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Lek Behavior of the Lesser Bird of Paradise 992 ShortCommunications [Auk,Vol. 100 at the mouth of the Rio Magdalenanear Barranquilla, siblethat armentimay occurnearer to Panamain suit- discovereda population of armentithere, and in both able spotsin the still little-worked sectionsof north- yearshe collecteda good number of specimens,thus ern C6rdobaand Antioquia provinces.The isolation corroboratingthe correctnessof the type locality and of armentiprobably resulted from the intervening casting doubt on Leticia as a locality of record. Dr. presence of a considerable area of humid forest in Toro-Garciafirst observedthe specieson 16 June1969, Panama and in northwestern Colombia. M. aeneus is in company with Shiny Cowbirds (Molothrusbonar- but one of a number of speciesof semi-arid,or at iensis)but saw no more until early November of that least fairly open, parts of Middle America that has year, when he noted flocksof them daily until early its nearestally on the Caribbeanslope of Colombia. December. On 25 March 1970 he collected a young Now, with increasingnumbers of man-madeclear- male but saw no others until 20 October, when flocks ings, M. aeneushas been extendingits range south- of 100-200 individuals were noted and additional ward in Panama on both slopes. specimens were taken. From then to 11 December With 12 specimensnow availablefor study in the flocks of 20-30 birds were seen almost daily. museumsof New York and Bogota,it becomesclear Dr. Toro-Garciareported seeing armenti chiefly on that armenticannot be looked upon as other than a the ground (in which respectit agreeswith aeneus), southernmostrace of the speciesM. aeneus.M. ar- perchingon small trees(Prosopis julifiora), or flying mentiagrees with aeneusin the emarginationof the 30-50 m from tree to tree as he approachedthem. So primaries in the adults. Inasmuch as no adult females far he has not been able to furnish any information of armentihave been describeduntil now, it may be abouttheir breeding habits.If armenti,like the closely put on record that in colorationthey resemble,but allied aeneus,is a brood parasite,which seemsalmost are not quite as glossyas, female M. aeneusassimilis. certain, it probably dispersesand no longer gathers The adult males of armenti resemble those of aeneus in conspicuousflocks during the main nestingseason but are lessbronzy, more brownish. The brown color of its presumedhost species,when it would be more of the head and body of armenti,while more irides- likely to be overlooked.In 1913,when the late M. A. cent, is not unlike that of the head of male ater (Parkes Carriker, Jr. visited the narrow sandy eastern end of and Blake 1965). The bill of adult armentiis somewhat Salamanca Island, he never saw armenti. Dr. Toro- shorter and slenderer than that of aeneus. Garciaeven venturedto surmisethat while breeding it may be absent from the island, as it is a bird of LITERATURE CITED partly open habitats,favoring lesshumid regionsor areasthat, becauseof clearingby man for agriculture, FRIEDMANN, H. 1933. Notes on Arment's Cowbird, have becomesemi-arid. In the stomachsof the spec- Tangaviusarmenti (Cabanis). Ibis 75: 492-494. imens he collectedhe found small, unidentified grass 1957. The rediscoveryof Tangaviusarmenti seedsand also many rice seeds.(There are somesmall (Cabanis). Auk 74: 497-498. rice plantations to the south of Salamanca Island, PARKES,K. C., & E. R. BLAKE.1965. Taxonomy and across the Carlo Clarin.) nomenclature of the Bronzed Cowbird. Field- The southernmost Panamanian locality where iana-Zoology 44: 207-216. aeneushas been found, Rio Chepo,on the Pacificslope of eastern Panama, is about 400 km (220 miles) from Received3 January1983, accepted 18 April 1983. Cartagena,where armentioccurs. It seemsquite pos- Lek Behavior of the Lesser Bird of Paradise BRUCE BEEHLER • Departmentof Biology,Princeton University, Princeton, New Jersey08544 USA The birds of paradise(Paradisaeidae) are promi- cited as textbookexamples of the promiscuouslife- nent among an array of vertebrate groups that have style,and the exaggerateddisplay plumes of the males becomethe focalpoint for biologistsstudying sexual of some paradisaeidspecies are consideredto be il- selectionand lek behavior (Emlen and Oring 1977, lustrativeof the extremeproducts of runawayselec- Foster 1977, Wittenberger 1979, Wrangham 1980, tion, which occurswhen only a small proportion of LeCroy et al. 1980,Bradbury 1981,LeCroy 1981,Dia- malesreproduce successfully in any breedingseason. mond 1981, Oring 1982). Birds of paradise are often This view is surprising,as there has been no defini- tive field study of these birds in their native habitat that documentseither polygynousmating or skewed Present address:National Museum of Natural History, MRC 114, mating successfor males of any speciesof bird of Smithsonian Institution, Washington, D.C. 20560 USA. paradise(Gilliard 1969, Dinsmore 1970, Cooperand October 1983] ShortCommunications 993 Forshaw 1977, LeCroy 1981). In this paper I present TABLE1. Lek attendanceand mating successin Less- the first conclusivefield observationsof a strongly er Birds of Paradise. skewedmating successamong