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Bird Predation on Periodical Cicadas in Ozark Forests: Ecological Release for Other Canopy Arthropods?

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Bird Predation on Periodical Cicadas in Ozark Forests: Ecological Release for Other Canopy Arthropods? Studies in Avian Biology No. 13:369-374, 1990. BIRD PREDATION ON PERIODICAL CICADAS IN OZARK FORESTS: ECOLOGICAL RELEASE FOR OTHER CANOPY ARTHROPODS? FREDERICK M. STEPHEN, GERALD W. WALLIS, AND KIMBERLY G. SMITH Abstract. Population dynamics of canopy arthropods were monitored in two upland forests in the Arkansas Ozarks during spring and summer of 1984-1986 to test whether the emergence of adult 13-year periodical cicadas on one site during late spring in 1985 would disrupt normal patterns of bird predation on canopy arthropods, resulting in ecological release for those prey populations. Canopy arthropods on foliage of oak, hickory, and eastern redcedar were sampled weekly beginning in June 1984, and April 1985 and 1986, and continuing through August in all years. We classed arthropods into four broad guilds based on foraging mode (chewers, suckers, spiders, and lepidopterous larvae) and expressed densities as number of individuals per kg of foliage sampled. Two-way analysis of variance revealed no significant treatment effects for densities of chewing, sucking, or lepidopteran larval guilds. A significant interaction of mean density between sites among years was detected for the spider guild, but not when cicadas were present, indicating that ecological release did not occur. We trace the development of the notion that bird populations are capable of affecting prey population levels, and discuss those ideas in light of our results, which suggest that birds have little impact upon their arthropod prey in Ozark forests. Kev Words: Arkansas: canonv arthronods: ecological release; guilds; insect sampling; Magicicada; Ozarks; periodical cicadas; predation. _ ’ One of the most predictable events in nature (Mugicicudu tredecim Walsh and Riley, M. tre- is emergence of periodical cicadas (Homoptera: decussini Alexander and Moore, and M. trede- Cicadidae: Magicicada). Unavailable to above- culu Alexander and Moore; Brood XIX, Simon ground consumers for long periods (either 13 or 1979) in northwestern Arkansas in 1985 afforded 17 years), they become superabundant for about us an opportunity to test this idea. We sampled 6 weeks as adults, while they reproduce, then die canopy arthropods during pre-emergence (1984) (Marlatt 1907). Densities may approach 3 mil- emergence (1985), and post-emergence (1986) lion/ha (Dybas and Davies 1962) and emer- summers on two study sites, one on which ci- gences are regarded as a classic example of pred- cadas emerged in 1985 and another nearby where ator swamping (Lloyd and Dybas 1966a, b). they did not. Periodical cicadas apparently contain no noxious compounds (Brown and Chippendale 1973) and STUDY AREAS AND METHODS have only limited anti-predatory behaviors We used anecdotal records of emergences of 13-year (Steward et al. 1988a) and during their emer- periodical cicadas from Brood XIX (Simon 1979) from gences become a highly desirable prey for many 1959 and 1972 in northwestern Arkansas to locate study bird species (Marlatt 1907, 1908; Forbush 1924; areas during 1984. Cicada chorusing had been notable Beamer 1931; Allard 1937; Howard 1937; Lcon- at certain farms adjacent to the White River, near Dur- ard 1964; Nolan and Thompson 1975; Anderson ham, Washington County (L. 0. Warren and R. Wat- 1977; Best 1977; Karban 1982; Murphy 1986; son, pers. comm.), and examination of favored tree Strehl and White 1986; Steward et al. 1988a; species revealed substantial twig scars (Marlatt 1907) from 1972. Digging among roots of scarred trees yield- Kellner et al., this volume). Indeed, predation is ed vertical pre-emergence tunnels, which cicada nymphs generally assumed to have been a driving force construct up to one year before emergence (Cory and in the evolution of the life cycle (Lloyd and Dy- Knight 1937). A 16 ha study site (Cassidy) was estab- bas 1966b, May 1979, Karban 1982). lished here as the cicada site. Another 16 ha site (Til- Given that most forest birds in the Ozarks eat lery) located 3.5 km to the east served as the control adult periodical cicadas when they are available, site. what is the effect of prey switching behavior by The Cassidy site was more xeric than the Tillery site birds on populations of the normal canopy ar- and was on a predominantly west-facing slope, meeting thropod prey? In particular, do normal prey items a pasture adjacent to the White River. Both locations were similar in tree species composition, with a wide experience a type of “ecological release,” where- variety of hardwoods characteristic of Ozark upland by populations expand greatly as a result of a forests (Moore 1972) including a predominance of oaks decrease in avian predation pressure owing to (Quercus spp.) and hickories (Curya spp.), plus abun- the appearance of a superabundant prey? dant eastern redcedar (Juniperusvirginiana L.). Those Emergence of 13-year periodical cicadas three taxa were sufficiently numerous and distributed 369 370 STUDIES IN