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 information 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 off-cycleto 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 or searching at locations brood and life cycle evolution. Any study making use of older maps, (with no vehicle engine running) for a minimum of 2 minutes. We or comparing older maps with newer maps, cannot dismiss the collected locality information by using handheld GPS units, writing - detailed location descriptions, or marking highly detailed road maps ing criteria for collecting and evaluating records (Marshall 2001). and digitizing the latter two types of record using GIS or geocoding possibility that apparent changes in distribution may reflect chang American Entomologist • Volume 55, Number 2 107 software packages (Arcview 3.0, Street Atlas, Google Earth). At some locations, we also recorded choruses using a Marantz digital recorder or a Sony DAT recorder sampling at 48 kHz. dpositiveo not attempt records to(light delineate green); brood verified boundaries negative records by drawing (dark closed gray); and unverified negative records (open circles). Like Marlatt, we - in rUnverifiedesponse to mediarecords solicitations consist of in observations the Washington, submitted DC, Cincinnati, by the areas around data points. Because we concentrated our efforts on publicOH, and through Allentown, specific PA, regions. Web sites, Records e-mails, submitted or other by communications the public con- only some locations, our verified records do not include many loca sisted of locality information (a street address, intersection, public stionsample where the range unverified of Brood records X. For suggest these reasons, cicadas this were map present. should It not is park, etc.) and comments. The comments sections of the records were bealso considered not possible exhaustive, to judge althoughhow completely it may be the used unverified to estimate records the examined, and records that described the emergences incorrectly brood’s boundaries. (e.g., wrong body colors, nocturnal choruses, wrong time of year) were discarded (~3% of all records). Locality information was used to geocode the remaining records are Severalopen to doubt,assumptions clusters are of recordsnecessary validate to interpret each other the and unverified suggest records in our map. First, even though individual unverified records to the Google Maps API geocoder (~7% of records were geocoded this saturation and the number of responses, it is unlikely that there were way).in the Recordsfollowing that manner: could notthe recordsbe geocoded were in first this submitted way were in geocoded batches alarge, higher unnoticed degree of populations confidence. nearLikewise, the metropolitan given the degree areas of wheremedia by hand, using Google Earth (~37% of records). Records that did media coverage was greatest. Thus, the total absence of records from not describe unique locations or that could not be geocoded (~8% a given area within one of these well-sampled regions suggests that of all records) were discarded. All geocoded records were placed as a data layer in an Arc GIS 9.2 map, and records that fell far outside make inferences about brood boundaries or range changes, but we the known boundaries of the brood were discarded, as were records haveno cicadas included were them present. because Unverified they provide records sampling should in not areas be thatused the to whose geocoded location fell outside the state or county described mapping teams could not investigate, and they will serve as a guide in the record submission (<5% of all records). All remaining records were entered into a database (http://hydrodictyon.eeb.uconn.edu/ Given these assumptions, this new map, like earlier maps, sug- projects/cicada/) and used as layers in an Arc GIS 9.2 map. gtoe stsfuture three mapping main divisionsefforts. in Brood X: a Midwestern division in western Ohio, southern Michigan, Indiana, and eastern Illinois; a Results southern division in eastern Tennessee, western North Carolina, and northern Georgia; and an eastern division concentrated in Washington, DC, Pennsylvania, northern Maryland, northern Dela- Our map (Fig. 2) includes field-verified positive records (dark green), in which we have a high degree of confidence; unverified

Fig. 2. Map of records collected during the 2004 emergence. Positive records collected or verified by the authors are shown in dark green; unverified positive records collected from the general public are shown in light green. Verified negative records are shown in dark gray and unverified negative records are shown as open circles. Base map is USGS National Land Cover Database (NLCD) 2001 30m resolution forest canopy (Homer et al. 2004, Homer et al. 2007)

108 American Entomologist • Summer 2009 Fig. 3. Verified positive records of low-density populations (blue symbols) and verified positive records (green symbols) found in Kentucky during the 2004 emergence. At each record location, on days and at times appropriate for chorusing activity, only single individuals or a handful of individuals were heard, and no choruses were heard. We made no attempt to survey the entire periphery, so other low-density populations may exist elsewhere.

