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Nepidae () of the United States and Canada

Article in Annals of the Entomological Society of America · January 1994 DOI: 10.1093/aesa/87.1.27

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Robert W. Sites University of Missouri

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Available from: Robert W. Sites Retrieved on: 26 October 2016 SYSTEMATICS (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 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- 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 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- 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 . 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 (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 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 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 in aquatic insects. imen of R. fusca take a small, live fly from the Although nepids exhibit a preference for lentic surface, and Packauskas & McPherson (1986) re- situations, habitat selection may be generalized ported R. fusca preying on water striders. The for each of the three North American nepid gen- major dietary item of R. montezuma in Monte- era. Curicta and Nepa typically occur along zuma Well is an endemic amphipod, and, because muddy shorelines of spring-fed bodies of water, fish are absent in this closed ecosystem, R. mon- where their dark-colored, flattened bodies are tezuma is the ultimate consumer with a high sec- particularly well adapted for concealment in ondary production rate (Runck & Blinn 1992). mud. Nepa also is found in sparse to dense veg- Weissmann (1986) proposed the use of Ranatra etation (Torre Bueno 1905b, Severin & Severin for mosquito control; Hoffmann (1930) reared R. January 1994 SITES & POLHEMUS: NEPIDAE OF UNITED STATES AND CANADA 29 chinensis entirely on a diet of mosquitoes, but approximately 110 species worldwide were ex- noted that large mosquito larvae sometimes kill amined. S. L. Keffer has (in progress) an exhaus- early instars. Evaluations of the predation effi- tive study of the phylogeny of Nepidae based on ciency of Nepidae on mosquito larvae have detailed examination of the male genitalia of al- been provided by Ouda et al. (1986) and Chock- most all species worldwide. Preliminary results alingam & Somasundaram (1987). Extensive pre- of that study support the separation of the above dation on carp eggs by R. linearis L. in some species (and others) into a group separate from British ponds resulted in the recommendation the type species of Ranatra sensu strictu, R. lin- that the ponds be drained and restocked with fish earis; however, several other equivalent groups (Anonymous 1878). There are many other records must also be recognized, including one with R. of predation on fish (e.g., Ormerod 1878, Rao kirkaldyi along with some South American spe- 1976, Bisht & Das 1981) and tadpoles (Hoffmann cies. We believe the recognition of genus-group 1930). names for any of these entities before the com- An eyeless Nepa species, recently discovered pletion of Keffer's study is premature, because it in a Romanian cave, is the first known troglobite would either require proposing genus-group among the aquatic (Sarbu & Popa names at this time for all of Keffer's groups or 1992). In that closed cave system, there is no result in paraphyletic clades. Thus, although vegetation, and Nepa is the ultimate consumer. Lee's (1991) proposal has merit, a possible end Curicta, Nepa, and Ranatra disperse by noc- result of Keffer's studies is the establishment of turnal flights and colonize temporary habitats, equivalent groups within Ranatra without nam- although actual observations of flight are rare. ing them subgenera and in that event, the genus- It was long thought that Nepa was incapable group name Syoitia would be placed as a syn- of flight because specimens were found with onym of Ranatra. For these reasons, we treat degenerated flight muscles; however, Larsen Ranatra in this work as a single genus without (1949b) stated that functional flight musculature subordinate taxa. does exist, summarized flight observations for The general morphology of Nepidae has been Nepa, and illustrated how flight occurs in Rana- adequately treated and illustrated by Hunger- tra (see also Torre Bueno [1906]). Ranatra will ford (1922), and the male genitalia were dis- fly at night from an uncovered aquarium if a stick cussed in detail by Lansbury (1974b). We gener- is available for it to leave the water. Near Lake ally have followed the structural nomenclature P6tzcuaro in Mexico, a hotel swimming pool was of Hungerford (1922), the last definitive revision recolonized on successive nights by Curicta of North American species; however, for male scorpio Stal (J.T.P., unpublished data). Nepa also genitalia we follow the more modern terminol- is apparently able to estivate, burying and encas- ogy of Lansbury (1974a). As a result, two male ing