Expression patterns of melanin-related genes are linked to crypsis and conspicuousness in a pumpkin toadlet.

Colour signals play pivotal roles in different communication systems, and the evolution of these characters has been associated with behavioural ecology, integumentary production processes and perceptual mechanisms of the species involved. Here, we present the first insight into the molecular and histological basis of skin colour polymorphism within a miniaturized species of pumpkin toadlet, potentially representing the lowest size threshold for colour polytypism in tetrapods. Brachycephalus actaeus exhibits a coloration ranging from cryptic green to conspicuous orange skin, and our findings suggest that colour morphs differ in their capability to be detected by potential predators. We also found that the distribution and abundance of chromatophores are variable in the different colour morphs. The expression pattern of coloration related genes was predominantly associated with melanin synthesis (including dct, edn1, mlana, oca2, pmel, slc24a5, tyrp1 and wnt9a). Up-regulation of melanin genes in grey, green and brown skin was associated with higher melanophore abundance than in orange skin, where xanthophores predominate. Our findings provide a significant foundation for comparing and understanding the diverse pathways that contribute to the evolution of pigment production in the skin of amphibians.

The role of historical biogeography in shaping colour morph diversity in the common wall lizard.

The maintenance of polymorphisms often depends on multiple selective forces, but less is known on the role of stochastic or historical processes in maintaining variation. The common wall lizard (Podarcis muralis) is a colour polymorphic species in which local colour morph frequencies are thought to be modulated by natural and sexual selection. Here, we used genome-wide single-nucleotide polymorphism data to investigate the relationships between morph composition and population biogeography at a regional scale, by comparing morph composition with patterns of genetic variation of 54 populations sampled across the Pyrenees. We found that genetic divergence was explained by geographic distance but not by environmental features. Differences in morph composition were associated with genetic and environmental differentiation, as well as differences in sex ratio. Thus, variation in colour morph frequencies could have arisen via historical events and/or differences in the permeability to gene flow, possibly shaped by the complex topography and environment. In agreement with this hypothesis, colour morph diversity was positively correlated with genetic diversity and rates of gene flow and inversely correlated with the likelihood of the occurrence of bottlenecks. Concurrently, we did not find conclusive evidence for selection in the two colour loci. As an illustration of these effects, we observed that populations with higher proportions of the rarer yellow and yellow-orange morphs had higher genetic diversity. Our results suggest that processes involving a decay in overall genetic diversity, such as reduced gene flow and/or bottleneck events have an important role in shaping population-specific morph composition via non-selective processes.

The Andes as a semi-permeable geographical barrier: Genetic connectivity between structured populations in a widespread spider.

Geographical barriers like mountain ranges impede genetic exchange among populations, promoting diversification. The effectiveness of these barriers in limiting gene flow varies between lineages due to each species' dispersal modes and capacities. Our understanding of how the Andes orogeny contributes to species diversification comes from well-studied vertebrates and a few arthropods and plants, neglecting organisms unable to fly or walk long distances. Some arachnids, such as Gasteracantha cancriformis, have been hypothesized to disperse long distances via ballooning (i.e. using their silk to interact with the wind). Yet, we do not know how the environment and geography shape its genetic diversity. Therefore, we tested whether the Andes contributed to the diversification of G. cancriformis acting as an absolute or semi-permeable barrier to genetic connectivity between populations of this spider at opposite sides of the mountain range. We sampled thousands of loci across the distribution of the species and implemented population genetics, phylogenetic, and landscape genetic analyses. We identified two genetically distinct groups structured by the Central Andes, and a third less structured group in the Northern Andes that shares ancestry with the previous two. This structure is largely explained by the altitude along the Andes, which decreases in some regions, possibly facilitating cross-Andean dispersal and gene flow. Our findings support that altitude in the Andes plays a major role in structuring populations in South America, but the strength of this barrier can be overcome by organisms with long-distance dispersal modes together with altitudinal depressions.

