Not so colourful after all: eggshell pigments constrain avian eggshell colour space

Birds'' eggshells are renowned for their striking colours and varied patterns. Although often considered exceptionally diverse, we report that avian eggshell coloration, sampled here across the full phylogenetic diversity of birds, occupies only 0.08–0.10% of the avian perceivable colour space. The concentrations of the two known tetrapyrrole eggshell pigments (protoporphyrin and biliverdin) are generally poor predictors of colour, both intra- and interspecifically. Here, we show that the constrained diversity of eggshell coloration can be accurately predicted by colour mixing models based on the relative contribution of both pigments and we demonstrate that the models'' predictions can be improved by accounting for the reflectance of the eggshell''s calcium carbonate matrix. The establishment of these proximate links between pigmentation and colour will enable future tests of hypotheses on the functions of perceived avian eggshell colours that depend on eggshell chemistry. More generally, colour mixing models are not limited to avian eggshell colours but apply to any natural colour. Our approach illustrates how modelling can aid the understanding of constraints on phenotypic diversity.     

Visual warning signals of the ladybird Harmonia axyridis: the avian predators’ point of view

Previous studies have suggested that spotted patterns are important in the protection of ladybirds against attack by avian predators. Nevertheless, these studies were based on the comparison of several ladybird species differing in colouration, but also in other traits (e.g., chemical protection). We presented natural as well as artificial colour modifications (using brown, red, and black paint) of the ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) – an invasive alien species for Europe – to an avian predator, the great tit, Parus major L. (Passeriformes: Paridae). All forms were considered to be equal in size, but differed in colouration and in the presence of spots. The chemical protection was equal except for one form. The birds displayed strong avoidance of all forms with red and black colouration; beetles with artificially removed red colouration (painted brown) were attacked more often. The beetles painted brown with black spots were slightly better protected than the painted beetles without spots. We can sum up that spots are of some importance in the protection of ladybirds; nevertheless, red and black colouration is the main part of the visual signal.     

Flower colours and pigments in Disa hybrid (Orchidaceae)

Flower colours and the composition of pigments in the perianths of five cultivars of Disa orchids were analyzed. Carotenoids were major pigment components in the orange-red flowers of ‘Dawn Angel’. We identified two types of pigment composition in the red flowered cultivars: ‘San Francisco’ contained more carotenoids and less anthocyanins, while ‘Marlene’ contained more anthocyanins than carotenoids. The red-purple flowered cultivars, only contained slight amounts of carotenoids, and the red-purple colour was attributed to the relatively high density of a cyanidin-based anthocyanin. The importance of the characterization of pigments in the perianths of orchid has been discussed in both breeding for flower colour improvement and chemotaxonomy.     

Structural coloration in a fossil feather

Investigation of feathers from the famous Middle Eocene Messel Oil Shale near Darmstadt, Germany shows that they are preserved as arrays of fossilized melanosomes, the surrounding beta-keratin having degraded. The majority of feathers are preserved as aligned rod-shaped eumelanosomes. In some, however, the barbules of the open pennaceous, distal portion of the feather vane are preserved as a continuous external layer of closely packed melanosomes enclosing loosely aligned melanosomes. This arrangement is similar to the single thin-film nanostructure that generates an iridescent, structurally coloured sheen on the surface of black feathers in many lineages of living birds. This is, to our knowledge, the first evidence of preservation of a colour-producing nanostructure in a fossil feather and confirms the potential for determining colour differences in ancient birds and other dinosaurs.     

Characterization of the porcine KIT ligand gene: expression analysis, genomic structure, polymorphism detection and association with coat colour traits

Kit ligand (KITLG) is the ligand for the type III receptor tyrosine kinase KIT. Studies of the KIT/KITLG pathway in a number of mammalian species have shown that it is important for the development of stem cell populations in haematopoietic tissues, germ cells in reproductive organs and the embryonic migrating melanoblasts that give rise to melanocytes. Consequently, mutations in the pathway may result in a range of defects including anaemia, sterility and de-pigmentation. The cDNA sequence of the porcine KITLG gene has been reported previously, and is an attractive candidate locus for moderating coat colour in pigs. In this paper we report the gene structure and physical mapping of the porcine gene. We also report the identification of polymorphisms in the gene, one of which was used to confirm linkage to chromosome 5. Preliminary RNA expression studies using a panel of tissues have shown that in addition to the known variant lacking exon 6, there is alternative splicing of exon 4. However, little evidence was found for the KITLG gene being linked to variation in colour in a Meishan x Large White cross.     

