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Vocal Repertoire of the Long-Tailed Manakin and Its Relation to Male-Male Cooperation’

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Vocal Repertoire of the Long-Tailed Manakin and Its Relation to Male-Male Cooperation’ THE CONDORDEC-61 A JOURNAL OF AVIAN BIOLOGY LIBRARY Volume 95 Number 4 November 1993 The Condor 95:769-78 I 0 The Cooper Ornithological Society 1993 VOCAL REPERTOIRE OF THE LONG-TAILED MANAKIN AND ITS RELATION TO MALE-MALE COOPERATION’ JILL M. TRAINER Department of Biology, Universityof Northern Iowa, Cedar Falls, IA 50614 DAVID B. MCDONALD ArchboldBiological Station, P.O. Box 2057, Lake Placid, FL 33852 Abstract. We examined the vocal repertoire of lek-mating Long-tailed Manakins (Chi- roxiphia linearis, Pipridae) in Monteverde, Costa Rica. Males in this genusare unusualin performing a cooperativecourtship display, including duet songsand coordinateddual-male dance displays. Males give at least 13 distinct vocalizations, several of which occur in clear behavioral contexts. By observing the behavioral context and the sequencein which calls were given, we found that the most frequent calls occurred during three types of activity: song bouts, dance, and noncourtship interactions. The responsesof males to playback of six vocalizations indicated that the calls function as much in mediating cooperative inter- actions as in expressingmale-male agonism. The evolution of the large vocal repertoire in Long-tailed Manakins may be associatedwith their unique social system based on long- term, cooperative relationshipsamong males. Key words: Vocalization;call function; Long-tailed Manakin; Chiroxiphia linearis; so- ciality; cooperation,lek. Resumen. Estudiamosel repertorio vocal de1Saltarin Toledo (Chiroxiphia linearis, Pi- pridae) en Monteverde, Costa Rica. Los machos de este genera se comportan muy parti- cularesen relacibn el cortejo cooperative, incluyendo cancionesa duo y danzascoordinadas de parejasde machos. Los machos emiten al menos 13 vocalizacionesdistintas, muchasde ellas con un context0 claro con respeto al comportamiento. Se encontrb, mediante obser- vacion de1 context0 de comportamiento y las secuenciasde sus vocalizaciones, que las llamadas m&s frecuentesse emiten durante tres tipos de actividades:el canto, el baile, y la interaction no-relacionada al cortejo. Las respuestasde 10s machos a seis vocalizaciones pre-grabadassugieren una funcion de interaction cooperativa tanto coma para expresar agresionentre machos. La evolution de las vocalizacionesdiversas en Chiroxiphia linearis puede ser asociadacon el sistema social unico fundado en relaciones cooperativasa largo plazo entre machos. INTRODUCTION most passerines in that males provide no re- Although much information exists on the use sourcesvaluable to breeding females other than and function of vocalizations in territorial song- gametes (Bradbury 1981). In addition, lek spe- birds, little such information exists about pas- cies lack persistent pairbonds and do not defend serines with other social systems. Here we de- exclusive, resource-basedterritories. Long-tailed scribethe diverse vocal repertoire of a lek-mating Manakins are especially unusual in that groups species, the Long-tailed Manakin (Chiroxiphia of males form complex networks of cooperative lineuris). Lek social systemsdiffer from those of alliances (McDonald 1989a). This cooperation involves a joint courtship display that consists of highly coordinated singing and dancing per- I Received 11 March 1993. Accepted 8 July 1993. formed by two males (Snow 1977, Foster 1981, [7591‘ 170 JILL M. TRAINER AND DAVID B. MCDONALD McDonald 1989b). Our results suggestthat the playing males tolerate the presence of other af- functions of Long-tailed Manakin calls reflect the filiates, and overt aggressionamong males is rare. importance of cooperative as well as aggressive In order to attract females to the perch-zone, interactions among males, in contrast to what is two male partners cooperate in performing the observed in territorial species. coordinated song and dance displays. The court- Chiroxiphia manakins are unusual among their ship display usually is performed by the alpha suboscinerelatives in possessinga large and var- and beta, although occasionally other combina- ied repertoire of vocalizations. Several speciesof tions of affiliates may display (McDonald 1989b). manakins have mechanical pops and snaps in Males begin by broadcastinglong bouts of unison the acoustic repertoires, but generally, manakin duet song, or toledos, from the lower branches vocalizations are few in number and simple in of a tree crown (subcanopy). If a female arrives, structure (Prum 1990b). The Long-tailed Mana- both males and the female descendto the dance kin’s repertoire of 13 distinct calls is rich, even perch, a horizontal branch or vine within 1.5 m when compared to the call repertoires of song- of the ground. The males begin a series of 20- birds (exclusive of primary songvariants). A ma- 100 backward leapfrog hops alternating with as jor factor contributing to the evolution of such much as 60 set of a special butterfyjlight char- a diverse repertoire may be the complex network acterized by deep wing beats. A dance bout may of long-term