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 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 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 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 developmentof 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). 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,, 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, 52040'W) from dawn to dusk for a total of 173 days (1900 h). Locations around a granite dome 100 km from the coast (Erard were recorded visually or by compass triangulation from points et al. 1989). Between January 1985 and July 1987 (23 placed every 20 m in the field. Fixes were systematicallytaken for months), I color-banded254 manakins of six species: each new 's location. Home-rangeswere computed for six or 75 white-crownedmanakins Pipra pipra pipra, 67 gold- seven days of continuous radio-tracking,using high and equivalent en-headed manakins P. numbers of fixes for each monitored individual (Table 2, mean+ erythrocephala erythrocephala, SD=384+21 2= Contour areas 35 white-frontedmanakins P. serena 32 white- fixes, 15.66, df=13, P>0.27). serena, were drawn on a map using the modified minimum-areapolygon bearded manakins Manacus manacus manacus, 23 white- method (Harvey and Barbour 1965), and measuredby planimetry. throated manakins Corapipogutturalis, and 22 thrush- I located on each female home-range map the areas where the like manakins Schiffornis turdinus wallacii. The family individual spent the highest percentageof time, and calculated the of status of Schiffornis turdinus is uncertain (Wetmore percentage female home-rangesize accounting for the top 75% Snow McKitrick and recenttaxonom- of the observations. C. gutturaliswas not equipped with transmit- 1972; 1975; 1985) ters because body weights are less than 10 g (Table 2). Therefore, ic studies,using proteinelectrophoresis, DNA-DNA hy- home-range size was estimated for eight individuals using as fixes bridizationor morphologicalanalysis, no longer include the locations of 8 captures, 19 recapturesand 179 visual sightings the genus Schiffornisin the Pipridae(Lanyon 1985; Sib- away from leks (Table 2). Locations of leks and distances between

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Table 1. Breeding system and numbers of banded individuals, recaptures,visual sightings and individuals monitored by radio-tracking for each species, age and sex of manakins studied

Species Age, sex NB NRb NVC NRTd Breedingsystem

Piprapipra Adult male 10 9 9 3* Exploded lek Immaturemale 27 11 4 2 Adult female 38 6 18 2 P. erythrocephala Adult male 19 18 7 3* Classical lek Immaturemale 14 6 2 1 Adult female 34 5 8 2 P. serena Adult male 15 14 10 3 Double-spot lek Immaturemale 6 4 4 1 Adult female 14 5 10 3* Manacusmanacus Adult male 18 2 14 2 Classicallek Immaturemale 5 1 3 2 Adult female 9 0 7 2 Corapipogutturalis Adult male 9 10 107 0 Exploded lek Immaturemale 6 5 38 0 Adult female 8 9 72 0 Schiffornisturdinus Adult male 12 7 8 2 Solitary display Adult female 10 3 6 1

254 115 327 28 a NB = number of banded individuals b NR = number of away from leks c recaptures NV = number of visual sightings away from leks d NRT = number of individualsmonitored by radio-tracking * Indicateswhen one individualwas monitored outside the breedingseason

Table2. Body weight, duration of radio-tracking,number of fixes, and home-range size of 25 manakins monitored during the breeding season, and area accounting for 75% of female traffic

Species CTa Home-range sizeb (NDC, NFd) Se (body weight) Adult males Immaturemales Adult females

P. pipra RT 2.1 (7, 361) 18.7 (6, 380) 8.8 (6, 370) 25 (11.0-12.8 g) 2.3 (7, 350) 20.9 (7, 357) 9.2 (6, 381) 29 P. erythrocephala RT 3.0 (6, 379) 22.7 (7, 413) 12.8 (6, 380) 28 (11.8-12.6 g) 3.4 (6, 390) 14.4 (7, 363) 30 P. serena RT 2.8 (6, 410) 24.1 (7, 365) 11.2 (6, 405) 22 (10.7-11.3 g) 3.2 (6, 387) 13.0 (7, 385) 28 M. manacus RT 2.1 (7, 355) 20.1 (6, 378) 13.0 (6, 420) 27 (16.0-17.5 g) 2.5 (6, 372) 20.3 (7, 350) 14.1 (6, 375) 29 C. gutturalis C 2.0-2.5 [123] ? 9.0-10.0 [83] ? (8.1-8.8 g) S. turdinus RT 14.3 (6, 416) ? 20.1 (7, 407) 68 (30.4-31.8 g) 16.5 (6, 434)

