Evolution and assessment of colour patterns in stream-resident and anadromous male threespine stickleback Gasterosteus aculeatus from three regions

We compared the colour patterns of free swimming, reproductively active male threespine stickleback Gasterosteus aculeatus of the anadromous and stream ecotypes from three geographically distinct regions. Consistent with the hypothesis of environmentally mediated selection, our results indicate ecologically replicated differences in G. aculeatus coloration between anadromous and stream-resident populations, and that G. aculeatus probably have the visual acuity to discriminate colour pattern differences between anadromous and stream-resident fish.     

Carotenoid-based bill colour as an indicator of immunocompetence and sperm performance in male mallards

Female mate choice is often based on exaggerated sexual traits, signals of male qualities that females cannot assess directly. Two such key qualities are male immune and/or sexual competence, whereby honesty in signalling could be maintained by physiological trade-offs. Carotenoid-based ornaments likely constitute such honest signals, as there is direct competition for (limited) carotenoids between ornament deposition and anti-oxidant support of immune or sperm functioning. Using spectrometry, we assessed the potential signalling function of the yellow, carotenoid-based colour of the bill of male mallards, a target of female mate choice. Here we demonstrate that bill reflectance varied with plasma carotenoid level, indicating antioxidant reserves. Moreover, lower relative UV reflectance during autumn pairing predicted immune responsiveness and correlated positively with sperm velocity during breeding, a trait that affects fertility. Our data provide support for current theories that females could use carotenoid-based sexual signals to detect immune vigour and fertilizing ability of prospective mates.     

Hybridization and transgressive exploration of colour pattern and wing morphology in Heliconius butterflies

Hybridization can generate novel phenotypes distinct from those of parental lineages, a phenomenon known as transgressive trait variation. Transgressive phenotypes might negatively or positively affect hybrid fitness, and increase available variation. Closely related species of Heliconius butterflies regularly produce hybrids in nature, and hybridization is thought to play a role in the diversification of novel wing colour patterns despite strong stabilizing selection due to interspecific mimicry. Here, we studied wing phenotypes in first‐ and second‐generation hybrids produced by controlled crosses between either two co‐mimetic species of Heliconius or between two nonmimetic species. We quantified wing size, shape and colour pattern variation and asked whether hybrids displayed transgressive wing phenotypes. Discrete traits underlain by major‐effect loci, such as the presence or absence of colour patches, generate novel phenotypes. For quantitative traits, such as wing shape or subtle colour pattern characters, hybrids only exceed the parental range in specific dimensions of the morphological space. Overall, our study addresses some of the challenges in defining and measuring phenotypic transgression for multivariate traits and our data suggest that the extent to which transgressive trait variation in hybrids contributes to phenotypic diversity depends on the complexity and the genetic architecture of the traits.     

Evolutionary significance of ontogenetic colour change in animals

Ontogenetic colour changes are non-reversible colour changes associated with normal progressive development of an individual of a species. This paper provides the first review of the evolutionary significance of this phenomenon in animals. Proximate mechanisms and environmental cues are briefly discussed and a conceptual framework for understanding the ultimate reasons for ontogenetic colour change is established. Changes in size, vulnerability, reproductive status, habitat and metabolism are often associated with ontogenetic colour change and can aid in understanding its adaptive significance. Neutral or non-adaptive ontogenetic colour changes due to phylogenetic inertia and developmental constraints are also considered. Existing studies of ontogenetic colour changes in marine invertebrates, terrestrial invertebrates, fish, amphibians, reptiles, birds and mammals are discussed within this framework. A need is identified for more experimental tests of hypotheses for the significance of ontogenetic colour change.     

Pollen colour morphs take different paths to fitness

Colour phenotypes are often involved in communication and are thus under selection by species interactions. However, selection may also act on colour through correlated traits or alternative functions of biochemical pigments. Such forms of selection are instrumental in maintaining petal colour diversity in plants. Pollen colour also varies markedly, but the maintenance of this variation is little understood. In Campanula americana, pollen ranges from white to dark purple, with darker morphs garnering more pollinator visits and exhibiting elevated pollen performance under heat stress. Here, we generate an F2 population segregating for pollen colour and measure correlations with floral traits, pollen attributes and plant‐level traits related to fitness. We determine the pigment biochemistry of colour variants and evaluate maternal and paternal fitness of light and dark morphs by crossing within and between morphs. Pollen colour was largely uncorrelated with floral traits (petal colour, size, nectar traits) suggesting it can evolve independently. Darker pollen grains were larger and had higher anthocyanin content (cyanidin and peonidin) which may explain why they outperform light pollen under heat stress. Overall, pollen‐related fitness metrics were greater for dark pollen, and dark pollen sires generated seeds with higher germination potential. Conversely, light pollen plants produce 61% more flowers than dark, and 18% more seeds per fruit, suggesting a seed production advantage. Results indicate that light and dark morphs may achieve fitness through different means—dark morphs appear to have a pollen advantage whereas light morphs have an ovule advantage—helping to explain the maintenance of pollen colour variation.     

