Plasticity Of Pigmentation Research Articles

The H3K79me3 methyl-transferase Grappa is involved in the establishment and thermal plasticity of abdominal pigmentation in Drosophila melanogaster females

Temperature sensitivity of abdominal pigmentation in Drosophila melanogaster females allows to investigate the mechanisms underlying phenotypic plasticity. Thermal plasticity of pigmentation is due to modulation of tan and yellow expression, encoding pigmentation enzymes. Furthermore, modulation of tan expression by temperature is correlated to the variation of the active histone mark H3K4me3 on its promoter. Here, we test the role of the DotCom complex, which methylates H3K79, another active mark, in establishment and plasticity of pigmentation. We show that several components of the DotCom complex are involved in the establishment of abdominal pigmentation. In particular, Grappa, the catalytic unit of this complex, plays opposite roles on pigmentation at distinct developmental stages. Indeed, its down-regulation from larval L2 to L3 stages increases female adult pigmentation, whereas its down-regulation during the second half of the pupal stage decreases adult pigmentation. These opposite effects are correlated to the regulation of distinct pigmentation genes by Grappa: yellow repression for the early role and tan activation for the late one. Lastly, reaction norms measuring pigmentation along temperature in mutants for subunits of the DotCom complex reveal that this complex is not only involved in the establishment of female abdominal pigmentation but also in its plasticity.

Genetic basis of variation in thermal developmental plasticity for Drosophila melanogaster body pigmentation.

Seasonal differences in insect pigmentation are attributed to the influence of ambient temperature on pigmentation development. This thermal plasticity is adaptive and heritable, and thereby capable of evolving. However, the specific genes contributing to the variation in plasticity that can drive its evolution remain largely unknown. To address this, we analysed pigmentation and pigmentation plasticity in Drosophila melanogaster. We measured two components of pigmentation in the thorax and abdomen: overall darkness and the proportion of length covered by darker pattern elements (a trident in the thorax and bands in the abdomen) in females from two developmental temperatures (17 or 28°C) and 191 genotypes. Using a GWAS approach to identify the genetic basis of variation in pigmentation and its response to temperature, we identified numerous dispersed QTLs, including some mapping to melanogenesis genes (yellow, ebony, and tan). Remarkably, we observed limited overlap between QTLs for variation within specific temperatures and those influencing thermal plasticity, as well as minimal overlap between plasticity QTLs across pigmentation components and across body parts. For most traits, consistent with selection favouring the retention of plasticity, we found that lower plasticity alleles were often at lower frequencies. The functional analysis of selected candidate QTLs and pigmentation genes largely confirmed their contributions to variation in pigmentation and/or pigmentation plasticity. Overall, our study reveals the existence and underlying basis of extensive and trait-specific genetic variation for pigmentation and pigmentation plasticity, offering a rich reservoir of raw material for natural selection to shape the evolution of these traits independently.

The adaptive role of melanin plasticity in thermally variable environments.

Understanding the evolution of adaptive plasticity is fundamental to our knowledge of how organisms interact with their environments and cope with environmental change. Plasticity in melanin pigmentation is common in response to variable environments, especially thermal environments. Yet, the adaptive significance of melanin plasticity in thermally variable environments is often assumed, but rarely explicitly tested. Furthermore, understanding the role of plasticity when a trait is responsive to multiple environmental stimuli and plays many functional roles remains poorly understood. We test the hypothesis that melanin plasticity is an adaptation for thermally variable environments using Hyles lineata, the white-lined sphinx moth, which shows plasticity in melanin pigmentation during the larval stage. Melanin pigmentation influences thermal traits in H. lineata, as melanic individuals had higher heating rates and reached higher body temperatures than non-melanic individuals. Importantly, melanin pigmentation has temperature specific fitness consequences. While melanic individuals had an advantage in cold temperatures, neither phenotype had a clear fitness advantage at warm temperatures. Thus, the costs associated with melanin production may be unrelated to thermal context. Our results highlight the importance of explicitly testing the adaptive role of plasticity and considering all the factors that influence costs and benefits of plastic phenotypes across environments.

