Plumage coloration and social context influence male investment in song.

Animals use multiple signals to attract mates, including elaborate song, brightly coloured ornaments and physical displays. Female birds often prefer both elaborate male song and intense carotenoid-based plumage coloration. This could lead less visually ornamented males to increase song production to maximize their attractiveness to females. We tested this possibility in the highly social and non-territorial house finch (Haemorhous mexicanus), in which females discriminate among males based on both song and on the intensity of red carotenoid-based plumage coloration. We manipulated male plumage coloration through carotenoid supplementation during moult, so that males were either red or yellow. Males were then housed under three social environments: (i) all red birds, (ii) all yellow birds or (iii) a mixture of red/yellow birds. We recorded song after presentation of a female. Red males produced more song than yellow males. But when yellow males were housed with red conspecifics, they produced more song relative to yellow males housed with equally unattractive yellow males. This study provides novel evidence that a male's plumage coloration and the plumage colour of his social competitors influence investment in song.

Independent sources of condition dependency and multiple pathways determine a composite trait: lessons from carotenoid-based plumage colouration.

Many colour ornaments are composite traits consisting of at least four components, which themselves may be more complex, determined by independent evolutionary pathways, and potentially being under different environmental control. To date, little evidence exists that several different components of colour elaboration are condition dependent and no direct evidence exists that different ornamental components are affected by different sources of variation. For example, in carotenoid-based plumage colouration, one of the best-known condition-dependent ornaments, colour elaboration stems from both condition-dependent pigment concentration and structural components. Some environmental flexibility of these components has been suggested, but specifically which and how they are affected remains unknown. Here, we tested whether multiple colour components may be condition dependent, by using a comprehensive 3 × 2 experimental design, in which we carotenoid supplemented and immune challenged great tit nestlings (Parus major) and quantified effects on different components of colouration. Plumage colouration was affected by an interaction between carotenoid availability and immune challenge. Path analyses showed that carotenoid supplementation increased plumage saturation via feather carotenoid concentration and via mechanisms unrelated to carotenoid deposition, while immune challenge affected feather length, but not carotenoid concentration. Thus, independent condition-dependent pathways, affected by different sources of variation, determine colour elaboration. This provides opportunities for the evolution of multiple signals within components of ornamental traits. This finding indicates that the selective forces shaping the evolution of different components of a composite trait and the trait's signal content may be more complex than believed so far, and that holistic approaches are required for drawing comprehensive evolutionary conclusions.

Differential ability of carotenoid C4-oxygenation in yellow and red bishop species (Euplectes spp.).

Male breeding plumages of African widowbirds and bishops (Euplectes spp.) show striking variation in carotenoid-based plumage coloration, with saturated yellow or orange-red patches of different size. Yet, from observations and experiments, agonistic signaling appears to have been a generalized sexual selection pressure for redness in the genus. Recent results show that yellow and red widowbird colors derive from distinctly different pigment profiles, and suggest that species vary in their ability to metabolize ingested carotenoids. We used reflectance spectrometry and High Performance Liquid Chromatography (HPLC) to describe the species-specific colors and plumage carotenoids of the congeneric yellow-crowned bishop (E. afer) and southern red bishop (E. orix). Results show that the yellow rump color of E. afer primarily derives from direct-deposited, dietary yellow pigments, i.e. lutein and zeaxanthin. In the red breast of E. orix, these are complemented by smaller amounts of derived red C4-keto-carotenoids: mainly alpha-doradexanthin, but also beta-doradexanthin, canthaxanthin, astaxanthin and adonirubin. We also performed a diet supplementation experiment to investigate the relative importance of nutritional and metabolic constraints in determining the differential occurrence of C4-keto-carotenoids, and thus red plumage color, in the two species. Our results indicate that E. orix, but not E. afer, can manufacture red C4-keto-carotenoids (alpha-doradexanthin and canthaxanthin) from yellow dietary precursors (lutein and beta-carotene).

Condition dependence of nestling mouth colour and the effect of supplementing carotenoids on parental behaviour in the hihi (Notiomystis cincta).

Carotenoids are integument pigments that often reflect foraging efficiency, disease resistance and body condition. In contrast to the widespread attention this relationship has received in adult birds, the condition dependence of nestling colouration remains an understudied component of animal communication. Here we assess the condition dependence of carotenoid pigmentation in nestling hihi (Notiomystis cincta, an endangered New Zealand bird) and examine the influence of carotenoid supplementation on nestling quality and parental visitation rates. Our results show that carotenoids provided to breeding adult hihi were transferred to their offspring and resulted in an intensified orange-yellow flange colour. After accounting for carotenoid supplementation the parameter that most consistently explained variation in nestling flange colour was nestling tarsus length at 23 days, indicating condition dependence of this trait. We did not, however, detect direct effects of carotenoid supplementation on nestling mass or immune response (or any other fitness parameter measured). Carotenoid supplementation did, however, result in an increased paternal provisioning rate.

Hypothalamic GnRH-I and its precursor during photorefractoriness onset in free-living male Dark-eyed Juncos (Junco hyemalis) of different year classes.

The termination of seasonal breeding in most photoperiodic passerines is heralded by spontaneous gonadal regression as a result of long day exposure. This phenomenon is termed absolute photorefractoriness and this insensitivity to long days may only be dissipated by short photoperiod exposure. The timing of the transition to a photorefractory stage in adult free-living male Dark-eyed Juncos is age-dependent. Second-year males (SY, entering their first breeding season) molt earlier, suggesting that they become refractory earlier, than older males (ASY). We determined whether the earlier onset of photorefractoriness in SY males concurs with decreased expression of hypothalamic GnRH-I or its precursor, pro-GnRH-GAP. Male SY and ASY juncos were caught at the end of the breeding season either during, or following, testicular regression. Immunoreactivity for GnRH-I and pro-GnRH-GAP was found in perikarya in the preoptic area, and fibers in the median eminence. The number and size of cells immunoreactive for GnRH-I and pro-GnRH-GAP did not significantly differ with age or time of year. In ASY juncos, there was a significant decrease in both GnRH-I and pro-GnRH-GAP staining density at the median eminence following testicular regression at a time of photorefractoriness onset. Density of pro-GnRH-GAP at the median eminence was also reduced following testicular regression in SY males but GnRH-I staining density did not change. Thus, age-class differences in the transition to photorefractoriness do not appear to result from differences in GnRH-I synthesis. Instead, the regulation of GnRH-I secretion by non-photoperiodic factors may determine photorefractoriness onset.