Patch Utilization by Migrating Birds: Resource Oriented?
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ORNISSCANDINAVICA 17: 165-174. Copenhagen1986 Patch utilization by migrating birds: resource oriented? Thomas E. Martin and James R. Karr Martin T. E. and Karr, J. R. 1986. Patch utilization by migrating birds: resource ori- ented? - Ornis Scand. 17: 165-174. Use of gap (created by tree falls) and non-gap forest understory sites by migrating birds in central Illinois was studied during spring and autumn for three years (1978- 1980). Fruit and understory foliage were concentrated in gaps. Birds that relied on these resources (foliage-gleaning insectivores, frugivores in autumn) used gaps more than non-gaps. Birds that fed on food other than fruit and foliage insects ("frugi- vores" in spring, other insectivores) did not use gaps more than non-gaps. Bird abundance varied markedly among gap and non-gap sites, potentially reflecting differences in site preferences. Site selection, as determined by bird abundances, was consistent (correlated) between years for birds that fed on items that were concen- trated in gaps but not for birds that did not rely on these patchy resources. Foliage density is a measure of foraging substrates for foliage-gleaning birds to search. Abun- dance of foliage-gleaning insectivores was highly correlated with foliage density in both spring and autumn. Frugivore abundance was highly correlated with fruiting fo- liage density during autumn when they are frugivorous, but not during spring when they are insectivorous. Insectivores not relying on foliage insects or fruit were un- correlated with either index of resource availability. These same relationships hold even when examining gap sites only. Thus, migrants can be consistent in their se- lection of foraging sites and this consistency appears to exist when resource densities are markedly different among sites (patchy) but not when resources are more disper- sed. T. E. Martin, Dept of Zoology, Arizona State University, Tempe, Arizona 85287, U.S.A. J. R. Karr, Smithsonian Tropical Research Institute, Box 2072, Balboa, Re- public of Panama. bers at the same time as their food and 1. Introduction peak (Graber Graber 1983). Migratory birds use considerable amounts of energy for When resources are patchy, foraging efficiency also is migration; they lose 1-4% of their gross body weight enhanced by selecting habitat patches with more abun- per hour of flying (Graber and Graber 1962, Hussell dant food resources (Charnov 1976, Krebs et al. 1978, 1969). As a result, migrants require periodical replen- Cowie and Krebs 1979, Martin 1985a). However, patch ishment of lost fat stores at stop-over sites to allow suc- selection by migrating birds is rarely studied. Instead, cessful completion of migration (Nisbet and Medway analyses of habitat selection during migration typically 1972, Berthold 1975). Individuals able to maximize have been restricted to general habitat patterns (e.g., their foraging efficiency (rate of food intake) at these Parnell 1969, Laursen 1976, Rappole and Warner 1976, stop-over sites increase their rate of fat deposition and Martin 1980). Only Willson et al. (1982) examined fine their chances of successful migration. One means of en- scale habitat selection patterns during migration. hancing foraging efficiency is to migrate when food is Willson et al. (1982) showed that migrants were more most abundant. Indeed, spring arrival of migrant war- abundant in understory of light gaps than in undisturbed blers in southern Illinois coincides with eruptions of forest understory. Light gaps are created by tree falls. their primary food (lepidopteran larvae); warbler num- More light reaches the forest floor in light gaps, causing Received 7 January1985 Accepted 20 June 1985 ? ORNIS SCANDINAVICA 11 ORNIS SCANDINAVICA 17:2 (1986) 165 increasedgrowth and colonization by understoryveg- 1978-1980. Autumn was divided into early (before 15 etation (Hartshorn 1978, Thompson 1980, Runkle September) and late (after 15 September) subseasons. 1982). As a result, understoryfruits and foliage insects Nets (30 or 36 mm mesh, 4 shelves, 12 m long) were are more concentratedin light gap patchesthan in non- paired between gap and non-gap sites in the forest un- gaps. Thus, preferences for light gaps may reflect se- derstory. Gap nets were placed at the edge of gaps lection of patches with abundantfood by migrants. rather then in the centres to minimize net visibility as a Yet, the density of plants, and associated food re- bias on capture rates. Similar methods have been used sources, differsamong gap sites (Runkle 1982). If birds by Schemske and Brokaw (1981) and Willson et al. are truly selecting sites based on availability of re- (1982) to examine gap and non-gap use by birds. A core sources, then two resultsare predicted:(1) Birdsshould of 10 (5 pairs) nets was placed in the same locationsin be more abundantsat sites with greateramounts of the all seasons and years to examine consistency of site pref- food