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The Distribution of Periodical Cicadas in Illinois

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The Distribution of Periodical Cicadas in Illinois TThehe DiDistributionstribution ooff PPeriodicaleriodical CicadaCicada John R. Cooley, Gene Kritsky, Marten J. Edwards, John D. Zyla, David C. Marshall Kathy B. R. Hill, Rachel Krauss, and Chris Simon eriodical cicadas (genus Magicicada) are found only in periodical cicada “broods,” or year-classes. Complicating matters, eastern North America and are notable for their long, prime- there are two life cycles (13 and 17 years), three species groups Pnumbered life cycles, precisely timed mass emergences, and (-decim, -cassini, and -decula), and seven recognized species of dense, multispecies choruses. Their uniqueness has given them a special appeal and cultural status. Members of the Onondaga Na- Moore 1962; Dybas and Lloyd 1962, 1974; Lloyd and Dybas 1966a; tion maintain the oral tradition of being rescued from famine by periodicalWhite 1980) cicada, (Marshall with slightand Cooley ecological 2000). differences Individual broods(Alexander usually and periodical cicadas (Cooley et al. 2004). Early European colonists contain multiple synchronized species of the same life cycle type. viewed periodical cicadas with a mixture of religious apprehen- Broods are one of the more puzzling aspects of periodical ci- sion and loathing (Kritsky 2004); and modern Americans maintain cada biology. On one hand, broods have a kind of cohesiveness in numerous Web sites to assist in planning weddings, graduations, which local populations are bound together by a reliance on high and other outdoor activities around Magicicada emergences (e.g., cicadamania.com). Periodical cicadas have attracted the attention satiation (Dybas and Lloyd 1962, Lloyd and Dybas 1966a, Karban 1982,population Williams densities and Simon (several 1995). million On perthe acre)other tohand, effect broods predator can species), and Darwin, who commented on their unusual life cycles fragment and give rise to other broods, so that small isolated popu- of(quoted such scientificin Simon etluminaries al. 2000). as In Linnaeusthe future, (who periodical named cicadas one of may the lations separated from the main body of a brood may be relicts of become important bioindicators of ecological health and climate a previously larger brood distribution, or they may have arisen change, both natural and human-mediated (Reding and Guttman 1991, Clark 1992, Cooley et al. 2003, Yang 2004, Heckel and Keener Moore 1962, White and Lloyd 1979, Simon and Lloyd 1982, Kritsky 2007). These diverse ways of understanding periodical cicadas are andindependently Simon 1996). from Detailed a different information brood about(Young brood 1958, ranges, Alexander isolated and united by one common theme: all rely on accurate information about populations, and brood overlap can help clarify their origins and emergence timing and location. To facilitate and promote future biological interactions. Our understanding of broods developed from emergent patterns and public information that we solicited via the Internet. in records accumulated by early naturalists. By the 19th century, study, we present a detailed map of Brood X based on field mapping enough information existed that several authors developed maps and Background nomenclatural schemes for keeping track of broods. For example, C. - V. Riley (Riley 1885) compiled periodical cicada distribution records and presented a series of maps and schedules that could account Periodical cicada emergences in different regions are not syn chronized;106 different populations comprise the 15 largely parapatric American Entomologist • Summer 2009 for past emergences and accurately predict future emergences. Riley named each brood by assigning it a Roman numeral, but his maps, including Stannard’s maps of the Illinois broods (Stannard nomenclatural scheme did not clearly separate 13- and 17-year pe- 1975),There haveKritsky’s been 1987 efforts map to of Broodmake entirely X in Ohio new (Kritsky periodical 1988), cicada and riodical cicadas. This situation quickly led to confusion because the Zyla’s map of Brood XIX in Maryland (Zyla 2004), but no projects progression of emergence sequences of 17- versus 13-year broods have been attempted on the scale of whole-brood distributions, and disordered the numbering scheme. To address this problem, C. L. Marlatt (1923) presented a series devices and georeferencing software. So far, only published maps of of distribution maps and proposed a nomenclature for the broods Broodfew such VII efforts (Cooley have et al. relied 2004) on and recent partial advances maps ofin Broodinexpensive III (Irwin GPS that assigned separate numbering systems to the 13- and 17-year and Coelho 2000) and Brood X (Edwards et al. 2005) consist entirely populations. Marlatt designated 17-year broods with Roman numer- of newly acquired, georeferenced records. als I–XVII, and 13-year broods XVIII–XXX. Marlatt’s nomenclature is In this article, we present maps of the 2004 emergence of periodi- the basis for all subsequent published periodical cicada brood maps cal cicada Brood X, among the largest, by geographical extent, of all (e.g., Simon 1988). 