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Nepidae (Hemiptera) of the United States and Canada

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Nepidae (Hemiptera) of the United States and Canada See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/233593939 Nepidae (Hemiptera) of the United States and Canada Article in Annals of the Entomological Society of America · January 1994 DOI: 10.1093/aesa/87.1.27 CITATIONS READS 11 279 2 authors, including: Robert W. Sites University of Missouri 167 PUBLICATIONS 643 CITATIONS SEE PROFILE Available from: Robert W. Sites Retrieved on: 26 October 2016 SYSTEMATICS Nepidae (Hemiptera) of the United States and Canada ROBERT W. SITES1 AND JOHN T. POLHEMUS2 Ann. Entomol. Soc. Am. 87(1): 027-042 (1994) ABSTRACT Seventy-two years have elapsed since the most recent synoptic treatment of North American Nepidae. This paper updates our present knowledge and brings together information on the family from various sources. Notes on the biology and ecology of Nepidae, a redescription of Ranatra texana Hungerford, and an illustrated key to the 13 species of Nepidae of the United States and Canada are provided. Range extensions, distribution maps, diagnoses, and synonymies also are given. KEY WORDS Nepidae, waterscorpion, key THE KNOWN WATERSCORPION fauna of the con- Curicta remained essentially unstudied until tinental United States and Canada comprises 13 Keffer (1991). extant species in three genera (Polhemus 1988), Since the first natural history of American ne- two fossil Ranatra species from the McKittrick pids (Uhler 1884), the literature concerning biol- Tar Pits (Pierce 1948, Miller 1983), and one fossil ogy and ecology of North American Nepidae has Nepa species from Florissant shale beds in Col- been summarized or reported by various authors orado (Hungerford 1932). Of the extant fauna, the (e.g., Hungerford 1920, Torre-Bueno 1923, Rad- genus Nepa is monotypic in North America and inovsky 1964, Bobb 1974, Weissmann 1986), and occurs throughout the eastern United States and Menke presented an excellent review in 1979. Canada and into the midwest; Curicta is repre- More recently, several life history papers (Pack- sented by two species in the south and south- auskas & McPherson 1986, McPherson & Pack- west; and Ranatra by 10 species with ranges that auskas 1987), a series of papers concerning eco- vary from an isolated limestone sink in Arizona logical aspects of R. montezuma (specifically (R. montezuma Polhemus) to transcontinental nocturnal planktonic behavior [Blinn et al. (R. fusca Palisot de Beauvois). 1982], parameters pertaining to the eggs [Blinn The Nepidae were among the first aquatic & Runck 1989], population dynamics and sec- bugs studied by early naturalists. The first fig- ondary production [Runck & Blinn 1990], forag- ures of nepids we have found are two excellent ing ecology [Runck & Blinn 1992], and the im- figures of Ranatra by Aldrovandi (1602, repeated pact of prey behavior and density on foraging in 1638) and crude woodcuts by Moufet (1634). strategy [Blinn et al. 1993]), and several distribu- Frisch (1728) provided good figures of both Nepa tion notes (e.g., DuBois 1978, Cochran et al. and Ranatra; however, Swammerdam (who died 1992) have been added to the literature. in 1680) gave us the first excellent anatomical Much of the literature dealing with North studies and drawings of these two genera in his American Nepidae has focused on taxonomy. posthumous Biblia Naturae (1737-1738). Since Hungerford (1922) contributed the first major then, the morphology of Ranatra has been stud- synoptic treatment of North American Nepidae, ied by several workers, most notably Dufour which included biological information, keys to (1821, 1833; both Nepa and Ranatra), Locy species, descriptions of new species, and correc- (1884), Marshall & Severin (1904), and Neiswan- tions of nomenclatural errors in Kirkaldy & Torre der (1925), and more specialized studies by Par- Bueno (1909). Kuitert (1947) revised the Ameri- sons (1972, 1974). The morphology of Nepa has can Nepidae, but only fragments of this disserta- been more intensively studied, with contribu- tion were published (Kuitert 1949a,b). Subse- tions by a number of scientists reviewed in the quently, several emendations to the North excellent work of Hamilton (1931). Other than American fauna have been published. Specifi- brief notes and several figures in Hungerford cally, R. annulipes Stal has been removed be- (1922) and Wiley (1922,1924), the morphology of cause it is neotropical (Polhemus 1988); and R. spatulata, described by Kuitert (1949a) from a Key West, FL, specimen, was considered by Pol- hemus (1976) to be a mislabeled African speci- 1 Wilbur R. Enns Entomology Museum, Department of En- men. Ranatra texana was described by Hunger- tomology, University of Missouri, Columbia, MO 65211. ford (1930) from