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Top-Down Cascade from a Bitrophic Predator in an Old-Field Community

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Top-Down Cascade from a Bitrophic Predator in an Old-Field Community Wright State University CORE Scholar Biological Sciences Faculty Publications Biological Sciences 1996 Top-Down Cascade from a Bitrophic Predator in an Old-Field Community Matthew D. Moran Thomas P. Rooney Wright State University - Main Campus, [email protected] L. E. Hurd Follow this and additional works at: https://corescholar.libraries.wright.edu/biology Part of the Biology Commons, Medical Sciences Commons, and the Systems Biology Commons Repository Citation Moran, M. D., Rooney, T. P., & Hurd, L. E. (1996). Top-Down Cascade from a Bitrophic Predator in an Old- Field Community. Ecology, 77 (7), 2219-2227. https://corescholar.libraries.wright.edu/biology/82 This Article is brought to you for free and open access by the Biological Sciences at CORE Scholar. It has been accepted for inclusion in Biological Sciences Faculty Publications by an authorized administrator of CORE Scholar. For more information, please contact [email protected]. Top-Down Cascade from a Bitrophic Predator in an Old-Field Community Author(s): Matthew D. Moran, Thomas P. Rooney and L. E. Hurd Source: Ecology, Vol. 77, No. 7 (Oct., 1996), pp. 2219-2227 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/2265715 . Accessed: 19/03/2013 16:01 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology. http://www.jstor.org This content downloaded from 130.108.169.201 on Tue, 19 Mar 2013 16:01:19 PM All use subject to JSTOR Terms and Conditions Ecology, 77(7), 1996, pp. 2219-2227 ( 1996 by the Ecological Society of America TOP-DOWN CASCADE FROM A BITROPHIC PREDATOR IN AN OLD-FIELD COMMUNITY' MATTHEW D. MORAN Ecology Program, Department of Biology, University of Delaware, Newark, Delaware 19716 USA THOMAS P. ROONEY Department of Biology, Indiana University of Pennsylvania, Indiana, Pennsylvania 15705 USA L. E. HURD2 Department of Biology, Washington and Lee University, Lexington, Virginia 24450 USA Abstract. We tested the hypothesis that a bitrophic (third and fourth level) arthropod predator can exert a cascading, top-down influence on other arthropods and plants in an early successional old field. First-stadium mantids, Tenodera sinensis, were added to rep- licated open-field plots in numbers corresponding to naturally occurring egg hatch density and allowed to remain for ;2 mo. Sticky-trap dispersal barriers around both control and mantid-addition plots allowed us to monitor emigration of arthropods continuously during the experiment. Biomass of herbivores, carnivores, and plants, and abundances of arthropod taxa within plots were determined at the beginning, middle, and end of the experiment. The impact of mantids on the community was a top-down trophic cascade, beginning at the fourth trophic level and evident at each of the lower three levels. Mantids induced marked behavioral responses in other predators, but interference among predators did not prevent the trophic cascade. The most common predators, cursorial spiders, emigrated from mantid addition plots in significantly greater numbers than from controls. This behavioral response may have resulted from avoidance of predation or competition. Mantids decreased biomass of herbivorous arthropods through predation, and this de- crease in turn increased biomass of plants. Therefore, these generalist predators were able to decrease herbivory enough to affect plant growth. This and other recent studies indicate that top-down effects can be important in structuring terrestrial communities. Ours is the first example of a top-down cascade by a generalist arthropod predator in a nonagricultural ecosystem and illustrates the importance of detecting behavioral responses in studies of trophic interactions. Key words: arthropod assemblages; behavioral responses; herbivore load; intraguild interac- tions; mantids; old fields; predator load; predators, bitrophic and generalist; Tenodera sinensis; top- down forces; trophic cascades. INTRODUCTION have a tendency to produce intraguild interactions such The importance of top-down vs. bottom-up forces in as predation (Polis et al. 1989, Hurd and Eisenberg trophic level interactions has been the topic of recent 1990a), exploitation competition (Spiller 1984, Hurd debate in ecology. No real agreement has been reached and Eisenberg 1990b), interference competition (Spil- on the importance of top-down and bottom-up forces ler 1984, Wise 1993), and predator avoidance behavior in terrestrial systems (Hunter and Price 1992, Power (Stamps 1983, Moran and Hurd 1994). These inter- 1992, Strong 1992). A strict dichotomy between these actions can result in both direct and indirect effects on two processes may be counterproductive, as it is likely arthropod assemblages (Risch and Carroll 1982, Hurd that both