-= (63, 3854) (40, 2243) (57, 3486) a Census Technique:RT = radio-tracking,C = netting and visual control b Home range size for each monitored individualin ha c ND =duration of radio-trackingin days d NF = correspondingnumber of fixes used in home-rangeestimate e S= % female home-rangesize accounting for the top 75% of female traffic In C. gutturalis, home-range size is estimated for four males and four females using locations of 8 captures, 19 recaptures and 179 visual controls (numbersof fixes are given in square brackets)

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neighboring males were measured directly in the field from topo- species,P. pipra (Fig. 1). For the solitarydisplaying spe- graphically known points. Because I worked alone for a limited cies, I presentthe activitiesand home-rangesof one male period, sample sizes per species were relatively small, particularly and one female duringthe breedingseason (S. turdinus, for home-range estimates. Despite this limitation and only five studied species, correlation coefficients between home-range sizes Fig. 2). and variations of male clusteringwill be presentedbecause patterns emergedclearly. Behavioralobservations were conducted at 29 leks Piprapipra. Each of the three banded adult males sam- and 7 bathing sites during 318 days (2813 h) between January 1985 pled for lek attendancewas on its territory75-88% of and July 1987, from blinds erected 5-10 m from these sites. In the day. Home-rangesof the two monitoredadult males order to measure attendance at the lek, three adult males of each contained only one lek and one bathing site (Fig. 1), lekking species were sampled continuously for 2 days in the middle and covered2.1 ha and 2.5 ha All of the breeding season. In C. gutturalis, supplementarystudies of respectively(Table 2). mating skew were conducted at one lek during 66 days (600 h) males most often displayedafter dawn and early in the between October and December 1990. afternoon:103 (70%) out of 147 displayswere observed before 0930 hours or between 1300 and 1600 hours. In P. pipra no was observed. Of 107 visits by Results females 53 (49.5%) were directedtowards one male in each group (mean number of males/group 3.3, range Activity and home-ranges during the breeding season 2-4, n= 6 groups).Between the burstsof display,move- ments of males from arenas were very short (2-4 min) For the lek-breeding species, I present one figure show- and they fed less than 60 m from their display sites. ing the typical activities and use of space for females Males visited a stream bathing site in the afternoon, and adult and immature males of the most abundant mostly between 1600 and 1730 hours (85% of manakin visits observed at seven bathing sites, n=265). Intra- and interspecificinteractions were frequent at bathing sites because males and females of different species bathed sequentiallyat the same place. When followed to bathing sites, the two monitored adult males ap- proachedseven banded femalespreviously seen at their display sites. Philopatryto a forest area was very strong - in adult males: 8 out of 10 were netted or visually re- (/n/ml(n.1 m.*y moioefoI6. 0,--*0" h. *= \ ? cordedin the same area duringsuccessive years. } . , ..... / Unlike adult males, adult females were very mobile and visiteddifferent leks and bathingsites (Fig. 1). They used areas 4 times larger than adult male home-ranges I j :. . .. ?/ (Table2). Femaleuse of space was very heterogeneous: * the two individualsmonitored pl , do * l. \ : / by radio-trackingspent Il\\/; / I * '...... / : / 75% of their time in 25-29% of their ranges. Female activities were concentratedat fruiting places around j ' . ! ..' leks and bathing sites (Fig. 1). Of 48 feeding bouts of females monitorednear leks, 18 (37%) ended in a visit .. * to mates. of two females V.".. . ' / potential Home-ranges trapped near the same lek showed 74% spatial overlap. At the beginning of the breeding season, monitored females I / . / \~i\ ~* * . .. / // visitedthree leks, frequentlyin groupsof 2-5 individuals. ' o *' / Laterin the season, they focused on one or two particu- lar leks, probably for mating. One monitored female, Fig.1. Home. rangesand space use in Pi resightedwith two dependentyoung, used a home-range 3 times smallerthan that used when she was monitored ---home ' range* * / beforenesting. Two immature males in transitionalplumage were banded and equippedwith transmittersat the same lek as one of the monitoredadult males (Fig. 1). Both juve- niles visited the same four and their roostingplace. Immature male (no. 11, monitored for 7 days): leks, home-ranges were 8-10 times largerthan those of adult males (Table 2). Immaturemales were very mobile and frequently Fig. 1. Home ranges and space use in Pipra pipra. Adult male visited four leks on the same day. All immaturemales (no. 1, monitored for 7 days): 0 display site,o -*- home range, in their first year of life had female-likeplumage and were toleratedon leks by adults(" visitor" social status). place, display sites of other adult males (no. .. Adult Duringtheir brief visits to leks, they displayedafter adult female (no. 10, monitored for 6 days): '*****.* home range,. I****range, males performances.Males in their second year exhib- ited intermediateplumage ("intruder" social status); they used peripheraldisplay courts on adults' leks, but

