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SAB 013 1988 P44-52 Quantifying Bird Predation of Arthropods In

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SAB 013 1988 P44-52 Quantifying Bird Predation of Arthropods In Studies in Avian Biology No. 13:44-52, 1990. QUANTIFYING BIRD PREDATION OF ARTHROPODS IN FORESTS DONALD L. DAHLSTEN, WILLIAM A. COPPER, DAVID L. ROWNEY, AND PAULA K. KLEINTJES Abstract. Sampling insects and other arthropods in forest environments is complicated becauseof the unique attributes of this ecosystem.Entomologists have used many techniquesto quantify forest arthropods, some of which are applicable for quantifying the impact of bird predation, as we illustrate in studies of several defoliators and bark beetles. We describe sampling methods for a defoliator, Douglas-fir tussockmoth (Orgyiapseudotsuguta),and a bark beetle,western pine beetle(Dendroctonus brevicomis).We discussthe types of information that can be obtained for insect populations from thesemethods, the time or cost for different levels of sample error, and the application of thesemethods for evaluating bird predation on arthropods. Key Words: Sampling; predation; defoliators; insectivorous birds; forests; conifers; western pine beetle; Scolytidae; Douglas-fir tussockmoth; Lymantriidae. Forest entomologists have struggled with the insects, particularly insectivorous cavity-nesting quantification ofarthropod abundance for many birds (Dahlsten and Copper 1979). In this paper years. Much work has been done by applied bi- we discuss the types of sampling we have used ologists interested in population dynamics of cer- to assess avian impact on insects on two sub- tain species, efficacy of treatments, or the impact strates in the forest, foliage and bark, and also of insects on resources. Quantitative studies are what it costs to obtain useful information. more complicated in forests than in other en- vironments where insects are of economic im- FOLIAGE SAMPLING portance (such as agriculture; Dahlsten 1976) because forests are vast, continuous regions com- LODGEPOLENEEDLE MINER posed of different tree species of different ages, The lodgepole needle miner (Coleotechnites and a mosaic of stocking (density) patterns milleri; Lepidoptera: Gelechiidae), because of its (Pschom-Walcher 1977). The advantage of for- cyclic availability, is a suitable species for study- est ecosystems is that they generally encounter ing the role of birds in its dynamics. The adult less perturbation than agricultural systems and moths appear only in alternate years and have a probably have a more stable arthropod com- short period of activity, whereas the larvae and munity. Outbreak species (those that reach very pupae are available for a long period. The insect high densities periodically) are relatively rare in has a discrete 2-year life cycle, passing the first forests (Berryman 1986). winter in an early larval instar and the second Most studies of forest insect populations deal winter in the fifth instar. As the birds feed only with single species;associated insects such as nat- on larger larvae, this food source is available only ural enemies, inquilines, and organisms in the in alternate years. In addition, because the insect same feeding guild are often ignored. Regional is a needle miner, the birds must open needles or forest-type arthropod faunistic or community to obtain the larvae, leaving evidence of their studies are rare and typically more qualitative feeding. Finally, the distribution of immature than quantitative. Yet, population information needle miners in the trees has been studied and gathered by entomologists may be useful in as- a sampling method developed (Stark 1952, Ste- sessing the impact of birds on a single insect vens and Stark 1962). The method is similar to species. that for tussock moths, discussed below, and in- Meanwhile, ornithologists desire quantitative volves sampling the tips of lodgepole pine population information about arthropod species branches. eaten by birds. Birds typically feed on several At a study site in the Inyo National Forest, different species and at different heights in the Telford and Herman (1963) found that Moun- foliage. To quantify an adequate number of prey tain Chickadees (Purus gambeli) concentrated items on several substrates is costly and time- their feeding efforts in alternate years on the consuming, however, so compromises and strat- needle miner larvae and that the chickadees ex- ifications are required. hibited a functional response to prey density. The Based on work by our laboratory, we believe chickadees peeled needles in a characteristic way, that better quantification of arthropod prey for leaving evidence of their feeding, and Cassins’ birds is possible. We have had a long-term in- Finches (Carpodacus’ cassinii) also fed on needle terest in the impact of natural enemies on forest miners by clipping the ends of the needles 44 PREDATION ON ARTHROPODS--D&z&en et al. 45 (Dahlsten and Herman 1965). Nest boxes were substance is probably distasteful (Eisner et al. later placed in areas infested and not infested 1974). with needle miners. Mountain Chickadees in- In studying