Sympatry drives colour and song evolution in wood-warblers (Parulidae)

Closely related species often exhibit similarities in appearance and behaviour, yet when related species exist in sympatry, signals may diverge to enhance species recognition. Prior comparative studies provided mixed support for this hypothesis, but the relationship between sympatry and signal divergence is likely nonlinear. Constraints on signal diversity may limit signal divergence, especially when large numbers of species are sympatric. We tested the effect of sympatric overlap on plumage colour and song divergence in wood-warblers (Parulidae), a speciose group with diverse visual and vocal signals. We also tested how number of sympatric species influences signal divergence. Allopatric species pairs had overall greater plumage and song divergence compared to sympatric species pairs. However, among sympatric species pairs, plumage divergence positively related to the degree of sympatric overlap in males and females, while male song bandwidth and syllable rate divergence negatively related to sympatric overlap. In addition, as the number of species in sympatry increased, average signal divergence among sympatric species decreased, which is likely due to constraints on warbler perceptual space and signal diversity. Our findings reveal that sympatry influences signal evolution in warblers, though not always as predicted, and that number of sympatric species can limit sympatry''s influence on signal evolution.     

Physiological stress links parasites to carotenoid‐based colour signals

Vertebrates commonly use carotenoid‐based traits as social signals. These can reliably advertise current nutritional status and health because carotenoids must be acquired through the diet and their allocation to ornaments is traded‐off against other self‐maintenance needs. We propose that the coloration more generally reveals an individual’s ability to cope with stressful conditions. We tested this idea by manipulating the nematode parasite infection in free‐living red grouse (Lagopus lagopus scoticus) and examining the effects on body mass, carotenoid‐based coloration of a main social signal and the amount of corticosterone deposited in feathers grown during the experiment. We show that parasites increase stress and reduce carotenoid‐based coloration, and that the impact of parasites on coloration was associated with changes in corticosterone, more than changes in body mass. Carotenoid‐based coloration appears linked to physiological stress and could therefore reveal an individual’s ability to cope with stressors.     

Condition-dependent traits as signals of the functionality of vital cellular processes

Condition is a nearly ubiquitous term in the behavioural, physiological and evolutionary ecology literature; however, existing definitions are incomplete or ambiguous. This poor conceptualization has led to confusion regarding what is being signalled by condition-dependent traits and how to interpret links between ornamentation and individual characteristics such as nutrient reserves, oxidative state and immunocompetence. I propose that the combined effects of the somatic state, epigenetic state and genotype of an organism determine condition. I define condition as the relative capacity to maintain optimal functionality of vital systems within the body. A condition-dependent trait is a conspicuous feature of an organism that enhances perception of condition. Ornament expression can link to system functionality in at least four ways: (1) resources are traded off between operation of physiological pathways and production of ornaments; (2) a regulatory agent necessary for ornament expression depresses a vital physiological process; (3) ornament production requires a product of a vital physiological process; and (4) pathways are shared between ornament production and vital physiological processes. If the honesty of ornamental traits derives from connections to vital cellular processes then there is no need to invoke a fitness cost of ornamentation to insure signal honesty.     