Background matching through fast and reversible melanin-based pigmentation plasticity in tadpoles comes with morphological and antioxidant changes

Facultative colour change is widespread in the animal kingdom, and has been documented in many distantly related amphibians. However, experimental data testing the extent of facultative colour change, and associated physiological and morphological implications are comparatively scarce. Background matching in the face of spatial and temporal environmental variation is thought to be an important proximate function of colour change in aquatic amphibian larvae. This is particularly relevant for species with long larval periods such as the western spadefoot toad, Pelobates cultripes, whose tadpoles spend up to six months developing in temporary waterbodies with temporally variable vegetation. By rearing tadpoles on different coloured backgrounds, we show that P. cultripes larvae can regulate pigmentation to track fine-grained differences in background brightness, but not hue or saturation. We found that colour change is rapid, reversible, and primarily achieved through changes in the quantity of eumelanin in the skin. We show that this increased eumelanin production and/or maintenance is also correlated with changes in morphology and oxidative stress, with more pigmented tadpoles growing larger tail fins and having an improved redox status.

Elevational divergence in pigmentation plasticity is associated with selection and pigment biochemistry.

Phenotypic plasticity is predicted to evolve in environmentally variable habitats, or those experiencing a high frequency of strong selection. The evolution of plasticity may however be constrained by costs or physiological limitations. In flowers, UV-absorbing pigmentation ameliorates UV damage to pollen, and is linked with elevated UV exposure. Whether plasticity contributes to this pattern remains unclear. Petals of Argentina anserina have larger UV-absorbing petal areas at high elevations where they experience higher and more variable UV exposure than low elevations. We measured UV-induced pigmentation plasticity in high- and low-elevation populations (hereafter, "high," "low"), and selection on pigmentation via male fitness. We dissected UV pigment biochemistry using metabolomics to explore biochemical mechanisms underlying plasticity. High displayed positive UV-induced pigmentation plasticity but low lacked plasticity. Selection favored elevated pigmentation under UV in high, supporting adaptive plasticity. In high, UV absorption was conferred by flavonoids produced in one flavonoid pathway branch. However, in low, UV absorption was associated with many compounds spanning multiple branches. Elevated plasticity was thus associated with reduced pigment diversity. These results are consistent with adaptive floral pigmentation plasticity in more extreme and variable environments. We discuss how biochemical underpinnings of pigmentation may permit or constrain the evolution of pigmentation plasticity.

Aspartate-β-alanine-NBAD pathway regulates pupal melanin pigmentation plasticity of ladybird Harmonia axyridis.

Phenotypic plasticity is observed in many animal species and it is effective for them to cope with many types of environmental threats. The multicolored Asian ladybird Harmonia axyridis shows a cuticular pigmentation plasticity that can be rapidly induced by temperature changes, and in the form of changeable melanin spot patterns to adjust heat-absorbing. Here, H. axyridis with thermal stimulation were selected for determining the molecular regulations behind it. First, we confirmed the melanin level changes of H. axyridis pupa could be induced by temperature, and then screened the efficient time window for thermal sensing of H. axyridis pre-pupa; it is suggested that the late stage of pre-pupa (late stage of 4th instar larva) is the critical period to sense thermal signals and adjust its pupal melanin spot area size to adapt to upcoming thermal conditions. The Ha-ADC (aspartate decarboxylase) and Ha-ebony (NBAD synthase) of aspartate-β-alanine-NBAD pathway were then proved in regulation of cuticular melanization for pupa through RNA interference experiments; knockdown of these two genes enlarged the melanin spot size. Finally, we designed a random injection of Ha-ADC at different pre-pupal stages, to further study the regulation window during this process. Combined with all evidence observed, we suggested the spot size determination can be regulated very close to the time point of pupation, and genes of the aspartate-β-alanine-NBAD pathway play an important role at the molecular level. In brief, H. axyridis exhibits a flexible active physiological regulation through transcriptional modification to thermal changes.