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 231 also disruptedadult males duringfemales' visits. Recap- 0930 hours or between 1300 and 1600 hours. In the tures and visual sightings of immature males showed morning, banded and monitored adult males displayed that "visitors" had the largest ranges, which progres- at the top of the slopes and daily moved furtherdown- sively reduced in size when they became "intruders" ward in the valleysat midday(Thery 1990b). Threecop- and settled on a particularlek. ulations were obtained by one male, and 25 (32%) out of 77 females visits were directedtowards one male in Each of the three adult males was Pipra erythrocephala. each 5-8, n=5 on its territory80-85% of the As in P. group (6 males/group, range groups). samplingdays. As in P. pipra and P. erythrocephala, adult males ate pipra, home-rangesof two adult males monitored by fruits near the leks and used the contained one lek and one very rapidly always radio-tracking only bathing same bathing site in the end of the afternoon. Around site. Although small in size, their ranges were 30-60% the the two monitored males interacted than those of P. adult males For bathing place, larger pipra (Table2). with three banded females previouslyseen at their dis- all males, the daily rhythmof courtshipdisplay was less sites. to sites was well-markedthan in P. 234 out of 360 play Philopatry display very strong pipra: (65%) in adult males: 14 out of 15 adult banded males (93%) displays occurred before 0930 hours or between 1300 were seen two successive at the same lek. and 1600 hours. Seven three males were years copulationsby Two females,banded and equippedwith transmitters observed at three different leks, and 42 (35%) of 121 near the leks of monitored visited the same two females visits were directed towards one male in each males, leks and four bathingsites. Theirhome-ranges were 3.5- 6-12, n=5 Be- group (9.6 males/group,range groups). 4.6 times larger than adult male ranges; they used 22- tween these display bouts, males fed in short sessions 28% of these areasin 75% of the concen- less than 70 m from the lek. Like P. day (Table2), (2-5 min) pipra trating around leks and bathing sites. There was 70% males, their furthestdaily movementwas to one bathing in home of these females. Of 52 where interactedwith five adult fe- overlap ranges feeding place they socially bouts of females monitored near leks 20 (38%) were males. Aggregation was evidently due to the limited followed a visit to mates. One of the moni- number of suitable sites the by potential bathing plus synchronized tored females,resighted with one dependentyoung, cov- activity rhythms of different sexes and species. Adult ered an area one-thirdthat of other females. male site fidelity was very high: 16 out of 19 banded One immature banded and monitored in the males were seen in two successive at the male, (84%) years same area as adults, was female-likein his visitations same lek. to the same two leks and four sites. His home- of two monitored adult females were bathing Home-ranges range was 7.5-8.6 times largerthan that of adult males 3-5 times than those of adult males (Table 2), larger (Table2). and contained two leks and two bathing sites. As in in less than 30% P. pipra, femalesspent 75% of the day Manacusmanacus. Each of three banded adult males was concen- of their ranges (Table2), and their activity was on its 78-86% of the bouts territory day. Home-ranges trated near leks and bathing sites. Of 39 feeding of two monitored 2.1 ha and 2.5 ha 19 ended in a males, covering (Ta- of females monitored near leks, (48%) ble were the smallest of all the studied For sites. Two 2), species. visit to potentialmates at neighboringdisplay all the of was well females near the same lek exhibited65% males, daily rhythm display very trapped spatial marked, with 103 (79%) out of 131 displays given to overlapin home-ranges. females before 0930 hours or between 1300 and in transitional was One immature male plumage 1600 hours. Five were realized two males at the same copulations by banded and equipped with a transmitter from different and 63 out of 132 female adult males. This leks, (48%) lek as one of the monitored juvenile visits were directedtowards one male in each and group (6.2 regularlyvisited two leks and three bathing sites, n=8 Two dominant than the adult males/group,range 4-13, groups). his home-range was 6.7 times larger removedfor 2 from their were "visitor" males, days leks, replaced male's range (Table2). Female-plumaged the first subdominant males. Remov- males brief visits to whereas"in- day by neighboring paid displayingadults, dominantmales increasedcourtship disruption (n = truders"in transitional more time at leks, ing plumagespent 12 disruptionsfor 131 courtshipsbefore male removal: restrictingtheir activity for extendedperiods to limited removal: "Vis- 9.2%; n=7/23 during 30.4%). areaswhere they attemptedto establishterritories. contained one lek and one formed leks Home-ranges only bathing itors" and "intruders"jointly permanent site used at the end of afternoon.Between The individual displaybouts, near the main fruiting places. composi- males fed in 1-5 less than 50 m from the lek. Six betweensuccessive min, tion of these leks was different years, femalesbanded at leks werefollowed monitoredadult but the location was the same. by males on their way to bathingsites. Of 18 bandedmales Pipra serena. Each of three banded males sampled for 17 (94%) were seen in two successiveyears at the same lek attendancewas on its territory78-86% of the day. lek. Home-rangesof two monitored adult males contained Home-rangesof two monitoredfemales were 5.2-6.7 one lek and two bathing sites, and covered 2.8 ha and times larger than those of adult males (Table2), and 3.2 ha respectively(Table 2). For all males, most court- contained two leks and two bathing sites. Female use ship displays were observed shortly after dawn or at of space was very heterogeneousand covered 27-29% the beginningof the afternoon:among 87 displaysgiven of the home-rangesin 75% of the time budgets(Table 2). in presence of females, 65 (75%) occured before Among 47 feedingbouts of femalesmonitored near leks,

