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Ecology of Herbivorous Arthropods in Urban Landscapes

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Ecology of Herbivorous Arthropods in Urban Landscapes ANRV397-EN55-02 ARI 2 November 2009 10:40 Ecology of Herbivorous Arthropods in Urban Landscapes Michael J. Raupp,1 Paula M. Shrewsbury,1 and Daniel A. Herms2 1Department of Entomology, University of Maryland, College Park, Maryland 20742; email: [email protected]; [email protected] 2Department of Entomology, The Ohio State University, Ohio Agricultural Research and Development Center, Wooster, Ohio 44691; email: [email protected] Annu. Rev. Entomol. 2010. 55:19–38 Key Words First published online as a Review in Advance on urbanization gradient, top-down, bottom-up, defense free space, October 5, 2009 biodiversity, ecosystem processes The Annual Review of Entomology is online at ento.annualreviews.org Abstract This article’s doi: Urbanization affects communities of herbivorous arthropods and pro- 10.1146/annurev-ento-112408-085351 vides opportunities for dramatic changes in their abundance and rich- Copyright c 2010 by Annual Reviews. ness. Underlying these changes are creation of impervious surfaces; All rights reserved variation in the density, diversity, and complexity of vegetation; and 0066-4170/10/0107-0019$20.00 maintenance practices including pulsed inputs of fertilizers, water, and Annu. Rev. Entomol. 2010.55:19-38. Downloaded from arjournals.annualreviews.org pesticides. A rich body of knowledge provides theoretical underpinnings for predicting and understanding impacts of urbanization on arthro- pods. However, relatively few studies have elucidated mechanisms that explain patterns of insect and mite abundance and diversity across ur- banization gradients. Published accounts suggest that responses to ur- by UNIVERSITY OF MARYLAND - COLLEGE PARK MCKELDIN LIBRARY on 12/05/09. For personal use only. banization are often taxon specific, highly variable, and linked to proper- ties of urbanization that weaken top-down and/or bottom-up processes, thereby destabilizing populations of herbivores and their natural ene- mies. In addition to revealing patterns in diversity and abundance of herbivores across urbanization gradients, a primary objective of this re- view is to examine mechanisms underlying these patterns and to identify potential hypotheses for future testing. 19 ANRV397-EN55-02 ARI 2 November 2009 10:40 INTRODUCTION associated with impervious surfaces and ele- vated temperatures, as well as escape from nat- Urbanization brings about dramatic changes in ural enemies. In most cases, however, the data Urbanization land use as croplands, old fields, forests, and are insufficient to establish a mechanistic basis gradient: an natural areas give way to cities and suburbs and for the observed patterns, and the population environmental their buildings, roads, airports, waterways, gar- gradient with less dynamics of arthropods in urban environments dens, and parks (32, 97). Biotic elements shift developed natural or remains poorly understood (27, 32, 37, 97, from communities of plants and animals at var- agricultural areas at 98, 123). one extreme and ious stages of succession to communities as- Several previous reviews provided concep- highly developed areas sociated with ornamental plants and remnants tual frameworks for understanding the ecology dominated by of natural habitats (32, 97). Urbanization cre- anthropogenic of urban biota in general (97, 127) and insects ates a unique ecological gradient along which structures and inputs in particular (27, 32, 37, 98, 130). This review patterns of richness, abundance, and ecosystem at the other extreme examines how features of urban landscapes gen- processes can be studied in a context of hu- Bottom-up: when erate properties that affect arthropod richness man culture, economics, and politics (27, 37, 97, host plants regulate and abundance and predispose plants to attack the structure and 98, 123). by herbivorous arthropods. In some studies, the population dynamics Dramatically elevated densities have been urbanization gradient was clear, ranging from of herbivores and documented frequently in urban forests by her- other organisms at natural areas and parks to residential areas and bivorous arthropods that rarely, if ever, reach higher trophic levels finally to cities. Other studies measured pat- high densities in natural forests (27, 37, 50, Top-down: when terns of herbivore abundance along key features 141, 149, 151). For example, the abundance natural enemies and that vary across urban gradients such as plant of tetranychid mites generally increased, often organisms at higher density, abundance, diversity, vegetational com- trophic levels regulate dramatically, with increasing levels of urban- plexity, relative frequency of native and exotic the structure and ization (6, 30, 78, 132, 