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Surprising Northern Disjunct Population of 13-Year Cicada Brood XXII (Hemiptera: Cicadidae, Magicicada)

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Surprising Northern Disjunct Population of 13-Year Cicada Brood XXII (Hemiptera: Cicadidae, Magicicada) Evolution and Geographic Extent of a Surprising Northern Disjunct Population of 13-Year Cicada Brood XXII (Hemiptera: Cicadidae, Magicicada) GENE KRITSKY, ROY TROUTMAN, DAN MOZGAI, CHRIS SIMON, STEPHEN M. CHISWELL, SATOSHI KAKISHIMA, TEIJI SOTA, JIN YOSHIMURA, AND JOHN R. COOLEY ntil now, periodical cicada Brood XXII was recog- along the lower Mississippi Valley in eastern Louisiana nized as having the most southern distribution of and southwestern Mississippi. Uthe three extant Magicicada 13-year broods (XIX, It was therefore quite surprising, so many years later, XXII, XXIII). Phares (1845) recorded the 1845 Brood XXII to discover an unrecognized large northern disjunct emergence in The Woodville Republican, a Mississippi population of Brood XXII (Kritsky 2004) in a populated newspaper; in addition, he had per- area along the Ohio River, separated by sonally witnessed the cicadas’ 1832 more than 1,000 km from the record- appearance and received verbal Magicicada Primer ed range of Brood XXII in the lower accounts from observers of the 1819 Species: Mississippi Valley. Geographic dis- and 1806 emergences. He also pub- Decim group: juncts are common in 17-year cicadas lished details of the 1858 emergence, Magicicada septendecim (L.) (Simon and Lloyd 1982, Cooley 2015, leaving no doubt as to the 13-year life M. tredecim (Walsh & Riley) Cooley et al. 2015) and they have also cycle of this brood (Kritsky 2004). M. neotredecim Marshall & Cooley been noted in 13-year cicadas (e.g., After the 1871 emergence of Brood Cassini group: populations of Brood XXIII in Dewitt XXII, Phares sent specimens to C. V. County Illinois separated by more M. cassini (Fisher) Riley (Riley 1885) along with a record M. tredecassini Alexander & Moore than 250 km from the main range of of the timing of emergence events. All the brood; Lloyd et al. 1983, Cooley et Decula group: of these 19th-century observations al. 2006), but none had been reported were summarized by Marlatt (1907), M. septendecula Alexander & Moore for Brood XXII. M. tredecula Alexander & Moore who mapped all 13- and 17-year peri- There are three prevailing hypothe- odical cicada broods, having previ- Distribution: ses for the formation of disjunct pop- ously developed the brood nomencla- Largely co-occurring, east of the ulations within a single brood year. ture now in use (Marlatt 1902; I-XVII U.S. Great Plains Disjunct populations may form via a for 17-year broods and XVIII-XXX for Broods: one-time life cycle shift in which cica- 13-year broods with numbers I and Periodical year classes numbered das from a given brood come out early XVIII chosen arbitrarily). Brood XXII’s sequentially and identified solely by or late, usually in increments of four distribution, as mapped by Marlatt year of emergence years, to form a new population on (1907), appeared small and centered a schedule matching, by chance, the American Entomologist • Volume 63, Number 4 E-15 Downloaded from https://academic.oup.com/ae/article-abstract/63/4/E15/4713030 by guest on 17 December 2017 Here we present an entirely new GIS-based map of Brood XXII during its 2001 and 2014 emergences, with detail on all three species found in the brood (M. trede- cassini, M. tredecim, M. tredecula). We present genetic data documenting that M. tredecim and M. tredecassini in the Ohio Valley disjunct of Brood XXII have mtDNA haplotypes in common with their respective species in the lower Mississippi Valley portion of Brood XXII; thus, these populations appear to be relicts of a once-larger distribution and could not have been derived recently from neighboring 17-year cicadas. M. tredecula in the Ohio Valley disjunct are of more ambiguous origin. Methods Verified records.We created a new map of Brood XXII by methods similar to those used in constructing maps of Broods I (Cooley 2015), II (Cooley et al. 2015), III (Cooley et al. 2013a), VII (Cooley et al. 2004), X (Cooley et al. 2009), XI (Cooley et al. 2013b), XIII (Cooley et al. 2016), and XIV (Cooley et al. 2011). We searched for active periodical Fig. 1. Verified 13-year cicada Brood XXII presence (red symbols) and absence (gray cicadas, using record collection methods and criteria symbols) records collected between 8 May and 15 June 2014. included in Cooley et al. (2013a). We collected locality information with handheld GPS units (Garmin, various models) and a custom datalogger described in Cooley et al. (2013a). Distributional information was organized and mapped using ArcGIS 9.3 (ESRI 2009). Crowdsourced records from the general public. In 2014, the website www.magicicada.org collected unveri- fied (“crowdsourced”) periodical cicada sightings from the general public. Not all seemed credible, so we assigned a weight to each