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17 (36%) ended in visits to potential mates. One female was monitored 4 days before the start of the breeding season. Althoughcaptured inside the primaryforest, this female intensively searched for fruits in light gaps on the granite slopes of the "inselberg des Nouragues", and spent 45% of her time near the forest edge. Follow- ing this female I found 12 hithertounknown abandoned display courts clusteredinto a classical lek 50 m away ee e ". /''A \ from her main feedingplace. At this time no adult male ...' could be seen around the lek or at feeding places. The first heavy rains came after 4 day of radio-tracking,and eight adult males then settled at this lek. The monitored female was still feeding at the forest edge and the only shift in her activitywas to visit adult males at the nearby : displaycourts. Each of two monitoredimmature males visited two / leks and two bathing sites, coveringareas 8.0-9.7 times larger than adult males' home-ranges(Table 2). "Visi- tors" frequently displayed with "residents" on their courts, whereas "intruders" were outsted from "resi- dents"' territoriesand settledon smallperipheral display bat600 aalw ol courts. Corapipogutturalis. Although no individualof this spe- cies was equippedwith a transmitter,8 captures, 19 re- capturesand 179 resightingsof eight adult males or fe- males allowed an estimation of size in C. oioe or6dya oatiiycnesdr home-range mae in saon(Tbe).Ulielekigmaakn, w gutturalis (Table 2). Each of threeadult males sampledfor lek attendance i n 0h0tdeeid.Thymvdaldylne tweent~~eding sites andused large home-ranges (Fig.2,0 was on its territory78-86% of the day. Among the five '!~ abl'1.-65h) ~ vnweog eehadi lekkingspecies, the daily rhythmof displaywas the least th oet ete fte oioe ae a er well-marked:202 (60%) out of 337 displaysoccured be- fore 0930 hours or between1300 and 1600 hours. Eleven copulationswere observedat two leks: two by different males in a lek of eight males and nine by one male in a lek of six males. Of 234 females visits 107 (46%) iy andHom eranges and spaces from the lekking species. weredirected towards one male in each group(6.4 males/ * 0 A n =5 Two dominant (20-04-1986),,~nig feeding places, bathing site, roosting place, group, range 5-8, groups). males, Bandedttindividualg places over twoThs successive years. Femaleae wr (no.moni-recaptured lot 2, etil aut n removed for 2 days from their leks, were replaced by exibtesiia civt n seo p orwoucs subdominantneighboring males. Male removal did not increase disruption, which was very low at that time (n=6/168 before removal:3.6%; n=0/42 dur- the As in other adult ing experiment). lekking species, ing season (Table1). Unlike leaking manakins, two males' home-rangeswere very small (Table 2: 2.0-2.5 ha), fruits were found less than 70 m from the lek, and males visited only one lek and one bathing site. All banded males were seen in two successiveyears at the same lek. Banded adult females exhibited home-ranges 4-5 more than 500 m homfr rainitialthe mist-nettingplace, netting times larger than those of adult males, and were ob- served at only two leks and two bathing sites. Banded placesover successivetwo inyears the same forest area. immaturemales were observedat two differentleks, but the numberof observationsaway from display logs was insufficientto estimatehome-range size. "Visitors"were toleratedon leks and assumedthe position of the female adult even if no was home-range size were minimal (Table 2). S. turdinusalso during displays, copulationattempt bathedbetween 1600 and 1700 hours, but alwayssolitar- observed(Thery 1990 b). "Intruders"often visited"resi- dent" males and in ily and in different places from the lekking species. displayedcompetitively groups;they Banded individualswere never resightedor recaptured displayederratically at peripheralcourts. more than 500 m from the initial mist-netting place Schiffornis turdinus. I color-banded 22 S. turdinus, and three were equippedwith transmittersduring the breed- successiveyears in the same forest area.