the population dynamics of a pine- creased in density in the infested areas, both dur- feeding sawfly in the Neodiprion jiilviceps com- ing the breeding and postbreeding periods. plex at Mt. Shasta, California, Dahlsten (1967) The needle miner-chickadee system has great watched Evening Grosbeaks (Coccothraustesves- potential for evaluating the impact of a bird on pertinus) feeding on their larvae. Ten trees, 2-4 a single insect species. Because the insect is cryp- m in height, were sampled in each of three study tic during the stage eaten, evidence of chickadee areas at different elevations in a plantation. All feeding can be easily detected. The system is ideal sawfly stages, starting with eggs, were counted. for studying the functional response of chicka- Drop cloths were placed beneath each sample dees, because the prey is available only in alter- tree. The cloths did not catch cocoons, but they nate years, and nest boxes and avian census tech- did catch head capsules and thoraxes of larvae, niques permit study of the numerical response which were discarded by grosbeaks. Some larval of the predator to its prey. remains were also stuck to foliage; counts on and beneath the trees showed a total of 166 sawflies- BUD-MINING SAWFLIES 10% of all the larvae on the study trees in one Bud-mining sawflies (Pleroneuru spp.; Hy- area (Dahlsten 1967). menoptera: Xyelidae) are also well suited for Because the birds were feeding on a known evaluating avian predation. Four species mine population, the portion taken was known, at least new buds on expanding shoots of white fir (Abies from the sample trees. Area-wide estimates can concolor) in California; three have been studied be made from such samples. This is a labor- in detail (Ohmart and Dahlsten 1977, 1978, intensive technique, limited to smaller trees where 1979). The species ofearly instar larvae and adults foliage-feeding insects could be counted and lar- can be distinguished, but the late larval instar val remains could be found on foliage or drop (the stage most likely to be eaten by birds) cannot cloths. be separated to species. The three species were treated as a single group DOUGLAS-FIR TUSSOCKMOTH (DFTM) in an analysis of within-crown distribution and the development of sampling methods at Blod- The tussock moth (Orgyiapseudotsugata;Lep- gett Forest, El Dorado County, California idoptera: Lymantriidae), because of its economic (Ohmart and Dahlsten 1978). Over 94% of the importance in western North America, has been infested buds occurred in the outer portion of the focus of many studies, including the role of the crown, coinciding with the foraging area of insectivorous birds in its population dynamics several birds at Blodgett, particularly the Moun- (Brooks et al. 1978; Torgersen et al., this vol- tain and Chestnut-backed (P. rufescens)Chick- ume). The tussock moth overwinters as eggs in adees. Also, the chickadee nesting period coin- masses on top of female cocoons. Both male and cided with the late larval instars of the sawflies, female cocoons are commonly spun on foliage, late May to early June (Ohmart and Dahlsten although cryptic sites such as cavities in trees are 1977). also used. The cocoons and egg masses, in par- We did not learn how birds open buds to re- ticular, are suitable for stocking studies. Egg move larvae, or if they leave characteristic evi- masses can be sampled and then examined for dence. However, mortality ofthe Pleroneuru fifth evidence of predation, or they can be stocked on larval instar was substantial (Ohmart and Dahl- branches or trunks of trees at different known sten 1977), seemingly because of avian preda- densities and predation evaluated (Dahlsten and tion, as chickadees were observed and photo- Copper 1979, Torgersen and Mason 1987). Pu- graphed by nest box camera units bringing pal stocking showed that most predation was due numerous Pleroneura larvae to their young. to birds, although some was due to ants (Dahl- sten and Copper 1979, Torgersen et al. 1983). PINE SAWFLIES Larvae that feed in the open, like sawflies, are SAMPLING ARTHROPODSON WHITE FIR often fed upon by birds, but no evidence is left This study illustrates how the distribution of on the foliage when they are removed. However, a community of organisms on a given tree species birds often remove sclerotized portions of in- can be determined. Sampling programs can then sects, such as the elytra of beetles and the head be developed for any species known to be eaten capsules of larvae, before eating them or feeding by birds. Relationships among sampling error, them to nestlings. Sawfly larvae, in particular, time spent sampling, and cost are shown, so that exude a brownish substance from their mouth the researcher can better manage available fi- when threatened by a parasitoid or predator. This nancial resources. 46 STUDIES IN AVIAN BIOLOGY NO. 13 caught in large canvas bags and beaten over a large canvas on the ground. All insects and spi- ders were recorded, as was the branch number, dimensions (for foliage area), and other charac- teristics. Some insects were retained for rearing or identification.
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