Molecular Plasticity in Animal Pigmentation: Emerging Processes Underlying Color Changes.

Animal coloration has been rigorously studied and has provided morphological implications for fitness with influences over social behavior, predator-prey interactions, and sexual selection. In vertebrates, its study has developed our understanding across diverse fields ranging from behavior to molecular biology. In the search for underlying molecular mechanisms, many have taken advantage of pedigree-based and genome-wide association screens to reveal the genetic architecture responsible for pattern variation that occurs in early development. However, genetic differences do not provide a full picture of the dynamic changes in coloration that are most prevalent across vertebrates at the molecular level. Changes in coloration that occur in adulthood via phenotypic plasticity rely on various social, visual, and dietary cues independent of genetic variation. Here, I will review the contributions of pigment cell biology to animal color changes and recent studies describing their molecular underpinnings and function. In this regard, conserved epigenetic processes such as DNA methylation play a role in lending plasticity to gene regulation as it relates to chromatophore function. Lastly, I will present African cichlids as emerging models for the study of pigmentation and molecular plasticity for animal color changes. I posit that these processes, in a dialog with environmental stimuli, are important regulators of variation and the selective advantages that accompany a change in coloration for vertebrate animals.

Phenotypic plasticity in insects

Insects represent 85% of the animals. They have adapted to many environments and play a major role in ecosystems. Many insect species exhibit phenotypic plasticity. We here report on the mechanisms involved in phenotypic plasticity of different insects (aphids, migratory locust, map butterfly, honeybee) and also on the nutritional size plasticity in Drosophila and the plasticity of the wing eye-spots of the butterfly Bicyclus anynana. We also describe in more detail our work concerning the thermal plasticity of pigmentation in Drosophila. We have shown that the expression of the tan, yellow and Ddc genes, encoding enzymes of the melanin synthesis pathway, is modulated by temperature and that it is a consequence, at least in part, of the temperature-sensitive expression of the bab locus genes that repress them.

Low-latitude zooplankton pigmentation plasticity in response to multiple threats.

Crustacean copepods in high-latitude lakes frequently alter their pigmentation facultatively to defend themselves against prevailing threats, such as solar ultraviolet radiation (UVR) and visually oriented predators. Strong seasonality in those environments promotes phenotypic plasticity. To date, no one has investigated whether low-latitude copepods, experiencing continuous stress from UVR and predation threats, exhibit similar inducible defences. We here investigated the pigmentation levels of Bahamian ‘blue hole’ copepods, addressing this deficit. Examining several populations varying in predation risk, we found the lowest levels of pigmentation in the population experiencing the highest predation pressure. In a laboratory experiment, we found that, in contrast with our predictions, copepods from these relatively constant environments did show some changes in pigmentation subsequent to the removal of UVR; however, exposure to water from different predation regimes induced minor and idiosyncratic pigmentation change. Our findings suggest that low-latitude zooplankton in inland environments may exhibit reduced, but non-zero, levels of phenotypic plasticity compared with their high-latitude counterparts.

The role of plasticity in the evolution of cryptic pigmentation in a freshwater isopod.

