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Cannibalism, Food Limitation, Intraspecific Competition See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/7437851 Cannibalism, food limitation, intraspecific competition, and the regulation of spider populations Article in Annual Review of Entomology · February 2006 DOI: 10.1146/annurev.ento.51.110104.150947 · Source: PubMed CITATIONS READS 213 593 1 author: David H. Wise University of Illinois at Chicago 118 PUBLICATIONS 5,094 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: RESTORE Project View project All content following this page was uploaded by David H. Wise on 09 April 2016. The user has requested enhancement of the downloaded file. 2Nov 2005 13:14 AR ANRV263-EN51-19.tex XMLPublishSM(2004/02/24) P1: KUV 10.1146/annurev.ento.51.110104.150947 Annu. Rev. Entomol. 2006. 51:441–65 doi: 10.1146/annurev.ento.51.110104.150947 Copyright c 2006 by Annual Reviews. All rights reserved First published online as a Review in Advance on September 22, 2005 CANNIBALISM,FOOD LIMITATION, INTRASPECIFIC COMPETITION, AND THE REGULATION OF SPIDER POPULATIONS David H. Wise Department of Entomology, University of Kentucky, Lexington, Kentucky 40546-0091; email: [email protected] KeyWords Araneae, intraguild predation, population regulation, trophic cascades ■ Abstract Cannibalism among generalist predators has implications for the dy- namics of terrestrial food webs. Spiders are common, ubiquitous arthropod generalist predators in most natural and managed terrestrial ecosystems. Thus, the relationship of spider cannibalism to food limitation, competition, and population regulation has direct bearing on basic ecological theory and applications such as biological control. This review first briefly treats the different types of spider cannibalism and then focuses in more depth on evidence relating cannibalism to population dynamics and food web interactions to address the following questions: Is cannibalism in spiders a foraging strategy that helps to overcome the effects of a limited supply of calories and/or nu- trients? Does cannibalism in spiders reduce competition for prey? Is cannibalism a significant density-dependent factor in spider population dynamics? Does cannibalism dampen spider-initiated trophic cascades? INTRODUCTION Terrestrial food webs have a high diversity of generalist predators (32, 102, 105), by UNIVERSITY OF ILLINOIS - CHICAGO on 08/23/07. For personal use only. among which spiders are abundant and ubiquitous (143). Cannibalism occurs in a Annu. Rev. Entomol. 2006.51:441-465. Downloaded from arjournals.annualreviews.org wide range of generalist predators (43, 100) and is perceived by many researchers to be common among spiders. Types of cannibalism can be classified according to the life-history stages and relatedness of cannibal and victim. Among spiders sexual cannibalism has been the most extensively studied, yet Elgar (31) points out that among invertebrates “...sexual cannibalism is surprisingly uncommon relative to the other types of cannibalism.” The current review focuses on the less- studied types of cannibalism in spiders. This emphasis is not meant to suggest a clear dichotomy, because some of the selective pressures molding both sexual and nonsexual cannibalism are similar. Thus, aspects of sexual cannibalism are reviewed briefly. Emphasizing research findings on nonsexual forms of spider cannibalism hopefully places in perspective what is currently known of the roles 0066-4170/06/0107-0441$20.00 441 2Nov 2005 13:14 AR ANRV263-EN51-19.tex XMLPublishSM(2004/02/24) P1: KUV 442 WISE that cannibalism plays in the ecology of a major generalist predator, with a primary goal of identifying research questions deserving future scrutiny. Spiders are frequently generalized as highly cannibalistic, which could mean that most spider species exhibit cannibalism and/or that killing and eating con- specifics occurs frequently in those species that are cannibalistic. Given the gaps in our knowledge about the frequency of cannibalism in nature, generalizations about rampant spider cannibalism are sometimes too glibly stated, but it is true that spiders and other arachnids display a wide range of cannibalistic behaviors. Elgar & Crespi (32) list five types based upon the life-history stage of cannibal and victim: adults cannibalizing adults, adults cannibalizing juveniles, juveniles can- nibalizing juveniles, adults cannibalizing eggs, and juveniles cannibalizing eggs. Among invertebrates only arachnids exhibit all five types, and all five occur in spi- ders (32). However, it does not follow that cannibalism occurs frequently in most spider species. Sexual cannibalism has been documented in many spider families; however, Elgar (31) points out that we have scant data on the actual frequency of sexual