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Table 3. Male dispersion and home-rangesize for five sympatic lekking manakins

Species na DMALb DLEK +SE HRAMd HRIM HRFE'

M. manacus 8 3 525+30 2.3 20.2 15.5 P. pipra 6 30 150 + 25 2.2 19.8 9.0 P. erythrocephala 5 5 530+ 30 3.2 22.7 13.6 P. serena 5 6 420+ 30 3.0 24.1 12.1 C. gutturalis 5 25 230+ 20 2.2 ? 9.5 a Number of studied leks d Mean home-rangesize of adult males in ha b Mean minimumdistance between neighboringmales in m e Mean home-rangesize of immaturemales in ha c f Mean distances between neighboringleks in m Mean home-rangesize of adult females in ha

Table4. Correlationcoefficients between home-rangesize and male Table5. Correlationcoefficients between home-range size and male clusteringvariables computed for four sympatriclekking manakins clusteringvariables computed for five sympatriclekking manakins

HRIMb HRFEc DLEKd DMALe HRFEb DLEKc DMALd

HRAMa 0.9019 0.5400 0.5494 -0.5666 HRAMa 0.6437 0.6398 -0.6600 * * * * * * * HRIM 1.0000 0.4007 0.4245 -0.5376 HRFE 1.0000 0.9964 -0.9739 * * * *** ** HRFE 1.0000 0.9993 -0.9655 DLEK 1.0000 -0.9792 *** ** **

DLEK 1.0000 -0.9744 a ** Mean home-rangesize of adult males b Mean home-rangesize of adult females c a Mean size of adult males Mean distance between neighbouring leks home-range d Mean minimumdistance between neighbouringmales b Mean home-rangesize of immaturemales * ** P<0.01; *** P<0.001 c Mean home-rangesize of adult females non-significant; d Mean distance between neighbouringleks Mean minimum distance between neighbouringmales * ** *** P<0.001 non-significant; P<0.05; home-ranges; (4) mean size of adult male home-ranges; and (5) mean size of immature male home-ranges. Three significant correlations were found between these five Activity and home-ranges during the nomadic period of variables (Table 4): a positive correlation between female lekking species home-range size and distance between neighboring leks, a negative correlation between female home-range size I monitored the activity of three individuals when leks and distance between neighboring males, and a negative were abandoned: two males (P. pipra and P. erythroce- correlation between male-male nearest neighbor dis- phala) for 5 successive days, and one female (P. serena) tances and lek-lek distances. Exploded leks of P. pipra for only 3 days. For all species, estimated ranges were were associated with small female home-ranges and extremely large and unstable: the monitored birds fed short distances between neighboring leks, whereas classi- for 2 or 3 days in the same canopy tree, then flew directly cal leks (M. manacus, P. erythrocephala)and double-spot 1-2 km away and found another feeding place. During leks (P. serena) exhibited large female home-ranges and the very short periods of radio-tracking, male ranges large distances between neighboring leks. covered 30 and 32 ha respectively, and the female's range For C. gutturalis, I also computed values of home- was 32 ha. If the monitoring period had been as long range size (estimated from recaptures and visual sight- as during the breeding season, each home-range would ings away from leks) and male clustering with the other probably have reached more than 100 ha. four lekking species. Correlations were very similar to that described in the first analysis (Table 5): large female home-ranges were associated with higher degrees of male Correlations of male clustering with home-range sizes clustering and distances between leks.

Because home-range size was not determined for every individual manakin whose dispersion was studied on the Discussion leks, leading to unequal numbers of home range and male clustering data, I computed for the four lekking Variationin home-range size species monitored by radio-tracking (Table 3): (1) mini- mum distances between neighboring males; (2) mean dis- Beehler and Foster (1988) expressed the difficulty of de- tances between neighboring leks; (3) mean size of female fining a female home-range because "for at least two