Las barreras geográficas como las cordilleras montañosas impiden el intercambio genético entre poblaciones, promoviendo la diversificación. La efectividad de estas barreras para limitar el flujo genético varía entre linajes debido a los modos y capacidades de dispersión de cada especie. Nuestra comprensión de cómo la orogenia de los Andes contribuye a la diversificación de especies proviene de vertebrados y algunos artrópodos y plantas bien estudiados, descuidando a los organismos incapaces de volar o caminar grandes distancias. Se ha hipotetizado que algunas arañas, como Gasteracantha cancriformis, se dispersan a grandes distancias mediante la técnica de "ballooning" (es decir, utilizando su seda para interactuar con el viento). Sin embargo, no sabemos cómo el entorno y la geografía han dado forma a su diversidad genética. Por lo tanto, probamos si los Andes contribuyeron a la diversificación de G. cancriformis actuando como una barrera absoluta o permeable para la conectividad genética entre poblaciones de esta araña en lados opuestos de la cordillera. Muestreamos miles de loci a través de la distribución de la especie e implementamos análisis de genética de poblaciones, filogenéticos y de genética del paisaje. Identificamos dos grupos genéticamente distintos estructurados por los Andes Centrales, y un tercer grupo menos estructurado en los Andes del Norte que comparte ascendencia con los dos anteriores. Esta estructura se explica en gran medida por la altitud a lo largo de los Andes, que disminuye en algunas regiones, posiblemente facilitando la dispersión y el flujo genético a través de los Andes. Nuestros hallazgos apoyan que la altitud en los Andes juega un papel importante en la estructuración de las poblaciones en América del Sur, pero la fuerza de esta barrera puede ser superada por organismos con modos de dispersión a larga distancia junto con depresiones altitudinales.

Habitat heterogeneity limits prey colour polymorphism maintained via negative frequency-dependent selection.

The persistence of non-neutral trait polymorphism is enigmatic because stabilizing selection is expected to deplete variation. In cryptically coloured prey, negative frequency-dependent selection due to search image formation by predators has been proposed to favour rare variants, promoting polymorphism. However, in a heterogeneous environment, locally varying disruptive selection favours patch type-specific optima, resulting in spatial segregation of colour variants. Here, we address whether negative frequency-dependent selection can overcome selection posed by habitat heterogeneity to promote local polymorphism using an individual-based model. In addition, we compare how prey and predator mobility may modify the outcome. Our model revealed that frequency-dependent predation could strongly promote local prey polymorphism, but only when differences between morphs in patch-specific fitness were small. The effect of frequency-dependent predation depended on the predator adjustment of search image and was hampered by the prey population structure. Gene flow due to prey movement counteracted local selection, promoted local polymorphism to some extent, and relaxed the conditions for polymorphism due to frequency-dependent predation. Importantly, abrupt spatial changes in morph frequencies decreased the probability that mobile frequency-dependent predators could maintain local prey polymorphism. Overall, our study suggests that in a spatially heterogeneous environment, negative frequency-dependent selection may help maintain local polymorphism but only under a limited range of conditions.

Genetic structure, UV-vision, wing coloration and size coincide with colour polymorphism in Fabriciana adippe butterflies.

Colour polymorphisms have long served as model systems in evolutionary studies and continue to inform about processes involved in the origin and dynamics of biodiversity. Modern sequencing tools allow for evaluating whether phenotypic differences between morphs reflect genetic differentiation rather than developmental plasticity, and for investigating whether polymorphisms represent intermediate stages of diversification towards speciation. We investigated phenotypic and genetic differentiation between two colour morphs of the butterfly Fabriciana adippe using a combination of ddRAD-sequencing and comparisons of body size, colour patterns and optical properties of bright wing spots. The silvery-spotted adippe form had larger and darker wings and reflected UV light, while the yellow cleodoxa form displayed more green scales and reflected very little UV, showcasing that they constitute distinct and alternative integrated phenotypes. Genomic analyses revealed genetic structuring according to source population, and to colour morph, suggesting that the phenotypic differentiation reflects evolutionary modifications. We report 17 outlier loci associated with colour morph, including ultraviolet-sensitive visual pigment (UVRh1), which is associated with intraspecific communication and mate choice in butterflies. Together with the demonstration that the wings of the adippe (but essentially not the cleodoxa) morph reflect UV light, that UV reflectance is higher in females than males and that morphs differ in wing size, this suggests that these colour morphs might represent genetically integrated phenotypes, possibly adapted to different microhabitats. We propose that non-random mating might contribute to the differentiation and maintenance of the polymorphism.