cannibalism in natural populations. It appears that sexual cannibalism is relatively rare among the therophosids (22, 61). Jackson (59) suggests that sex- ual cannibalism may not be as widespread in spiders as commonly believed and that many of the courtship behaviors attributed to the avoidance of sexual can- nibalism have alternative explanations. Elgar (31), however, suggests that sexual cannibalism in spiders is widespread. Relatively few investigators have attempted to quantify the frequency of sexual cannibalism among spiders in nature. We do not have a good idea of the extent to which nonsexual cannibalism is a major spider foraging behavior. Predation by spiders on other spider species has been documented for numerous families in both nature and laboratory (143), and authors (30, 41, 44) frequently echo Bristowe’s (14) assessment that puts “spiders easily at the top of the list of spider enemies.” Even if it is true, widespread predation by spiders on other spider species does not necessarily implicate cannibalism as a major factor in spider ecology. Furthermore, it does not necessarily follow that cannibalism is frequent in nature if conspecific individuals confined in the laboratory kill and eat each other. A review of the evidence indicates that, despite the absence of extensive data on the frequency of cannibalism in many families in by UNIVERSITY OF ILLINOIS - CHICAGO on 08/23/07. For personal use only. nature, available data from field surveys, from accumulating evidence from field Annu. Rev. Entomol. 2006.51:441-465. Downloaded from arjournals.annualreviews.org experiments, and from laboratory studies in which habitat structure and densities of conspecifics and alternative prey are similar to natural conditions suggest that nonsexual cannibalism may be a significant foraging behavior in some spider families, especially the families of wandering spiders, and among them, the wolf spiders (Lycosidae) in particular. THEORETICAL CONSIDERATIONS Cannibalism and Intraguild Predation The ecological roles of spider cannibalism in population and food web dynamics are understood best when compared with intraguild predation (IGP) because both 2Nov 2005 13:14 AR ANRV263-EN51-19.tex XMLPublishSM(2004/02/24) P1: KUV CANNIBALISM IN SPIDER POPULATIONS 443 involve generalist predators feeding on other predators that share one or more non- predacious prey species. Such a comparison also highlights how costs and benefits interact in shaping the evolution of cannibalistic behaviors in spiders. Polis (101) defined IGP as “the killing and eating of other species that use the same resources and are therefore potential exploitative competitors.” Many authors view cannibal- ism as a form of IGP, but the definition proposed by Polis excludes cannibalism. Separating IGP and cannibalism recognizes a crucial difference between the two behaviors, i.e., that costs of the two can be different because cannibalism can incur direct genetic costs that are impossible in IGP. On the other hand, their bene- fits overlap extensively, including the advantage gained by eliminating a potential competitor, an indirect effect that should be even greater for cannibalism than for IGP. The benefits ascribed to cannibalism are acquisition of a high-quality food source and elimination of a potential exploitative competitor. The possible costs are injury or death, contracting pathogens or parasites, lowering of inclusive fitness by killing relatives, and loss of sperm. Asymmetries, Ecological Factors, and Population Consequences One can make several generalizations about factors that should influence rates of cannibalism (28, 32, 43, 100). Certain asymmetries influence the risk involved: Cannibals should prey on smaller conspecifics in order to avoid the risk of retalia- tion; they should avoid killing kin; and they should avoid killing a mate unless the energy or nutrients acquired outweigh the cost of losing sperm. Because cannibal- ism is most frequently a predator-prey interaction, its frequency should respond to the ecological factors of cannibal density, alternative prey (both abundance and relative food quality), and habitat structure, all of which can act separately or together to determine the frequencies of encounters with potential cannibals or potential prey (conspecific and heterospecific), the spectrum of foraging choices, hunger level, and the strength of exploitative competition. Many of these general attributes lead to the prediction that certain types of cannibalism should act as a strong density-dependent factor regulating population density, which has conse- by UNIVERSITY OF ILLINOIS - CHICAGO on 08/23/07. For personal use only. quences for spider-initiated trophic cascades. Annu. Rev. Entomol. 2006.51:441-465. Downloaded
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