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A Generalized Female Bias for Long Tails in a Short-Tailed Widowbird

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A Generalized Female Bias for Long Tails in a Short-Tailed Widowbird Received 7 May 2002 Accepted 8 July 2002 Publishedonline 25September 2002 Ageneralizedfemale biasfor long tailsin a short-tailedwidowbird Sarah R.Pryke * and Staffan Andersson AnimalEcology, Department of Zoology,Go ¨teborgUniversity, Box463, SE-405 30 Go ¨teborg,Sweden Tail elongation in thepolygynous widowbirds( Euplectes spp.)has evokedboth adaptive andnon-adaptive explanations. Female choicehas beenshown in thethree longest tailed species(20– 50 cm),whereas an agonistic functionwas proposed for amedium-tailed (10 cm) widowbird.To testthe generality anddirec- tionality ofsexual selectionon tail length in widowbirds,we experimentally investigated selectionin the relatively short-tailed (7cm)red-shouldered widowbirds ( E. axillaris ).Prior toterritory establishment, males wereassigned to four tail-treatment groups;control, short, long andsupernormal (similar toa sympatric long-tailed congener).No effects on male competition weredetected as the groups wereequally successfulin acquiring territories ofsimilar sizeand quality. However,mating successamong the92 territorial males wasstrongly skewedin favour ofsupernormal-tailed males (62% ofactive nests;5.2 ± 1.3 nestsper territory). Long-tailedmales also acquired more nests(1.9 ± 0.7) than control(0.7 ± 0.5) and short-tailed (0.5 ± 0.3) males,while thelatter twogroups didnot differ signi cantly. Theseresults support ageneral, open-endedfemale preferencefor long tails in widowbirdsand may representa receiver bias that aroseearly in their divergencefrom theshort-tailed weaverbirds (Ploceinae). Keywords: sexual selection;tail length; receiver bias; Euplectesaxillaris ;widowbirds 1. INTRODUCTION To differentiatebetween the debated functions of tail elongation, weexamined theselection pressures on tail The extreme variation in tail elongation among themale length in oneof the shortest-tailed species, the red- breeding plumages ofthe highly polygynous andecologi- shoulderedwidowbird (tail ca.7cm).Like other widow- cally similar African widowbirds Euplectes spp. (Craig birds,the male has pronouncedsexual andseasonal 1980; Andersson& Andersson1994) is astriking example dimorphism, andmoults from acryptic sparrow-brown ofenigmatic diversity in ornamentexpression. Male tail non-breedingplumage intoa black nuptial plumage with a length varies from therelatively short tails (7cm) ofyel- shorttail andred carotenoid-based epaulettes (lesser wing low-rumped( E. capensis)andred-shouldered widowbirds coverts).During thebreeding season,these birds are terri- (E.axillaris )to50 cm in thelong-tailed widowbird torial andinhabit marsh andgrassland areas in central and E. progne.The functionof this signal has evokedboth southernAfrica. In ight display, themale has aslow, adaptive andnon-adaptive explanations (Savalli 1993, bouncy‘ rowing’action with exaggerated wing beatsand 1995; Andersson& Andersson1994; Craig &Villet afoldedtail. On their territories, males build numerous 1998), butmany ofthe basic assumptionsremain incon- coarsenest frames (‘cock’s nests’), which thefemales line clusively tested.One of the main questionsconcerns the with nergrass, and then incubate and feed the nes- generality ofdirectional female preferencesfor elongated tlings alone. tails. Female choiceof long tails wasinitially demonstrated In this study,we experimentally manipulated male red- in long-tailed widowbirds(Andersson 1982) andhas sub- shoulderedwidowbird tails both within thenatural range sequentlybeen shown in thelekking Jackson’s widowbird (i.e.short, average andlong) andfar beyond,creating a E. jacksoni (Andersson1989, 1992 (tail 20 cm)) andterri- supernormal tail similar in length tothat ofthesympatric torial red-collared widowbirds E. ardens (Pryke et al. 2001 red-collared widowbirds.As described by Tinbergen (tail 22 cm)).By contrast,tail length in themedium-tailed (1948) in early ethological literature, andrecently (10 cm) yellow-shoulderedwidowbird ( E.macrourus ) explored in articial neural networks(e.g. Enquist & Arak seemedto function in male competition for territories 1998), supernormal stimuli oftenelicit stronger receiver rather than female choice(Savalli 1994 a,b).Furthermore, responsesthan thenatural range ofstimuli towhich the other selectiveforces, such as stabilizing selection,may receiver is genetically or phenotypically adapted.For have contributedto the diverse tail variation among the example, if tail length is subjectto sexual selection,males widowbirds.For example, Savalli (1995) suggestedthat displaying supernormal tails may bemore dominantin tail length might beconstrained by speciesrecognition in male–male contestsor attract more females.Furthermore, areas ofsympatry with