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.

Regional variation in climate change alters the range-wide distribution of colour polymorphism in a wild bird.

According to Gloger's rule, animal colouration is expected to be darker in wetter and warmer climates. Such environmental clines are predicted to occur in colour polymorphic species and to be shaped by selection if colour morphs represent adaptations to different environments. We studied if the distribution of the colour polymorphic tawny owl (Strix aluco) morphs (a pheomelanic brown and a pale grey) across Europe follow the predictions of Gloger's rule and if there is a temporal change in the geographical patterns corresponding to regional variations in climate change. We used data on tawny owl museum skin specimen collections. First, we investigated long-term spatiotemporal variation in the probability of observing the colour morphs in different climate zones. Second, we studied if the probability of observing the colour morphs was associated with general climatic conditions. Third, we studied if weather fluctuations prior to the finding year of an owl explain colour morph in each climate zone. The brown tawny owl morph was historically more common than the grey morph in every studied climate zone. Over time, the brown morph has become rarer in the temperate and Mediterranean zone, whereas it has first become rarer but then again more common in the boreal zone. Based on general climatic conditions, winter and summer temperatures were positively and negatively associated with the proportion of brown morph, respectively. Winter precipitation was negatively associated with the proportion of brown morph. The effects of 5-year means of weather on the probability to observe a brown morph differed between climate zones, indicating region-dependent effect of climate change and weather on tawny owl colouration. To conclude, tawny owl colouration does not explicitly follow Gloger's rule, implying a time and space-dependent complex system shaped by many factors. We provide novel insights into how the geographic distribution of pheomelanin-based colour polymorphism is changing.

Does thermal biology differ between two colour pattern morphs of a widespread Australian lizard?

Alternative phenotypes allow individuals to pursue different adaptive pathways in response to the same selective challenge. Colour polymorphic species with geographically varying morph frequencies may reflect multiple adaptations to spatial variables such as temperature and climate. We examined whether thermal biology differed between colour morphs of an Australian lizard, the delicate skink, Lampropholis delicata. The delicate skink has two colour pattern morphs, with frequencies varying across latitude and sex: plain (darker, more common at temperate latitudes, more common in males) or striped (lighter, more common at lower latitudes, more common in females). We tested heating and cooling rate, sprint speed, thermal preference, field body temperature and metabolic rate in both morphs and sexes to determine any link between colour and morph frequency distribution. Plain individuals heated more quickly, but other thermal traits showed little variation among morphs. Lampropholis delicata colour influences rates of heat exchange, but the relationship does not appear to be adaptive, suggesting that behavioural thermoregulation homogenises body temperature in the field. While we find no substantial evidence of thermal differences between the two colour morphs, morph-specific behaviour may buffer against differences in heat exchange. Latitudinal variation in species colour may be driven by selection pressures other than temperature.

Spatial patterns of flower colour variation in native and introduced ranges of Convolvulus arvensis (Convolvulaceae) revealed by citizen-science data and machine learning.

Flower colour polymorphism refers to the presence of multiple colour variants in plant populations. Investigation of this phenomenon led to multiple discoveries, including the principles of heredity and the foundations of population genetics. I examined flower colour variation across native and introduced ranges of Convolvulus arvensis, which exhibits flower colour polymorphism (individuals have white or pink petals). To study flower colour variation of this species throughout large geographic scale, I used observations gathered from the iNaturalist platform. To handle a large amount of data, I trained a neural network to classify the plants' morphs based on photographs. After which I performed spatial analyses to examine the patterns of the colour frequency, also in relation to environmental factors. The results show that flower colours are polymorphic across the whole species range, but the frequency of pink versus white flowers varies. In the Palearctic, I observed geographic clines of colour morph frequencies: a higher frequency of the pink morph in populations from Northwest Europe, whereas in South and East Europe, towards the eastern edge of the range, the white morph was dominant. In contrast, pattern of colour distribution in North America (where the species is invasive) seems random, but the model indicates a link between higher proportions of pink morphs in mild and humid climates. The mechanisms behind the observed patterns remain largely unknown, as changes in a morphs' frequency are not strongly linked to abiotic factors. To understand the spatial pattern, a detailed investigation, accounting for the species' phylogeography is needed. This study provides another example of how the general public may collect data relevant to ecological studies, even when the data are not collected for a specific project.

