Is the later arrival of young male red-breasted flycatchers (Ficedula parva) related to their physical condition?

Intraspecific variation in the arrival time of migratory birds to breeding grounds is common. Although this phenomenon has been explained in various ways, the condition-dependency of arrival is often evoked. I analyzed the arrival time of male red-breasted flycatchers Ficedula parva, a long-distance migratory passerine, in relation to age, body size and body condition. Data were collected over the course of 5 years in the primeval Białowieża Forest of northeast Poland. Each of these years, young males arrived at the breeding grounds significantly later in the year than did older males. Young males also exhibited significantly shorter wings and lower body condition than older males. Settlement speed was significantly related to age, but not to wing length or body condition. Only the arrival time of old males was related to the body mass and condition. Later arrival of young males could be explained by the lack of experience or by avoidance of both aggression and competition from older males, and both explanations are thought to conserve energy for breeding. Young male red-breasted flycatchers exhibit delayed plumage maturation, and this duller plumage supports the strategy of restraint in the arrival time of young males in this species.     

Plumage color as a status signal in male–male interaction in the red-flanked bushrobin, Tarsiger cyanurus

Recent studies have suggested that structural-based coloration is an honest signal of male genetic and/or conditional quality in sexual selection. However, whether structural coloration functions in intrasexual competition is unknown. We examined whether plumage color functions as a status signal during intrasexual interactions in the red-flanked bushrobin Tarsiger cyanurus; adult males have many blue plumes as structural coloration whereas yearling males and females are olive brown with few blue plumages. Blue males did not always dominate olive-brown males. The number of interactions did not differ with the colors of the two birds involved. The interactions of a blue male and an olive-brown male were less aggressive than those of two blue or of two olive-brown males. In this study, we found that structural plumage coloration may serve as a signal of aggressive intent and lower the escalation level of an aggressive interaction in a manner consistent with hypotheses regarding the evolution of delayed plumage maturation.     

Seasonal colour change by moult or by the abrasion of feather tips: a comparative study

Many birds undergo seasonal changes in plumage coloration by prebreeding moult, abrasion of cryptic feather tips, or both. Seasonal dichromatism is thought to result from optimizing coloration to the conflicting demands of different life-cycle periods, sexual selection for conspicuousness being substantial during the mating season, whereas selection for camouflage and for social signals may act in all seasons. Furthermore, energetic and time demands may constrain the extent of moult, thereby limiting colour change. We investigated the relative importance of several factors in shaping this variation in a songbird clade using phylogenetic comparative methods. We found that prebreeding moult relates most strongly to breeding onset and winter diet, demonstrating that both time and food availability constrain feather replacement. Feather abrasion was best predicted by winter flocking behaviour, and secondarily by open habitats, implying that exposure to predators and the simultaneous need for social signalling may favour the expression of partially obscured ornaments in the non-breeding season. The combined occurrence of prebreeding moult and feather abrasion was associated with the polygynous mating system, suggesting that species under strong sexual selection may employ both strategies of colour change to ensure the full expression of breeding coloration.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 711–721.     

Components of phenotypic variation in avian ornamental and non-ornamental feathers

Phenotypic variation, measured as the coefficient of variation (CV), is usually larger in secondary sexual characters than in ordinary morphological traits. We tested if intraspecific differences in the CV between ornamental and non-ornamental feather traits in 67 evolutionary events of feather ornamentation in birds were due to differences in (1) the allometric pattern (slope of the regression line when regressing trait size on an indicator of body size), or (2) the dispersion of observations around the regression line. We found that only dispersion of observations around the regression line contributed significantly to total variation. A large dispersion of observations around the regression line for ornamental feathers is consistent with these characters showing condition-dependence, supporting indicator models of sexual selection more strongly than a pure Fisher process. Ornamental feathers in males demonstrated negative allometry when regressed on tarsus length, which is a measure of skeletal body size. This finding is consistent with ornamental feather traits being subject to directional selection due to female mate preferences, where large body size is less important than in male–male competition. This pattern of phenotypic variation for avian secondary sexual characters contrasts with patterns of variation for insect genitalia, supposedly subject to sexual selection, since the latter traits only differ from ordinary morphology traits in allometry coefficient. The prevailing regime of selection (directional or stabilizing) and the effects of environmental factors are proposed to account for these differences among traits.     

Signal evolution and morphological complexity in hummingbirds (Aves: Trochilidae)

Understanding how animal signals are produced is critical for understanding their evolution because complexity and modularity in the underlying morphology can affect evolutionary patterns. Hummingbird feathers show some of the brightest and most iridescent colors in nature. These are produced by optically complex stacks of hollow, platelet-shaped organelles called melanosomes. Neither how these morphologies produce colors nor their evolution has been systematically studied. We first used nanoscale morphological measurements and optical modeling to identify the physical basis of color production in 34 hummingbird species. We found that, in general, the melanosome stacks function as multilayer reflectors, with platelet thickness and air space size explaining variation in hue (color) and saturation (color purity). Additionally, light rays reflected from the outer keratin surface interact with those reflected by small, superficial melanosomes to cause secondary reflectance peaks, primarily in short (blue) wavelengths. We then compared variation of both the morphological components and the colors they produce. The outer keratin cortex evolves independently and is more variable than other morphological traits, possibly due to functional constraints on melanosome packing. Intriguingly, shorter wavelength colors evolve faster than longer wavelength colors, perhaps due to developmental processes that enables greater lability of the shapes of small melanosomes. Together, these data indicate that increased structural complexity of feather tissues is associated with greater variation in morphology and iridescent coloration.     