Texas on the basis of only one 2 University of Colorado Museum, 3115 S. York, Englevvood, CO 80010. male specimen, and R. montezuma was de- 0013-8746/94/0027-0042302.00/0 © 1994 Entomological Society of America 28 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 87, no. 1 scribed by Polhemus (1976) from Arizona. More 1911a, Lindeman 1941) and under leaves or recently, Stys & Jansson (1988) presented a stones in shallow water (Uhler 1884, Torre- world checklist of all Nepomorpha (at and above Bueno 1923); DuBois (1978) found Curicta the generic level), and Polhemus (1988) pre- clinging to the side of a submerged log in a sented all synonymies, provided a brief discus- stream. Long series of both genera have been sion of biology, and listed published state and collected, often by hand, along the very shallow province records for the Nepidae of the United edges of streams and ponds, where the bugs States and Canada. blend with leaf litter. Specimens of Curicta and Presented herein are notes on the biology and Nepa are difficult to collect using traditional ecology of Nepidae; a redescription of R. texana; aquatic insect net techniques. Although Torre an annotated list of species; distributions includ- Bueno (1905b) recommended sweeping a net ing range extensions; and an updated, illustrated over these insects several times to disturb and key to the United States and Canadian species of make them float into it, moving a stick laterally, Nepidae. held lengthwise against the mud substratum, will usually dislodge the waterscorpions, and they are then easily noticed as they attempt to Biology and Ecology regain their original position. The immature stages of Nepa and Ranatra Ranatra is found among aquatic vegetation, have been the subject of many studies: Hinton debris, and overhanging vegetation that extends (1961, 1970) reported on egg structure and func- into the water, where it is most easily collected tion, Davis (1961) described the hatching pro- by sweeping an aquatic net vigorously through cess, Holmes (1907) reported on various aspects the vegetation. Ranatra sometimes occurs in of Ranatra nymphs, Hoffmann (1925) showed deep water, such as swimming pools or large that Nepa has only four instars, and Cloarec stock tanks, and has been seen swimming awk- (1976) demonstrated the importance of mechano- wardly. If emergent vegetation such as reeds or reception in predation behavior. Packauskas & cattails is present, Ranatra clings to it, moving McPherson (1986) and McPherson & Packauskas down the stems to hide during the day and re- (1987) provided descriptions for R. fusca and N. turning to the surface during the night. Nepids apiculata, respectively; the latter oviposits in possess "static sense organs" that keep them ori- mud. Oviposition is endophytic for many species ented correctly in the water (Baunacke 1912, of Ranatra, including R. montezuma (Blinn & Thorpe & Crisp 1947). Because nepids exhibit Runck 1989) and R. quadridentata Stal (Holmes death-feigning, they appear very sticklike and 1907), although Hoffmann (1930) stated that R. can be difficult to detect when collecting. The chinensis Mayr oviposits in mud banks, and propensity for nepids to undergo death-feigning Packauskas & McPherson (1986) reported that R. is well documented for both Nepa (Hamilton fusca oviposits in mud banks devoid of vegeta- 1931) and Ranatra (Holmes 1906; Severin & tion. Severin 1911a,b; Abbott 1940; Larson 1949a,b). Adults of most New World species of Ranatra After several minutes in a dry situation, they possess a well-developed stridulatory mecha- begin moving in an apparent attempt to locate nism, consisting of serrations on the fore coxal water and are then most easily separated from cavity that contact coxal ridges, which appear to plant debris. Alternatively, by placing the be sclerotized setae (Torre Bueno 1905a). The debris back into the water, much plant material only New World species known to lack stridula- will sink, whereas specimens of Ranatra usually tory structures are R. texana, R. kirkaldyi Torre will float and begin to move, and are easily de- tected. Bueno, R. signoreti Montandon, and R. parvula Kuitert. Further, stridulatory structures are ab- Cloarec (1986 and references therein) exten- sent in all Old World species examined. Nymphs sively studied many aspects of Ranatra develop- of Ranatra also possess the stridulatory mecha- ment, behavior, and prey capture. Blois & Cloarec nism, a rare occurrence in aquatic insects (Aiken (1983) demonstrated a density-dependent prefer- 1985) and unknown in any other aquatic het- ence for prey size. Prey may be taken from the eropteran. Indeed, the possession of a stridula- water surface (Radinovsky 1964, Weissmann tory apparatus in both nymphs and adults may be 1986); J.T.P. (unpublished data) observed a spec- unique
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