Cryptic pigmentation of prey is often thought to evolve in response to predator-mediated selection, but pigmentation traits can also be plastic, and change with respect to both abiotic and biotic environmental conditions. In such cases, identifying the presence of, and drivers of trait plasticity is useful for understanding the evolution of crypsis. Previous work suggests that cryptic pigmentation of freshwater isopods (Asellus aquaticus) has evolved in response to predation pressure by fish in habitats with varying macrophyte cover and coloration. However, macrophytes can potentially influence the distribution of pigmentation by altering not only habitat-specific predation susceptibility, but also dietary resources and abiotic conditions. The goals of this study were to experimentally test how two putative agents of selection, namely macrophytes and fish, affect the pigmentation of A.aquaticus, and to assess whether pigmentation is plastic, using a diet manipulation in a common garden. We performed two experiments: (a) in an outdoor mesocosm experiment, we investigated how different densities of predatory fish (0/30/60 three-spined stickleback [Gasterosteus aculeatus] per mesocosm) and macrophytes (presence/absence) affected the abundance, pigmentation and body size structure of isopod populations. (b) In a subsequent laboratory experiment, we reared isopods in a common garden experiment on two different food sources (high/low protein content) to test whether variation in pigmentation of isopods can be explained by diet-based developmental plasticity. We found that fish presence strongly reduced isopod densities, particularly in the absence of macrophytes, but had no effect on pigmentation or size structure of the populations. However, we found that isopods showed consistently higher pigmentation in the presence of macrophytes, regardless of fish presence or absence. Our laboratory experiment, in which we manipulated the protein content of the isopods' diet, revealed strong plasticity of pigmentation and weak plasticity of growth rate. The combined results of both experiments suggest that pigmentation of A.aquaticus is a developmentally plastic trait and that multiple environmental factors (e.g. macrophytes, diet and predation) might jointly influence the evolution of cryptic pigmentation of A.aquaticus in nature on relatively short time-scales.

Bric à brac (bab), a central player in the gene regulatory network that mediates thermal plasticity of pigmentation in Drosophila melanogaster.

Drosophila body pigmentation has emerged as a major Evo-Devo model. Using two Drosophila melanogaster lines, Dark and Pale, selected from a natural population, we analyse here the interaction between genetic variation and environmental factors to produce this complex trait. Indeed, pigmentation varies with genotype in natural populations and is sensitive to temperature during development. We demonstrate that the bric à brac (bab) genes, that are differentially expressed between the two lines and whose expression levels vary with temperature, participate in the pigmentation difference between the Dark and Pale lines. The two lines differ in a bab regulatory sequence, the dimorphic element (called here bDE). Both bDE alleles are temperature-sensitive, but the activity of the bDE allele from the Dark line is lower than that of the bDE allele from the Pale line. Our results suggest that this difference could partly be due to differential regulation by AbdB. bab has been previously reported to be a repressor of abdominal pigmentation. We show here that one of its targets in this process is the pigmentation gene tan (t), regulated via the tan abdominal enhancer (t_MSE). Furthermore, t expression is strongly modulated by temperature in the two lines. Thus, temperature sensitivity of t expression is at least partly a consequence of bab thermal transcriptional plasticity. We therefore propose that a gene regulatory network integrating both genetic variation and temperature sensitivity modulates female abdominal pigmentation. Interestingly, both bDE and t_MSE were previously shown to have been recurrently involved in abdominal pigmentation evolution in drosophilids. We propose that the environmental sensitivity of these enhancers has turned them into evolutionary hotspots.

Thermal Physiology and Developmental Plasticity of Pigmentation in the Harlequin Bug (Hemiptera: Pentatomidae).

Traits that promote the maintenance of body temperatures within an optimal range provide advantages to ectothermic species. Pigmentation plasticity is found in many insects and enhances thermoregulatory potential as increased melanization can result in greater heat retention. The thermal melanism hypothesis predicts that species with developmental plasticity will have darker pigmentation in colder environments, which can be an important adaptation for temperate species experiencing seasonal variation in climate. The harlequin bug (Murgantia histrionica, Hemiptera: Pentatomidae, Hahn 1834) is a widespread invasive crop pest with variable patterning where developmental plasticity in melanization could affect performance. To investigate the impact of temperature and photoperiod on melanization and size, nymphs were reared under two temperatures and two photoperiods simulating summer and fall seasons. The size and degree of melanization of adults were quantified using digital imagery. To assess the effect of coloration on the amount of heat absorption, we monitored the temperature of adults in a heating experiment. Overall, our results supported the thermal melanism hypothesis and temperature had a comparatively larger effect on coloration and size than photoperiod. When heated, the body temperature of individuals with darker pigmentation increased more relative to the ambient air temperature than individuals with lighter pigmentation. These results suggest that colder temperatures experienced late in the season can induce developmental plasticity for a phenotype that improves thermoregulation in this species. Our work highlights environmental signals and consequences for individual performance due to thermal melanism in a common invasive species, where capacity to respond to changing environments is likely contributing to its spread.

