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Use of Floral Nectar in Heliconia Stilesii Daniels by Three Species of Hermit Hummingbirds

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Use of Floral Nectar in Heliconia Stilesii Daniels by Three Species of Hermit Hummingbirds The Condor89~779-787 0 The Cooper OrnithologicalSociety 1987 ECOLOGICAL FITTING: USE OF FLORAL NECTAR IN HELICONIA STILESII DANIELS BY THREE SPECIES OF HERMIT HUMMINGBIRDS FRANK B. GILL The Academyof Natural Sciences,Philadelphia, PA 19103 Abstract. Three speciesof hermit hummingbirds-a specialist(Eutoxeres aquikz), a gen- eralist (Phaethornissuperciliosus), and a thief (Threnetesruckerz]-visited the nectar-rich flowers of Heliconia stilesii Daniels at a lowland study site on the Osa Peninsula of Costa Rica. Unlike H. pogonanthaCufodontis, a related Caribbean lowland specieswith a less specialized flower, H. stilesii may not realize its full reproductive potential at this site, becauseit cannot retain the services of alternative pollinators such as Phaethornis.The flowers of H. stilesii appear adapted for pollination by Eutoxeres, but this hummingbird rarely visited them at this site. Lek male Phaethornisvisited the flowers frequently in late May and early June, but then abandonedthis nectar sourcein favor of other flowers offering more accessiblenectar. The strong curvature of the perianth prevents accessby Phaethornis to the main nectar chamber; instead they obtain only small amounts of nectar that leaks anteriorly into the belly of the flower. Key words: Hummingbird; pollination; mutualism;foraging; Heliconia stilesii; nectar. INTRODUCTION Ultimately affectedare the hummingbird’s choice Species that expand their distribution following of flowers and patterns of competition among speciationenter novel ecologicalassociations un- hummingbird species for nectar (Stiles 1975, related to previous evolutionary history and face 1978; Wolf et al. 1976; Feinsinger 1978; Gill the challenges of adjustment to new settings, 1978). Use of specificHeliconia flowers as sources called “ecological fitting” (Janzen 1985a). In the of nectar by particular species of hermit hum- case of mutualistic species, such as plants and mingbirds, however, varies seasonally and geo- their pollinators, new ecological settingsmay in- graphically (Stiles 1975). Comparative studies of clude new arrays of speciesvarying in ability to the foragingecology of hermit hummingbirds and function as partners. Bird-pollinated plants in a the pollination biology of Heliconiaflowers could new setting, for example, will face new selection help us to understand the loosening and tight- pressureson the form of floral display, the ac- ening of mutualistic relationships in different cessibility of nectar, and the phenology of flow- ecological settings. ering, all of which affect ability to compete for In this paper I examine the use of nectar in the services of hummingbirds (Brown and Ko- flowers of Heliconia stilesii Daniels by three dric-Brown 1919, Kodric-Brown and Brown speciesof hermit hummingbirds at one locality 1979, Stiles 1980). in the Pacific lowlands of southern Costa Rica. Hermit hummingbirds (Trochilidae, Phae- Belonging to different genera, the three species thorninae) and Heliconiaflowers (Zingiberales: of hermit hummingbirds differ strikingly in bill Heliconiaceae) provide striking examples of spe- form: Phaethornissuperciliosus (Long-tailed cialized pollination mutualisms (Snow and Snow Hermit) has a long (38 to 39 mm) decurved bill; 1972, 1980; Stiles 1975, 1979; Feinsinger 1983; Threnetesruckeri (Band-tailed Barbthroat) has a Dobkin 1984). Promoting the parallel evolution shorter (28 to 29 mm) nearly straight, sharp- of bills and flowers is the effect of the precise fit tipped bill; Eutoxeresaquilu (White-tipped Sick- between the two on the hummingbird’s rate of lebill) has a sharply bent, stout bill (photos in nectar extraction and the associatedprobability Stiles 1975). The differencesin bill form affected of pollen transfer (Wolf et al. 1972, Stiles 1980). their abilities to extract nectar from H. stilesii flowers, which were abundant next to a large lek ’ ’ Received24 October 1986. Final acceptance 13 of P. superciliosus,and thus an obvious potential May 1987. sourceof energy for their breeding efforts.Nectar [7791 780 FRANK B. GILL TABLE 1. Floral characteristicsof two speciesof Hel- iconia. Character H. rmbricata ’ H. stilesii Flower (perianth) A c Length (cm) 2.5-3.0 5.5 I I Curvature slight strong ,0”l?l Nectar Concentration f 1 SD (O/asucrose equivalents) 22.4 t 2.1 29.1 k 5.4 Energy content f 1 SD (J/pi) 3.6 f 0.3 4.9 * 0.9 Production’ (&hr) 19 18 I Ratesof nectarproduction declined during the day (seeStiles 1975; Gill, m press).The valuespresented here are averageworking estimates for early to midmorning. FIGURE 1. Mid-longitudinal section of Heliconia imbricata(upper) and H. stilesii(lower) flowers. A = into Panama (Daniels and Stiles 1979). It was main nectar chamber; B = belly of flower where nectar accumulatesafter overflowing from the main chamber; common at several forest edge localities at Si- C = site of bill insertion by Phaethornissuperciliosus rena. It was