphisms: viewing the ‘alba’ female polymorphism through the visual system of male Colias butterflies

Summary Although colour polymorphisms are a widespread and conspicuous component of extant biodiversity, the selective pressures that act to maintain multiple morphs within populations remain poorly understood in most cases. In particular, the role that visual system limitations may play in maintaining multiple colour morphs is not well explored. We used a female‐limited colour polymorphism common to the butterfly genus Colias, called the ‘alba’ polymorphism, to investigate the hypotheses that mate‐searching males may struggle to discriminate pale ‘alba’ females from co‐occurring heterospecific white butterflies and/or heterospecific ‘alba’ females, or that ‘alba’ females may be more difficult to detect than non‘alba’ females in natural scenes. Such perceptual limitations may influence the relative mating rates of ‘alba’ versus non‘alba’ females, contributing to the evolutionary persistence of both morphs. Based on receptor‐noise‐limited modelling of the male Colias visual system, we find that ‘alba’ females exhibit chromatic and luminance contrasts against background foliage that are most similar to the ‘alba’ females of other co‐occurring Colias species and females of the co‐occurring white butterfly Pieris rapae. When compared to other co‐flying butterflies including non‘alba’ females, ‘alba’ females are consistently lower in chromatic contrast against background, but higher in luminance contrast. When viewed side‐by‐side, we estimate that male Colias should be able to discriminate ‘alba’ females from other co‐occurring heterospecific butterflies, including heterospecific ‘albas’. However, under field conditions that involve larger distances in space or time, males are likely to face challenges discriminating between conspecific ‘alba’ females and co‐occurring heterospecific white butterflies, particularly heterospecific ‘alba’ females. Our results suggest that constraints arising from male visual function may be involved in the maintenance of this colour polymorphism, particularly in populations that co‐occur with other ‘alba’‐polymorphic Colias species. We argue that such visual system constraints may play a larger role in the maintenance of colour polymorphism than has been empirically appreciated to date.

Multiple Paternity Increases Phenotypic Diversity inTetrix subulataPygmy Grasshoppers

Multiple paternity within clutches has been recorded among a variety of organisms. The degree of genetic similarity between parents may influence the number and viability of offspring. Females may therefore mate with several males as an insurance against sterile, low quality or genetically incompatible mates, but also to obtain half sibling offspring that are genetically and phenotypically more diverse. We examine the links between polyandry, multiple paternity and offspring phenotypic diversity in the color polymorphic pygmy grasshopper Tetrix subulata. By experimentally mating virgin females and genotyping the resulting offspring using microsatellite markers, we demonstrate that polyandrous females can produce offspring sired by different males. Analyses of microsatellite data and color patterns of captive reared families produced by wild caught females that were not mated in the laboratory, confirmed that multiple paternity occurs in the wild, and that it may increase color morph diversity among half-siblings. Polyandrous mating behavior may thus influence the evolutionary dynamics and maintenance of color polymorphism in this species.

Abiotic factors may explain the geographical distribution of flower colour morphs and the maintenance of colour polymorphism in the scarlet pimpernel

SummaryFlower colour polymorphism is traditionally attributed to pollinator selection although other factors, such as indirect selection on correlated traits, can play an important role.Lysimachia arvensisis a widespread annual species with two colour morphs differing in anthocyanin composition. We explored the hypothesis that colour polymorphism is maintained by selection related to environmental heterogeneity.Morph frequencies and environmental traits were recorded in 51 populations along a wide geographical range. To explore the existence of morph‐by‐environment interactions, we conducted an experimental study comparing the two morphs under treatments differing in water and light availability.A geographical pattern was found with a negative association between blue frequencies and latitude. The proportion of the blue morph increased with temperature and sunshine hours, but decreased with precipitation. Flowering onset and flower size differed between morphs and scarcely varied across treatments. In contrast, several fitness components such as germination, seedling survival, seedling mass and flower production showed important morph‐by‐environment interactions. The blue morph showed higher overall male and female fitness in all the treatment combinations excepting in sun‐wet conditions where the red morph had higher fitness.Synthesis. Our results indicate that the mechanism of selection on flower colour seems to be related to differences in fitness of both morphs due to abiotic factors. These differences could explain the geographical distribution of flower colour morphs and the maintenance of the colour polymorphism. The marked difference in flowering time between morphs leaves open the potential for assortative mating and speciation inLysimachia arvensis.

A potential role for parasites in the maintenance of color polymorphism in urban birds

Urbanization is a major challenge for biodiversity conservation, yet the evolutionary processes taking place in urbanized areas remain poorly known. Human activities in cities set new selective forces in motion which need to be investigated to predict the evolutionary responses of animal species living in urban areas. In this study, we investigated the role of urbanization and parasites in the maintenance of melanin-based color polymorphism in the feral pigeon Columba livia. Using a correlative approach, we tested whether differently colored genotypes displayed alternative phenotypic responses to urbanization, by comparing body condition, blood parasite prevalence and parasite load between colored morphs along an urbanization gradient. Body condition did not vary with urbanization, but paler individuals had a higher body condition than darker individuals. Moreover, paler morphs were less often parasitized than darker morphs in moderately urbanized habitats, but their parasite prevalence increased with urbanization. In contrast, darker morphs had similar parasite prevalence along the urbanization gradient. This suggests that paler morphs did better than darker morphs in moderately urbanized environments but were negatively affected by increasing urbanization, while darker morphs performed equally in all environments. Thus, differently colored individuals were distributed non-randomly across the urban habitat and suffered different parasite risk according to their location (a gene-by-environment interaction). This suggests that melanin-based coloration might reflect alternative strategies to cope with urbanization via different exposure or susceptibility to parasites. Spatial variability of parasite pressures linked with urbanization may, thus, play a central role in the maintenance of plumage color polymorphism in this urban species.