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Male Courtship Vocalizations As Cues for Mate Choice in the Satin Bowerbird (Ptilonorhynchus Violaceus)

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Male Courtship Vocalizations As Cues for Mate Choice in the Satin Bowerbird (Ptilonorhynchus Violaceus) MALE COURTSHIP VOCALIZATIONS AS CUES FOR MATE CHOICE IN THE SATIN BOWERBIRD (PTILONORHYNCHUS VIOLACEUS) CHRISTOPHER A. LOFFREDO AND GERALD BORGIA Departmentof Zoology,University of Maryland,College Park, Maryland 20742 USA AI3STRACT.--MaleSatin Bowerbirds(Ptilonorhynchus violaceus) court femalesat specialized structurescalled bowers. Courtship includes a complexpattern of vocalizationsin which a broad-band,mechanical-sounding song is followed by interspecificmimicry. We studiedthe effect of male courtshipdisplays on male mating successin Satin Bowerbirds.Data from 2 yearsof field researchshowed low between-maledifferences in mechanicalcomponents of courtshipsong and high variability betweenmales in mimeticsinging. Older malessang longer and higher-qualitybouts of mimicry than did youngermales. In one year, courtship song featureswere correlatedwith male mating success.The resultssuggest that female SatinBowerbirds use male courtship vocalizations in their mate-choicedecisions. We discuss hypothesesabout assessment of male age and dominancefrom courtshipvocalizations and suggestthat thesesongs have evolvedas a result of selectionfor male displaycharacteristics that provide femaleswith information about the relative quality of prospectivemates. Re- ceived27 June1985, accepted20 September1985. MALESatin Bowerbirds(Ptilonorhynchus vio- ingnessto copulateor by flying away. Given laceus)build specializedstructures called bow- the complexity of male display and the atten- ers that are used as sites for courting females tion femalespay to displayingmales, it is likely and for mating (Vellenga 1970,Donaghey 1981, that male displayshave an important effect on Borgia 1985). Femalesraise their young unas- female mating decisions,yet there have been sistedby males,and malesdo not associatewith no detailedstudies of displayin any bowerbird females after mating. Males decorate their species. bowers with a variety of natural objectsand The relationshipbetween male display and attempt to steal decorationsand destroy the female choice in polygynous species,such as bowers of other males. bowerbirds, in which males do not provide ma- Much effort has been focused on under- terial benefitsto femalesor their offspring,has standingthe role of the bower and its decora- been the subject of considerable speculation tions in female mating decisions.Borgia (1985) (Fisher 1930, Zahavi 1975, Davis and O'Donald demonstrateda skewed distribution of matings 1976, Emlen and Oring 1977, Halliday 1978, among male Satin Bowerbirds, differences Borgia 1979, West-Eberhard 1979, Bradbury among males in bower decoration and con- 1981, Hamilton and Zuk 1982, Bradbury and struction, and a consistent pattern of female Gibson 1983, Kodric-Brown and Brown 1984, preference for males with well-constructedand Borgia et al. 1985). One view of the sexual se- well-decorated bowers. In addition, male mat- lection process is that females choose mates ing successis influenced negatively by bower based on characters that indicate the relative destruction and decoration-stealing (Borgia ability of malesto sire fit (e.g. viable) offspring 1986a,Borgia and Gore 1986). (Zahavi 1975,Halliday 1978,Borgia 1979, Ham- Male courtship vocalizations are another ilton and Zuk 1982, Thornhill 1980, Borgia et component of male sexual display that may al. 1985). An alternative model suggeststhat have important effectson female mating deci- male sexualdisplays are arbitrary resultsof the sions. Male courtship is intense, involving runaway processof sexual selection (Fisher dance movements and postures that are coor- 1930).This implies that femalepreferences have dinated with the simultaneousproduction of evolved that have no effect on enhancingthe complex vocalizations. The female crouches viability of the offspring(Lande 1981,Kirkpat- within the bower, closely observing the male rick 1982, Arnold 1983). during the display, and controls the outcome Developing hypothesesthat separate these of the courtshipeither by signaling her will- modelshas proven very difficult. One method 189 The Auk 103:189-195. January 1986 190 LOFFREDOAND BORGIA [Auk, Vol. 103 is to consider the evolution of male display Analyzer (Model 4512). A Kay ElemetricsDigital (Borgia 1986a).Models that predict gainsin off- Sono-Graphwas used to prepare spectrograms. spring viability require that male displayspro- Analysis-of-variance tests (one-way ANOVA), vide cues that allow females to assess characters Spearman rank correlations (r.), and the Student- importantto demonstratingthe quality of males Newman-Keuls (SNK) procedure for multiple comparisonsof meanswere used for statisticalcom- as