Differential regulation of the Tor gene homolog drives the red/green pigmentation phenotype in the aphid Myzuspersicae.

In some aphid species, intraspecific variation in body colour is caused by differential carotenoid content: whilst green aphids contain only yellow carotenoids (ß-, γ-, and ß,γ-carotenes), red aphids additionally possess red carotenoids (torulene and 3,4-didehydrolycopene). Unusually, within animals who typically obtain carotenoids from their diet, ancestral horizontal gene transfer of carotenoid biosynthetic genes from fungi (followed by gene duplication), have imbued aphids with the intrinsic gene repertoire necessary to biosynthesise carotenoids. In the pea aphid, Acyrthosiphon pisum a lycopene (phytoene) desaturase gene (Tor) underpins the red/green phenotype, with this locus present in heterozygous form in red individuals but absent in green aphids, resulting in them being unable to convert lycopene into the red compounds 3,4-didehydrolycopene and torulene. The green peach aphid, Myzus persicae, separated from the pea aphid for ≈45MY also exists as distinct colour variable morphs, with both red and green individuals present. Here, we examined genomic data for both red and green morphs of M. persicae and identified an enlarged (compared to A. pisum) repertoire of 16 carotenoid biosynthetic genes (11 carotenoid desaturases and five carotenoid cyclase/synthase genes). From these, we identify the homolog of A. pisum Tor (here called carotene desaturase 2 or CDE-2) and show through 3D modelling that this homolog can accommodate the torulene precursor lycopene and, through RNA knockdown feeding experiments, demonstrate that disabling CDE-2 expression in red M. persicae clones results in green-coloured offspring. Unlike in A. pisum, we show that functional CDE-2 is present in the genomes of both red and green aphids. However, expression differences between the two colour morphs (350-700 fold CDE-2 overexpression in red clones), potentially driven by variants identified in upstream putative regulatory elements, underpin this phenotype. Thus, whilst aphids have a common origin of their carotenoid biosynthetic pathway, two aphid species separated for over 40MY have evolved very different drivers of intraspecific colour variation.

Integrative Approaches Establish Colour Polymorphism in the Bamboo-Feeding Leafhopper Mukaria splendida Distant (Hemiptera: Cicadellidae) from India.

The leafhopper species, Mukaria splendida Distant, is economically important due to itsstatus as a pest on bamboo and was recently reported to bea vector for phytoplasmal disease. Morphological identification is often difficult and requires a high level of taxonomic expertise, with misidentifications causing problematic false-positive/negative results. In this paper, colour polymorphism was recorded in the bamboo-feedingleafhopper M. splendida (Distant, 1908), which is a major insect pest in the bamboo ecosystem based on explorations conducted in different locations of India. Ten morphs were identified for each sex of M. splendida Distant based on the colour pattern on the pronotum and forewings. However, in view of the economic importance of the species, the morphological studies were integrated with the molecular data for the accurate identification of the species. The morphological characteristics and sequence results of the amplified product of the mitochondrial cytochrome oxidase subunit I (mtCOI) gene confirmed that all the morphs were M. splendida and the pairwise distance matrix showed a negligible genetic distance in the COI mtDNA gene. Simulated future predictions, along with detailed notes on the colour polymorphic forms with illustrations, and additional distribution records as well as thebiology of M. splendida were discussed in light of the available literature, all of which will aid the authentic identification of this species.

Beyond morphs: Inter-individual colour variation despite strong genetic determinism of colour morphs in a wild bird.