Modulation of yellow expression contributes to thermal plasticity of female abdominal pigmentation in Drosophila melanogaster

Phenotypic plasticity describes the ability of a given genotype to produce distinct phenotypes in different environments. We use the temperature sensitivity of abdominal pigmentation in Drosophila melanogaster females as a model to analyse the effect of the environment on development. We reported previously that thermal plasticity of abdominal pigmentation in females involves the pigmentation gene tan (t). However, the expression of the pigmentation gene yellow (y) was also modulated by temperature in the abdominal epidermis of pharate females. We investigate here the contribution of y to female abdominal pigmentation plasticity. First, we show that y is required for the production of black Dopamine-melanin. Then, using in situ hybridization, we show that the expression of y is strongly modulated by temperature in the abdominal epidermis of pharate females but not in bristles. Interestingly, these two expression patterns are known to be controlled by distinct enhancers. However, the activity of the y-wing-body epidermal enhancer only partially mediates the effect of temperature suggesting that additional regulatory sequences are involved. In addition, we show that y and t co-expression is needed to induce strong black pigmentation indicating that y contributes to female abdominal pigmentation plasticity.

Pigmentation plasticity enhances crypsis in larval newts: associated metabolic cost and background choice behaviour

In heterogeneous environments, the capacity for colour change can be a valuable adaptation enhancing crypsis against predators. Alternatively, organisms might achieve concealment by evolving preferences for backgrounds that match their visual traits, thus avoiding the costs of plasticity. Here we examined the degree of plasticity in pigmentation of newt larvae (Lissotriton boscai) in relation to predation risk. Furthermore, we tested for associated metabolic costs and pigmentation-dependent background choice behaviour. Newt larvae expressed substantial changes in pigmentation so that light, high-reflecting environment induced depigmentation whereas dark, low-reflecting environment induced pigmentation in just three days of exposure. Induced pigmentation was completely reversible upon switching microhabitats. Predator cues, however, did not enhance cryptic phenotypes, suggesting that environmental albedo induces changes in pigmentation improving concealment regardless of the perceived predation risk. Metabolic rate was higher in heavily pigmented individuals from dark environments, indicating a high energetic requirement of pigmentation that could impose a constraint to larval camouflage in dim habitats. Finally, we found partial evidence for larvae selecting backgrounds matching their induced phenotypes. However, in the presence of predator cues, larvae increased the time spent in light environments, which may reflect a escape response towards shallow waters rather than an attempt at increasing crypsis.

Phenotypic Plasticity through Transcriptional Regulation of the Evolutionary Hotspot Gene tan in Drosophila melanogaster.

