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Fruits and the Ecology of Resplendent Quetzals

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Fruits and the Ecology of Resplendent Quetzals FRUITS AND THE ECOLOGY OF RESPLENDENT QUETZALS NATHANIEL T. WHEELWRIGHT 1 Departmentof Zoology,University of Washington,Seattle, Washington 98195 USA ABsTR^CT.--ResplendentQuetzals (Pharomachrus rnocinno) are typicallytermed "special- ized" fruit-eatingbirds, althoughthere are few data describingthe breadthof their diet or the characteristicsof the fruits they select.In fact, there is no generalconsensus about the meaning or consequencesof being a fruit specialist. In the lower montane forests at Monteverde,Costa Rica, quetzalsfeed on a minimum of 12-18 speciesof fruits at most timesof the year and on an annualtotal of at least41 species.Although their diet includes the watery,small-seeded berries of many second-growthplants, they depend mostly on the largedrupes of about18 speciesin the laurelfamily (Lauraceae).The phenologiesand habitat distributionsof the Lauraceaeappear to dictatethe timing and directionof seasonalmove- mentsby quetzals.Mutual dependenceand, possibly,general coevolution between quetzals and the lauraceoustrees whose seeds they disperseare suggestedby the birds' morphology, distribution, behavior, and life history. Nestling quetzalsare broughtentire fruits as early as the secondday after hatching. Thereafter,they consumegradually increasing amounts of fruit, but, even immediately beforethey fledge,most of their diet consistsof insects,snails, and lizards.Brooding drops off rapidly by the time chicksare 9 days old. Considerablevariation in broodingduration, parentalsex roles, and nestlingdiet existsbetween nests, however, and apparentlybetween dutches. Adults take far less time to deliver fruits to nestlingsthan to deliver insectsor lizards, which reflectsthe relativeease of "capturing"ripe fruits (asopposed to animal prey) during the breeding season.The male parent delivered significantlymore insectsand food items in generalthan did the female at a first-dutchnest but not at a second-clutchnest. SeveralCentral Americanmontane reserves have been establishedto protectpopulations of quetzals,the nationalsymbol of Guatemalaand an importanttourist attraction throughout the Isthmus.Unfortunately, the reservestend to be too small and to include only a limited representationof criticalhabitats. If other CentralAmerican quetzal populationsare similar to Monteverde's,the birds must migrate to different habitatsas the availability of ripe fruits fluctuatesbetween seasonsor years. Once reservesbecome isolated by deforestation,they will fail to prevent local extinctionof quetzals. Received9 March 1982, accepted4 October 1982. A SMALLgroup of tropical fruit-eating birds dispersersand with which they are believed to has come to be recognizedas "different." The have coevolved(McKey 1975).There have been diet, behavior, and morphologyof thesebirds, few long-termstudies adequate to test the spe- which include oilbirds, fruit-pigeons, cotin- cialist-generalistdichotomy and the predic- gas, and toucans,seem to set them apart from tions ot McKey's influential model, however, other fruit-eating birds, such as flycatchers, let alone its assumptionsabout differences in thrushes, and finches (Snow 1971, 1981). The seed dispersalquality and behavior between distinctionsbetween the two groupsprompted specialistsand generalists(Wheelwright and McKey (1975) to introduce the terms "special- Orians 1982).A key questionis whether, or in ized" and "generalized" frugivores. Special- what ways, specialistsare ecologicallysimilar ists' diets are thought to be comprisedmainly to each other. Does coevolution between fruit- of the nutritious fruits of only a few plant eating birds and plants result in similar pat- species,for which the birds serveas major seed terns within different habitats and taxa? Or do specialists,unified only by their distinctness from birds that are lessdependent upon a fruit Present address: Department of Zoology, Uni- diet, differ from one another as well? versity of Florida, Gainesville, Florida 32611USA. The difficulties of identifying specialists' 286 TheAuk 100:286-301. April 1983 April 1983] Fruitsand Quetzal Ecology 287 unique traits lie partly in the shortage of nat- 33,000 ha) surrounds the Reserve on the Atlantic ural history information.In an attempt to ad- slope.Bordering the Reserveon the Pacificslope, the dressthe need for suchinformation, this paper Monteverde community itself consistsof scattered describesthe ecologyof ResplendentQuetzals cattle pasturesand extensivewood lots. The rainy (Pharomachrusmocinno), neotropical members seasonusually begins in earlyMay and extends,with a brief dry spell in July, until mid-December. The of the Trogonidaethat are typicallyconsidered abundanceand diversity of fruits vary between