itself in mud (Wood 1884, Larimore et al. genitalia terms of Hungerford (intromittent 1959). organ and clasper) are herein referred to as posterior diverticulum and paramere, respec- Taxonomy tively. A number of attempts have been made to key De Carlo (1967) divided Nepidae into two fam- at least part of the waterscorpion fauna of North ilies, the Nepidae and Ranatridae, and under the America (e.g., Blatchley 1926, Herring 1951b, latter proposed the subfamilies Curictinae and Usinger 1974), but none addressed the entire Ranatrinae (De Carlo 1968). This concept was fauna including R. texana and R. montezuma. De rejected by Menke & Stange (1964), Lansbury Carlo (1972) contributed a key to the New World (1974a), and Stys & Jansson (1988). We concur species of Ranatra (including R. texana but be- with Stys & Jansson that there is one family, fore the description of R. montezuma) (English Nepidae, with two subfamilies, Nepinae and translation of the couplets available from Ranatrinae, the former containing the tribes Cu- R.W.S.); however, the key is difficult and in some rictini and Nepini. Further, Keffer & McPherson instances uses relative characteristics. Lans- (1993) reported the copulatory posture of Curicta bury's (1972) key to Oriental Ranatra was pro- and concluded that nepid copulatory posture posed by Menke (1979) as a model to follow; may be of subfamilial significance. however, those characters have not been consis- Lee (1991) proposed Miyamotoa as a subgenus tently appropriate for the North American spe- of Ranatra based on characters of the male gen- cies. Other than for eggs (Hinton 1962), keys to italia; this is a preoccupied name, since replaced immature stages are lacking. by Syoitia (Lee 1992). The subgenus was pro- Waterscorpions are notorious for much in- posed to include three North American and one traspecific variability, particularly with regard to Mexican species ( Hungerford several key characters. For example, the lengths [type species], R. fusca [as americana Montan- of the metaxyphus and the projection of the pen- don], R. quadridentata, and R. operculata Kuit- ultimate antennal segment are quite variable. ert). Although Lee's study included both Old This variability led Bennett (1987) to suggest World and New World species, only seven of the hybridization, and he showed character clines 30 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 87, no. 1 for several species. Because of this variability, vated as tylus (Fig. 4); Arizona, Mexico occasional individuals will not key correctly; Curicta pronotata Kuitert thus, when possible, series of specimens should 3'. Juga slightly convex, not as elevated as be collected for many reasons, including to facil- tylus (Fig. 5); Louisiana, Texas, Mexico, itate characterization of the population and for a Mesoamerica Curicta scorpio Stal more confident use of the key. 4. Prothorax with midventral hollow groove Distinction between the parapatric R. brev- (Fig. 6); southeastern United States icollis Montandon and R. fusca may be quite Hungerford difficult. The suite of characters historically used 4'. Prothorax without midventral hollow to distinguish these species is generally unreli- groove, but may be ventrally flattened or able because of the high level of intraspecific have paired ventrolateral longitudinal de- morphological variability in both species. Usu- pressed lines 5 ally, the morphology of the penultimate antennal 5. Profemur with midventral narrow, curved, segment is diagnostic (see couplet 11); however, elongate spine, height of spine subequal nepid antennae are variable. Nonetheless, where to width of femur at that point (Fig. 22); the ranges of the species overlap (i.e., mid- to male with laterotergites of terminal ab- southern California), morphological divergence dominal segment wrapped around sterna is evident in the structure of the metaxyphus. (Fig. 19); penultimate antennal segment Elsewhere in the range of R. fusca, the expres- with lateral projection ca. V2 length of ter- sion of the metaxyphus is quite variable. A key to minal antennal segment (Fig. 20); Texas, identify California specimens was presented by Mexico, Mesoamerica Menke (1979); those characters have not been Ranatra texana Hungerford incorporated into the key presented here, but 5'. Profemur with midventral spine at most are discussed in the Diagnosis section for each of slightly longer than wide, usually no the two species. In distinguishing between R. longer than % width of femur at that point quadridentata and R. fusca, the principal char- (Figs. 7-11); male with laterotergites of acter is the degree of excavation of the profemo- terminal abdominal segment not wrapped ral apical notch. When examining specimens for around sterna; penultimate antennal seg- this feature, it is important to disregard the ves- ment with lateral projection variable ... 6 titure lining the ventral margin of the femur and 6. Penultimate antennal segment with lateral to focus on the cuticle. projection absent or