top-down and bottom-up forces interact to and Eisenberg 1984b, 1990a, Fagan and Hurd 1991, structure all communities (Menge 1992). However, the 1994, Riechert and Bishop 1990). How interactions question remains: in which communities and under within the generalist predator guild influence plant as- what conditions are top-down or bottom-up forces im- semblages is poorly understood, and may not be portant? straightforward. For example, spiders eat pollinating The effects of generalist predators on prey assem- insects, which may negatively affect plants, but also blages is complicated by the fact that they occupy two eat seed predators, which may benefit plants (Louda trophic levels (third and fourth) at the same time. Such 1982). bitrophic predators (sensu Hurd and Eisenberg 1990a) In spite of the argument by Price et al. (1980) that enemies of herbivores are mutualists of plants, there ' Manuscript received 7 July 21 1995; revised November have been relatively few studies investigating the ef- 1995; accepted 9 December 1995; final version received 25 January 1996. fects of predators on plants in terrestrial systems. Stud- 2 Address reprint requests to this author. ies detecting top-down effects in terrestrial commu- 2219 This content downloaded from 130.108.169.201 on Tue, 19 Mar 2013 16:01:19 PM All use subject to JSTOR Terms and Conditions 2220 MATTHEW D. MORAN ET AL. Ecology, Vol. 77, No. 7 nities have focused on arthropod specialists such as inant members of the generalist arthropod predator hymenopteran parasitoids (Gomez and Zamora 1994). guild in such systems. Studies with parasitoids typically involve only one or a few host insects and therefore have little chance of MATERIALS AND METHODS detecting effects on very diverse systems such as ter- Study site restrial insect communities. Other top-down studies The study site was a large field located in northern have shown effects of vertebrate predators on plant New Castle County, Delaware. It had been periodically growth and reproduction (Altegrim 1989, Spiller and cut for hay in recent years (last time in fall 1993) so Schoener 1990, 1994, Marquis and Whelan 1994, that at the beginning of the study aboveground plant McLaren and Peterson 1994, Dial and Roughgarden biomass was low. The major plant species present ini- 1995). Generalist arthropod predators are less well tially were Trifolium pratense, Poa sp., Viola papil- studied, although it is clear that they can exert impor- ionacea, Phleum sp., Lynchis alba, and Allium sp. tant influences on arthropod assemblages, including There was not an established population of mantids at herbivorous arthropods that can be important to plant the beginning of the study, but a diverse assemblage succession (Brown 1985). We have found no experi- of insects and other arthropods was present (M. D. mental evidence in the literature that generalist arthro- Moran, personal observation from previous year). Ap- pod predators affect primary producers in complex nat- proximately 30 m from the south side of experimental ural communities; however, Riechert and Bishop plots was a line of trees, and the other three sides of (1990) found that introduced spider predation de- the plots were on the edges of large open fields. creased leaf damage in a garden test system. Possible mechanisms by which trophic cascades Experimental design function have not been well investigated in terrestrial During April of 1994 12 6 X 6 m plots were estab- systems, although the role of behavior has been noted lished in pairs, resulting in six groups composed of two in several aquatic studies (Werner et al. 1983, Douglas plots each. Each treatment and control plot within a et al. 1994, Hill and Lodge 1994). Behavior is an im- pair was separated by 3 m while each pair of plots was portant component of predator-prey interactions (Lima separated from other pairs by 20 m. Treatment plots and Dill 1990, Anholt and Werner 1995) and thus (receiving mantids) and control plots (no mantids) were should not be ignored in design of cascade experiments. systematically interspersed (i.e., alternating) within the The most abundant generalist predators in old-field pairs of plots (Hurlbert 1984). Each plot was bounded communities in our study area are mantids and spiders. by an 0.5 m wide black plastic barrier that was stapled The most common mantid in the mid-Atlantic region to furring strips placed around the outside edge of the of the United States is the Chinese mantid (Tenodera plot. The day before introduction of mantids, a 6-cm sinensis), which often reaches high densities upon band of Tangletrap (Tanglefoot Company, Grand Rap- emergence in the spring (Hurd and Eisenberg 1984a, ids, Michigan) was painted in a broad stripe down the Eisenberg and Hurd 1990). This is a semelparous uni- middle of the barrier to trap emigrating mantids and voltine insect that hatches in the spring, matures late spiders.
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