Foraging Behavior of American Redstarts in Breeding and Wintering Habitats: Implications for Relative Food Availability
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Dartmouth College Dartmouth Digital Commons Open Dartmouth: Peer-reviewed articles by Dartmouth faculty Faculty Work 8-1-1995 Foraging Behavior of American Redstarts in Breeding and Wintering Habitats: Implications for Relative Food Availability Irby J. Lovette Dartmouth College Richard T. Holmes Dartmouth College Follow this and additional works at: https://digitalcommons.dartmouth.edu/facoa Part of the Ecology and Evolutionary Biology Commons Dartmouth Digital Commons Citation Lovette, Irby J. and Holmes, Richard T., "Foraging Behavior of American Redstarts in Breeding and Wintering Habitats: Implications for Relative Food Availability" (1995). Open Dartmouth: Peer-reviewed articles by Dartmouth faculty. 3902. https://digitalcommons.dartmouth.edu/facoa/3902 This Article is brought to you for free and open access by the Faculty Work at Dartmouth Digital Commons. It has been accepted for inclusion in Open Dartmouth: Peer-reviewed articles by Dartmouth faculty by an authorized administrator of Dartmouth Digital Commons. For more information, please contact [email protected]. The Condor97:X32-791 0 The CooperOrnithological Society 1995 FORAGING BEHAVIOR OF AMERICAN REDSTARTS IN BREEDING AND WINTERING HABITATS: IMPLICATIONS FOR RELATIVE FOOD AVAILABILITY l IRRY J. LOVETTE~ AND RICHARD T. HOLMES Department of Biological Sciences,Dartmouth College,Hanover, NH 03755 Abstract. We investigatedfood availability for a long-distancemigrant species,the Amer- ican Redstart (Setophagaruticilla), in both its summer breeding habitat in New Hampshire and in its winter habitat in Jamaica. We used four components of foraging behavior (prey attack rate, foraging speed, time spent foraging, and foraging maneuver use) as indicators of the relative availability of prey in the two seasons. Redstartsattacked prey at a significantlygreater rate in summer than in winter, indicating that foragingbirds encounteredprey more frequently in summer. The winter prey-encounter rate was low even though redstarts moved almost twice as fast while foraging in winter as in summer. Male redstartsalso spent more time foraging in winter (85%) than in summer (43-65%), possiblyto balancethe low rate at which they encounteredprey. In winter, redstarts used more foraging maneuvers that were directed towards small flying prey, whereas in summer they usedmaneuvers that resultedin the capture of relatively large and presumably energy-richprey suchas lepidopteranlarvae. That wintering redstartsforaged faster, attacked prey lessoften, and spent more time foragingthan thosein summer indicatesthat the winter is a period of relative food scarcity for this species,whereas the breeding seasonis a period of greater resourceabundance. Key words: Foraging behavior;American Redstart;Setophaga ruticilla; Neotropicalmi- grant; prey availability; Jamaica. INTRODUCTION small, mainly insectivorous species. No direct Food is often considered the resourcemost likely method of estimating prey availability is free from to limit bird populations (Lack 1966, Wiens potential biases (Cooper and Whitmore 1990) 1989) and has been implicated as an important and these biases may be exacerbatedwhen com- limiting resourcefor passerinetaxa as diverse as parisons are made between seasons,habitats, and corvids (Hogstedt 1980, 1981), sparrows (Pul- prey communities. Furthermore, insectivorous liam and Dunning 1987, Arceseand Smith 1988), birds usually consume a variety of prey types, warblers (Rodenhouse and Holmes 1992) and which often differ in motility, crypsis, size and finches (Smith et al. 1978, Schluter 1982). Mi- palatability (Sherry 1984, Holmes and Schultz gratory birds could experience food limitation in 1988, Hutto 1990). Prey availability is affected winter (Alerstam and Hogstedt 1982, Fretwell by all of thesefactors, as well as the overall abun- 1986), in summer (Probst 1986, Holmes et al. dance of prey items. Thus, no prey sampling or 1986, Martin 1987, Sherry and Holmes 1992), census method directly assessesthe prey avail- or during both periods (Cox 1985, Sherry and ability actually experienced by foraging birds Holmes, in pressa). To date, however, there have (Wolda 1990). been few attempts to compare summer and win- An alternative way to obtain information on ter food availability for any long-distance mi- food availability is to examine how the birds gratory species. themselves searchfor and capture food, because The lack of information on food availability foraging behavior often reflects in predictable for migrant songbirds largely results from the ways the types and abundances of available prey difficulties of measuring prey abundance for these (Hutto 1990). At least four