17 Figs. 1-17. (1) Abdominal sterna of Ranatra. (2) Abdominal sterna of Nepa. (3) Abdominal sterna of Curicta. (4) Head of Curicta pronotata. (5) Head of Curicta scorpio. (6) Prosternal groove of Ranatra buenoi. (7) Prothoracic femur and tibia of Ranatra nigra. (8) Prothoracic femur and tibia of Ranatra kirkaldyi. (9) Prothoracic femur of Ranatra australis. (10) Prothoracic femur of Ranatra quadridentata. (11) Prothoracic femur of . (12) Left antenna of Ranatra nigra. (13) Left antenna of Ranatra kirkaldyi. (14) Left antenna of Ranatra brevicollis. (15) Left antenna of Ranatra fusca. (16) Sterna of Ranatra quadridentata showing metaxyphus. (17) Sterna of Ranatra fusca showing metaxyphus.

piratory siphons ^50 mm; penultimate southeastern seaboard antennal segment with lateral projection Ranatra drakei Hungerford V2-V5 length of terminal antennal seg- 10' .Pronotum with anterior portion 1.5-2.5 ment 10 times length of posterior portion when 10. Pronotum with anterior portion 3 times measured at midline; width of each eye length of posterior portion when mea- subequal to or less than interocular dis- sured at midline; each compound eye tance 11 slightly wider than interocular distance; 11. Penultimate antennal segment with lateral 32 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 87, no. 1

projection V2—% length of terminal anten- Distribution. This species is an inhabitant of nal segment when measured from base of pools in mountain streams. In the United States, bifurcation (Fig. 14); southern California C. pronotata is restricted to Arizona (Fig. 24), north to =s40° latitude, Mexico where it has been collected only a few times in Ranatra brevicollis Montandon the Huachuca Mountains and Sabino Canyon 11'. Penultimate antennal segment with lateral near Tucson. In Mexico, it is locally common in projection 3A-Vs length of terminal anten- the Sierra de los Ajos and Sierra Tarahumara of nal segment when measured from base of Sonora and occurs also in western Chihuahua, bifurcation (Fig. 15) 12 Sinaloa, Nayarit, and Jalisco (Keffer 1991). 12. Profemoral apical notch usually semicircu- lar and ^2 times as wide as deep (Fig. Curicta scorpio Stal 10), distal limit of notch may be poorly defined; metaxyphus digitate, narrow, ex- Curicta scorpio Stal, 1862: 203; Keffer, 1993. tending to posterior margin of hind coxae Nepoidea montandoni Martin, 1898: 68 (syn- (Fig. 16); southwestern United States, onymized by Champion, 1901: 353). Mexico, Mesoamerica Curicta howardi Montandon, 1910a: 181; Pol- Ranatra quadridentata Stal hemus, 1988: 529 (synonymized by Keffer, 1993). 12'. Profemoral apical notch broadly exca- Curicta drakei Hungerford, 1922: 432 (synon- vated, >2 times as wide as deep (Fig. 11); ymized by Kuitert, 1949b: 68). metaxyphus variable, ranging from as Diagnosis. The slightly smaller average size, above to broader, extending to middle of unswollen tylus (Fig. 5), and prominent longitu- hind coxae (Fig. 17); widespread across dinal carinae on the last abdominal tergum sep- northern states and southern Canada; arate this species from C. pronotata. northern California south to —36° latitude Distribution. This species is found in ponds Ranatra fusca Palisot de Beauvois and slow streams in the southern United States; however, it has been collected in rocky hill The following is an annotated list of the spe- stream pools in southern Mexico. In the James cies of Nepidae of the United States and Canada. River (Mason County, Texas), many specimens The synonymies list only works that present have been taken on the shaded, muddy bank important taxonomic or nomenclatural informa- among grasses completely out of the water tion. Localities listed in Material Examined for (R.W.S., unpublished data). In the United States, most species present new state or province C. scorpio occurs in southern Texas and Louisi- records (i.e., not listed in Polhemus [1988]) or ana (Fig. 24), where it sometimes is abundant. In significant range information. Because R. texana Mexico, it occurs in the eastern states of Oaxaca, is redescribed, localities in Material Examined San Luis Potosi, and Veracruz and extends across are reported for all specimens examined re- central Mexico in the states of Colima, Morelos, gardless of range significance. For more general Jalisco, Michoac^n, Nayarit, and Guerrero to distribution information for all species, see overlap the range of C. pronotata. It occurs in the distribution maps. Abbreviations for collec- Mesoamerica as far south as Guatemala, Hondu- tions listed in the Material Examined sections ras, and Nicaragua (Keffer 1991). are as follows: CAS (California Academy of Sciences), CNC (Canadian National Collec- tion), CSU (Colorado State University), NCSU Genus Nepa (North Carolina State University), JTPC (John T. Polhemus Collection), TAM (Texas A&M Uni- Nepa apiculata Uhler versity), UF (University of Florida), UMC Nepa apiculata Uhler, 1862: 12, pi. I, fig. 1; (University of Missouri-Columbia), USNM Polhemus, 1988: 530. (United States National Museum), USU (Utah Ferrari, 1888: 189, in part. State University), WSU (Washington State Uni- Diagnosis. The short, wide habitus, a charac- versity). teristic of the genus, readily distinguishes N. api- culata from all other North American nepids. Distribution. This species is found in the Genus Curicta northeastern United States as far west as North Dakota, Iowa, southwestern Kansas, and Okla- homa, with one isolated record in Georgia (Fig. Curicta pronotata Kuitert 24). An early instar housed at the USNM is la- Curicta pronotata Kuitert, 1949b: 66; Polhe- beled: Florida: Cape Sable. Because the identity mus, 1988: 529. of this specimen is tenuous and the record is Diagnosis. The slightly larger average size, widely disjunct from the remainder of the range, bulging tylus (Fig. 4), and slender longitudinal it is considered suspect and requires confirma- carinae on the last abdominal tergum separate tion; therefore, it is not included in the range this species from C. scorpio. presented here. In Canada, N. apiculata occurs January 1994 SITES & POLHEMUS: NEPIDAE OF UNITED STATES AND CANADA 33