17-year periodical cicada broods. Earlier maps show that this brood Published periodical cicada distribution maps have uses beyond is divided into three main regions and bordered by several other simply cataloguing and predicting emergences. For example, peri- broods (Fig. 1). Our maps are based exclusively on more than 8,000 odical cicada distributions allow some inferences about Pleistocene positive (present) and negative (absent) georeferenced records that glacial cycles because these insects are so closely associated with are available to the public (searchable database: http://hydrodictyon. eastern deciduous forests (Alexander and Moore 1962, Lloyd and eeb.uconn.edu/projects/cicada/). These records are of two types: Dybas 1966b, Cox and Carlton 1988, Cox 1992, Marshall et al. 2003). records (~45% of records) were collected by the authors and their distribution maps; some brood ranges have contracted noticeably research“field-verified groups. records” We made and no “unverifiedattempt to search records.” the entireField-verified possible Contemporarywithin historical climate times (Youngand habitat 1958, change Simon may1988, also Cooley be reflected et al. 2004, in distribution of Brood X uniformly. Rather, each research group con- Nelson 2004). At least two small broods have become extinct: Brood centrated on mapping either general distributions or brood edges in XXI in the 19th century (Marlatt 1923), and Brood XI between 1954 and 1971 (Manter 1974). Maps may help reveal the evolutionary implications of disjunct specific areas. The Edwards group collected distribution records in brood patches, such as V, IX, and X on Long Island; XIV in New Eng- be independently derived “parallel broods” not related by a common land;temporal VI in originWisconsin; to the or main XXIII inbody DeWitt of the County, brood IL—all (Simon of whichand Lloyd may 1982). Maps have revealed aspects of periodical cicada biology that bear on processes of speciation in the group, such as the pattern of reproductive character displacement between M. neotredecim and M. tredecim (Marshall and Cooley 2000). Finally, distributional infor- mation provides important tests of hypotheses about the formation of broods and species. Some of these hypotheses invoke patterns of spatial and temporal separation of broods of the same life cycle (Lloyd and Aspinwall 1975), or possible replacement of 17-year cicadas by 13-yearand Dybas cicadas 1966b), (Lloyd overlap and Whiteof broods 1976, of Coxdifferent and Carlton life cycles 1991). (Bryce For all their uses, Marlatt’s maps and their derivatives have Fig. 1. Composite map of all extant periodical cicada broods, adapted limitations. Most records used in these maps were summarized from maps published in Marlatt (1923) and Simon (1988). Base map is the original Marlatt base map. Individual brood records in Marlatt and and mapped by county, which limits the resolution of brood edges. Simon maps were traced on this map and then modified by removing or Other records used in these maps appear to be incorrect, resulting adding records on the basis of unpublished data. For clarity, dot size has from confusion with morphologically similar cicadas of the genus been reduced. Okanagana (e.g., records in Maine or Canada). Marlatt’s maps are also cross-generational: the records for all of the years pertaining Pennsylvania (Edwards et al. 2005); the Kritsky group surveyed dis- to a given brood (e.g., 2008, 1991, 1974) are combined in a single tributions in southern Ohio; the UConn group mapped brood edges in southern Illinois and Indiana; and the Zyla group mapped the limits of Brood X in Delaware, Maryland, Virginia, and West Virginia. To col- bmap,rood making (Marshall it difficult 2001). to For identify all of these records reasons, that are Marlatt’s best explained maps tend by lect records, we searched within the known distribution of the brood tooff-cycle overestimate (“straggler”) periodical emergences cicada brood from ranges an adjacent (Maier 1985,or overlapping Marshall and in adjacent areas for physical evidence of cicadas (emerging 2001). Recent revisions have helped correct some of these problems nymphs, cast skins, adults, etc.). We also listened for singing cicadas (Simon 1988), but even so, uncertainties associated with these maps by driving slowly (<40 mph) along roads with car windows open. hamper resolution of important questions about periodical cicada We obtained negative records by listening
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