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The Evolution of Leks Through Female Choice: Differential Clustering And BehavEcol Sociobiol (1992) 30:227-237 Behavioral Ecology and Sociobiology ? Springer-Verlag1992 The evolutionof leks throughfemale choice: differentialclustering and space utilizationin six sympatricmanakins Marc Thery C.N.R.S. - U.R.A. 1183, Mus6umNational d'Histoire Naturelle, Ecologie Generale,4 avenue du Petit-Chateau, F-91800 Brunoy, France Received October 1, 1990 / Accepted October 2, 1991 Summary.The degreeto which lekkingand non-lekking the increasingfemale home-rangesize could have led male manakinsselect display sites in order to maximise to the evolution of classicalleks. proximityto femaleswas examinedby contrastingmove- ments of females with male dispersion.Data on female visitingpatterns, male courtshipdisruption, and mating skew were also collected over three successivebreeding Introduction seasons. For the five lek-breedingspecies, female home- rangeswere 3-7 times largerthan those of adult males. Lek matingsystems, where males congregatefor the sole Femalemovements were concentrated around leks, fruit- purpose of attractingand courtingfemales, are typified ing places and streambathing sites. None of the females by a heavilybiased operational sex ratio and the inability monitoredby radio-trackingexpanded her normalrange of individualmales to economicallycontrol or monopo- in order to visit males on leks. On the contrary,feeding lize the resourcesessential for femaleacquisition (Emlen bouts of femalesfrequently preceded a visit to potential and Oring 1977). Across lek mating systems,male clus- mates at neighboringleks. Despite small sample sizes, teringis highly variableand can be dividedinto "classi- significant correlations were found between female cal leks" (Bradbury1977, 1981) also called "true leks" home-rangesize and male clustering(distances between (Oring 1982), where males are densely grouped within neighboring leks and distances between neighboring sight of each other, and "explodedleks" (Gilliard1963; males), as predicted by the female choice model and Snow 1970; Bradbury 1981; Foster 1983) or "quasi- the hotspot model. Adult and immature male home- leks" (Oring 1982), where males display from courts range sizes were not significantlycorrelated with male which are clumpedwhen consideringstatistical patterns dispersionor female ranges. On the other hand, males of dispersion. Beehler and Foster (1988) consider leks and females of the only non-lekking species exhibited to be only those court-based systems in which males similar use of space and home-rangesize. Male settle- are clusteredin space; non-clusteredsite-based systems ment at sites with high levels of female traffic showed are treated as non-lek "court" systems. After several that the hotspot model is adequateto explaindifferences attemptsto explain the evolution of leks (see Bradbury in male dispersion among sympatric lekking species. 1981; Oring 1982; Bradburyand Gibson 1983; Arak Comparisonswith other studies suggest that apparent 1984; Beehlerand Foster 1988; H6glundand Robertson female choice could be overidden by past and present 1990a), two sets of hypotheses presentlyinfluence the male-maleinteractions or female mate-comparisontact- theoreticalperception of lek systems: ics. In fact, both the hotspot model and the attractive- 1. The "hotspot model" where clumpingresults be- ness hypothesisappear to shapemale dispersionon leks: cause males settle at sites where the largest number of males appear to settle under hotspot conditions with females are likely to pass (Lill 1976; Emlen and Oring "despotic"rules generated through bias in femalechoice 1977; Payne and Payne 1977; Bradburyand Gibson or male-maleinterference. It is proposedthat the evolu- 1983). Leks can be located less than one female home- tion of leks is ecologicallymotivated by the spatio-tem- range apart and most females visit more than one lek poral distributionof trophic resources,initially leading before mating. Female range size should be positively to a dispersedmale-advertisement polygyny. Following correlatedwith male clumping and hence the distance this, a foragingecology that promotes high mobility by betweenleks. Despotic settlementrules, arisingthrough femalesand the magneticeffect of matingskew in partic- female preferencesfor particularmales or asymmetric ular malesmay have favoredclustering on explodedleks. interferencebetween neighboring males, may help to ex- Later, the development of male-male interferenceand plain why males are less clumpedthan expected(Brad- This content downloaded from 134.153.184.170 on Tue, 13 Jan 2015 08:19:15 AM All use subject to JSTOR Terms and Conditions 228 bury et al. 1986). Hotspots could be discriminatedfrom ley and Ahlquist 1985; Sibley et al. 1988; Prum 1990; the "female