crowdsourced record using methods described in Cooley (2015). This weighting method assigns greater import to crowdsourced records that are near verified records or that are clustered. Genetic data. Mitochondrial cytochrome oxidase sub- unit I (COI) sequences of specimens from Ohio and Ken- tucky were determined following the methods described in Sota et al. (2013) and compared to existing haplotype information (ibid.). Details of specimens and accession numbers of the haplotype sequences are given in Table 1. Newly discovered haplotypes are deposited at DNA Data Bank of Japan (DDBJ; accession no. LC331667 - LC331670). Fig. 2. Records submitted to www.magiciada.org between 8 May and 15 June 2014 (orange symbols) scaled by size as described in (Cooley 2015). Red symbols indicate Results verified Brood XXII presence records. The year 2014 was also a 17-year cicada Brood III emergence year, and web records reflect the emergence of the brood in the upper Between 8 May and 15 June 2014, our field teams collected Midwest, although record weights are relatively low due to distance from verified 5,216 verified presence and absence records of Brood XXII 13-year cicada Brood XXII records. using automatic data loggers or hand-held GPS (Fig. 1); in 1975, 1988, and 2001, we mapped individual Brood schedule of an existing brood located elsewhere (e.g., Long XXII localities in Southern Mississippi and southeastern Island populations of Broods I, V, and IX; Simon and Lloyd Louisiana that also provide verification (unpublished 1982). Alternatively, disjuncts may be relicts of once-larg- notes). Between 8 May and 15 June 2014, the website er distributions (e.g., Maryland populations of Brood XIX; www.magicicada.org collected 992 records that were Marshall and Cooley 2000). A third possibility is that pop- located within the continental U.S. and that could be ulations of one life cycle may be relatively recently derived resolved to location (Fig. 2). In general, the crowdsourced from populations of the other life cycle via permanent records follow the distribution of Brood XXII indicated life-cycle shifts (e.g., Magicicada neotredecim, a 13-year by the verified records; however, since 2014 was a year in species found in northern parts of Broods XIX and XXIII which 17-year Brood III emerged in the upper Midwest that originated after the last glacial maximum from 17-year (Cooley et al. 2016), many of the crowdsourced records M. septendecim; Martin and Simon 1988, 1990; Marshall clearly belong to that brood. and Cooley 2000; Simon et al. 2000; Cooley et al. 2001). Our records show that Brood XXII consists of two E-16 American Entomologist • Winter 2017 Downloaded from https://academic.oup.com/ae/article-abstract/63/4/E15/4713030 by guest on 17 December 2017 Fig. 3. Detail of lower Mississippi Valley portion of 13-year cicada Fig. 4. Detail of Ohio River portion of 13-year cicada Brood XXII. Brood XXII. Brood XXII presence noted by colored symbols; Brood XXII presence (red symbols) and absence (gray symbols). black: Magicicada tredecassini; orange: M. tredecim; red: M. tre- decula. Absence records are shown by gray symbols. distinct parts: a southern part, in the lower Mississippi River Valley (Fig. 3), and a northern disjunct along the Ohio River southeast of Cincinnati, along the southern margins of Clermont and Brown Counties, OH, and in Campbell, Bracken, and Pendleton Counties, KY, extend- ing into Kenton, Grant, Harrison, Robertson, and Mason Counties, KY (Fig. 4). Three species, Magicicada tredecas- sini, M. tredecim, and M. tredecula are found in both por- tions of Brood XXII, though M. tredecassini appear to be the most abundant species in the Ohio Valley disjunct and the rarest species in the Lower Mississippi Valley portion of Brood XXII. The Ohio Valley disjunct occupies roughly 3,000 km2 or 1/5 the area of the main portion of Brood XXII, estimated at 16,778 km2 (Fig. 4); for comparison, this area is slightly larger than the 17-year cicada Brood I disjunct (2,600 km2) described in 2012 (Cooley 2015). The Ohio Valley disjunct of Brood XXII is remarkable in that it is entirely surrounded by 17-year Broods X and XIV, and it also substantially overlaps Brood XIV (Fig. 5). We obtained COI sequences for nine M. tredecim, 10 M. tredecassini, and one M. tredecula specimens (Table 1). All M. tredecim sequences belong to lineage B, which is the unique haplotype group of M. tredecim (Sota et al. 2013). All sequences from M. tredecassini specimens Fig. 5. Detail of Ohio River portion of Broods X, XIV, and XXII. belong to lineage Cw, which is shared by all Cassini-group Symbols indicate presence of 13-year cicada Brood XXII (red), 13-year cicadas and the western 17-year Cassini-group 17-year cicada Brood XIV (green), and Brood X (blue). Absence cicadas. The only M. tredecula sequence was identical to records for each brood are shown by gray symbols. the haplotype De (eastern Decula) and not Dm (middle Decula), as found in southern Brood XXII M.
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