Categorizing individuals into discrete forms in colour polymorphic species can overlook more subtle patterns in coloration that can be of functional significance. Thus, quantifying inter-individual variation in these species at both within- and between-morph levels is critical to understand the evolution of colour polymorphisms. Here we present analyses of inter-individual colour variation in the Reunion grey white-eye (Zosterops borbonicus), a colour polymorphic wild bird endemic to the island of Reunion in which all highland populations contain two sympatric colour morphs, with birds showing predominantly grey or brown plumage, respectively. We first quantified colour variation across multiple body areas by using a continuous plumage colour score to assess variation in brown-grey coloration as well as smaller scale variation in light patches. To examine the possible causes of among-individual variation, we tested if colour variation in plumage component elements could be explained by genotypes at two markers near a major-effect locus previously related to back coloration in this species, and by other factors such as age, sex and body condition. Overall, grey-brown coloration was largely determined by genetic factors and was best described by three distinct clusters that were associated to genotypic classes (homozygotes and heterozygote), with no effect of age or sex, whereas variation in smaller light patches was primarily related to age and sex. Our results highlight the importance of characterizing subtle plumage variation beyond morph categories that are readily observable since multiple patterns of colour variation may be driven by different mechanisms, have different functions and will likely respond in different ways to selection.

Cardiac performance and heat shock response variation related to shell colour morphs in the mudflat snail Batillaria attramentaria.

Gastropods exhibit remarkable variation in shell colour within and among populations, but the function of shell colour is often not clear. In the present study, body temperature in the field and physiological and transcriptomic responses to thermal stress were investigated in different shell colour morphs of the mudflat snail Batillaria attramentaria. Using biomimetic models, we found that the body temperature of snails with a dark unbanded shell (D-type morph) was slightly higher than that of snails with a white line on the upper side of each whorl (UL-type morph) when exposed to sunlight. Despite no differences in upper lethal temperature among shell colour morphs, their Arrhenius breakpoint temperature (ABT) for cardiac thermal performance differed significantly, and the ABT of snails with the D-type morph was higher than that of snails with the UL-type morph. Transcriptomic analysis showed that D-type snails exhibit higher levels of four heat shock proteins (HSPs) than UL-type snails at control temperature. The unfolded protein response was activated in UL-type snails but not in D-type snails under moderate thermal stress. And 11 HSPs showed an increase in UL-type snails in contrast to 1 HSP in D-type snails, suggesting a 'preparative defence' strategy of the heat shock response in D-type snails under moderate thermal stress. When exposed to sublethal temperature, eight molecular chaperones were uniquely upregulated in D-type snails, suggesting these genes may allow D-type snails to improve their cardiac thermal tolerance. Our results suggest that the preparative defence strategies and higher ABT for cardiac thermal performance may allow the dark shell snails to adapt to rapid and stronger thermal stress in the field.

Colour Variation in the Crocodile Lizard (Shinisaurus crocodilurus) and Its Relationship to Individual Quality.

Colour plays a key role in animal social communication including as an indicator of individual quality. Using spectrophotometry, we examined colour variation in the throat and venter of the crocodile lizard (Shinisaurus crocodilurus), an endangered species native to southern China and northern Vietnam. We detected two broad colour variants, individuals with and without red, for each body region and each sex. A cluster analysis of spectral colour measurements (hue, chroma, luminance) revealed discrete throat and ventral morphs when measured in a single snapshot in time. However, photographic evidence revealed that the amount of red relative to body size increased as they got older. Individuals with red were equally likely to be male or female and throat colour was unrelated to ventral colour. Therefore, it is premature to claim that crocodile lizards have discrete colour morphs. We used visual modelling to show that the throat and venter were easily discriminable to a lizard visual system, suggesting they function in social communication. We also asked whether colour variation signalled individual quality. Females with red throats had greater bite force while males with red throats were older. In addition, females with red venters had larger heads. We also detected differences in morphology linked to colour. Females with red throats had slender bodies and longer tails, while individuals lacking red on their throats were stouter and had shorter tails. Finally, throat and ventral colour were unrelated to reproductive output (litter size and mass) in females. Males with greater ventral luminance contrast sired offspring from litters with greater litter mass (including stillborns), while males with greater ventral chromatic contrast sired offspring whose collective live mass (excluding stillborns) was greater. Males with greater luminance contrast also sired more live offspring (excluding stillborns). Collectively, these results suggest that male ventral colour signals individual quality in males. Conservation initiatives should take colour variation into account when planning future captive breeding and release programs for this endangered species.