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 234 species of birds that display on courts, data indicate and predictablefruit shortage limits breeding to well- that females travel some distancesoutside their normal defined seasons (Worthington 1982). The shift from ranges in order to visit males on their courts." On the lekkingto nomadismis also relatedto seasonalchanges other hand, Higlund and Robertson(1990b), studying in fruit dispersion and in fruit availability near leks spacing of leks in relation to female ranges in the great (Snow 1962a, 1962b; Bradbury 1981; Worthington snipe Gallinagomedia, used only 5-12 fixes to calculate 1982; Thery 1990a). Beehler and Pruett-Jones(1983), home-rangesizes and did not include lek visits by fe- comparingdiets and dispersionin nine species of birds malesin home-rangeestimates. In this study,based upon of paradise,found a significantrelationship between the radio-trackingduring the breedingseason, none of the extent of frugivoryand the breakdownof widely-spaced monitoredfemales expanded her normal range in order territorialbehavior. Lek matingis correlatedwith a for- to visit males on their courts. On the contrary, every aging ecology that promotes high mobility by females, monitored female regularly visited different leks, and and yet permitsforaging from a fixed point by the males leks were always situated near peaks of female feeding restrictedto the lek for more than 6 months a year activity.Female home-ranges were fixed in space at least (Beehler1987). Although population density may consti- during several stages of the breeding cycle (i.e. when tute a proximatefactor of social organization(Brosset visiting males on leks, or when feeding dependent 1982), the spatio-temporaldistribution of trophic re- young). Data collected during annual cycles for four sources appears to be the major force motivating the lekking manakin species showed that home-rangesize shift to communal display (Snow 1962a, 1971, 1976; changedwith the stage of the breedingcycle (very large Lill 1974c; Emlen and Oring 1977; Willis et al. 1978; home ranges outside the breeding season), the sex of Snow and Snow 1979; Bradbury1981; Brosset 1981a, the individual(home-ranges 3-7 times largerin females) b, c, 1982; Oring1982; Worthington 1982; Beehler1983, and the social status of males (larger home-rangesin 1987). juvenile than in adult residentmales). Significantcorrelations were found between female home-range sizes and male dispersions (distances be- tween neighboringmales and distances between neigh- Evaluating hypotheses of lek evolution boring leks). As predicted by Bradburyet al. (1986), larger female home-rangesize was associated with in- Hotspot model. In this paper, theoretical implications creased male clustering. Adult and immature male are restrictedgiven the circumstantialnature of the ra- home-rangesizes were not significantlycorrelated with dio-trackingdata which are obviously limited in terms male dispersionor female ranging.Unlike lekkingman- of number of individuals per species and duration of akins, male and female S. turdinusexhibited similar use monitoring.However, individualranging patterns were of spaceand home-rangesize. In P. serena,males seemed very repetitive,more than 350 fixes were used for each to initiatethe processof clumpingand to use two court- of 25 home-rangesize estimates,and intersexualdiffer- ship localitiesin orderto meet as many femalesas possi- ences in rangesize were more pronouncedthan intrasex- ble, supporting the hotspot model of lek evolution ual or interspecificdifferences. Male spacingand female (Thery 1990b). In M. manacus, the onset of nesting de- ranging were consistent with the female choice model pends upon and follows quickly after the first heavy (Bradbury 1981) and the hotspot model suggested by rainfall (Snow 1962a, present study). For example, on Bradburyand Gibson (1983)and simulatedby Bradbury the slopes of the "inselbergdes Nouragues",displaying et al. (1986). First, as predictedby these authors,ranges males depended on the presence of an intermittent of differentfemales visiting the same lek had high spatial streambathing site near the lek; femalesbuilt their nests overlaps,and males settled near peaks of female traffic down in the valley, using fruiting panicles of Nespera (i.e. clumpedfruiting resources). Among manakins,large aquaticaavailable during the rainyseason (Snow 1962a). overlappingfemale ranges includingdifferent leks have With the onset of rainfall,males joined the lek and set- also been found in Manacusmanacus (Lill 1974b), Pipra tled accordingto female use of space. In contradiction erythrocephala (Lill 1976), and Chiroxiphia linearis with Beehlerand Foster's(1988) hypothesis,limited ra- (McDonald 1989a). Femaleuse of space was very heter- dio-trackingof a female M. manacussuggested that fe- ogeneous and 75% of female traffic was restrictedto males show preferencefor these sites in the absence of 25-30% of their ranges around leks, fruiting sites and males. In this case female rangingmay have influenced bathing places. Of 186 feeding bouts of females moni- location of lek sites, and not the reverse. tored near leks 74 (40%) ended in a visit to potential This study presents the first set of data showing a mates. continuumof variationin male aggregationwith chang- During the breeding season, males aggregatednear ing femalerange-size among sympatricspecies. On finer the most important fruiting resourcesexploited by fe- scales, the role of dispersion and positional arrange- males; adult males used small home-rangesand could ments of display sites in determiningmating skew has feed in short sessions from display sites located in the been the focus of a continuingcontroversy. Correlations vicinity of these rich fruit patches. The breedingseason between position of male territorieson the arena and of manakins,and thereforelek behavior,seems adapted mating successhave been found in the ruff Philomachus to seasonal fluctuations in food supply (Snow 1962a, pugnax (Hogan-Warburg1966), Uganda kob Adenota 1962b; Snow and Snow 1964). Fruit scarcity reduces kob (Buechnerand Schloeth 1965; Buechneret al. 1966; male and female reproductivesuccess (Foster 1977a), Buechner and Roth 1974; Floody and Arnold 1975),

