Genetic and Ecological Characterisation of Colour Dimorphism in a Coral Reef Fish

Synopsis Many recognised species of coral reef fishes exhibit two or more colour variants, but it is unknown whether these represent genetically identical phenotypes, genetic polymorphisms or closely related species. We tested between these alternatives for two colour morphs of the coral reef fish, Pseudochromis fuscus, from Lizard Island (Great Barrier Reef). A molecular analysis using mtDNA did not detect any genetic differentiation between co-occurring ‘yellow’ and ‘brown’ colour morphs. A previous study proposed that these two colour morphs are aggressive mimics of yellow and brown damselfishes. Here, a manipulative field experiment was used to evaluate whether the colour dimorphism in P. fuscus is a phenotypic response to the presence of two different model species. Colonies of either yellow or brown damselfish species were established on different patch reefs, and each of the two different P. fuscus morphs was then placed on the different reefs. Contrary to expectations, all yellow individuals that stayed on the reefs changed to brown, regardless of the model species. No brown individuals changed to the yellow colouration. However, P. fuscus were more likely to emigrate from, or suffer higher mortality on, patch reefs where they were not matched with similarly coloured models. Clearly, yellow and brown P. fuscus are members of a single species with sufficient phenotypic plasticity to switch from yellow to brown colouration.     

Phylogeography of colour polymorphism in the coral reef fish Pseudochromis fuscus, from Papua New Guinea and the Great Barrier Reef

Body colour has played a significant role in the evolution of coral reef fishes, but the phylogenetic level at which colour variation is expressed and the evolutionary processes driving the development and persistence of different colour patterns are often poorly understood. The aim of this study was to examine the genetic relationships between multiple colour morphs of Pseudochromis fuscus (family Pseudochromidae), both within and among geographic locations. Pseudochromis fuscus is currently described as a single species, but exhibits at least six discrete colour morphs throughout its range. In this study, P. fuscus from Papua New Guinea (PNG) and the Great Barrier Reef (GBR), Australia, formed three genetically distinct clades based on mitochondrial DNA (control region) sequence data: (1) yellow and brown morphs from the GBR and southern PNG, as well as an orange morph from southern PNG; (2) a pink morph from southern PNG; and (3) all three morphs (pink, orange and grey) found in Kimbe Bay, northern PNG. The three groups showed deep levels of divergence (d=14.6–25.4%), suggesting that P. fuscus is a complex of polychromatic species, rather than a single widespread species with many different colour morphs. Population genetic analyses indicate that the three clades have experienced unique evolutionary histories, possibly from differential effects of sea level fluctuations, barriers to gene flow and historical biogeography.     

Comparative life history and parasitism of a new colour morph of the walnut aphid in California

1 The walnut aphid Chromaphis juglandicola is a yellow aphid. In 2003, however, a white colour morph was discovered in the Sacramento Valley of California. The colour dimorphism occurs between clone lines and, when white morphs are present, they occur in mixed colour morph colonies on the underside of walnut leaves. 2 Laboratory experiments were undertaken to evaluate the thermal requirements for development, adult longevity and progeny production of the two colour morphs. Host instar preference of Trioxys pallidus, a parasitoid responsible for the successful biological control of the walnut aphid in California, was examined separately for each colour morph, and host colour preference was investigated for the preferred instar. 3 No differences in thermal requirements for development, adult size or mean longevity were detected between yellow and white colour morphs. A small difference in early reproduction was detected: white colour morphs produced more progeny on each of the two first days of adult reproduction than yellow colour morphs. 4 Trioxys pallidus showed a slight preference for the fourth instar of the yellow morph over the second‐ and third‐, but equal preference for second, third and fourth instars of the white morph. When offered equal numbers of fourth instars of the two colour morphs, T. pallidus did not show any colour preference. 5 The differences in early aphid reproduction and host instar preference by T. pallidus were combined in a stage‐structured matrix model. Model analysis showed a greater potential for population growth of the white morph over the yellow morph, with early reproduction having a greater influence than host instar preference.     

