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Effects of Trophic Morphology and Behavior on Foraging Rate of Three Hawaiian Honeycreepers

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Effects of Trophic Morphology and Behavior on Foraging Rate of Three Hawaiian Honeycreepers Oecologia (Beri) (1982) 55:157-159 CJ?COlOg?? ? Springer-Verlag 1982 Effects of Trophic Morphology and Behavior on Foraging Rates of Three Hawaiian Honeycreepers John H. Carothers Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA Summary. The effects of trophic morphology and behavior rates is possible. Tongue morphology can have an impor- of three Hawaiian honeycreepers (Fringillidae: Drepani- tant effect upon nectar gathering efficiency (e.g. Hains- dinae) upon their foraging rates on the flowers of Vaccinium worth 1973), and these birds show differences in tongue calycinum are examined. The Maui Creeper (Paroreomyza morphology with respect to diet (Raikow 1977; Richards montana), the Amakihi (Hemianathus virens), and the Fiwi and Bock 1973). Thus, included in the analyses is discussion (Vestiario cocc?nea) show shifts from a short straight bill of the role of tongue morphology in affecting the observed to a long decurved bill, from a tongue adapted for insect patterns. feeding to one specialized for nectar, and diets ranging from primarily probing for insects to primarily nectarivorous. This diversity is examined feeding on the simple straight Materials and Methods tubular corollas of the Vaccinium. Significant differences (P< 0.001, Mests) exist among the birds with respect to Study Area foraging rates on these flowers. The Tiwi, with its greatly The work was in the Ko'olau Forest Reserve decurved (64?) bill feeds the quickest (2.09 s/flower) while performed on the north of Haleakala Maui, from 23 the Maui Creeper with its rather straight (18?) bill feeds slope volcano, to 27 December 1981. The site was elevation the slowest (3.87 s/flower). These differences are seen to study high rainforest, with a formed of ohias be the result of differing abilities of tongues to extract nectar (2,000 m) canopy (Metro- sideros collina) 13 m in The as well as differing behavioral tactics of floral exploration averaging height. understory was of six Pelea and nectar extraction. This suggests that predictions of for- composed major plant species, clusiaefolia, tameiameiae, lessertiana, Ilex anomola, aging efficiency based solely on bill morphology are not Styphelia Myrsine Rubus hawaiiensis, and Vaccinium All exhibit a necessarily valid, and that other factors (tongue morpholo- calycinum. form and 3 m in gy, foraging maneuvers, and typical food spectrum) must bushy growth average height. also be considered. The Flower During much of the year the often abundant blooms of canopy ohias provide the main food source for nectar- Carothers in Introduction feeding drepanidids (Baldwin 1953; prep.; Carpenter 1976). At various times some understory species The trophic morphology of an animal has a great influence bloom in sufficient quantity to become major nectar sources on the type of food utilized and the success in using the to at least a few of the bird species. Vaccinium calycinum, items. It has been observed that the proboscis length in a member of the blueberry (Ericaceae) family, is a most bees (Inouye 1980; Real 1981) and bill shape in birds (Gill important nectar source in winter (Carothers pers. obs.). and Wolf 1978) has a pronounced effect on foraging effi- More component species of the Maui drepanidid assem- ciencies upon nectar sources. Flower morphology is often blage were observed feeding on this plant than any other coevolved with the trophic morphology of potential pollina- species, including ohia. tors to insure maximal outcrossing through pollinator fidel- The flowers of Vaccinium were the focus of observations ity (review in Gilbert and Raven 1975). on foraging rates. This species produces from 5 to 15 + The Hawaiian honeycreepers (Fringillidae: Drepani- umbelliform flowers arising from leaf nodes along branches. dinae) show a wide diversification in bill morphology, The pale green corollas average 15 mm in length and 5 mm ranging from parrot-like to warbler-like to radically elon- in width. The corolla is tubular, straight, with a slightly gated and decurved bills (Raikow 1977). In this study a flared opening, and has no internal subdivisions (see Fig. 1). subgroup of drepanidids was examined to determine forag- Nectar is produced basally within easy access of the birds. ing efficiencies on a single species of plant. Because of the During the study period the flowers were in profusion, morphological and behavioral variance in bill morphology averaging 50/m3/plant. Nectar volume from 161 flowers (from short straight to long decurved) and degree of nectar sampled ranged from 0-56 ??). The nectar is a hexose-rich dependence (very weakly to strongly nectarivorous), an ex- type typically used by passerine nectar feeders (Irene Baker, amination of the consequences of this variation in foraging pers. comm.). 