sea- specialized fruit-eating birds (McKey 1975, sons and years (G. Frankie and W. Haber pers. Snow 1981). Data on adult and nestling diets, comm.; pers. obs.). morphology, life history, and predation in a Two times every month from June 1980 until Au- Costa Rican population are presented in the gust 1981 (exceptmid-November), I censusedquet- contextof currenttheory about avian frugivory zals by recordingsightings and callsover a route of and diet specialization. about 7 km from the cloud forest to the dry forest, Despite their designation as the national spending about 6 h in each of four habitat types, in order to determine diets and seasonal movements. symbol of Guatemala and a reputation as "the General field observations(over 2,000 h), including most spectacularbird in the New World" (Pe- periodic censuseson the Atlantic slope, added in- tersonand Chalif 1973), quetzalshave attracted formation about migrations,foraging behavior, and little scientificstudy. Acrossmuch of their range natural history. Even during the nonbreeding sea- (southernMexico to westernPanama) they are son, when quetzalsrarely vocalize, their noisy flight, scarce,and their montane habitat is relatively large size, and regurgitationof bulky seedsreveal inaccessible(Hanson 1982).Skutch's (1944) pa- their presence.I spent more than 200 h watching at per remains the most thorough descriptive ac- fruiting trees,especially 23 speciesin the Lauraceae count of the natural history of quetzals. Over (laurel, or avocado, family). Over 120 h were spent the course of one year, Skutch documented monitoring food deliveries to nestlings at three nesting chronology, behavior, vocalizations, quetzaI nests. I used a 15•50x spotting scopeand binoculars to observe from a blind at a distance of plumage, and nestling developmentat a Costa about 35 m, from which ! had little difficulty iden- Rican site. Drawing from observationsin Gua- tifying most itemsbrought to the nestby the parents. temala during the breeding season,LaBastille ! made observations at a total of 11 active nests. and others (Bowes and Allen 1969, LaBastille Quetzalshabitually perch on a particularlow branch et al. 1972, LaBastille 1973) supplemented when arriving at or departing from their nest. Be- Skutch's work while emphasizing the conser- neath these perches I placed "seed traps" (cf. Snow vation of quetzals. 1970)of dark, fine-mesh, i m e nylon netting elevated 0.5 m above the ground. Although pocketmice (Het- STUDY SITE AND METHODS eromys demerestianus),the major vertebrate seed predatorsat Monteverde, can climb (Eisenberg1963), Between 1979 and 1982, I studied quetzalsand the no seeds were removed from control seed traps in reproductive biology of the plants whose fruits which a known number of seeds had been placed. quetzals eat in the lower montane forests of Mon- At 5-day intervals I identified and countedseeds re- teverde, Costa Rica (10ø18'N, 84ø48'W; Holdridge gurgitated or defecatedby perching quetzals. Most 1967).The 18-monthstudy period 0une-August 1979, or all of the seedsin the traps were believed to have June 1980-July1981, March 1982) included one entire been dropped by members of the pair nesting near- annual cycle and parts of breeding seasonsduring by, becauseseed traps were positioned directly be- four different years. Monteverde lies on a plateau low perchesthat the territorial quetzals used exclu- straddlingthe continentaldivide in the Cordillera de sivelyduring more than 120h of observation.Control Tilaran. Strong northeast Trade Winds blow across seed traps located a short distance away collected0- the divide at all times of the year, especiallyduring 2 seeds/weekversus 20-60 seeds/weekbelow quetza• the dry season, when winds often carry sheets of perches. The accuracy of seed-trap sampling was mist. Westward from the divide, in the lee of the confirmed by independent sightings of quetzals prevailing winds, habitats change rapidly along a feeding in the field on the samefruit speciesin ap- marked moisture gradient. The forests range from proximately the same proportions. In an attempt to epiphyte-laden"elfin" cloudforest at the crestof the identify individual birds, ! banded one adult male cordillera (elevation 1,550 m) to the relatively dry, quetzal, but its short legs and loosebelly plumage taller forest of the edge of the plateau 4 km to the completelyobscured the leg bands. I could often dis- west (1,350m) (seeLawton and Dryer 1980for a more tinguish birds, however, by differencesin tail co- completedescription of forest types). The study site verts (the long paired covertsthat extend over the included the 2,700-ha Monteverde Cloud Forest Re- tails of males are frequently broken or lost) or bill serve. The undisturbed Arehal National Forest (ca. color (females'bills vary from slate to yellow). 288 NATHANIELT. WHEELWRICHT [Auk, Vol.
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