Evidence for reproductive isolation between two colour morphs of cavity nesting honey bees (Apis) in south India

In south India there are two distinct colour morphs of cavity nesting honey bees: the yellow ‘Plain’ morph and the black ‘Hill’ morph which are collectively known as Apis cerana. We show that the Hill morph is associated with a widely distributed mitochondrial haplotype that is present throughout mainland populations of south east Asian A. cerana. In contrast, the Plain morph, which is apparently confined to low to moderate elevations in India and Sri Lanka, is associated with a unique mitochondrial haplotype that is not present in other cavity nesting honey bees. We further show that in a region of sympatry (Bangalore, Karnataka State) the drone mating flight times of the two colour morphs barely overlap. Combined, drone flight data and the complete separation of mitochondrial haplotypes suggest that the two morphs are reproductively isolated. The Plain morph is distinguished from the Hill morph by the first three abdominal tergites of the worker, which are completely yellow in the Plain morph, whereas in the Hill morph they are black or black with yellow patches. Although the two morphs are generally distinguishable in the field by overall colouration, microscopic examination of the first 3 abdominal tergites is preferred. Received 16 February 2006; revised 11 May 2006; accepted 23 May 2006.     

Colour morph related performance in the meadow grasshopper Chorthippus parallelus (Orthoptera,Acrididae)

Abstract 1. Polymorphism has been described for a number of herbivorous insects, but little is known about whether differences in body colour cause fitness differences. In Chorthippus parallelus, three main colour morphs occur, namely brown, green, and dorsally striped. 2. The present study examined colour morph abundances and morph‐related differences in body size, oviposition rate, and offspring numbers in females of C. parallelus collected in 15 montane grasslands. The study also examined the effect of plant species richness, composition, community productivity, and solar radiation on colour morph frequency and fitness. 3. The relative frequencies of the three colour morphs was 31.7% (brown), 33.1% (green), and 35.2% (dorsally striped), but the morphs were not evenly distributed across the 15 sites. 4. There was no effect of the habitat variables on the distribution of the green and the striped morph in the study sites, however 80% of the variation in the abundances of the brown morph was explained by plant species richness and composition. 5. Grasshopper size was equal among the morphs. Brown females laid significantly more egg pods than the green and dorsally striped morphs. There were no significant differences in offspring numbers among the colour morphs. 6. Body colour in C. parallelus seems to be a fitness‐relevant trait, raising the question of the evolutionary maintenance of polymorphism.     

Substrate preference in a colonial spider: is substrate choice affected by color morph?

Animals can show preference for a particular background as a way of decreasing visibility. Species with color polymorphism may have morph‐dependent background preference. I test this hypothesis on the orb‐weaving spider Parawixia bistriata. Adult females of P. bistriata present two distinct morphs characterized by brown and yellow opisthosomata. This nocturnal spider can be found in its retreat on the vegetation during the day. In order to examine whether females exhibit substrate preference dependent on their color morph, I first recorded the distribution of color morphs on different substrates (leaf and branch) and then performed a mark and release experiment. Field censuses indicated that the yellow morph was associated with leaves while the brown morph was found on either substrate type. The results of a mark and reciprocal release experiment agreed with the censuses and suggest that the two morphs differ in their association to substrate type: yellow females were associated with the leaf substrate, while brown females showed no association to a particular substrate type. Possible forces behind these differences in substrate choice are discussed.     