AVIAN BIOLOGY NO. 13 within both sites so that they were selectedas sample as small (~6 mm), medium (~6 and < 19 mm), or trees for the canopy arthropod study. large (2 19 mm). Specimenswere categorizedinto four guilds based loosely on feeding behavior: (1) chewing Arthropod sampling insect guild, containing all families in orders Lepidop- Each site was divided into a grid consistingof 100, tera and Orthoptera, sawfly families in the order Hy- 0.16 ha (40 x 40 m) subplots. Data were collected menoptera, and families Chrysomelidae, Scarabaeidae weekly from 1984 through 1986. Sampling began in and Curculionidaein the Coleoptera;(2) suckinginsect June 1984 and in April of 1985 and 1986, approxi- guild, containing all families in orders Homoptera and mately coincident with appearanceof new foliage, and Thysanoptera, plus families Aradidae, Berytidae, Co- continued through August in all three years of study. reidae, Miridae, Pentatomidae, Scutelleridae, Tingi- The sampling regime varied slightly each year, as dae, and Thyreocoridae in the order Hemiptera; (3) refinementswere made to optimize efficiency. In 1984, spider guild, containing all spiders;and (4) medium to eight subplotswere randomly selectedon each site:four large lepidopterous larval guild, a subgroup of the alongthe perimeter of each site and four in the interior. chewing insect guild. Within those subplots,two crown heights (< 10 m and Not all taxa collected are included in our guilds. > 10 m) were sampled on each of the three tree taxa. Choices for guild membership were made to incor- Each week, 48 samples (8 subplots x 3 tree taxa x 2 poratethe two main phytophagousinsect feeding types, crown heights) were taken, resulting in 1008 samples. plus the large,entirely entomophagous,arachnid group. In subsequentyears, high and low crown heights were Medium to largelepidopteran larvae (a relatively large- merged into one lower mid-crown sample, as we de- sized and flightlessfood resource)were examined sep- tected no significant differences in arthropod popula- arately, as they form a common food for insectivorous tions as a function of height in 1984. Analysesof 1984 birds of the forest canopy (Holmes et al. 1979~).Other data in this paper used only the 504 lower crown sam- authors, working with similar canopy arthropod data, ples, as they were most similar in height to all subse- have demonstrated that choice of taxa in guild for- quent samples. mation can significantlyinfluence results (Stork 1987; The number of subplotswas increasedin 1985 to 16 Cooper et al., this volume). We are aware that the at the control and 24 at the cicada site for a total of feeding guilds we specified are broad, but their taxo- 2400 samples. Sampling in 1986 was similar to the nomic composition remained relatively constant be- previous year, but one week shorter, resultingin 2244 tween cicada and non-cicada sites. samples. All samplingwas conductedin morning to minimize Data analyses variation in insect movement within the crown (e.g., We summarized data in two ways: (1) numbers of Holmes et al. 1978). Each sample unit consisted of individuals/sample unit; and (2) numbers of arthro- three terminal branches from a tree cut with a pole pods standardized by kg of tree foliage sampled, cal- pruner, and dropped into an attached muslin bag. We culated by dividing number of individuals by weight attempted to standardizeeach branch cutting to sample of foliage.Exploratory data analysisindicated that most similar amounts of foliage for each species through variableswere not normally distributed and that mean/ time. Foliage consisted of leaves, petioles, and small variance ratios were not stable. Becausemost stan- twigs (larger stems were clipped and discarded). All dardized variables approximated a negative binomial foliage was immediately placed into a paper bag (ca. or Poisson distribution, for all analysesof variance a 30 x 17 x 30 cm), which was folded and stapledclosed. squareroot transformationwas usedafter 3/8 was added Bags were returned to the laboratory, weighed to de- to each value, a processthat stabilizes the variance of termine wet weight of foliage biomass, then frozen a Poisson distribution regardlessof the mean (Ans- overnight to kill all arthropods. The next day, foliage combe 1948). Standardizeddata reported in tables are was shaken, and all stem and leaf surfacescarefully untransformed means, presentedwith associatedsam- examined. Total arthropod wet weight was measured ple sizes and standard errors of the mean. and specimenswere held for identification in 70% ethyl The General Linear Models procedure in SAS (SAS alcohol. 1982) was used for analysisof-variance, with Tukey’s Multiple methods of sampling may be required to mean separation tests (P 5 0.05) where appropriate. properly sample entire canopy arthropod communities Two-way analysisof variance was used to test for dif- (Cooper 1989, Morrison et al. 1989). Methods such as ferences in mean densities between sites and among pole pruning miss actively flying insects, while tech- years, and, more importantly, to determine if any sig- niques that do catch fliers (e.g., sticky traps) usuallydo nificant site-year interactions existed.
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