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 235 black grouse Lyrurustetrix (Kruijt and Hogan 1967; speciesexhibiting cooperative displays (Snow and Good- Kruijt et al. 1972), sage grouse Centrocercus urophasian- win 1974; Schodde and Mason 1974; Foster 1977b; us (Wiley 1973) and fallow deer Dama dama (Clutton- LeCroyet al. 1980). The hypothesisof strongerbias due Brock et al. 1988). Opposite conclusions were reached to male dominancein small clustershas been expressed in Manacus manacus (Lill 1974a, b), Pipra erythrocepha- by Bradburyet al. (1986). In C. linearismale dominance la (Lill 1976), Centrocercus urophasianus(Hartzler 1972; relationshipsdetermined access to females in each lek Gibson and Bradbury 1985), buff-breastedsandpiper (Foster 1977b; McDonald 1989a, b), but femalescould Tryngites subruficollis (Pruett-Jones 1988), Dama dama choose between differentleks (McDonald 1989a). Sec- (Clutton-Brocket al. 1989),Lawes' parotia Parotia lawe- ond, male disruptionof courtship may influence mate sii (Pruett-Jonesand Pruett-Jones1990) and Gallinago choice in explodedleks (swallow-tailedmanakin Chirox- media(H6glund and Robertson1990a). Position effects, iphiacaudata: Foster 1981), or in classicalleks Philoma- consideredas a consequencerather than a cause of mat- chuspugnax: Hogan-Warburg 1966; L. tetrix: Kruijt and ing skew (H6glund and Robertson 1990a), appear to Hogan 1967; Centrocercus urophasianus: Hartzler 1972; act in conjunctionwith other cues of female choice or Wiley 1973;greater prairie chicken Tympanuchus cupido: to reflectother processeslike male relocation(Bradbury Robel and Ballard 1974; M. manacus:Lill 1974a; P. and Gibson 1983). In C. urophasianusGibson and Brad- erythrocephala: Lill 1976; T. subruficollis: Myers 1979; bury (1985) reanalysed the issue of mate choice cues cock-of-the-rockRupicola rupicola: Trail 1985; Trailand and found, like Hartzler (1972), phenotypic correlates Adams 1989).As predictedby Foster(1983) and Beehler of male mating success. In a recentstudy of cooperative and Foster (1988), I found lower levels of disruption displays in C. linearis,McDonald (1989a, b) described in explodedleks (3.6% in C. gutturalis)than in classical behavioral and age-related correlates of male mating leks (9.2% in M. manacus).However, disruptionrates success. Ryan (1983), H6glund and Lundberg (1987), can vary betweenleks, and I recentlyobserved 32 disrup- Clutton-Brocket al. (1988), Andersson(1989), Clutton- tions for 131 female visits in Corapipo(24.4%). In con- Brock et al. (1989), Trail and Adams (1989), H6glund tradictionto Bradburyand Gibson (1983)and Bradbury and Robertson (1990a), H6glund et al. (1990), Pruett- et al. (1985), male interferenceduring female visits can Jones and Pruett-Jones(1990) also found male charac- bias mate choice; there is convincing evidence that fe- ters correlatedwith reproductivesuccess. In the present males may change their preferencesfor a given male study whether females do copy the choice of other fe- as a result of experiencingdisrupted copulations (Trail males, use a prior mating site, or employed a "pool- 1985), and that apparentfemale choice could be overri- comparison tactic" (Wittenberger 1983; Trail and den by past and presentmale-male interactions (Foster Adams 1989) instead of having a true preferencefor 1981, 1983; Beehlerand Foster 1988; McDonald 1989a, a particularterritory, is still unclear. However, female b). In their evaluation of the hotspot model, Beehler choice among maleswithin leks is not criticalto discrimi- and Foster(1988) ignored the "despotic"settlement rule nating between models of lek evolution. Even when fe- examinedby Bradburyet al. (1986).Despotic settlement, male choice plays an importantrole in , generatedthrough bias of female choice or male-male it may be unrelated to the process of lek initiation interference,led to reducedclustering of males. Lek sys- (Pruett-Jonesand Pruett-Jones1990). My conclusions tems are expected to stabilizeat intermediatevalues of and those reached by Bradburyet al. (1989a, b) and bias and male clustering(Bradbury et al. 1986). In their Pruett-Jones(1985, 1988) are consistent with the "hot- recentsimulation study, Gibson et al. (1990)have shown spot model" of lek evolution because males were clus- that female preferenceeither for largerleks or for male tered over peaks of female traffic. Male spacing and qualitymay causemales to aggregate.In fact, male-male female ranging are also consistent with the "female interactions, female preference and mate-comparison choice" model of lek evolution (Bradburyet al. 1986; tactics all appear to act synergisticallyto shape male Gibson et al. 1990), but to test this model one needs dispersionon leks. For example, H6glund and Robert- to show that females of every species studied prefered son (1990b) found a hotspot settlementeven if the at- largeclusters of males. tractivenessof particularmales and the relocation of unsuccessful ones may explain why they were more clumpedthan expected.Hotspot settlementmodels may Complementaryhypotheses also be adequate to explain male dispersionon coarse scales, but habitat preferencesor male tendencies for Beehlerand Foster (1988) stressedthat two of the basic clusteringmay be invoked to explain specific lek siting assumptionsof Bradburyet al. (1985) may be false: (1) (Bradburyet al. 1989b). that "the major determinantof the distributionof mat- ings among males is female choice" and (2) that "male interferencedid not bias mate choice". In fact, females Lek developmentand maintenance of some lekking species do not exert any effectivemate choice within leks, although they may do so between Althoughbehavioral and location effectson mate choice leks. First, behavior or location effects describedabove may also be explained by the hotspot model (Beehler could be correlatedwith male age, dominanceor mating and Foster 1988), correlationsof male clusteringwith success (Beehler and Foster 1988). Male interactions female home range size are predictionsof the hotspot prior to female visits determinemating success only in model (Bradburyet al. 1986), confirmed here for the