ware, northern Virginia, and eastern West Virginia (Fig. 2). Brood X includes several metropolitan areas, such as Princeton, NJ; Wash- some limited predator immunity. ington, DC; Philadelphia; Indianapolis; Cincinnati; and a scattering becauseThis phenomenon their synchronization of low-density, with an peripheral existing broodpopulations affords may them be of records on Long Island, NY. We received no reports from west of the Mississippi River. al. 1994). It may also be a source of confusion about brood boundaries In the eastern United States, the 2004 Brood X map (Fig. 2) gener- anbecause important stragglers avenue from of gene one flowbrood between may be two mistaken broods for (Heliövaara low-density et ally resembles maps such as those published by Marlatt and Simon populations of another brood (Edwards et al. 2005). There is no way (Fig. 1), although the new map suggests a smaller range for Brood yet known to determine the age of an adult periodical cicada, nor X in the East. Brood X does not appear to reach as far north in most of Pennsylvania as suggested by Marlatt’s county-level dots, nor is cycle, so that where two broods are adjacent, unambiguously as- the coverage of New Jersey as extensive as suggested by the earlier aresigning there brood known membership genetic differences to some border among populations broods of the may same not lifebe records. In at least one area, the new map suggests a larger range possible. We include records of low-density populations in our map than was reported historically. The southwestern edge of Brood X of the 2004 Brood X emergence with the caveat that brood member- in Pennsylvania was comparatively well sampled with positive and ship of low-density, peripheral populations is an open question, and negative records, and it appears to extend farther west than the old with the hope that these records will be carefully examined during records, into central Somerset County. The records from Garrett upcoming emergences of Broods XIV, II, V, and VI or other brood County, MD, also appear to extend the range, or at least resolve it pairs with shared boundaries (e.g., I/V, II/VI, V/IX). Older brood maps (Fig. 1) suggest that many broods inhabit Armstrong County, PA, should be checked in future emergences. - with better precision. The unverified public report from southern lapping, discounting the possibility that broods are fragmented, inter- range changes or are artifacts of map-making techniques (e.g., we well-defineddigitating, or regions.that some Older enclose maps isolated also tend populations to portray of broods other asbroods. over Whetherused smaller the dotsdifferences on our map between and did old not and lump new records maps byare county) due to is open to question. - deThus,tailed the records spatial are relationships needed to determineof many neighboring whether these broods—such broods are ingly low density for Magicicada, sometimes so low that they consist fragmented,as II, VI, V, VI, whether IX, X and or XIV—are not they notoverlap well locallyunderstood in some (Fig. regions, 1). More or of scatteredSome of the individuals field-verified rather records than full are choruses of populations (Fig. 3). at Somesurpris of whether the apparent confusion is mainly a consequence of accu- these records, such as those in northern Kentucky, are found in mulated records of stragglers (Lloyd and White 1976, Maier 1982, locations where Brood X and XIV border each other, and some are Marshall 2001). Detailed, Internet-based surveys of the public, com- fully within the known range of Brood XIV (Lloyd and White 1976; Cooley et al. unpublished data). Such low-density populations could rare, isolated populations and resolve complex brood boundaries. In belong to either Brood X or XIV because periodical cicadas occasion- thebined Baltimore–Washington, with efforts to verify DC,key region, records, the have public the records potential from to 2004detect they show that the closely spaced county-level dots from earlier maps tend to do so in increments of 1 or 4 years, with 4-year advance correspond to an almost continuous band of populations, saturated emergencesally emerge off-schedule.especially common When among periodical 17-year cicadas species accelerate, (Lloyd and even at the scale of Fig. 4. Similarly, the complete absence of negative Dybas 1966b, Simon et al. 1981, Simon and Lloyd 1982, Kritsky public records from large areas of south-central Pennsylvania and 1987). In some circumstances, repeated instances of straggling may central Maryland, combined with evenly spaced positive records, allow low-density populations to exist at the periphery of a brood. suggests that Brood X is also widespread in that region and not inter- For example, where two adjacent broods are separated by 4 years, rupted by populations of other broods. At the same time, clusters of early stragglers from the later brood will be synchronized with negative records suggest that populations of other broods may be the earlier brood, and straggler populations may be able to persist isolated within the range of Brood X (e.g., York and Adams Counties,

American Entomologist • Volume 55, Number 2 109 Fig. 4. Detail of 2004 Brood X emergence in the eastern United States. Verified positive records are shown in dark green; unverified positive records collected from the public are shown in light green. Verified negative records are shown in dark gray, and unverified negative records are shown as open circles.