GENETIC AND COLOR INTERACTIONS AT A CONTACT ZONE OF ACANTHOCHROMIS POLYACANTHUS: A MARINE FISH LACKING PELAGIC LARVAE

Abstract. — Acanthochromis polyacanthus is an unusual tropical marine damselfish that uniquely lacks pelagic larvae and has lost the capacity for broad-scale dispersal among coral reefs. Different color morphs exist in different regions of the Great Barrier Reef, and morphs from northern and southern regions are genetically distinct. In the Hydrographers Passage area, which is a large break through the reef matrix in the central Great Barrier Reef that may have acted as a bottleneck on the migration of these animals during sea level rise, three morphs recognized from other regions were found on neighboring reefs. The transition between them is abrupt with three loci (AAT-2*, GPI-1*, and PGM*) showing allelic frequency patterns close to fixation between opposite alleles within a few kilometers. On two reefs (Hyde, Bebe), a pair of morphs was found to coexist and exhibited a habitat partitioning pattern with each morph restricted to one side on the reef and steep transitions in between. Outside these transition zones, phenotypes and genotypes matched those on surrounding reefs without coexistence and were little changed from reefs several hundred kilometers away. An electrophoretic survey across one transition zone on Hyde Reef showed steep genetic gradients along one kilometer of reef slope. Significant linkage disequilibria in samples collected in Hyde Reef as a result of dispersal of parental combinations of alleles into the center or because parental combinations of alleles confer greater fitness, allowed us to estimate the dispersal rate (189 m/generation) and the selection pressure on the marker loci (0.411). Finally, we investigated models that could lead to such a steep transition in genotypic and phenotypic combinations. Both contact zones on each side of Hyde Reef were associated with geomorphological discontinuities in the reef structure. We suggest that assortative mating may be a proximal mechanism for maintaining isolated each color morph, which could be reinforced by selective predation against hybrids outside the zone of their formation (i.e., the frequency-dependent selection model of Mallet and Barton (1989). Acanthochromis is a midwater planktivore and, when in coexistence, the two morphs forage in different habitats amid multispecific flocks of other damselfishes of matching colors.     

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.     

Genetically distinct colour morphs of European perch Perca fluviatilis in Lake Constance differ in susceptibility to macroparasites

The unusual yellow‐finned morph of European perch Perca fluviatilis found in Lake Constance suffers more severely from macroparasite infections, including the tapeworm Triaenophorus nodulosus and the gill worm Ancyrocephalus percae, than conspecifics elsewhere. Microsatellite analysis of yellow‐finned P. fluviatilis and red‐finned variant recently discovered in Lake Constance revealed significant genetic differentiation. Red‐finned P. fluviatilis and fish with mixed fin colour, suggested backcrosses between red and yellow‐finned colour morphs, exhibit better resilience to parasite infection, suggesting that the inability of the yellow‐finned morph to reject macroparasites may have a genetic basis.     

Intraspecific variation of flower colour and its distribution within a sea lavender, Limonium wrightii (Plumbaginaceae), in the northwestern Pacific Islands

Differentiation of flower colour is thought to be one of the most important factors promoting plant speciation. We describe the intraspecific variation of flower colour and its distribution in Limonium wrightii. We conducted a survey on 36 islands in the northwestern Pacific and discriminated six morphs of flower colour variation. Two flower colour morphs, pink and yellow, were most frequently observed, and their geographical distributions were basically allopatric. These two morphs were in contact in a narrow zone on Okinoerabu Island, located in the middle region of the Ryukyu Archipelago. In addition, orange, white, and ivory flower morphs were also found in this zone. The geographical distribution of pink and yellow morphs showed a “leapfrog” pattern; the distribution of pink flowers was divided into two areas, intercalated by the distribution of the yellow flower morph. The orange morph may have resulted from hybridization between the pink and yellow flower morphs.     

Influence of shell colour on the mortality rate of the land snail Arianta arbustorum under different microclimatic regimes

The mortality of phenotypic shell colour morphs and age classes of the snail Arianta arbustorum in several microclimatic conditions was recorded. An analysis of variance was performed on five factors: adaptation temperature, relative humidity, shell colour, age class and test temperature. There were no significant differences in the mortality between different adaptation temperatures or relative humidities, but the interaction of these two factors was highly significant. There were significant differences in mortality rate between test temperature and age class. The mortality of the brown morph was higher than the yellow one at all adaptation temperatures (overall, P < 0.05). There were no significant differences between the mortality rates of the two morphs at different relative humidities. The mortality of the brown morph was higher than that of the yellow at six out of seven test temperatures. Juvenile snails survived significantly better than adults at all test temperatures.     

