Bower Decorations Attract Females but Provoke Other Male Spotted Bowerbirds
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Bower Decorations Attract Females but Provoke Other Male Spotted Bowerbirds: Bower Owners Resolve This Trade-off Author(s): Joah Robert Madden Source: Proceedings: Biological Sciences, Vol. 269, No. 1498 (Jul. 7, 2002), pp. 1347-1351 Published by: The Royal Society Stable URL: http://www.jstor.org/stable/3067697 Accessed: 29/09/2009 19:19 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=rsl. 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The Royal Society is collaborating with JSTOR to digitize, preserve and extend access to Proceedings: Biological Sciences. http://www.jstor.org Received 5 December 2001 E THE ROYAL Accepted 13 February 2002 SOCIETY Published online 11 June 2002 Bower decorations attract females but provoke other male spotted bowerbirds: bower owners resolve this trade-off Joah Robert Madden* Department of Animal and Plant Sciences, University of Sheffield, Sheffield, S10 2TN, UK ([email protected]) Elaborate secondary sexual traits offset the costs that they impose on their bearer by facilitating repro- ductive benefits, through increased success in intrasexual contests or increased attractiveness to choosy mates. Some traits enhance both strategies. Conversely, I show that spotted bowerbirds Chlamydera mac- ulata may face a trade-off. The trait that best predicts their mating success, numbers of Solanum berries exhibited on a bower, also provokes increased intrasexual aggression in the form of bower destructions by neighbouring bower owners, which reduce the quality of the male's bower. At natural berry numbers, levels of mating success in the population are skewed, but levels of destruction do not vary with berry number. When berry numbers are artificially exaggerated, increased levels of destructions occur, but mat- ing success does not increase. When offered excess berries, either to add to the bower or artificially placed on the bower, bower owners preferred to use numbers of berries related to the number that they displayed naturally. This decision is made without direct experience of the attendant changes in destruction or mating success. This indicates that bower owners may assess their own social standing in relation to their neighbours and modulate their display accordingly. Keywords: bowerbird; male competition; female choice; condition dependence 1. INTRODUCTION were dominated by duller males during the non-breeding season when for access to food. The evolution of elaborate sexual traits competing secondary puzzled Numerous studies have demonstrated that males exhib- Darwin who that the costs of males exhi- (1871) suggested the most elaborate sexual traits within a such traits could be offset benefits. iting secondary biting by reproductive the levels of success Elaborate traits enhance a male's chance of success population enjoy highest mating (for may reviews see Andersson Kotiaho in intrasexual 1994; Johnstone 1995; competition. Alternatively, they may few studies then on to enhance his attractiveness to females. In some 2001). However, relatively go choosy these correlates in order to test how such traits cases, traits function in both intrasexualcontest and inter- manipulate drive variation in mating success. Only one-third of the sexual choice. et al. (1996) review 48 cases of Berglund 232 studies of selected traits reviewed Anders- traits that function as both ornaments in intersexual sexually by (used son combine field observations with and armaments in intrasexual (1994) confirmatory choice) (used contests), One reason for the of such chemical and electrical experiments. paucity experi- including visual, acoustic, signals. ments is that a trait correlated with These two selective forces, inter- and intrasexualselection, altering morphological an organism's mating success is likely to have other effects usually operate in the same direction. For example, outside the context of sexual selection. For example, male Bisazza & Marconato (1988) showed that female bull- golden-headed cisticolas C. exilis with artificially shortened heads Cottus gobio preferred large males and that large tails-a trait that increases their mating success-showed males did better in intrasexualcompetitions. Male golden- reduced aerodynamic performance during slow-speed for- headed cisticolas Cisticolaexilis with artificiallyshortened aging flights (Balmford et al. 2000). tails were able to defend higher quality territories against Male bowerbirds build and decorate bowers, which act other males and also achieve increased reproductive suc- as the target of both intersexual choice by females (Borgia cess independent of territory quality, possibly due to 1985b; Borgia & Mueller 1992; Lenz 1994; Uy & Borgia enhanced aerial displays (Balmford et al. 2000). In cases 2000; Madden and intrasexual with such as these, a successful male fitness benefits via 2001a), contest, gains males the bower structure and both inter- and intrasexualselection neighbouring destroying through exaggeration decorations & Gore of the same trait. stealing (Borgia 1986; Borgia & Mueller 1992; Lenz This reduces the In some cases however, traits favoured female choice 1994). possibility by that such traits will confound success do not enhance the male's status. Work on house finches manipulating mating by altering other aspects of male behaviour. Despite this Carpodacusmexicanus (McGraw & Hill 2000) showed that apparently ideal situation, to my knowledge, only a single males bearing bright plumage (a trait favouredby females) previous study (Borgia 1985b) has attempted to determine whether widely reported correlations between numbers of decorations and male mating success reflect causal *Address correspondence to: 25 Church Lane, Deanshanger, Milton relationships. This experiment, which involved removing Keynes MK19 6HF, UK. all decorations except for three yellow leaves from satin Proc. R. Soc. Lond. B (2002) 269, 1347-1351 1 347 ? 2002 The Royal Society DOI 10.1098/rspb.2002.1988 1348 J. R. Madden Bowerbirdsresolve a trade-off Ptilonorhynchus violaceus bowers, was only able to conclude these 2 years. Data used in this analysis, relating to natural levels that decorations per se enhance mating success, rather than of destruction, were confined to those collected during the specifying which were of particular significance. breeding seasons of 1998 and 1999. This allowed me to com- At Taunton National Park (Central Queensland, pare average destruction rates from these 2 years with the rates Australia), spotted bowerbirds Chlamydera maculata dis- recorded during the experimental period, which covered the playing large numbers of Solanum berries were observed breeding season in 2000. Attendance rates in 1998 and 1999 to obtain increased mating success in 2 consecutive years were also collected from the videos in order to investigate (Madden 2001 a). I tested whether these berries were whether destruction rate was related to attendance at the bower. responsible for variation in levels of mating success by experimentally manipulating berry numbers at bowers. (c) Do males exhibit the maximum numbers of During the experiment, I observed that the manipulations Solanum berries available? that I was making provoked a surprising side-effect: alter- Two experiments were conducted to address this question. ing levels of bower destruction. First, a 'harvesting'(Diamond 1987) experiment was conducted This presents a situation where males may face a trade- at 19 bowers. All berries were removed from the avenue and off between whether to increase numbers of berries and placed in a cache 1.5 m from the avenue entrance. An equal enjoy increased mating success, or reduce berry numbers number of new berries, or 30 berries in total (equating to the and hence avoid intrasexual destructions. I tested this mean number of berries; see above) if there were few (less than trade-off experimentally with a series of cache experi- 15) berries naturallyon the bower, were added to the cache and ments, investigating whether males offered large numbers thoroughly shuffled, thereby providing easy access to at least of berries forego the opportunity to use them, despite the twice as many Solanum berries as were initially present on the potential for increased mating success. bower. Old berries were marked with a small cross using a per- manent pen, while new berries were marked with a circle. The cache was left for 2 and when I returned I recorded the 2. METHODS days, number of berries placed in the avenue. A second 'weeding' (a) Do numbers of Solanum berries cause