PA). Revised maps of Broods XIV and II may show that these other Maryland localities for 13-year Brood XIX (Fig. 2; Zyla 2004). In broods completely surround isolated populations of Brood X. Once southern Indiana, Broods X and XXIII have a complex microparapatric relationship, including at least one area of slight overlap in the same events or other processes that lead to temporal isolation and brood woods; in Illinois, Brood X does not currently contact Broods XIX or formationclarified, these in periodical patterns cicadas. may be used to understand the climatic XXIII, nor does it contact 17-year Brood XIII (unpublished data). Our data add some information to ongoing discussions of contact Our map is a single-generation map of Brood X as it emerged in 2004. We urge caution in making inferences about range changes major mitochondrial lineages in the “decim” species of periodical - cicada,zones between and contact broods between with different these lineages life cycles. is of There particular are two interest known though range contraction could explain some discrepancies between to theories of life cycle switching and species formation (Cox and ourbased map on and differences earlier maps, between other our explanations map and other include published the incomplete maps. Al Carlton 1988, 2003; Lloyd et al. 1983; Marshall and Cooley 2000; sampling of our map, the lower precision of earlier maps, and the Cooley et al. 2001; Simon et al. 2000 Marshall et al. 2003). Lineage tendency of cross-generational data sets to accumulate erroneous “A” is found in M. neotredecim and M. septendecim, and lineage “B” is records (Marshall 2001). Although this map, by itself, cannot resolve found only in M. tredecim (Martin and Simon 1988, 1990; Cooley et past questions about range change, it will be useful as a baseline al. 2001). Where M. neotredecim and M. tredecim co-occur in Broods map for future emergences of Brood X and all adjacent broods. Any XIX and XXIII, these lineages are in contact and display positive assortative mating and a striking pattern of reproductive character displacement in M. neotredecim calling song pitch (Marshall and bodifferencesundary with between positive a andmap negativebased on records,the 2021 would emergence be strong and evi-this Cooley 2000, Simon et al. 2000, Cooley et al. 2006). d2004ence map, for range especially changes. in areas where we have identified the brood This pattern suggests that these lineages have a history of selection against interbreeding. The two mitochondrial lineages also come into Acknowledgments contact wherever M. septendecim in Brood X (or Broods II, VI, IX, or Steve Chiswell, Richard Crist III, Amy Faivre, Andrew Martin, XIV) is in contact with M. tredecim (Broods XIX or XXIII), although co- emergences occur only once every 221 years. One hypothesis is that there are undiscovered patterns of reproductive character displace- WMargarete also th Pfiester,ank the Michael owners Sitvarin, of our researchJessee Smith, sites. Paige This Swientisky, work was ment along this contact zone; however, it is also possible that co-emer- madeJennifer possible Webb, byand a NationalZachary ZylaGeographic provided grant assistance to JC, a NASAin the Grantfield. NAG5-12416 to Muhlenberg College, Grants to GK from The College signals. Where we mapped Illinois and Indiana in detail, we found of Mount St. Joseph, and National Science Foundation Grants. DEB gencesno evidence are too of reproductiverare to cause charactersufficient divergentdisplacement selection within on Brood mating X; 99-82039, NSF DEB 04-22386, DEB 05-29679, DEB 06-19012 (REU), across its range, M. septendecim chorus pitch from 2004 Brood X was between 1.29 and 1.37, well within the 1.25–1.50 kHz range reported recommendations expressed in this material are those of the authors for this species (Fig. 5; Marshall and Cooley 2000). Our unpublished and DEB 07-20664 to CS. Any opinions, findings, and conclusions or data suggest that overlap between Brood X and the 13-year broods dedicate this paper to the memory of her father Walter W. Simon in is minimal. In Maryland, no Brood X emergences were reported in St. andrecognition do not necessarily for his help reflect in collecting the views many of broodsthe NSF. of CS cicadas would and like for to Mary’s County, which contains 17-year Brood II and the only known Magicicada when she was 12 years old.