Biology and ecology of the Oriental flower-breeding Drosophila elegans and related species.

Animals adapt to their environments in the course of evolution. One effective approach to elucidate mechanisms of adaptive evolution is to compare closely related species with model organisms in which knowledge of the molecular and physiological bases of various traits has been accumulated. Drosophila elegans and its close relatives, belonging to the same species group as the model organism D. melanogaster, exhibit various unique characteristics such as flower-breeding habit, courtship display, territoriality, sexual dimorphism, and colour polymorphism. Their ease of culturing and availability of genomic information makes them a useful model for understanding mechanisms of adaptive evolution. Here, we review the morphology, distribution, and phylogenetic relationships of D. elegans and related species, as well as their characteristic flower-dependent biology, food habits, and life-history traits. We also describe their unique mating and territorial behaviours and note their distinctive karyotype and the genetic mechanisms of morphological diversity that have recently been revealed.

Multispecies colour polymorphisms associated with contrasting microhabitats in two Mediterranean wrasse radiations.

Intraspecific colour polymorphisms (CPs) present unique opportunities to study fundamental evolutionary questions, such as the link between ecology and phenotype, mechanisms maintaining genetic diversity and their putative role in speciation. Wrasses are highly diverse in ecology and morphology and harbour a variety of colour-polymorphic species. In the Mediterranean Sea, wrasses of the tribe Labrini evolved two species radiations each harbouring several species with a brown and a green morph. The colour morphs occur in complete sympatry in mosaic habitats with rocky outcrops and Neptune grass patches. Morph-specific differences had not been characterized yet and the evolutionary forces maintaining them remained unknown. With genome-wide data for almost all Labrini species, we show that species with CPs are distributed across the phylogeny, but show evidence of hybridization. This suggests that the colour morphs are either ancient and have been lost repeatedly, that they have evolved repeatedly or have been shared via hybridization. Focusing on two polymorphic species, we find that each colour morph is more common in the microhabitat providing the best colour match and that the morphs exhibit additional behavioural and morphological differences further improving crypsis in their respective microhabitats. We find little evidence for genetic differentiation between the morphs in either species. Therefore, we propose that these colour morphs represent a multi-niche polymorphism as an adaptation to the highly heterogeneous habitat. Our study highlights how colour polymorphism (CP) can be advantageous in mosaic habitats and that Mediterranean wrasses are an ideal system to study trans-species polymorphisms, i.e. polymorphisms maintained across several species, in adaptive radiations.

Intraspecific diversity alters the relationship between climate change and parasitism in a polymorphic ectotherm.

Climate-modulated parasitism is driven by a range of factors, yet the spatial and temporal variability of this relationship has received scant attention in wild vertebrate hosts. Moreover, most prior studies overlooked the intraspecific differences across host morphotypes, which impedes a full understanding of the climate-parasitism relationship. In the common lizard (Zootoca vivipara), females exhibit three colour morphs: yellow (Y-females), orange (O-females) and mixed (mixture of yellow and orange, M-females). Zootoca vivipara is also infested with an ectoparasite (Ophionyssus mites). We therefore used this model system to examine the intraspecific response of hosts to parasitism under climate change. We found infestation probability to differ across colour morphs at both spatial (10 sites) and temporal (20 years) scales: M-females had lower parasite infestations than Y- and O-females at lower temperatures, but became more susceptible to parasites as temperature increased. The advantage of M-females at low temperatures was counterbalanced by their higher mortality rates thereafter, which suggests a morph-dependent trade-off between resistance to parasites and host survival. Furthermore, significant interactions between colour morphs and temperature indicate that the relationship between parasite infestations and climate warming was contingent on host morphotypes. Parasite infestations increased with temperature for most morphs, but displayed morph-specific rates. Finally, infested M-females had higher reductions in survival rates than infested Y- or O-females, which implies a potential loss of intraspecific diversity within populations as parasitism and temperatures rise. Overall, we found parasitism increases with warming temperatures, but this relationship is modulated by host morphotypes and an interaction with temperature. We suggest that epidemiological models incorporate intraspecific diversity within species for better understanding the dynamics of wildlife diseases under climate warming.