Color pan traps often catch less when there are more flowers around

When assessing changes in populations of species, it is essential that the methods used to collect data have some level of precision and preferably also good accuracy. One commonly used method to collect pollinators is colour pan traps, but this method has been suggested to be biased by the abundance of surrounding flowers. The present study evaluated the relationship between pan trap catches and the frequency of flowers on small (25 m²) and large (2–6 ha) spatial scales. If pan traps work well, one should assume a positive relationship, that is, more insects caught when they have more food. However, in contrast, we found that catches in pan traps were often negatively affected by flower frequency. Among the six taxa evaluated, the negative bias was largest in Vespoidea and Lepturinae, while there was no bias in solitary Apoidea (Cetoniidae, Syrphidae and social Apoidea were intermediate). Furthermore, red flowers seemed to contribute most to the negative bias. There was also a tendency that the negative bias differed within the flight season and that it was higher when considering the large spatial scale compared to the small one. To conclude, pan trap catches may suffer from a negative bias due to surrounding flower frequency and color. The occurrence and magnitude of the negative bias were context and taxon dependent, and therefore difficult to adjust for. Thus, pan traps seem less suited to evaluate differences between sites and the effect of restoration, when gradients in flower density are large. Instead, it seems better suited to monitor population changes within sites, and when gradients are small.     

The seven ways eukaryotes produce repeated colour motifs on external tissues

The external tissues of numerous eukaryote species show repeated colour patterns, usually characterized by units that are present at least twice on the body. These dotted, striped or more complex phenotypes carry out crucial biological functions, such as partner recognition, aposematism or camouflage. Very diverse mechanisms explaining the formation of repeated colour patterns in eukaryotes have been identified and described, and it is timely to review this field from an evolutionary and developmental biology perspective. We propose a novel classification consisting of seven families of primary mechanisms: Turing(-like), cellular automaton, multi-induction, physical cracking, random, neuromuscular and printing. In addition, we report six pattern modifiers, acting synergistically with these primary mechanisms to enhance the spectrum of repeated colour patterns. We discuss the limitations of our classification in light of currently unexplored extant diversity. As repeated colour patterns require both the production of a repetitive structure and colouration, we also discuss the nature of the links between these two processes. A more complete understanding of the formation of repeated colour patterns in eukaryotes will require (i) a deeper exploration of biological diversity, tackling the issue of pattern elaboration during the development of non-model taxa, and (ii) exploring some of the most promising ways to discover new families of mechanisms. Good starting points include evaluating the role of mechanisms known to produce non-repeated colour patterns and that of mechanisms responsible for repeated spatial patterns lacking colouration.     

The evolution and genetics of sexually dimorphic ‘dual’ mimicry in the butterfly Elymnias hypermnestra

Sexual dimorphism is a major component of morphological variation across the tree of life, but the mechanisms underlying phenotypic differences between sexes of a single species are poorly understood. We examined the population genomics and biogeography of the common palmfly Elymnias hypermnestra, a dual mimic in which female wing colour patterns are either dark brown (melanic) or bright orange, mimicking toxic Euploea and Danaus species, respectively. As males always have a melanic wing colour pattern, this makes E. hypermnestra a fascinating model organism in which populations vary in sexual dimorphism. Population structure analysis revealed that there were three genetically distinct E. hypermnestra populations, which we further validated by creating a phylogenomic species tree and inferring historical barriers to gene flow. This species tree demonstrated that multiple lineages with orange females do not form a monophyletic group, and the same is true of clades with melanic females. We identified two single nucleotide polymorphisms (SNPs) near the colour patterning gene WntA that were significantly associated with the female colour pattern polymorphism, suggesting that this gene affects sexual dimorphism. Given WntA''s role in colour patterning across Nymphalidae, E. hypermnestra females demonstrate the repeatability of the evolution of sexual dimorphism.     

Association between population structure and allele frequencies of the glycogen synthase 1 mutation in the Austrian Noriker draft horse

The aim of this study was to determine the allele frequency of the glycogen synthase 1 (GYS1) mutation associated with polysaccharide storage myopathy type 1 in the Austrian Noriker horse. Furthermore, we examined the influence of population substructures on the allele distribution. The study was based upon a comprehensive population sample (208 breeding stallions and 309 mares) and a complete cohort of unselected offspring from the year 2014 (1553 foals). The mean proportion of GYS1 carrier animals in the foal cohort was 33%, ranging from 15% to 50% according to population substructures based on coat colours. In 517 mature breeding horses the mutation carrier frequency reached 34%, ranging on a wider scale from 4% to 62% within genetic substructures. We could show that the occurrence of the mutated GYS1 allele is influenced by coat colour; genetic bottlenecks; and assortative, rotating and random mating strategies. Highest GYS1 carrier frequencies were observed in the chestnut sample comprising 50% in foals, 54% in mares and 62% in breeding stallions. The mean inbreeding of homozygous carrier animals reached 4.10%, whereas non‐carrier horses were characterized by an inbreeding coefficient of 3.48%. Lowest GYS1 carrier frequencies were observed in the leopard spotted Noriker subpopulation. Here the mean carrier frequency reached 15% in foals, 17% in mares and 4% in stallions and inbreeding decreased from 3.28% in homozygous non‐carrier horses to 2.70% in heterozygous horses and 0.94% in homozygous carriers. This study illustrates that lineage breeding and specified mating strategies result in genetic substructures, which affect the frequencies of the GYS1 gene mutation.