relationships among males. include as many as 10 setsof hops and butterfly Here we describe the functions of Long-tailed flight. Dances leading to copulation include a Manakin vocalizations, and examine the com- solo butterflyflight by the alpha male. At times, plexity of the vocal repertoire in the context of two or more males, often including predefinitive sexual selection and the social structure. We em- males, gather on a dance perch in the absenceof ployed three approaches in our study. (1) We a female and perform a variation of the hops observed the social and behavioral context of (popcorns),in which males jump straight up and each call. (2) We analyzed the sequencein which do not leapfrog over the partner(s). calls were given. These approachesallowed us to identify three general contexts in which the ma- METHODS jority of calls occurred. (3) We recorded the re- The study area, in Monteverde, Costa Rica sponsesof manakins to experimental playback (1 0”18’N, 84”48’W), is 80 ha of premontane trop- of taped calls in the field. ical moist forest (Holdridge 1966) at an elevation of 1,300 m. McDonald uniquely color-banded NATURAL HISTORY AND DISPLAY 270 manakins between 198 1 and 1987. The sex A Long-tailed Manakin lek consists of a core of individuals with green plumage was deter- partnership of an alpha and a beta male and as mined by subsequent observations of behavior. many as 11 male affiliates (McDonald 1989b). Vocalizations were recorded with a Sony WM These males regularly attend a traditional display D6C cassetterecorder and a Sennheiser ME 80 area called a perch-zone or lek arena, which con- directional microphone. In addition, behavior tains one or two major perches for the dance sequenceswere recordedin the field using a Tandy display. Perch-zonesare separatedby 75-300 m, 102 personal computer programmed as an event and each is occupied by a different alpha male. recorder. Sonagramswere made using a Kay Ele- From a particular perch-zone, one can some- metrics Model 5500 Sonagraph and grey scale times hear the singingof manakins in other zones, printer. A synoptic tape of the Long-tailed Man- but cannot see them. akin vocal repertoire is on file at the Library of Plumage maturation occurs over a four-year Natural Sounds at the Cornell Laboratory of Or- period with distinct, age-specific, predefinitive nithology (Accession #49097). stages(McDonald 1989b, 1993a). Affiliates of a The terms “call” and “vocalization” are used perch-zone may include both definitive and pre- interchangeably to denote an individual utter- definitive males. Affiliations among males de- ance of sound. “Song” refers only to the toledo velop over their lifetimes of at least 13 years call, which is given in unison as a duet by singing (McDonald 1993b) and result in the establish- partners. “Call type” refers to one of the 13 ment of a dominance hierarchy. The alpha male acoustically distinguishable kinds of manakin is highest ranking, and with very rare exceptions calls. The term “dance bout” refers to a display only he mates with female visitors. Pairs of dis- sequenceincluding one or more of the following VOCAL REPERTOIRE OF LONG-TAILED MANAKIN 111 elements: a variable number of sets of leapfrog Chatfield 1971). Therefore we excluded transi- dance hops, dual-male butterflydisplay, and solo tions between identical call types when calculat- display by the alpha male. A “dance bout” is, ing the expected values. Expected values were therefore, essentially synonymous with a single calculated by iteration (Goodman 1968, Trainer continuous visit to the dance perch by one or 1988) becausesimply using row and column to- more females, because males almost invariably tals is not valid for an incomplete matrix. display throughout female visits. Comparing the observed and expected fre- quenciesofeach call type transition revealedthose BEHAVIORAL CONTEXT pairs of call types that tended to occur or not to occur together in sequence.To obtain a consis- Ad libitum observations of vocal behavior and tent criterion for identifying major departures responses of male and female manakins were from expected values, we collapsed the contin- made at perch-zones.In addition, observerscon- gency table around each matrix cell, combining ducted scheduled two-hour observations from frequenciesfrom other call types to form 42 two- blinds 8-12 m from display perches between by-two matrices (Slater 1973). We tested the re- 06:00-l 5:O0. During the scheduled observation sulting chi-square values at the 0.05 level. The periods, an observer recorded the number of to- chi-square analyses of the two-by-two matrices ledo and teeamoo calls occurring in each 5-min were not intended as significance tests; rather, block, and the number of female and male man- they indicate those transitions that contribute akins present in the perch-zone during the first most to the nonrandom sequence of call types min of each 5-min block. The details of this be- overall. havioral sampling protocol were described in McDonald (1989a). PLAYBACK EXPERIMENT The vocal responsesto each of five manakin call SEQUENCE ANALYSIS types were tested by broadcasting recorded calls Thirty real-time samples of vocalizations were through an Aiwa SC-A2 speaker.
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