scarceelsewhere in Corcovado Na- and Eutoxeresaquila. tional Park. H. stilesiiflowers throughout the year at Sirena with a general peak of bloom during from the quite similar H. pognonantha Cufo- the rainy seasonstarting in late May. Each flower dontis fuels reproduction by P. superciliosusat lasts only half a day, wilting conspicuously by La Selva in the Caribbean lowlands on the op- early afternoon. The flowers are long and sharply posite side of Costa Rica (Stiles and Wolf 1979). bent, making accessto the main nectar chamber Contrary to expectation, however, the flowers of extremely difficult. A tight passagewayat the an- H. stilesii served only as a temporary resource terior end of the chamber compounds the chal- for P. superciliosusin our study area. lenge of nonlinear accessto the distant nectar chamber. MATERIALS AND METHODS H. imbricata is an abundant, widespread I conducted this study of color-marked hum- speciesin both the Caribbean and Pacific low- mingbirds (see Stiles and Wolf 1973 for proce- lands of Central America, and was one of the dures) in 1980 and 198 1 in Corcovado National dominant plants in the wet second growth hab- Park on the Osa Peninsula of Costa Rica. Allen itat at Sirena. The dark red, compact, vertical (1956)andHartshorne(l983: 132-136)describe inflorescence produces short, slightly curved the forests of this region. Our study site was lo- flowers, which allow direct accessto the nectar cated on the edge of rain forest near the park chamber by a straight bill or capillary tube. The headquartersat Sirena at the base of a small ridge anterior opening to the nectar chamber of H. next to the park headquarters and adjacent to a imbricata flowers allows easy passage. Both lek of P. superciliosus.Both Phaethornis and straight-billed hummingbirds, such as Thalura- Threneteswere common at this locality, but Eu- nia fircata and Amazilia decora, and hermit toxereswas rare, as reflectedin the relative abun- hummingbirds can reach the floral nectar cham- dance of captures during our study, namely 155 ber. Phaethornis, 76 Threnetes,and 7 Eutoxeres. The ease of measuring floral nectar contents Table 1 summarizes the floral and nectar char- facilitates study of energetic rewards available to acteristics of H. stilesii, the principal botanical hummingbirds. Nectar concentrations in H. sti- subjectof this study, and of H. imbricata (Kuntze) lesii flowers were measured in terms of percent Baker, the main alternative sourceof nectar used sucroseequivalents with a temperature-compen- by Phaethornis in our study area in May to July. sated hand refractometer and converted to H. stilesii is found on the Pacific side of Central J/flower based on grams of solute per 100 ml America up to 1,000 m elevation from Par&a, (Bolten et al. 1979). The nectar contained fiuc- Costa Rica south through the Golfo Dulce region tose, sucrose, glucose, and unidentified amino THREE HERMITS 781 0 0 . 07,. I,, I I I. I I 20 24 29 1 5 9 13 17 21 25 29 3 7 11 15 19 23 27 31 4 9 MAY JUNE JULY AUG. FIGURE 2. Standingcrops of nectar energy in Heliconiastilesii (squares) and in H. imbricata(closed circles). Black squares= mean total nectar per H. stilesiiflower at 06:OO;white squares= mean total nectar per H. stilesii flower at 10:OO;black circles = mean total nectar per H. imbricataflower * 1 SD indicated for values after mid- July (deviations in May and June were similar but omitted to simplify figure).Nectar energyavailable as overflow in the bellv of H. stilesiiflowers is shown at the bottom left of the figure:horizontal bars = mean; vertical black bars = i 1 SD. acids (Gill, unpubl. data). Nectar volumes in studies.One lek male, color-marked Pink-White- plucked flowers were measured using 100 ~1 cap- Red (PWR), accounted for 83% of the visits to illary tubes, first from the belly of the flower “I” in 1981. anterior to the staminode and then separately To determine the patterns of nectar removal from the main chamber (Fig. 1). The presence from H. stilesiiflowers by Phaethornis,we bagged of insect larvae or ants in the nectar chamber bracts with new flowers at dawn and then re- was noted. Most flowers also contained floral moved the netting in midmorning to await hum- mites (see Colwell 1973, Dobkin 1984). In this mingbird visits. Baggedflowers remaining on an paper the term “standing crop” refers to nectar inflorescence served as controls. Flowers with present in flowers open to visitors of all kinds. beetle or fly larvae were excluded from the anal- Nectar production was estimated as the accu- ysis of nectar removal; suchflowers typically were mulation in flowers bagged with mesh cloth be- rejected by Phaethornis. Brief or aborted flower fore dawn. visits (less than 5% of total) also were discarded To establish the temporal patterns of flower to insure that the flower contained substantial visitation, we undertook continuous vigils at nectar and that the hummingbird fed without stands (= presumed clones) of H. stilesii. We interference from larvae inside the flower or from monitored all visits by hummingbirds to these Trigona bees. flowers from 07:OO to 12:00, and into the late afternoon on some days.
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