choice" model (Bradbury1981) where fe- Prum and Lanyon 1991; R.O. Prum in prep.). I captured male preferencefor males in larger groups causes leks only one tiny tyrant-manakin Tyranneutesvirescens, and to be spaced at least as far apart as an averagefemale will therefore not consider this species in the results. range diameter, although Gibson et al. (1990) have In French Guiana, as in other countries, M. manacus shown that some versions of the female choice model and P. erythrocephala form classical leks (Snow 1962a, can be consistent with a female's range including a 1962b, Lill 1974 a, b, 1976; Thery 1990 a), P. pipra forms numberof differentleks. exploded leks (Snow 1961; Thery 1990a), and S. turdinus 2. The "hotshot model" stresses the importanceof males sing from widely dispersed territories (Skutch male-male dominance interactions and attributes less 1969, 1981; Thery 1990a). On the other hand, P. serena significanceto female choice in the development and and C. gutturalis show geographical variation in lek be- maintenanceof leks (Foster 1983; Beehler and Foster havior. P. serena forms exploded leks in Suriname (Prum 1988). This hypothesis suggests that certain males, be- 1985), and "double-spot leks" in French Guiana: cause of behavioralor morphologicalattributes, are ex- groups of males, densely clustered on classical leks, tremely successful at attractingmates. Very similar to which move every day from one spot to another during the hotshot model, but weighingmore heavily the driv- set display hours (Thery 1990b). C. gutturalis forms ex- ing force of female preferences,is the "attractiveness ploded leks in French Guiana (Thery 1990b), and "mo- hypothesis" where aggregatingsubordinate males gain bile leks" in Surinam: males abandon exclusive display advantagesby exploitingthe ability of dominantsto at- territories for the competitive group display at a series tract potential mates (Waltz 1982; Arak 1983, 1984; of sites (Prum 1986). Hoglund and Robertson 1990a). Spatial position of males on leks should thereforebe consideredas a conse- quencerather than a causeof differencesin male attracti- Methods veness and dominance(H6glund and Robertson1990a). The attractivenesshypothesis makes no firm prediction A total of 83 adult males, 58 immaturemales and 114 adult females about female movements and thus may not constitute of all species combined were caught in mist nests 0-2 m from the an alternativebut a to the more ground during 125 days (1312 h) in January-December1985, Feb- complement hypotheses 1986 and 1987 councerned with female ruary-July February-July (Table 1). Birds were directly home-range patterns banded with permanentnumbered bands and individualcombina- (H6glundand Robertson1990a). tions of coloured plastic bands. The movements of 12 adult males, In the manakins(Passeriformes: Pipridae) lek mating 6 immature males in transitional plumage and 10 adult females is the rule, but sympatricspecies show differentpatterns were monitored with "SS1" 1.5-g radio transmitters from Bio- of clusteringon classical or explored leks. To test the track, England (Table 1). Twenty-fivebirds were studied for 6 or model of lek I followed 7 days during the breeding seasons when leks were fully active hotspot evolution, suggestions and three individuals were of et al. and McDon- copulations observed; monitored for Bradbury(1981), Bradbury (1986) 3 or 5 days during the nomadic period when leks were abandoned ald (1989a) by examiningmale clusteringand intersexual by adult males. For all lekking species, monitored females were use of space for sympatricspecies. I also presentinfor- known to be adults because they remained in green plumage for mation about female visiting patterns,mate choice, dis- more than one breedingseason, and monitoredmales where known ruptionand mating skew in the same species. to be immature when they were in transitional plumage or did not remain in green plumage for more than one season. In S. turdinus,the breeding season was considered as the period when males' were heard. In absence of sexual area and songs regularly dimorph- Study populations ism, S. turdinusindividuals were sexed using Payne's(1984) biomet- ric methods. Transmitterswere attached to the dorsum, after sec- My observationswere made at two locations in French tioning the feathers 1-2 mm from their base, with cyanoacrylate Guiana: the "piste de St Elie" site (5?04'N, 53018'W) surgicalglue (Aron alpha S2, Taogosei ChemicalIndustries, Japan). 12 km inside the tropical moist forest (Lescure et al. The transmittersdid not noticeably affect the birds' behavior. The Lescureand Boulet Sabatier and receiver, Yaesu FT 290R modified for 148 MHz, was used with 1983; 1985; 1985), the a directional antenna. Birds were monitored des Yagi continuously "inselberg Nouragues" site (4005'N,
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