110 first introducing her to American Entomologist • Summer 2009 Fig. 5. Verified positive records where M. septendecim chorus recordings were taken and analyzed. At each location, M. septendecim choruses were recorded with a Marantz digital recorder or a Sony DAT recorder sampling at 48 kHz. Recordings were not filtered, but background noise (pops, traffic, voices, etc.) was edited out, and 1–2-min samples were analyzed using Canary 1.2 (Cornell Bioacoustics Laboratory, Ithaca, NY). The dominant frequencies (pitch- es) of representative locations are shown in kHz.

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American Entomologist • Volume 55, Number 2 111 Martin, A., and C. Simon. 1988. Anomalous distribution of nuclear and mi- White, J., and M. Lloyd. 1979. 17-Year cicadas emerging after 18 years: a tochondrial DNA markers in periodical cicadas. Nature 336: 237–239. new brood? Evolution 33: 1193–1199. Martin, A., and C. Simon. 1990. Williams, K. S., and C. Simon. 1995. The ecology, behavior, and evolution divergence in the mitochondrial DNA of periodical cicadas related to of periodical cicadas. Annu. Rev. Entomol. 40: 269–295. historical biogeography. Evolution Differing 44: 1066–1080. levels of among-population Yang, L. H. 2004. Periodical cicadas as resource pulses in North American Nelson, B. 2004. Fearing the worst for cicada brood. Pp. A2. Newsday, forests. Science 306: 1565–1567. New York. Young, F. N. 1958. Some facts and theories about the broods and periodicity Reding, M. E., and S. I. Guttman. 1991. Radionuclide contamination as an of the periodical cicadas. Proc. Ind. Acad. Sci. 68: 164–170. Zyla, J. D. 2004. First report of the 13-year periodical cicada, Magicicada (Magicicada cassini) populations. Am. Midl. Nat. 126: 322–337. tredecim (Walsh and Riley) (Hemiptera: Cicadidae) in Maryland. Proc. Riley,influence C. V. 1885. on the The morphology periodical andcicada. genetic An account structure of Cicadaof periodical septendecim cicada Entomol. Soc. Wash. 106: 485–487. and its tredecim race, with a chronology of all broods known. Bull. U.S. Dept. Agric. Div. Entomol. 8: 1–46. Simon, C. 1988. Evolution of 13- and 17-year periodical cicadas. Bull. John R. Cooley, Lecturer at Yale University, studies the behavior and evolu- Entomol. Soc. Am. 34: 163–176. tion of cicadas. Current information about the cicada mapping project may Simon, C., and M. Lloyd. 1982. Disjunct synchronic population of 17-year be found at www.magicicada.org. Gene Kritsky, a Professor at the College periodical cicadas: Relicts or evidence of polyphyly? J. N.Y. Entomol. of Mount St. Joseph, has written a number of books and papers about pe- Soc. 110: 275–301. riodical cicadas. Marten Edwards is a mosquito physiologist and teaches Simon, C., R. Karban, and M. Lloyd. 1981. Patchiness, density, and aggrega- entomology as an Associate Professor at Muhlenberg College. John D. Zyla tive behavior in sympatric allochronic populations of 17-year cicadas. studies both periodical and annual cicada distribution in the Mid Atlantic Ecology 62: 1525–1535. states and runs the Mid-Atlantic Cicadas website (www.cicadas.info). David Simon, C., J. Tang, S. Dalwadi, G. Staley, J. Deniega, and T. R. Unnasch. C. Marshall is a postdoctoral associate in the Simon Lab at the University of 2000. Genetic evidence for assortative mating between 13-year cicadas Connecticut (UConn) studying the behavior and evolution of singing insects. and sympatric “17-year cicadas with 13-year life cycles” provides sup- Kathy B. R. Hill studies phylogenetic relationships of world cicada species port for allochronic speciation. Evolution 54: 1326–1336. and cicada behaviur in the Simon Lab at UConn. Rachel Krauss studied Stannard, L. J. 1975. The distribution of periodical cicadas in Illinois. Ill. cicadas in the Simon Lab as part of her BSMS degree at UConn. Chris Simon, Nat. Hist. Surv. Biol. Notes 91: 3–12. a Professor at UConn, studies molecular systematics and evolution of cicadas White, J. 1980. Resource partitioning by ovipositing cicadas. Am. Nat. worldwide and uses cicadas as model organisms to study the origin, spread, 115: 1–28. and maintenance of biodiversity.

