Multiple Sexual Ornaments in Satin Bowerbirds: Ultraviolet Plumage and Bowers Signal Different Aspects of Male Quality
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Behavioral Ecology Vol. 14 No. 4: 503–509 Multiple sexual ornaments in satin bowerbirds: ultraviolet plumage and bowers signal different aspects of male quality Ste´phanie M. Doucet and Robert Montgomerie Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6, Canada Much attention has been devoted to understanding the evolution of elaborate male ornaments and how they may signal male quality. However, the evolution of multicomponent sexual signals remains poorly understood, and past research on this type of signaling has been largely theoretical. Satin bowerbirds, Ptilonorhynchus violaceus, are polygynous, are sexually dichromatic, and construct sexually selected display structures (bowers): a model system for investigating the evolution and signal function of multiple sexual signals. We studied the interrelationship between bower features, plumage coloration, and indicators of male quality in this species. To do this, we located the bowers of male satin bowerbirds in rainforest in Queensland, Australia, and quantified bower quality. We captured the male bower owners and used reflectance spectrometry to objectively measure the plumage coloration of several body regions. We measured various indicators of male health and condition, including the intensity of infection from ectoparasites and blood parasites. Bower quality and male ultraviolet plumage coloration were significantly correlated. By using multiple regression analyses, we show that bower quality predicts ectoparasite load and body size, whereas ultraviolet plumage coloration predicts the intensity of infection from blood parasites, feather growth rate, and body size. Our findings support the multiple messages hypothesis of multicomponent signals: Female satin bowerbirds should assess both male and bower features to choose the highest quality mates. Key words: bowerbirds, bowers, honest advertisement, multiple ornaments, parasites, plumage color, sexual selection, structural colors. [Behav Ecol 14:503–509 (2003)] laborate, secondary sex traits are often presumed to on different time scales (daily for bowers and yearly for E communicate aspects of male quality to discriminating plumage color), these multicomponent sexual signals may females (Andersson, 1994). In many species, elaborate sexual reveal different aspects of male quality (Johnstone, 1995b; displays involve several distinct signals such as plumage Møller and Pomiankowski, 1993), and females may thus ornamentation, song, and display behavior. Some theoretical obtain useful information from evaluating all components of models have shown that, under certain circumstances, species male display. could evolve multiple quality-revealing sexual ornaments In satin bowerbirds, individual variation in the quality of ( Johnstone, 1995a), but other models propose that honest bower construction, the numbers and types of decorations advertising should favor a single most-revealing signal at the adorning the bower platform, and age-correlated features of expense of others ( Johnstone, 1996; Schluter and Price, 1993). courtship song are known to influence female choice of Despite some theoretical consideration ( Johnstone 1995a, copulation partners, as measured by male mating success 1996; Schluter and Price, 1993), the evolution of multiple (Borgia, 1985b; Lofredo and Borgia, 1986). In contrast to quality-indicating sexual ornaments has received little empir- other features of the sexual display of male satin bowerbirds, ical support (for exceptions, see Johnstone, 1995b). In many the structurally based iridescent plumage of males has species, the evolution of more than one sexual signal can be received less attention, even though this species is highly more accurately attributed to Fisherian arbitrary selection sexually dichromatic and plumage ornamentation is a prom- (Fisher, 1958; Møller and Pomiankowski, 1993; Pomian- inent feature of male courtship display. Recent studies of kowski and Iwasa, 1993), signal amplification (Hasson, other bird species have shown that structural plumage 1989), enhanced signal detection (Rowe, 1999), or to distinct coloration can honestly signal condition (Doucet, 2002; signals intended for different receivers (Pryke et al., 2001a,b). Keyser and Hill, 1999), male quality (Keyser and Hill, 2000), In satin bowerbirds, Ptilonorhynchus violaceus, males clear and viability (Sheldon et al., 1999). Thus plumage coloration display courts, build bowers (elaborate structures constructed may be a quality-indicating mechanism in male satin bower- from twigs), and decorate the bower platform with a variety of birds and may function as a key component of a complex, colorful natural and human-manufactured objects (see multimodal sexual signal of male quality. In the present study, Marshall, 1954). Females of this species observe male court- we investigated how multicomponent sexual signals can ship from within the bower avenue, and the sexual display of potentially reveal male quality in this species by studying the courting males incorporates song mimicry, ritualized pranc- relationship between male plumage coloration, bower fea- ing, and brilliant plumage coloration. Because different tures, and various indicators of male health and condition. components of the male satin bowerbird’s display will vary METHODS Address correspondence to S.M. Doucet, who is now at the This study was conducted from September–December 2000 in Department of Biological Sciences, 331 Funchess Hall, Auburn a 700-ha study area in Mount Baldy State Forest, Queensland, University, Auburn, AL 36849, USA. E-mail: [email protected]. Australia (17°309 S, 145°309 E). The dominant habitat was Received 3 April 2002; revised 3 September 2002; accepted 21 rainforest with a sharp transition to wet sclerophyll forest at September 2002. the northeastern edge of our study area. We located 12 satin Ó 2003 International Society for Behavioral Ecology bowerbird bowers in our study area by listening for male 504 Behavioral Ecology Vol. 14 No. 4 dicular to the feather surface. All spectral measurements are expressed as the proportion of light reflected relative to that reflected from a Spectralon white standard, an almost perfect reflector. We measured plumage reflectance on four body regions for each individual: wing coverts, breast, mantle, and rump, taking five readings from each region and moving the probe by at least 5 mm between readings. We restricted spectral analyses to wavelengths from 300–700 nm, as most birds are sensitive to ultraviolet (UV) wavelengths (300–400 nm; Cuthill et al., 2000), and 700 nm is likely the upper limit of the vertebrate visual spectrum (Jacobs, 1981). We summarized reflectance data by calculating five color variables: total brightness, UVV-brightness, UVV-chroma, con- trast, and hue. We calculated total brightness as the mean of reflectance values, in 1-nm intervals, from 300–700 nm (Andersson, 1999; Endler, 1990); this measurement thus gives the mean proportion of incident light that is reflected from the feather surface in the bird-visible range of wavelengths. Similarly, we calculated UVV-brightness as the mean of reflectance values in the UV–violet region of the spectrum (300–420 nm; Andersson et al., 1998), in which peak Figure 1 reflectance occurred in all males. As a measure of spectral sat- Representative reflectance spectrum (average of rump, mantle, breast, and wing coverts) for one adult male satin bowerbird. This uration, we calculated UVV-chroma as the proportion of total individual was chosen because its plumage color was closest to the reflectance occurring in the UV–violet region of the spec- mean for the color variables we calculated across all individuals trum. We calculated contrast as the difference between the sampled. maximum and minimum reflectance across the 300–700-nm range, such that higher contrast indicates a more intense color (Keyser and Hill, 1999, 2000). Given the simple, advertisement calls, usually given near bowers, and systemat- unimodal shape of adult male satin bowerbird reflectance ically searching for active display sites. We assessed bower spectra (Figure 1), contrast is likely a reliable estimate of color quality by evaluating bower construction and quantifying the intensity in this species. Finally, to approximate hue, we used number of decorations used. Two observers independently the wavelength at which maximum reflectance was reached, evaluated four features of bower construction—overall sym- a commonly used index of spectral location (see Keyser and metry of the structure, stick size, stick density, and overall Hill, 1999, 2000; Sheldon et al., 1999). quality of construction—each on a scale from one (poor) to We also calculated an overall ornamental color score using four (excellent). We calculated a bower quality score for each principal components analysis (PCA). We first calculated the bower as the sum of these measures averaged between the two mean of each color variable for each body region on each observers. Similar indices of bower quality have been shown to male. We then performed a PCA by using the mean total correlate with mating success in several species of bowerbird brightness, UVV-chroma, contrast, and hue averaged over the (Borgia, 1985b; Borgia and Mueller, 1992; Lenz, 1994; Uy and four regions of each male studied. The first principal Borgia, 2000). To assess the extent of bower decoration, we component (PC1) from this analysis explained 46% of the regularly visited bowers from mid October to late November variation in color among males, with total brightness,