components of for- aging behavior can provide information on food availability. First, prey attack rates are an indi- cation of how often prey are encountered by for- ’ ’ Received 12 December 1994. Accepted 24 April 1995. aging individuals (Davies and Houston 198 1, z Present address:Department of Biology, Univer- Price 198 1, Blancher and Robertson 1987, sity of Pennsylvania, Philadelphia, PA 19104. Thiollay 1988). This relationship has been ver- [7821 REDSTART FORAGING BEHAVIOR IN SUMMER AND WINTER 783 ified by experimental work with insectivorous imately 13 km west of Black River, St. Elizabeth warblers, which demonstrated that attack rates Parish, Jamaica (Holmes et al. 1989). This study increase with increasing prey abundance (Hutto area was located within a monospecific black 1990). mangrove (Avicennia germinans) woodland that Second, the rate at which birds move while had a dense, virtually contiguous canopy be- foraging (“foraging speed”) can be used as an tween 8 and 15 m above the ground, but little indicator of the intensity of foraging effort (Rob- vegetation at lower strata (Parrish and Sherry inson and Holmes 1982). By foragingfaster, birds 1994). The site was submergedby up to one me- searcha greateramount of substrate,and thereby ter of water in October-November, but was increase their prey encounter rate. Third, the mostly dry in March. Although the leafy canopy amount of time allocatedto active foragingshould cover within the mangroves decreasedin the late increase both during times of high energy de- winter dry season(Parrish and Sherry 1994) the mand and during periods of low food availabil- mangroves retained substantial green foliage ity. Estimates of the time spent foraging should throughout the year. Previous work has dem- thus provide an indirect measure of prey avail- onstrated that mangroves contain high densities ability. Fourth, birds use different classesof at- ofadult male American Redstartsrelative to oth- tack maneuvers to catch different types of prey: er Jamaican habitat types (Holmes et al. 1989, for example, many warblers use sallies to capture Sliwa 1991) and thus our winter study site was flying prey such as adult Diptera, whereasgleans presumably located within a high quality win- are often used to obtain sedentary prey such as tering habitat (Sherry and Holmes, in pressb). lepidopteran larvae (Bennett 1980, Robinson and Redstarts at both Hubbard Brook and Luana Holmes 1982). The proportional use of different Point were individually color-banded and their types of foraging maneuver thus gives informa- territories mapped. To reduce variance due to tion about the types of prey taken. between-sex differences in redstart foraging be- In this study we examined the foraging behav- havior (Holmes 1986) we report observations ior of the American Redstart, a long-distance only on male redstarts. Similar proportions of migrant, paruline warbler, in its temperate adult and yearling males were observed in all breeding habitat in New Hampshire and in a sampling periods. Although yearling male red- tropical wintering area in Jamaica. We reasoned starts are female-like in appearance, these indi- that if we found seasonal differences in redstart viduals could often be identified as male by foraging behavior, we could identify periods of plumage characteristics(Pyle et al. 1987) or by relatively low food availability by a conjunction song behavior during the breeding season. Data of low attack rates, high foraging speed, greater from the few female-plumaged birds that could time spent foraging, and low prey quality. While not be visually sexed in the field were excluded none of these behavioral parameters alone is a from analyses. sufficiently reliable index of resourcelevels, when Each of our observations were assignedto one considered in concert they provide insight into of five sampling periods, “early winter” (late Oc- differencesin the relative availability of food re- tober-early November) and “late winter” sourcesbetween seasons. (March), and three summer periods, “prebreed- ing, ” “incubation” (including the nest building METHODS and laying periods), and “nestling” periods. Data Summer observations were made from 20 May on summer males whose breeding statuswas not to 6 July 1990 on a 180-ha plot in the Hubbard known were not included in analyses. Brook Experimental Forest, West Thornton, New To obtain foraging and time budget behavior Hampshire (Sherry and Holmes 1985). The veg- sequences,we moved systematicallythrough each etation at the Hubbard Brook site consisted of study plot until we encountered a male redstart. mature second-growth northern hardwoods for- We then followed that individual for as long as est dominated by sugarmaple (Acer saccharum), it remained in sight, and dictated a constant nar- American beech (Fugus grandzjbfia), and yellow rative