Phenotypic plasticity is the ability of a given genotype to produce different phenotypes in response to distinct environmental conditions. Phenotypic plasticity can be adaptive. Furthermore, it is thought to facilitate evolution. Although phenotypic plasticity is a widespread phenomenon, its molecular mechanisms are only beginning to be unravelled. Environmental conditions can affect gene expression through modification of chromatin structure, mainly via histone modifications, nucleosome remodelling or DNA methylation, suggesting that phenotypic plasticity might partly be due to chromatin plasticity. As a model of phenotypic plasticity, we study abdominal pigmentation of Drosophila melanogaster females, which is temperature sensitive. Abdominal pigmentation is indeed darker in females grown at 18°C than at 29°C. This phenomenon is thought to be adaptive as the dark pigmentation produced at lower temperature increases body temperature. We show here that temperature modulates the expression of tan (t), a pigmentation gene involved in melanin production. t is expressed 7 times more at 18°C than at 29°C in female abdominal epidermis. Genetic experiments show that modulation of t expression by temperature is essential for female abdominal pigmentation plasticity. Temperature modulates the activity of an enhancer of t without modifying compaction of its chromatin or level of the active histone mark H3K27ac. By contrast, the active mark H3K4me3 on the t promoter is strongly modulated by temperature. The H3K4 methyl-transferase involved in this process is likely Trithorax, as we show that it regulates t expression and the H3K4me3 level on the t promoter and also participates in female pigmentation and its plasticity. Interestingly, t was previously shown to be involved in inter-individual variation of female abdominal pigmentation in Drosophila melanogaster, and in abdominal pigmentation divergence between Drosophila species. Sensitivity of t expression to environmental conditions might therefore give more substrate for selection, explaining why this gene has frequently been involved in evolution of pigmentation.

The phenotypic plasticity of developmental modules.

BackgroundOrganisms develop and evolve in a modular fashion, but how individual modules interact with the environment remains poorly understood. Phenotypically plastic traits are often under selection, and studies are needed to address how traits respond to the environment in a modular fashion. In this study, tissue-specific plasticity of melanic spots was examined in the large milkweed bug, Oncopeltus fasciatus.ResultsAlthough the size of the abdominal melanic bands varied according to rearing temperatures, wing melanic bands were more robust. To explore the regulation of abdominal pigmentation plasticity, candidate genes involved in abdominal melanic spot patterning and biosynthesis of melanin were analyzed. While the knockdown of dopa decarboxylase (Ddc) led to lighter pigmentation in both the wings and the abdomen, the shape of the melanic elements remained unaffected. Although the knockdown of Abdominal-B (Abd-B) partially phenocopied the low-temperature phenotype, the abdominal bands were still sensitive to temperature shifts. These observations suggest that regulators downstream of Abd-B but upstream of DDC are responsible for the temperature response of the abdomen. Ablation of wings led to the regeneration of a smaller wing with reduced melanic bands that were shifted proximally. In addition, the knockdown of the Wnt signaling nuclear effector genes, armadillo1 and armadillo 2, altered both the melanic bands and the wing shape. Thus, the pleiotropic effects of Wnt signaling may constrain the amount of plasticity in wing melanic bands.ConclusionsWe propose that when traits are regulated by distinct pre-patterning mechanisms, they can respond to the environment in a modular fashion, whereas when the environment impacts developmental regulators that are shared between different modules, phenotypic plasticity can manifest as a developmentally integrated system.Electronic supplementary materialThe online version of this article (doi:10.1186/s13227-016-0053-7) contains supplementary material, which is available to authorized users.

The costs of colour: plasticity of melanin pigmentation in an outbreaking polymorphic forest moth

AbstractOutside the context of industrial melanism, little is known about the physiological and ecological importance of genetic melanic polymorphisms in moths. Melanin pigments are synthesized from amino acid precursors and should therefore be costly to produce in nitrogen‐limited insects. A genetic melanic polymorphism is present in adult Malacosoma disstria Hübner (Lepidoptera: Lasiocampidae), a widespread forest moth with outbreaking population dynamics. We test the hypotheses that melanin‐based colouration is physiologically costly in M. disstria, that expression of melanin‐based colouration is a plastic trait which varies with population density and nutrition, and that the genetically based melanic phenotype is disadvantaged under nutritionally poor conditions. Two experiments were used to test these hypotheses. A field study compared pigmentation and phenotypic frequencies in moths collected from high‐ and low‐density populations. A laboratory experiment investigated the effects of larval nitrogen availability on adult pigmentation and phenotypic frequencies. High population density and nitrogen limitation reduced pigmentation and size of all moths, but phenotypic frequencies were not affected in either experiment. The effects of diet on both pigmentation and size were stronger for melanic moths than for typical moths. Our results show that adult melanism in M. disstria is physiologically costly, that colour expression is plastic despite its genetic component, and that the melanic phenotype may be disadvantaged under poor conditions but favoured under good conditions. We suggest that temporal variation in selection and trait plasticity help maintain polymorphism stability.