sires. For example, female preference for parisons(Helwig and Council 1982). Means are ex- older males may produceoffspring with high pressedas g + 1 SD. survivorship becausethese males have dem- Becauseof the low amplitude of mimetic sounds onstratedhigh viability through their own sur- (seebelow), it wasnot possibleto accuratelymeasure vival (Trivers 1972, Howard 1974, Halliday frequencies.However, it was apparentthat quality 1978, Wilbur et al. 1978). If such a model is of mimetic songsdiffered between males,and so a operating, then we expect that cues exist in subjectivemethod was used to produce an index of malesthat indicate male age and that these are qualitativedifferences among mimetic songsof dif- correlated with male mating success.Alterna- ferent males.All mimetic songswere given a score tively, females may prefer dominant (Borgia (1, 2, or 3) for "structuralintegrity": high-scoredmi- meticsongs were of relativelypure tonal structure, 1979) or disease-resistantmales (Hamilton and free from broad-bandand other atonalsounds pro- Zuk 1982). By contrast,runaway models make duced by the singer, and were free from acoustical no predictions about the nature of male dis- distortionsresulting from productionby the singer plays and are untestableon that basis. of other soundsduring mimetic singing.Figures 2 Our objectiveshere are to describeintrapop- and 3 represent several high- and low-scored mi- ulational variation in the courtship-display metic songs. vocalizations of Satin Bowerbirds, to look for To validate subjectivescoring of mimetic songs, correlates between male vocal patterns and two volunteerswere each given the same random other male characteristicssuch as age and bow- sampleof spectrogramstaken from the analyzeddata er quality, and to usethese results to determine and were askedto sort the spectrogramson the basis if the minimum conditions for male assessment of this three-level subjectivescoring system. There was completeagreement between the resultsof the models have been met. two independenttrials and the resultsobtained by the authorfor the samesample (r. = 1.00,P < 0.001). METHODS Field researchwas carried out at Wallaby Creek, RESULTS BeauryState Forest, New SouthWales, Australia, from August through December in 1982 and 1983. The Descriptionof the male courtshipsong.--The study site is characterizedby open-canopy,subtrop- song used by male Satin Bowerbirdsduring ical eucalyptusscrub forest and patchesof rain forest. courtship(and in other specificbehavioral set- Recordingsof maleSatin Bowerbird courtship songs tings; see below) consistsof a long seriesof were obtained with Uher 4000L and Marantz PMD harsh-soundingbuzzes and pulses followed 220 tape recorders,using small condensermicro- immediatelyby a boutof interspecificmimicry. phonessuspended in the foliage1 m from the bower The two major portions of the courtshipsong platform. Only sexualdisplay songs at or near the henceforth will be termed the "mechanical" and bower were recorded,although male Satin Bower- birds use a variety of calls in other behavioralset- "mimetic" portions. tings and at placesaway from the bower. Observa- The courtshipsong lasts24.0 + 8.2 s. Within tions of focal displaying maleswere made by field the mechanicalportion there are 2-6 (œ= 4.2) observers from blinds stationed within 10 m of the wide-band "long buzzes"(Fig. 1), eachof which bower. In addition, a camera system was used to is 0.35-1.50 s long with a frequency range of monitoractivities continuously at bowersand to ob- 0.06 --- 0.01 to 7.39 + 0.02 kHz. Each long buzz tain recordsof male mating success(see Borgia 1985). is separatedfrom the next by one or more "short We recorded vocalizations in 1983 for 22 individ- buzzes" (harsh soundswith a frequency range ual males(20 adultsand 2 juveniles).Recordings from of 4.00 +__0.03 to 7.20 ___0.05 kHz, each of which 1982 for 2 additional juveniles and 11 of the same lasts for less than 0.20 s) and several short adultswere includedin the analysis,for a totalof 24 individuals.A meanof 12.0vocal samples were ana- (<0.10 s) pulselike sounds(Fig. 1). The me- lyzedfor eachmale, with a totalof 288samples. Spec- chanicalportion of the song lasts 17.0 + 3.2 s. trographic sound analysiswas performed using a For eachmechanical song analyzed, the follow- PrincetonApplied ResearchFFT Real Time Spectrum ing variables were measured:the duration of January1986] BowerbirdCourtship Vocalizations 191 16- 0 • • 4 Time (sec) Fig. 1. Componentsof the mechanicalportion of male Satin Bowerbird courtshipsong. Time (sec) Fig. 3. Lewin'sHoneyeater mimetic song: (A) high the mechanical song; the numbers of long and (B) low scoresfor structuralintegrity. The sounds buzzes,short buzzes,and pulses;and the peak between 4.0 and 6.0 kHz are cicada calls. frequency(kHz), low frequency(kHz), and du- ration of each long buzz. ic calls are given, the kookaburra call always
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