title: How to reduce the costs of ornaments without reducing their effectiveness? An example of a mechanism from carotenoid-based plumage.
abstract: Carotenoid-based ornaments are often considered to be honest indicators of individual quality assessed by potential mates. However, males can use a variety of strategies that minimize the amount of costly carotenoids used while retaining the effectiveness of color signaling. Birds could do this by altering pigment intake, metabolism, or its presentation to a potential signal receiver. Here, we propose a new mechanism of lowering the costs of carotenoid displays in birds: differential allocation of pigments within single feathers. We studied the coloration of the yellow terminal tail bands of rectrices of male Bohemian waxwings. Using reflectance spectrometry, we show that the two central rectrices are most intensively colored compared to other rectrices. More detailed analyses reveal that these differences result from feather-specific patterns of rectrices coloration. The outer feather vanes of the outermost rectrices are more intensively colored compared to the inner vanes. However, the central rectrices have equally colored vanes that are, on average, more intensively pigmented than the outermost rectrices. When the waxwing tail is folded, the outermost rectrices are covered by other feathers, except for the narrow, outer vane. Central rectrices, however, form the outermost layers which are not obscured by other tail feathers. Thus, the feather vanes that are the most visible to potential viewers are also the most pigmented. These results support the occurrence of a previously overlooked mechanism to reduce the costs of carotenoid-based ornaments: precise pigment distribution to maximize efficiency of signals within single feathers.
  
  
  CONCLUSIONS
  Males of many bird species use bright carotenoid-based plumage coloration to attract females. These traits are physiologically expensive such that only individuals in prime condition can develop the most vivid colors. Males often "cheat" to obtain attractive appearances at lower costs. We showed that this goal could be achieved by differential deposition of pigments into the most conspicuous feather regions. Bohemian waxwing males have yellow tips on their rectrices of which the outer vanes are more brightly colored compared to the inner vanes. These inner feather vanes are usually covered by other feathers and are, thus, less visible to conspecifics. The only exception is the pair of central rectrices that are fully exposed, and both feather vanes are equally colored. In this species, males minimize the use of costly carotenoid pigments while maintaining elaborate ornamentation of plumage regions that are most visible to potential mates.
authors.affiliation: Department of Avian Biology and Ecology, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland.
authors.forename: Adrian
authors.initials: A
authors.lastname: Surmacki
authors.affiliation: Department of Avian Biology and Ecology, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland.
authors.forename: Anastazja
authors.initials: A
authors.lastname: Ragan
authors.affiliation: Department of Avian Biology and Ecology, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland.
authors.forename: Ziemowit
authors.initials: Z
authors.lastname: Kosiński
authors.affiliation: Institute of Zoology, Poznań University of Life Sciences, Wojska Polskiego 71C, 60-625 Poznań, Poland.
authors.forename: Marcin
authors.initials: M
authors.lastname: Tobółka
authors.affiliation: Department of Avian Biology and Ecology, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland.
authors.forename: Paweł
authors.initials: P
authors.lastname: Podkowa
keywords: Bombycilla garrulus; Color expression; Courtship behavior; Individual condition; Male display; Sexual secondary traits