types they eat. (2) Birds shouldbe more abundant erences by birds. The gaps used for these locations were at the same sites each year, assuming differences in all old and well established with dense shrub under- qualityamong sites varies only a little among years. To story. In each season an additional gap:non-gapnet examine these predictions, we established permanent pairing was placed in a new gap (< 1 year old) that had gap and non-gapsites and monitoreduse of the under- not developed a shrub understory. story of these sites by migratingbirds during spring and A total of 5,212 mist net hours (MNH) was accrued autumn for three years. We focus on three questions. over the six seasons with 458, 476, and 765 MNH during First, do migratingbirds of differingforaging habits dif- springs, and 1205, 1344, and 964 MNH for autumns of fer in their abundanceat gap versusnon-gap understory 1978, 1979, and 1980, respectively. Nets were opened 30 sites? Second, are birdsconsistent in their site selection min before to 30 min after sunrise and left open for 4-6 (i.e., more abundantin the same sites) each year? Fi- hours in all seasons. nally, are differences in bird abundancesamong sites Vegetation cover was sampled during autumn 1979, correlatedwith differences in indexed resource abun- when plants were fruiting, using the point sample dance? method of Karr (1971) in which presence/absence of vegetation was noted at the following height intervals: G (ground), < 0.25, 0.25-0.5, 0.5-1, 1-2, 2-3, 3-5, 5- 2. Study area and methods 7.5, 7.5-10, 10-12.5, 12.5-15, 15-20, and > 20 m. Num- The study site was Trelease Woods, a 22-ha woodlot lo- ber of points in which vegetation was present relative to cated northeast of Urbana, Illinois, USA. Woodlots and total number of points sampled provided percent veg- other forest islandsare importantsources of forest habi- etation cover in each height interval. A total of 30 tat for migratingand breedingbirds in the mid-western points was sampled on three transects at each net. Two U.S. (Martin 1980, 1981, Blake 1983). Principaltree transects ran parallel to the net at a distance of 3-4 m species in TreleaseWoods includedoaks Quercusspp., from the net on each side with points taken every 1 m. sugar maple Acer saccharum, elm Ulmus, white ash The third transect ran perpendicular to and bisected the Fraxinus americana, basswood Tilia americana, and net with points taken every 2 m. Foliage of individual hackberry Celtis occidentalis. The understory included plants that actually bore bird-dispersed fruits (males paw paw Asimina triloba, spicebush Lindera benzoin, and some female individuals of some plants species did grape Vitis spp., moonseed Menispermum canadensis, not bear fruit) was tabulated separately to provide fo- pokeweed Phytolacca americana, Virginia creeper Par- liage cover of fruiting plants. Each site was ranked rela- thenocissus, and poison ivy Toxicodendron radicans. tive to total foliage cover and fruiting foliage cover in Most census proceduresused in studies of birdswere the understory (< 3 m). Analyses based on ranks are designedfor use duringbreeding season when most spe- preferred because true availabilities of resources to ani- cies are relativelysedentary and vocal. Studies during mals are difficult to measure accurately (Johnson 1980). other seasons and in regions such as tropical forest Abundance of birds at each net site were ranked where assumptionsof more classicalprocedures are not based on capture rates (birds/100 MNH) and compared met requiredifferent methods (Karr1979, 1981). Birds among seasons and years and with foliage and fruit during migration are highly transient and often are not cover rankings using a Spearman rank correlation. vocal, especially in autumn. Thus, we used mist-nets to Comparisons among seasons and years allow examin- measure avian use of patches because: (1) We were in- ation of whether birds are consistently more abundant terested in avian use of forest understory patches from 0 at the same net sites. Comparisons with fruit and foliage to 3 m above the ground and mist-nets sample the un- cover rankings allow examination of whether birds are derstory, (2) mist-netsdo not depend on vocalizations more abundant at sites with more fruit or foliage cover. or sedentary birds, and (3) mist-nets allow examination Correlations were based only on the 10 sites netted in all of avian use of small areas, which is a central focus of 6 seasons. this study. Comparisons of capture rates among net sites, micro- Patchuse by birdswas sampledduring spring (15 Ap- habitats, seasons and years were made using the Fisher ril - 29 May) and autumn (24 August - 18 October) of binomial probability test when sample size was small (n 166 ORNIS SCANDINAVICA 17:2 (1986) < 35) and by X2analysis for large samples. Food habit Tab. 1. Capturerates (birds/100mist-net hours) for birdscap- followed Willson et al. for tured in light gaps and undisturbed(non-gap) understoryin assignments (1982) except - 14 and late Warbler and American Redstart.