a lek-displayinggroup of male birds of paradise. In addition, I comment on Timed the importanceof female choicevs. male-male com- attendance petition in the mating habits of this species. Bird' (min) b Copulations The Lesser Bird of Paradise (Paradisaeaminor) in- 1 960 25 habits rainforest of the lowlands and hills of north- 2 720 1 western New Guinea (Gilliard 1969, Cooper and For- 3 390 0 shaw 1977). Morphologically and behaviorally, it is 4 390 0 5 360 0 very similar to other membersof the Paradisaeaapoda 6 195 0 superspecies,all of which are notable for the re- 7 585 0 c markable nuptial plumage of the males,extreme sex- 8 105 0 ual dimorphism,and lek courtshipdisplay (LeCroy ' This linear sequenceis not equivalent to any perceived "mating 1981). ! observed a lek of Lesser Birds of Paradise at hierarchy." The numbers simply refer to the perch number that the the National Bird of ParadiseSanctuary, Baiyer Riv- bird attended. bTotal recorded attendancetimes for these birds during a period of er, Western Highlands Province,Papua New Guinea timed observationsduring Juneand July 1979. (5ø31'S,144ø71'E, 1,200 m elevation) on 18 days from c Female solicited from this male for more than 3 min, but the male 2 June to 4 July 1979, for a total of 49 h. During part never mounted. of this period, I timed attendanceby the birds, and at other times I monitored behavioral interactions. This lek has been used for at least 12 yr by an un- becausethe outer perches were occupied less faith- disturbed population of LesserBirds of Paradise(R. fully. During my study, ! documentedno exchange D. Mackaypers. comm.). Aided by a 15X telescope,I of perches, temporary or otherwise. I assignednum- observedthe birds from the ground, 45 m from the bers to perches;the males that held particular perch- lek. Althoughindividuals at the lek were not marked, es were given the same number as the perch they after a few daysof preliminary observationI wasable frequented. to identify the four males that occupiedthe central In addition to the regular attendant males, an ad- perches of the lek individually by physical differ- ditional numberof males,most in subadultplumage, encesin pectoralplumes, tail wires, and head mark- visited the lek but did not own perches.These birds ings. I monitored attendanceat the lek, visits by fe- spent most of the time out of the lek, calling from males,copulations, and activitiesamong males. My perches dispersedthrough the surrounding forest. observationsof the physical organization of the lek They would visit the lek when courtshipactivities of this speciesare in accord with those of Gilliard reacheda peak. These birds observedthe courtship (1969: 235), although elsewhere (Gilliard MS, as not- activitiesbut never actually displayed,nor did I ob- ed in LeCroy 1981: 10) Gilliard reported instancesof serve them to copulate with females at the lek. significantlocal variation in lek size and male orga- When a female entered the lek to mate, the perch- nization in this species.Here, I confine discussionto holding malesbecame visibly agitated,called loudly, activities in a single lek. erected their yellow pectoral plumes, and often The Baiyerlek was confinedto a singletree (32 m hoppedabout from limb to limb, flapping their wings tall) at the edgeof a clearing.Males held individually and shivering their plumes (see LeCroy 1981). After owned perchesclustered in a portion of the tree this brief flurry of motion and calling, the males as- where severallarge branchesheld up a tangle of epi- sumed their perches and quietly displayed their phytic vines. Individual malesoccupied perches on erected pectoral plumes while the female moved the horizontalor gently slopingparts of the branch- about the lek, often visiting the limbs of severaldif- es. The same percheswere occupiedday after day. ferent males in display. I never saw a male attempt Interperch distancewas as small as 50 cm. ! had un- to mount a female without a seriesof precopulatory obstructed views of the males because of the birds' behaviors that included explicit solicitation by the habit of plucking all green vegetationfrom the im- female (for details see LeCroy 1981).! observedmat- mediatevicinity of their perches.From the following ing to occur only with males at their particular observationsI determined that the male birds of par- perches. adise "owned" the perches that they attended and Males were present at the lek from 0600 to 0900 maintained. In the lek I was able to identify eight and 1430 to 1715. I recorded females visiting from perchesthat were usedby maleson a daily basis.The 0630 to 0730 and 1445 to 1715; most copulations, four central percheswere owned by positively iden- however, occurred from 0645 to 0715 and 1500 to tified males. The outer four perches were attended 1530.During the 18 days of observations,! recorded by males whose individual identity was less certain 99 visits by females and 26 copulations,which in- to me, in part becausein this short study I had to volved only two males. focus on the central (most active) birds, and in part Male 1 performed 25 of 26 copulationsthat I re- 994 ShortCommunications [Auk, Vol.
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