Trait‐specific consequences of inbreeding on adaptive phenotypic plasticity

Environmental changes may stress organisms and stimulate an adaptive phenotypic response. Effects of inbreeding often interact with the environment and can decrease fitness of inbred individuals exposed to stress more so than that of outbred individuals. Such an interaction may stem from a reduced ability of inbred individuals to respond plastically to environmental stress; however, this hypothesis has rarely been tested. In this study, we mimicked the genetic constitution of natural inbred populations by rearing replicate Drosophila melanogaster populations for 25 generations at a reduced population size (10 individuals). The replicate inbred populations, as well as control populations reared at a population size of 500, were exposed to a benign developmental temperature and two developmental temperatures at the lower and upper margins of their viable range. Flies developed at the three temperatures were assessed for traits known to vary across temperatures, namely abdominal pigmentation, wing size, and wing shape. We found no significant difference in phenotypic plasticity in pigmentation or in wing size between inbred and control populations, but a significantly higher plasticity in wing shape across temperatures in inbred compared to control populations. Given that the norms of reaction for the noninbred control populations are adaptive, we conclude that a reduced ability to induce an adaptive phenotypic response to temperature changes is not a general consequence of inbreeding and thus not a general explanation of inbreeding–environment interaction effects on fitness components.

Phenotypic and genetic variation within the Cricotopus sylvestris species-group (Diptera, Chironomidae), across a Nearctic - Palaearctic gradient

Intraspecific variation sometimes obscures species boundaries and makes identification of certain Chironomidae difficult. This is true for many species in the genus Cricotopus. We used DNA barcode data and multivariate statistical analyses to investigate which phenotypic characters in populations of the Cricotopus sylvestris species-group are useful for species identification. Specimens collected across a broad latitudinal range from the Southwest United States through subarctic Canada to northern Norway formed nine distinct barcode clusters. Body size of adult flies decreased by 51% from the northern to southernmost populations. Meristic characters in wings and legs were strongly related to overall body size, and related morphometric ratios were not species specific. Antennal ratio increased significantly with body size, thus limiting its value in species delimitation. Non-metric ordinations of setal and coloration pattern data were characteristic for most species in the sylvestris-group. DNA barcode data worked well in separating morphologically different populations, except for the case of C. (I.) sylvestris and C. (I.) trifasciatus, which were distinguished by ordination of color pattern, but not by barcoding data. These two species appeared closely related, and we conclude that sequence data from neutral nuclear markers will be necessary to determine if these are genetically distinct species, or whether there is merely a high level of environmental plasticity in pigmentation within this geographically widespread barcode cluster.doi: 10.5324/fn.v31i0.1417.Published online: 17 October 2012.

Zooplankton vertical migration and plasticity of pigmentation arising from simultaneous UV and predation threats

We assessed how zooplankton (copepods) handle the simultaneous threats of predators and ultraviolet (UV) radiation and whether they respond with changes in pigmentation, vertical migration, or both. We found weak vertical migration among copepods in response to UV stress, and this response was not apparently influenced by predation risk. Exposure to high levels of UV radiation caused copepods to retain pigments in the absence of a predation threat. When exposed to predation threat, they reduced their pigmentation regardless of UV level. Thus, they ranked predation as a threat more severe than UV radiation. Reducing the protective pigment level in response to predation in a situation in which UV radiation is high may, however, lead to higher mortality.