0029-8549/82/0055/0157/SOl .00 158 were timed after the from the initiation lOO-i entering plants, of foraging until cessation or until it was interrupted by movement to another part of the bush. The time used by the birds in short movements was negligible and roughly equal for all species. Number of flowers visited during the foraging interval and interval length were recorded. Obser- vations in which presence of a dominant species could po- tentially lower the foraging rate of a subordinate species were not included. In addition to sampling for average nectar content, notes were made on presence/absence of insects within the flowers. Pierces at the bases of corollas, indicating nectar thievary, were also noted. On several occasions flowers were examined for residual nectar content following foraging by the birds. Results Significant differences existed among foraging rates of the three species (P<0.001, 2-tailed /-tests: see Fig. 1). The Vaccinium m Maui Creeper had the slowest foraging rate, at 3.87 s/flow- calycinum JJ er. The Amakihi was intermediate in foraging rate, taking 2.18 s/flower. The Fiwi was the most efficient at nectar feeding, with a visitation rate of 2.09 s/flower. p? Nectar extraction by each species appeared complete, 40 20 30 probably aided by the simple morphology of the corolla. Number of flowers Thus, the foraging lengths reflect the time required to locate Fig. 1. Bill shapes and regression analyses of foraging rates. All a flower, insert the bill, and remove the approximately 10 ?? paired comparisons, both among slopes and among y-intercepts of nectar therein. (?-tests, 2-tailed), show significance at ? < 0.001 level. Correlation Differences were observed in the capabilities of the birds coefficients for each are as follows: Maui (r2) species Creeper to perform these different tasks. While both the Amakihi Amakihi Fimi -0.88 -0.86; -0.86; and the Fiwi located flowers with relative ease, the length of time required to remove the nectar appeared shorter for the for the visitation rate. The Table 1. Physical dimensions of bills (N = sample size). Curvature Fiwi, accounting higher was somewhat slower at is angle between tangents from base and tip of bill Maui Creeper locating flowers, but the large increase in foraging time was due to the inordi- ? Length Width Depth Curvature nate amount of time obtaining nectar from flowers. This species was observed to feed on flowers in several ways. 18? Maui Creeper 15 11.2 3.7 4.2 Simple bill insertion at the distal corolla opening with sub- Amakihi 3 15.4 4.2 4.7 40? sequent nectar extraction was observed to take a much lon- Fiwi 5 27.2 5.8 6.4 64? ger time. Other foraging maneuvers not used by the other species were also observed. Piercing of the corolla base was another method utilized. Indeed, 45% of the flowers exam- ined had at least one Other maneuvers The Drepanidids (N =250) puncture. included grasping the flower with the foot and stepping Three species in two tribes were observed feeding on the on it to hold it in place while probing it. Biting off and flowers: the Fiwi (Vestiaria cocc?nea) (tribe Drepanidini), mastication of the corolla to extract nectar were also ob- and the Amakihi (Hemignathus virens) and the Maui served. Bill insertion itself was not always a smooth, single Creeper (Paroreomyza montana) (tribe Psittirostrinae). motion but often represented a repeated pecking motion. Members of the first tribe are nectarivorous, while those None of the 250 flowers examined contained insects, ruling of the latter are usually more insectivorous. Indeed, prior out insectivory as a cause of these inappropriate maneuvers. to this study all foraging events of the Maui Creeper ob- served by myself and my coworkers were insect gleans and - probes nectar feeding was totally unexpected (but see Discussion Perkins 1903). Bill length and decurvature increase with Both morphology and behavior can be seen to determine nectarivory in the group (see Fig. 1 and Table increasing effectiveness at foraging on Vaccinium calycinum flowers. 1), adaptations for feeding on decurved corollas of endemic A long trophic apparatus does not seem to impose con- lobeliads Measurements were taken from (Berger 1981). straints on the use of short corolla flowers (Brian 1957; at the Bernice P. Bishop Museum. specimens this study). However, because of their short straight bills the Amakihi and the Maui Creeper are likely at an advan- Observations tage foraging on these flowers when compared to the Fiwi Observations were conducted on two bushes with the same with its long, decurved bill which should require greater flower densities, each being used by all three species. Birds skill to manipulate relative to a short bill. In a study of 159 bees Inouye (1980) suggests that a short proboscis allows meda dolei) indicate that this is indeed correct. Further greater efficiency when foraging on short corollas. comparisons of this nectar feeding assemblage can clarify Despite its long, decurved bill the Fiwi proved to be the constraints imposed by morphology in foraging rates.
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