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first time among sympatric lekking species. However, Bradbury JW, Gibson RM, McCarthy CE, Vehrencamp SL even Beehler and Foster (1988) agree "that inter-arena (1989b) Dispersion of displayingmale sage grouse. II. The role of female Behav distance is probably some function of size of female dispersion. Ecol Sociobiol 24:15-24 Brosset A (1981a) Occupationdu milieu et structured'une home-ranges". To explain lek formation in manakins, popula- tion du bulbul forestier Andropaduslatirostris (Pycnotidae). I suggest that environmental circumstances, especially L'oiseau et RFO 51:115-126 the nature and spatio-temporal distribution of food re- Brosset A (1981b) Evolution divergente des comportements chez sources, initially enhanced the likelihood of male eman- deux bulbuls sympatriques(Pycnotidae). Alauda 49:94-111 cipation from parental duties (Lill 1974a, 1976; Snow Brosset A (1981c) La periodicitede la reproductionchez un bulbul de fort - 1976; Emlen and Oring 1977; Wittenberger 1978; Snow equatoriale africaine Andropaduslatirostris ses inci- dences Rev Ecol 35:109-129 and Snow 1979; 1981; 1982; Beehler demographiques. (TerreVie) Bradbury Oring Brosset A (1982) The social life of the african forest to a male-advertisement yellow- 1983, 1987), leading dispersed whiskered greenbul Andropadus latirostris. Z Tierpsychol polygyny. Further, an ecologically determined increase 60:239-255 in female home-range size (Bradbury 1981) and the mag- Buechner HK, Roth HD (1974) The lek system in Uganda kob netic effect of mating skew in only a percentage of avail- antelope. Am Zool 14:145-162 able males (Arak 1983, 1984; Beehler and Foster 1988; Buechner HK, Schloeth R (1965) Ceremonial mating behaviour and Robertson have in Uganda kob (Adenotakob thomasiNeumann). Z Tierpsychol Hoglund 1990a, 1990b), would 22:209-255 an initial increase in male on court- promoted clustering Buechner HK, Morrison JA, Leuthold W (1966) Reproduction based systems or exploded leks, either with competitive in Uganda kob with special referenceto behavior. Symp Zool or cooperative displays. Later, the development of male- Soc Lond 15:69-88 male interactions, coupled with an increase in female Clutton-Brock TH, Green D, Hiraiwa-HasegawaM, Albon SD home range-size, could have led to the existence of classi- (1988) Passing the buck: resource defense, lek breeding and cal leks. mate choice in fallow deer. Behav Ecol Sociobiol 23:281-296 Clutton-Brock TH, Hiraiwa-Hasegawa M, Robertson A (1989) Mate choice in fallow deer leks. Nature 340:463-465 I am to J. C. Acknowledgements. very grateful Bradbury, Erard, Emlen ST, Oring LW (1977) Ecology, sexual selection, and the P. and three reviewersfor Charles-Dominique anonymous helpful evolution of mating systems. 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