140). Economically im- plants, patterns of land use, proportion of im- population dynamics portant infestations of scale insects have been of organisms at lower pervious surface, and frequency and intensity of documented more frequently in urbanized ar- levels common maintenance practices such as pulsed eas than in natural areas (28, 50, 141, 149, Heat islands: inputs of fertilizers, water, and pesticides. Prop- 151). By contrast, urbanization can also lead elevated air erties of urban environments that create op- to decreased abundance and diversity of phy- temperatures in portunities for insects and mites to increase in- middle and high tophagous arthropods. For example, destruc- clude changes in host quality and accessibility latitudinal cities tion and deterioration of habitats associated (bottom-up factors), natural enemy abundance compared with those with urbanization led to the disappearance of in nearby rural areas and diversity (top-down factors), microhabitats almost half of the indigenous species of butter- related to impervious including creation of heat islands, and matrixes flies in the San Francisco Bay Area, and local surfaces and other that disrupt movement and colonization of her- anthropogenic effects extinctions are common as buildings and roads bivores and natural enemies (58, 71, 137, 147). replaced forests and grasslands (46, 112). Annu. Rev. Entomol. 2010.55:19-38. Downloaded from arjournals.annualreviews.org Wefocus on assemblages of arthropods asso- In some cases, but not all, elevated densi- ciated with shade trees and woody ornamental ties of herbivores were called outbreaks. We plants rather than arthropods of turfgrass (re- consider outbreaks to be dramatic increases in viewed in Reference 109), structures, or those the abundance of arthropods that occur in rela- deemed as medically important (reviewed in tively short periods (9) and are often associated by UNIVERSITY OF MARYLAND - COLLEGE PARK MCKELDIN LIBRARY on 12/05/09. For personal use only. Reference 37). We found that most published with feeding injury to plants that exceeds eco- accounts of arthropods in urban settings dealt nomic or aesthetic thresholds. In many cases, with species associated with woody rather than herbivores were associated with plants of eco- herbaceous plants. We review empirical studies nomic or aesthetic value and hence were con- on the diversity and abundance of herbivores sidered pests. Herbivore outbreaks in urban and their natural enemies along urban gradi- environments have been linked to improved ents and explore how features and properties host quality resulting from environmental stress 20 Raupp · Shrewsbury · Herms ANRV397-EN55-02 ARI 2 November 2009 10:40 of urbanization affect arthropod abundance and may have fewer plant genera than sites never diversity, and ecosystem processes in general, as farmed, suggesting that legacy effects of agri- they relate to herbivore populations. culture on plant diversity can be long-lasting. Vegetational texture: Many studies speculated about causes of Plant diversity was inversely related to the age attributes of plant variation in abundance, but most did not ex- of housing in urban sites, with newly estab- communities that amine features of urbanization independently lished urban plantings more diverse than older influence the and variables were often confounded. In very ones (62). Studies in metropolitan New York distribution and few studies were mechanisms underlying pat- City (3) and Washington, DC (39, 61) docu- abundance of arthropods terns convincingly documented. Generally, it mented dramatic reductions in tree diversity is difficult or impossible to separate processes and density with increasing levels of urban- associated with urbanization, such as deterio- ization. For example, 30 species of trees lined ration in habitat quality, from those associated streets in a city in Maryland at a density of ap- with habitat loss, fragmentation, and isolation proximately 48 trees km−2, whereas nearby res- (105, 123). A primary objective of this review idential landscapes supported more than 100 is to propose mechanistic explanations as hy- species of woody plants at a density of about potheses for testing. In doing so, we hope to 69,000 plants km−2 (39, 61). advance the study of arthropod ecology in ur- Urbanization does not always correlate with ban environments. a reduction in floristic diversity (27, 107, 112). Extensive tree planting and landscaping, such as occurs in urban areas that lacked trees prior IMPACT OF URBANIZATION to development (e.g., cities built in deserts and ON ARTHROPOD DIVERSITY prairies) and those with rich horticultural tradi- AND ABUNDANCE tions, enjoy increased floristic richness and as- sociated biodiversity (27, 32, 62, 107). A garden Vegetational Texture Along in Leicester, United Kingdom, contained more the Urbanization
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