HISTORICAL EXAMINATION OF DELAYED PLUMAGE MATURATION IN THE SHOREBIRDS (AVES: CHARADRIIFORMES)

Delayed plumage maturation refers to the presence of nonadultlike immature plumages (juvenal plumage excluded). It is usually considered the result of selection for distinctive first-winter or first-summer appearance. In the present study, evolution of delayed plumage maturation is examined in the shorebirds: the sandpipers, plovers, gulls, and their allies. Nine plumage-maturation characters were identified, and their states were superimposed onto topologies generated during two recent investigations of shorebird relationships (Sibley and Ahlquist; revised Strauch). The characters were then optimized so as to assign character states to interior nodes of the trees in the most parsimonious way. Reconstructions of character evolution on six of the shortest revised Strauch trees were ambiguous with respect to delayed plumage maturation in the hypothetical ancestral shorebird. If plumage maturation was not delayed in the shorebird ancestor, optimization indicated that delay appeared when nonadultlike juvenal feathers were acquired. In contrast, on the single Sibley and Ahlquist tree, absence of delayed plumage maturation in the shorebird ancestor was indicated unambiguously, with three evolutionary novelties (nonadultlike juvenal feathers, seasonal plumage change, and a reduced first-spring molt) implicated in its acquisition. Optimization indicated that delayed plumage maturation in shorebirds can be explained plausibly without invoking selection for distinctive first-winter or first-summer appearance. Two of the novel conditions generating delayed plumage maturation (modified juvenal feathers and seasonal plumage change) did so only because they were acquired in a taxon possessing restricted first-year molts, which are primitive. Given these observations, it seems simplest to explain the delay in plumage maturation as an incidental consequence of the phylogenetic inertia of shorebird molts. The third novelty that generates delayed plumage maturation, a reduced first-spring molt, may have been acquired to reduce molt-associated energetic demands in young birds.     

SPECIATION AND FEATHER ORNAMENTATION IN BIRDS

The hypothesis that sexual selection promotes speciation has rarely been tested. We identified 70 evolutionarily independent events of feather ornaments in birds. For each focal species we noted the number of ornamented and nonornamented species belonging to its genus and its number of subspecies, as well as its mating system and the extent of its geographic range. For purposes of comparison, we randomly chose a second, nonornamented species for which we obtained information on the number of subspecies, and in cases in which the nonornamented species was in the same genus, we chose a third, nonornamented species in a related genus and obtained the same information. We then noted the number of species in each genus and the difference in numbers of species, or species richness, between paired genera. For the genera of the focal ornamented species, we regressed number of ornamented species on number of nonornamented species and found a positive relationship. As number of species per genus rose, number of ornamented species per genus rose more rapidly, indicating that more speciose genera have a higher proportion of ornamented species than less speciose genera. We then took the deviations from this regression, the residual number of species, and regressed them on the differences in species richness between the paired genera. This relationship was positive indicating that ornamented genera with more than the expected number of ornamented species were more speciose with respect to their paired genera than were genera with fewer than the expected number of ornamented species. Finally, we compared the deviations from this regression, the residual number of ornamented species, with species' mating system and found a greater residual number of ornamented species among species whose mating system is associated with greater skew in male mating success and thus more intense sexual selection. Ornamented species had more subspecies than nonornamented species, even when controlling for geographic range, suggesting an association between subspeciation and ornaments.     

Corticosterone regulates multiple colour traits in Lacerta [Zootoca] vivipara males

Ornamental colours usually evolve as honest signals of quality, which is supported by the fact that they frequently depend on individual condition. It has generally been suggested that some, but not all types of ornamental colours are condition dependent, indicating that different evolutionary mechanisms underlie the evolution of multiple types of ornamental colours even when these are exhibited by the same species. Stress hormones, which negatively affect condition, have been shown to affect colour traits based on different pigments and structures, suggesting that they mediate condition dependence of multiple ornament types both among and within individuals. However, studies investigating effects of stress hormones on different ornament types within individuals are lacking, and thus, evidence for this hypothesis is scant. Here, we investigated whether corticosterone mediates condition dependence of multiple ornaments by manipulating corticosterone levels and body condition (via food availability) using a two‐factorial design and by assessing their effect on multiple colour traits in male common lizards. Corticosterone negatively affected ventral melanin‐ and carotenoid‐based coloration, whereas food availability did not affect coloration, despite its significant effect on body condition. The corticosterone effect on melanin‐ and carotenoid‐based coloration demonstrates the condition dependence of both ornaments. Moreover, corticosterone affected ventral coloration and had no effect on the nonsexually selected dorsal coloration, showing specific effects of corticosterone on ornamental ventral colours. This suggests that corticosterone simultaneously mediates condition dependence of multiple colour traits and that it therefore accounts for covariation among them, which may influence their evolution via correlational selection.