Effects of local density and flower colour polymorphism on pollination and reproduction in the rewardless orchid Dactylorhiza sambucina (L.) Soò

The rewardless orchid Dactylorhiza sambucina shows a stable flower colour polymorphism, with both yellow- and red-flowered morphs growing sympatrically. Pollination biology and breeding system were investigated to examine the effects of density of plants, colour polymorphism, inflorescence dimension, and flower position within inflorescence on male and female reproductive success in three natural populations of D. sambucina. There were significant differences among sites in the number of pollinia removed and in fruit set per inflorescence. Number of removed pollinia and capsule production in D. sambucina were independent from flower and inflorescence size or flower position. As a whole, the red morphs showed the highest number of capsules produced, while the yellow morphs had the greatest male success. The relative male and female reproductive success were independent from plant density but were significantly correlated with the yellow morph frequency at the population level. Overall, our findings show that the contribution to the total reproductive success deriving from the two colour morphs does not conform with the predictions of negative frequency-dependent selection.     

Pollen competition between morphs in a pollen‐color dimorphic herb and the loss of phenotypic polymorphism within populations

Flower color polymorphism is relatively uncommon in natural flowering plants, suggesting that maintenance of different color morphs within populations is difficult. To address the selective mechanisms shaping pollen‐color dimorphism, pollinator preferences and reproductive performance were studied over three years in Epimedium pubescens in which some populations had plants with either green or yellow pollen (and anthers). Visitation rate and pollen removal and receipt by the bee pollinator (Andrena emeishanica) did not differ between the two color morphs. Compared to the green morph, siring success of the yellow morph's pollen was lower, but that of mixtures of pollen from green and yellow morphs was lowest. This difference, corresponding to in vivo and ex vivo experiments on pollen performance, indicated that pollen germination, rather than tube growth, of the green morph was higher than that of the yellow morph and was seriously constrained in both morphs if a pollen competitor was present. A rare green morph may invade a yellow‐morph population, but the coexistence of pollen color variants is complicated by the reduced siring success of mixed pollinations. Potential pollen competition between morphs may have discouraged the maintenance of multiple phenotypes within populations, a cryptic mechanism of competitive exclusion.     

Contents of ultraviolet-absorbing substances in two color morphs of the photosymbiotic ascidian Didemnum molle

The contents of mycosporine-like amino acids (MAAs) were compared in the two color morphs (dark-gray and brown colonies) of the tropical ascidian Didemnum molle (Herdman, 1886), which harbors the photosymbiotic prokaryote Prochloron. The colonies of each color morph were exclusively distributed in shallow reef lagoons at the different sites. Spectroscopic and chromatographic analyses showed that the Prochloron cell density and MAA concentration in the dark-gray colonies were an estimated 1.4 and 2.4 times higher, respectively, than in the brown colonies. The significant difference in MAA contents between the color morphs was primarily due to the difference in shinorine contents (p < 0.01, Mann–Whitney U-test). The high concentration of MAAs in the dark-gray colonies may provide better conditions for Prochloron cells, compared to the brown colonies with lower MAA concentrations.     

Shell colour polymorphism in a mangrove snail Littorina sp. (Prosobranchia: Littorinidae)

Shell colour polymorphism was examined in populations of a mangrove snail Littorina sp. in Queensland, Australia. Three morphs were recognized, yellow, red and brown, and morph frequencies varied both between widely spaced populations and between islands less than 1 km apart. Morph frequencies also varied with time of year. There was a relationship between shell colour and position on the tree, with yellow snails more often occurring amongst the foliage and brown snails more often on trunks and branches. In some populations yellow snails appeared to survive better than the other morphs, while in other populations there was no difference. The evidence for the maintenance of the polymorphism by natural selection is discussed.     