longer tailed congeners.However, if supernormal stimuli greatly exceed(by several s.d.)the only oneof the two investigated speciespairs showed natural range, astrong responseindicates that generalized character divergenceand only in apositive direction(i.e. preferencesand receiver biasesalone might besuf cient oneof the widowbirds had longer tails in sympatry than tocause and maintain signal exaggeration (e.g.Enquist & in allopatry). Arak 1998), i.e.without invoking Fisherian or ‘good genes’coevolution between the preference and signal. By contrast,females are also understrong selectionto *Authorfor correspondence ([email protected]). avoid heterospecic mating. If tail length is an essential Proc.R. Soc.Lond. B (2002) 269, 2141–2146 2141 Ó 2002 TheRoyal Society DOI10.1098/ rspb.2002.2131 2142S. R.Prykeand S. Andersson Generalfemale bias for long tails 7 120 y c n 6 e u q 80 s e t r s f e 5 l i n a t e 40 l v i a t r c 4 u a t a f n o 0 r e 3 b m u n 2 n a e m 1 0 246781012 14 16 18 20 22 manipulated tail length cm) Figure 1. Theaverage ( ± s.d.) reproductive success of tail-manipulated malesthat established territories; the6 cm manipulations represent theshort-tailed ( n = 25), 7cm thecontrol-tailed ( n = 23), 8cm thelong-tailed ( n = 23) and 22 cm the supernormal-tailed ( n = 21) males. Theinner histogram showsthe natural distribution of breeding male tail length in the population ( n = 252), corresponding to theaxis of experimental tail lengths. componentof species recognition, and if ‘supernormal ’ (b) Colorimetrics tail length in onespecies is ‘normal’ toanother species Spectral re ectance(at 2nmresolution) from the redepaul- with which hybrid matings might occur,females should ettewas measuredwith an S2000spectrometer (Ocean Optics, avoid or at least beindifferent to asupernormal tail com- Dunedin,USA), using an HL2000halogen light sourceand a pared with trait values within orcloserto the conspeci c bre optic re ectanceprobe. The probe was heldperpendicu- variation. By simultaneously,and under natural signal larlyagainst the plumageand vescans fromthe centreof the conditions(i.e. a eldexperiment), exploring the epaulettewere taken and averaged for eachindividual. Using responsesto both ‘normal’ and ‘supernormal ’ tail the C-specsoftware (Ancal,Las Vegas, USA)re ectance was manipulations (resembling thesympatric red-collared measuredas the proportion of light re ectedin relationto aWS- widowbird),this studyprovides adirecttest of stabilizing 2white standard (morethan 98%re ectanceacross the measur- constraintsarising from speciesrecognition versusdirec- ing range). tional sexual selectionfor elaborate male tails. Objectiveindices of the threemain dimensions of colourper- ception(spectral intensity, location and purity)were computed fromthe raw spectralre ectancedata and then averaged for 2. METHODS eachindividual. ‘Brightness’ (spectralintensity) was estimated (a) Morphologicalanalysis by R350–700,the sumof re ectancefrom 350 –700 nm. ‘Hue’ The experimentwas carriedout ona population of red- (spectralintensity) was estimatedas l(R50),the wavelength at shoulderedwidowbirds inthe Balgowan district,KwaZulu- which re ectanceis halfway betweenits minimum( Rmin) and Natal, South Africa,between early December 2001 and March maximum (Rmax). ‘Chroma’ (spectralpurity) is based on several 2002.Males were captured with mistnets at night roosts and aspects, such as re ectanceslope height and steepness.To markedwith an aluminiumring and threeunique colour bands. incorporateboth of these spectralshape aspects, we useda seg- Measurementsof wing chord(to the nearest0.5 mm), culmen ment-based estimatethat comparesthe re ectanceabove and (tothe nearest0.1 mm), tarsus (tothe nearest0.1 mm), tail below l(R50),divided by Rav (average re ectance)to control length (tothe nearest0.1 mm) before and after manipulation, for brightness, i.e. C[R50] = (R[l(R50) – 700] – R[350 – l(R50)])/Rav. and mass (tothe nearest0.5 g) weretaken. Becausethe three Detailson further methods and analysis of re ectancedata are linearmeasures of body size(tarsus, culmenand wing) were providedin Pryke et al. (2001). intercorrelated,the rst componentsof aprincipalcomponents analysis,explaining 81.7% of the variance,were used to extract (c) Behaviouraland territory characteristics an independentmeasure of body size.An index of body con- Territorialmales were observed for 30minweekly over a ditionwas estimatedfrom the residualsof the linearregression 13weekperiod when the malesdefended their breeding territor- of log(body mass)on 3log(tarsus length)(S. Andersson 1994). ies.Observationswere randomized and allactivities and their Epaulette sizewas calculated(to the nearest0.1 mm) as the durations werecontinuously recorded with the timegiven to the product of the maximumlength and the average of three nearest5 s.Data onthe timespent on the territory(min
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