Buzzwords (Continued from page 69) 15,000 Colorado potato beetles onto German References Askew, R.R. 1971. Parasitic Insects. New York: discussion, but the consensus is that stridu- supplies (leading to the establishment of a American Elsevier Publishing Company. Elmore, J.B. 1908. Love Among the Mistletoe, lation is a defense against predators. At least potatoKartoffelkaferabwehrdienst fields to destabilize ( PGermanotato Beetle food and Poems. Alamo (IN): J.B. Elmore. one competing hypothesis, however, led to Defence Service) in Kruft), and by the gov- Garrett. B.C. 1996. The Colorado potato beetle yet another appearance of kissing bugs in ernment of the German Democratic Republic goes to war. Historical note no. 2. Chemical popular culture (Askew 1971). According in 1950 of actually dropping thousands of Weapons Convention Bulletin 33: 2-3. to a front page story in the U.K. Guardian Colorado potato beetles in the southwest Howard, L.O. 1899. Spider bites and kissing bugs. Popular Science Monthly 56: 31-42. that ran on June 7 1966, the United States part of the country to demoralize a nation Howard, L.O. 1899. Insects to Which the Name Pentagon was reported to be “planning to fond of its potato dumplings (Garrett 1996). Kissing Bug Became Applied in the Sum- send bed-bugs [sic] to help to win the war For one thing, most estimates of the distance mer of 1899, United States Department of in Vietnam...Their plans are based on the from which blood-sucking bugs can detect Agriculture Division of Entomology Bulletin fact that bed-bugs scream with excitement a human blood meal are in the range of 10 22: 24-30. Lyman, H.H. 1899. The President’s annual to 20 feet, not 200 yards; moreover, it’s address. Thirtieth Annual Report of the unclear how or why bugs would ignore a Entomological Society of Ontario, Toronto: enat ablethe prospect the GI, sweating of feasting through on human the jungles flesh potential blood meal almost underfoot in Warwick Pros & Rutter, pp. 21-31. October of[sic!] South ...a soundVietnam, amplification to hear the system anticipatory would 11-12 1899. squeals of a captive bed-bug as it detects the away, and there’s no evidence that bugs Moncayo, A. 2003. Chagas disease: current preference to a meal two football fields epidemiological trends after the interruption Vietcong lying in ambush ahead. Tests have of vectorial and transfusional transmission apparently shown that a large and hungry forces. Finally, what adaptive value there in the Southern Cone countries. Mem. Inst. bed-bug will appropriately register the pres- canmight differentiate be to a bug of between announcing allies its and presence hostile Oswaldo Cruz 98: 577-591. ence of a man some two hundred yards to by squealing in anticipation while stalking Schofield, C.J. 1977. Sound production in some its front or side, while ignoring the person its prey is a mystery; it would seem such triatomine bugs. Physiol. Entomol. 2: 43-52. carrying it in a specially devised capsule” squeals are completely inconsistent with May Berenbaum is a pro- (The Guardian 7 June 1966). the stealth strategy displayed by the group fessor and head of the De- There are so many biological improb- as a whole. If nothing else, any Viet Cong partment of Entomology, abilities in the Guardian account that it’s within earshot would know of the presence University of Illinois, 320 hard to believe that such a proposal was ever of an American soldier immediately upon Morrill Hall, 505 South seriously considered by the Department of hearing the squeal. It would seem, then, that Goodwin Avenue, Urbana, IL 61801. Currently, she is studying the chemical as- same War Department was accused by the instead be a signal for the American soldier pects of interaction between herbivorous insects Defense—although it must be noted that this the amplified scream of a kissing bug could Nazi government in 1942 of plotting to drop to kiss his butt goodbye. 7 and their hosts.

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