Shades of grey: host phenotype dependent effect of urbanization on the bacterial microbiome of a wild mammal.

BACKGROUND: Host-associated microbiota are integral to the ecology of their host and may help wildlife species cope with rapid environmental change. Urbanization is a globally replicated form of severe environmental change which we can leverage to better understand wildlife microbiomes. Does the colonization of separate cities result in parallel changes in the intestinal microbiome of wildlife, and if so, does within-city habitat heterogeneity matter? Using 16S rRNA gene amplicon sequencing, we quantified the effect of urbanization (across three cities) on the microbiome of eastern grey squirrels (Sciurus carolinensis). Grey squirrels are ubiquitous in rural and urban environments throughout their native range, across which they display an apparent coat colour polymorphism (agouti, black, intermediate). RESULTS: Grey squirrel microbiomes differed between rural and city environments; however, comparable variation was explained by habitat heterogeneity within cities. Our analyses suggest that operational taxonomic unit (OTU) community structure was more strongly influenced by local environmental conditions (rural and city forests versus human built habitats) than urbanization of the broader landscape (city versus rural). The bacterial genera characterizing the microbiomes of built-environment squirrels are thought to specialize on host-derived products and have been linked in previous research to low fibre diets. However, despite an effect of urbanization at fine spatial scales, phylogenetic patterns in the microbiome were coat colour phenotype dependent. City and built-environment agouti squirrels displayed greater phylogenetic beta-dispersion than those in rural or forest environments, and null modelling results indicated that the phylogenetic structure of urban agouti squirrels did not differ greatly from stochastic expectations. CONCLUSIONS: Squirrel microbiomes differed between city and rural environments, but differences of comparable magnitude were observed between land classes at a within-city scale. We did not observe strong evidence that inter-environmental differences were the result of disparate selective pressures. Rather, our results suggest that microbiota dispersal and ecological drift are integral to shaping the inter-environmental differences we observed. However, these processes were partly mediated by squirrel coat colour phenotype. Given a well-known urban cline in squirrel coat colour melanism, grey squirrels provide a useful free-living system with which to study how host genetics mediate environment x microbiome interactions.

The complete mitochondrial genome of the wood tiger moth (Arctia plantaginis) and phylogenetic analyses within Arctiinae.

We report the assembly and annotation of the complete mitochondrial genome of the warningly-coloured wood tiger moth (Arctia plantaginis) and investigate its phylogenetic position within Arctiinae. The A.plantaginis mitogenome is 15,479 bp long with 13 protein-coding genes, 22 transfer RNAs, 2 ribosomal RNA genes, and an A + T-rich region (D-loop). The phylogenetic analyses based on 13 protein-coding genes showed A.plantaginis clustering within a clade of species with white wings and yellow or red bodies. This result can be useful in understanding the evolution of coloration in Arctiid moths.

Genetic mechanisms and correlational selection structure trait variation in a coral snake mimic.

Covariation among traits shapes both phenotypic evolution and ecological interactions across space and time. However, rampant geographical variation in the strength and direction of such correlations can be particularly difficult to explain through generalized mechanisms. By integrating population genomics, surveys of natural history collections and spatially explicit analyses, we tested multiple drivers of trait correlations in a coral snake mimic that exhibits remarkable polymorphism in mimetic and non-mimetic colour traits. We found that although such traits co-occur extensively across space, correlations were best explained by a mixture of genetic architecture and correlational selection, rather than by any single mechanism. Our findings suggest that spatially complex trait distributions may be driven more by the simple interaction between multiple processes than by complex variation in one mechanism alone. These interactions are particularly important in mimicry systems, which frequently generate striking geographical variation and genetic correlations among colour pattern traits.