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Extending the Resource Concentration Hypothesis to Plant Communities: Effects of Litter and Herbivores Author(S): Zachary T

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Extending the Resource Concentration Hypothesis to Plant Communities: Effects of Litter and Herbivores Author(S): Zachary T Extending the Resource Concentration Hypothesis to Plant Communities: Effects of Litter and Herbivores Author(s): Zachary T. Long, Charles L. Mohler, Walter P. Carson Source: Ecology, Vol. 84, No. 3 (Mar., 2003), pp. 652-665 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/3107860 . Accessed: 04/03/2011 10:46 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=esa. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. 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 Ecology, 84(3), 2003, pp. 652-665 X 2003 by the Ecological Society of America EXTENDING THE RESOURCE CONCENTRATION HYPOTHESIS TO PLANT COMMUNITIES: EFFECTS OF LITTER AND HERBIVORES ZACHARY T. LONG,"3 CHARLES L. MOHLER,24 AND WALTER P. CARSON' 'Department of Biological Sciences, University of Pittsburgh, Pittsburgh, Pennsylvania 15260 USA 2Department of Ecology and Evolutionary Biology, Corson Hall, Cornell University, Ithaca, New York 14853 USA Abstract. We extend the resource concentration hypothesis (herbivorous insects are more likely to find and stay in more dense and less diverse patches of their host plants) to plant communities. Specifically, whenever superior plant competitors spread to form dense stands, they will be found and attacked by their specialist insect enemies. This will decrease host plant abundance, causing a reduction in standing crop biomass, which will indirectly increase subordinate competitors and plant species richness. In this study, we found that a native, specialist chrysomelid beetle (Trirhabda virgata) in an old-field community de- creased total standing crop biomass, leading to an increase in plant species richness. This reduction in biomass was due solely to a reduction in the biomass of the beetle's host plant, meadow goldenrod (Solidago altissima), which was the dominant plant species in this community. Our results demonstrate that when a superior competitor increases in density, the per-stem impact of herbivores increases due to a buildup of these herbivores in high- density host patches. Specifically, we found that as goldenrod density increased, the per- stem abundance of Trirhabda virgata also increased. In turn, species richness increased as the negative effect of insects on goldenrod biomass increased. Recent research suggests that litter accumulation could negate or cancel the effect of herbivorous insects on plant communities because litter accumulation increases with stand- ing crop biomass, causing a decline in species richness. The litter accumulation hypothesis states that, in productive communities, the increase in the abundance of the superior com- petitor will lead to a dense accumulation of plant litter, causing a decline in species richness. Consistent with this hypothesis, we found that as the biomass of the dominant plant species increased, litter mass also increased. In turn, species richness decreased as the negative effects of litter on stem density increased. Interestingly, the effect of litter on stem density depended on whether insects were present. Our results suggest the potential for a general rule: specialist insect herbivores will function as classic keystone species in plant com- munities whenever host species form large, dense aggregations if host plants are dominant species. Key words: goldenrod; herbivory; insects; litter; old-field community; resource concentration hypothesis;Solidago altissima;Trirhabda virgata. INTRODUCTION fects of herbivorous insects on plant communities. Un- Examples of herbivorous insects regulating terres- fortunately, few empirical data directly test this asser- trial plant communities are scarce, possibly because tion (Crawley 1997). Recent evidence suggests that predators and parasites usually keep insect abundance there are at least three conditions that strengthen the low, or because plants are well defended, or both links between insects and plants. These conditions fre- (Strong et al. 1984, Strong 1992, Hairston and Hairston quently occur during outbreaks (Carson and Root 1993, Schmitz 1994, Carter and Rypstra 1995, Polis 2000), whenever insects can muster an effective anti- and Strong 1996, Price 1997, Uriarte and Schmitz predator defense (Carson and Root 1999), and poten- 1998). Polis and Strong (1996) argued that the retic- tially when there are strong feedbacks between host ulate and diffuse nature of terrestrial food webs weaken plant density and per-stem rates of damage by insects trophic links and typically prevent strong top-down ef- (Root 1973, Andow 1991, Carson and Root 2000). The resource concentration hypothesis (Root 1973) Manuscript received 27 September 2001; revised 4 August describes a viable insect-plant feedback mechanism 2002; accepted 7 August 2002. Corresponding Editor: D. A. that may typically operate in plant communities. To Spiller. 3Present address: Department of Ecology, Evolution, and date, this mechanism has not been applied at the scale Natural Resources, Cook College, Rutgers University, 14 of entire plant communities. The resource concentra- College Farm Road, New Brunswick, New Jersey 08901-8551 tion hypothesis states that "herbivores are more likely USA. E-mail: [email protected] to find and remain on hosts that are growing in dense 4Present address: Department of Crop and Soil Sciences, 907 Bradfield Hall, Cornell University, Ithaca, New York or nearly pure stands" (Root 1973). Numerous popu- 14853 USA. lation-level studies lend strong support to this hypoth- 652 March 2003 HERBIVORY, LITTER, AND PLANT DIVERSITY 653 Insect | Resource } herbivores Dominant Plant species concentration: increase in plant richness is ceonnraton inrdensehot biomass is indirectly F 1 patch a h reducedc pe increased |Dominant plant|| species increases in abundance R . , t Al L~~~~~~~ittern ~ ~ Totalstem | |Plant species| Titter ~~~~~accumulaton I HI density A Jrichness accumulation: ||increase WI| declines | | declines | FIG. 1. The resourceconcentration model andthe litteraccumulation hypothesis make contrasting predictions. The resource concentration model predicts that an increase in host plant density leads to an increase in specialist insect herbivores and a subsequent decrease in host plant biomass. Insects indirectly increase plant species richness by reducing host plant biomass if the host plant is a dominant competitor. The litter accumulation hypothesis predicts that plant litter accumulation causes a decrease in total stem density, leading to a decrease in species richness. esis (e.g., Root 1973, Risch et al. 1983, Andow 1991, herbivory, litter accumulation, and plant communities. Coll and Bottrell 1994, Schellhorn and Sork 1997). The goldenrod Solidago altissima is the dominant com- Furthermore, Bach (1980) found that the increased petitor in numerous old fields throughout the north- abundance of insects in host plant monocultures de- eastern United States (Bazazz 1996). This species often creased the growth of the host plant. Oddly, the logical forms dense, nearly monospecific stands with corre- extension of this hypothesis to entire communities has spondingly high net primary production and standing never been made. Here, for the first time, we extend crop biomass (Carson and Barrett 1988, Bazazz 1996, the resource concentration hypothesis to plant com- Carson and Root 2000); these stands result in com- munities. Specifically, if specialist herbivorous insects munities of low diversity (Bazazz 1996, Carson and are more likely to reduce the vigor and abundance of Root 2000). We tested if the density of the dominant host plants that form dense aggregations, and host plant species determined the abundance of herbivorous plants are dominant species in the community, then insects and their effect on plant diversity. We also test- specialist herbivorous insects could act as keystone ed whether dominant or subordinate plants primarily species, reducing the total standing crop biomass and determined litter mass and the effects of litter on plant increasing the diversity and fecundity of subordinate species richness. The factorial design of the experiment species. allowed us to integrate insect and litter effects. Litter accumulation, however, can have the opposite effect of herbivorous insects in plant communities. In- METHODS creases in plant community biomass are often associ- Study site ated with higher levels of litter accumulation (Carson and Barrett 1988, Tilman 1993, Foster and Gross 1997), We conducted this experiment in an abandoned
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