Peak shift discrimination learning as a mechanism of signal evolution

"Peak shift" is a behavioral response bias arising from discrimination learning in which animals display a directional, but limited, preference for or avoidance of unusual stimuli. Its hypothesized evolutionary relevance has been primarily in the realm of aposematic coloration and limited sexual dimorphism. Here, we develop a novel functional approach to peak shift, based on signal detection theory, which characterizes the response bias as arising from uncertainty about stimulus appearance, frequency, and quality. This approach allows the influence of peak shift to be generalized to the evolution of signals in a variety of domains and sensory modalities. The approach is illustrated with a bumblebee (Bombus impatiens) discrimination learning experiment. Bees exhibited peak shift while foraging in an artificial Batesian mimicry system. Changes in flower abundance, color distribution, and visitation reward induced bees to preferentially visit novel flower colors that reduced the risk of flower-type misidentification. Under conditions of signal uncertainty, peak shift results in visitation to rarer, but more easily distinguished, morphological variants of rewarding species in preference to their average morphology. Peak shift is a common and taxonomically widespread phenomenon. This example of the possible role of peak shift in signal evolution can be generalized to other systems in which a signal receiver learns to make choices in situations in which signal variation is linked to the sender's reproductive success.     

Noise affects the shape of female preference functions for acoustic signals

The shape of female mate preference functions influences the speed and direction of sexual signal evolution. However, the expression of female preferences is modulated by interactions between environmental conditions and the female's sensory processing system. Noise is an especially relevant environmental condition because it interferes directly with the neural processing of signals. Although noise is therefore likely a significant force in the evolution of communication systems, little is known about its effects on preference function shape. In the grasshopper Chorthippus biguttulus, female preferences for male calling song characteristics are likely to be affected by noise because its auditory system is sensitive to fine temporal details of songs. We measured female preference functions for variation in male song characteristics in several levels of masking noise and found strong effects of noise on preference function shape. The overall responsiveness to signals in noise generally decreased. Preference strength increased for some signal characteristics and decreased for others, largely corresponding to expectations based on neurophysiological studies of acoustic signal processing. These results suggest that different signal characteristics will be favored under different noise conditions, and thus that signal evolution may proceed differently depending on the extent and temporal patterning of environmental noise.     

Courtship in long-legged flies (Diptera: Dolichopodidae): function and evolution of signals

Analyzing the courtship behavior of long-legged flies (Diptera:Dolichopodidae), we focus on the evolutionary development ofcourtship signals. Long-legged flies may serve as a model forthis evolutionary process, because males of some species presentsexually dimorphic badges during courtship, whereas others donot exhibit such conspicuous signals but present lavish courtshipbehavior, including dynamic flight maneuvers. A comparison ofthese two groups within a single taxonomic family provides insightinto the evolution of courtship signals and the correspondingbehavior. Males of the closely related Empididae do not possesssuch badges. Within the super-family Empidoidea, we proposean evolutionary shift from dynamically courting and mating onthe wing (in Empididae) to courting and mating on ground (inDolichopodidae), accompanied by signaling through badge-waving.By comparing previously published data and observations on courtshipbehavior in Dolichopodidae, we present the hypothesis that thelatter replaced the former energetically expensive behavioras a case of automimicry and sensory trap.     

The evolution of novel animal signals: silk decorations as a model system

Contemporary animal signals may derive from an elaboration of existing forms or novel non-signalling traits. Unravelling the evolution of the latter is challenging because experiments investigating the maintenance of the signal may provide little insight into its early evolution. The web decorations, or stabilimenta of some orb web spiders represent an intriguing model system to investigate novel animal signals. For over 100 years, biologists have struggled to explain why spiders decorate their webs with additional threads of silk, producing a conspicuous signal on a construction whose function is to entangle unsuspecting prey. The numerous explanations for the maintenance of this behaviour starkly contrast with the absence of a plausible explanation for its evolutionary origin. Our review highlights the difficulties in resolving both the evolution and maintenance of animal signalling, and inferring the causative arrow-even from experimental studies. Drawing on recent research that focuses on physiological processes, we provide a model of the evolutionary progression of web-decorating behaviour.     

Influence of ambient light on the evolution of colour signals: comparative analysis of a Neotropical rainforest bird community

Rainforests offer two contrasted light environments: a bright canopy rich in blue and UV and a dark understorey, rich in green and orange. Therefore, natural selection for crypsis should favour dark brown signals in understorey and bright green signals in canopy, whereas sexual selection for conspicuousness should favour bright yellow‐red signals in understorey and dark blue and UV signals in canopy. Using spectrometry and comparative analyses, we examined the relationship between ambient light and colour signals in a bird community of French Guiana. It appears that brightness and hue are mostly naturally selected, while UV content of plumage is more likely sexually selected. At each height, both sexes present similar coloration but males display more conspicuous sexually selected patterns than females. These results show that ambient light drives the evolution of colour signals at community scale, and should be considered when studying signalling in other communities and light‐contrasted ecosystems.