Interaction between female mating preferences and predation may explain the maintenance of rare males in the pentamorphic fish Poecilia parae

Variation in mating preferences coupled with selective predation may allow for the maintenance of alternative mating strategies. Males of the South American live‐bearing fish Poecilia parae fall in one of five discrete morphs: red, yellow, blue, stripe‐coloured tail (parae) and female mimic (immaculata). Field surveys indicate that the red and yellow morphs are the rarest and that their rarity is consistent across years. We explored the role of variable female mating preference and selective predation by visual predators in explaining the rarity of red and yellow males, and more generally, the maintenance of this extreme colour polymorphism. We presented wild‐caught P. parae females and Aequidens tetramerus, the most common cichlid predator, with the five male colour morphs in separate trials to determine mating and prey preferences, respectively. We found that a large proportion of females shared a strong preference for the rare carotenoid‐based red and yellow males, but a distinct group also preferred the blue and parae morphs. The cichlid predator strongly preferred red and yellow males as prey. Together, these results suggest that the interaction between premating sexual selection favouring and predation acting against the red and yellow morphs may explain their rarity in the wild. The trade‐off between sexual and natural selection, accompanied by variation in female mating preferences, may therefore facilitate the maintenance of the striking colour polymorphism in P. parae.     

Endocrine differences among colour morphs in a lizard with alternative behavioural strategies

Alternative behavioural strategies of colour morphs are expected to associate with endocrine differences and to correspond to differences in physical performance (e.g. movement speed, bite force in lizards); yet the nature of correlated physiological and performance traits in colour polymorphic species varies widely. Colour morphs of male tawny dragon lizards Ctenophorus decresii have previously been found to differ in aggressive and anti-predator behaviours. We tested whether known behavioural differences correspond to differences in circulating baseline and post-capture stress levels of androgen and corticosterone, as well as bite force (an indicator of aggressive performance) and field body temperature. Immediately after capture, the aggressive orange morph had higher circulating androgen than the grey morph or the yellow morph. Furthermore, the orange morph maintained high androgen following acute stress (30 min of capture); whereas androgen increased in the grey and yellow morphs. This may reflect the previously defined behavioural differences among morphs as the aggressive response of the yellow morph is conditional on the colour of the competitor and the grey morph shows consistently low aggression. In contrast, all morphs showed an increase in corticosterone concentration after capture stress and morphs did not differ in levels of corticosterone stress magnitude (CSM). Morphs did not differ in size- and temperature-corrected bite force but did in body temperature at capture. Differences in circulating androgen and body temperature are consistent with morph-specific behavioural strategies in C. decresii but our results indicate a complex relationship between hormones, behaviour, temperature and bite force within and between colour morphs.     

Pollination ecology ofOxalis violacea (Oxalidaceae) following a controlled grass fire

Vernal grass fires may encourage profuse flowering in clonal, colonies ofOxalis violacea. Long-styled colonies appear to be more floriferous than short-styled colonies and set a greater number of capsules. Individual flowers of both morphs live one or two days, change position on their respective pedicels and advertise nectar concealed at the base of the floral throat. AlthoughDiptera, Hymenoptera, andLepidoptera forage for nectar, bees (Andrenidae,Anthophoridae, Halictidae, andMegachilidae) probably make the only effective pollen transfers between the two morphs. Both male and female bees may transport pollen of both morphs and short-tongued bees (e.g.,Augochlorella spp.,Dialictus spp.) may be more common but as effective as pollinators as long-tongued bees (e.g.,Calliopsis andreniformis andHoplitis spp.). The conversion rate of flowers into capsules is only 13–17%. The spreading style in the short-styled morph is interpreted as an adaptation restricting insect-mediated, self-pollination but encouraging bee-stigma contact during nectar foraging.