Territorial Behavior and Courtship of the Male Three-Wattled Bellbird

---The Three-wattled Bellbird (Procnias tricarunculata) was studied for 7V2 weeks (April-June 1974) at Monteverde, Costa Rica. In the study area, measuring approximately 1,400 by 3,200 m, 13 adult males held territories from which they advertised themselves by loud calls for 83-93% of the daylight hours. The majority had a repertoire of three different calls. Evidence from a tape recording from Panama and descriptions from elsewhere in Costa Rica show that the Monteverde dialect is distinctive. Two individuals had part or all of their repertoire different from other Monteverde males but matching vocalizations from elsewhere. Males call from exposed perches above the canopy and from a special broken-off branch, the visiting perch, beneath the canopy. Calling males perform two displays involving flight, each preceded by a characteristic call. These displays and a silent wattle-shaking display are mainly performed when another bellbird visits a calling male. The visitors were usually females or immature males but occasionally adult males. At the climax of the visit, the territow-holding male leans over his visitor, perched at the broken-off end of the visiting perch, and utters some extremely loud calls into its ear. This usually makes the visitor leave. Both sexes receive the same treatment. During May and June, females were watched coming to the visiting perches of calling males on 20 different occasions, but none of these visits culminated in mating. The male's wattles are fully extended when he is calling in his territory, but are usually retracted when he leaves his territory to feed. During encounters between closely matched males, first one and then the other may extend the wattles and call. Bellbirds were seen feeding only on fruits, mainly drupes of the family Lauraceae but also other drupes, particularly a species of Rutaceae. An individual male feeding exclusively on the latter fruit took an average of 9.0 g of pericarp per hour. A comparison is made of the calls and displays of the four Procnias species, and it is suggested how these may be related to habitat. ---Old Forge, Wingrave, Aylesbury, Buckinghamshire, England. Accepted 3 May 1976. IN an attempt to discover more about the breeding behavior of the Three-wattled Bellbird (Procnias tricarunculata) for comparison with the Bearded Bellbird (P. averano) and the White Bellbird (P. alba) previously studied in Trinidad and Guyana (B. K. Snow 1970, 1973), I stayed at Monteverde in the Cordillera de Tilaran of Costa Rica from 25 April to 17 June 1974. The Three-wattled Bellbird in adult male plumage is the most distinctive of the four bellbird species that make up the genus Procnias. Unlike the other three species in which the male is mainly white or very pale grey, the adult male is a bright rufous brown except for the head, neck, and upper breast which are white. One of its three pendant black wattles grows from the base of the upper mandible, the other two from either corner of the gape. The female and birds in juvenal plumage have no wattles and, as in the other species, are cryptically colored, olive-green above and yellow streaked with olive-green below. The size difference between the sexes is 623 The Auk 94: 623-645. October 1977 624 BARBARA K. SNOW [Auk, Vol. 94 tad •' ' . ;' "'. ' . ' :'"• . " • '" ; . ,•oo. •/Dk'"' . :' '. • . '::•// ß -. c io• . " _/,. ' #'. , _ % •,•? ,. •. ,"'11 ß ß . 2&&7 ". ftio . ' ". t ": I i . •' • . " '. ' . "\ , \\ ". :, /. ' ,. ,. :: •. " . : :' ". : I I %. • i r) :. ': . ; ! l ': •. ' . • ',. ". :\'•',. ". ,. "' --"". ::. -'"' 71 . :::. ,,. ,,. --:. ,-. ':':::: . Fig. 1. Map of the Monteverde study area showing approximate positions of the advertising territories of the 13 adult males. The continental divide is at the right-hand edge of the map; contour heights in