along the southern border from Quebec as far Ranatra buenoi Hungerford west as Manitoba. Ranatra buenoi Hungerford, 1922: 442; Polhe- Material Examined. North Dakota: Richland mus, 1988: 530. County: Mirror Pool, 23-29-VI-66, G. Aarhus, Diagnosis. This species is easily distinguish- USNM. Missouri: Platte City, 27-IV-36, F. W. able from the other North American species by Forbes, USNM. the very pronounced deep sulcus on the proster- num (Fig. 6). Distribution. The distribution of this species is Genus Ranatra similar to that of R. australis, the southeastern United States (Fig. 27). Material Examined. Illinois: Union County: 1 Ranatra australis Hungerford 8, Pine Hills Recreation Area, 5-V-57, R. Dim- mick, UMC. Kentucky: Butler County: Mud Ranatra australis Hungerford, 1922: 449; River, 37°07'24" N, 86°54'03" W, JTPC; Caldwell Lansbury, 1974b: 25, figs. 17-18; Polhemus, County: Land Branch wetland, 37°20'43" N, 1988: 530. 87°50'33" W, JTPC; Daviess County: Panther Diagnosis. This species is distinguishable Creek, Old Channel, 37°42'20"N, 87°12'54"W, from similar U.S. and Canadian species by the JTPC; Henderson County: Cypress Slough, lack of a distal femoral tooth, coupled with the 37°53'55" N, 87°29'30" W, JTPC; Hopkins Coun- well developed penultimate antennal segment, ty: Long Pond, 37°14'08" N, 87°22'20" W, JTPC; which distinguishes it from R. kirkaldyi and R. Ohio County: Muddy Creek, 37°26'05" N, nigra. Additionally, the lack of a prominent pros- 86°53'10" W, JTPC. Texas: Kaufman County: 1 ternal sulcus distinguishes it from R. buenoi, and 8,12.9 km, S Kemp, 15-X-88, B. A. Baugh, UMC. a short profemoral median tooth distinguishes it from R. texana. Distribution. Common in the southeastern Ranatra drakei Hungerford United States (Fig. 25). Material Examined. Illinois: Jackson County: Ranatra drakei Hungerford, 1922: 451; Lans- 1 6, Carbondale, farm pond, 8-VIII-87, R. W. bury, 1974b: 25, figs. 20-21; Polhemus, 1988: Sites, UMC; Union County: 1 9, Pine Hills, 531. 2-VIII-84, UMC. Kentucky: Ohio County: Rock Diagnosis. This species is clearly distinguish- House Slough, 37°28'38" N, 86°53'25" W, JTPC. able from the other United States and Canadian species by the very long, slender pronotum and large eyes; see key. Distribution. Ranatra drakei is a warm- Ranatra brevicollis Montandon adapted species restricted to the lower eastern Ranatra brevicollis Montandon, 1910a: 184; seaboard and extreme southeastern states (Fig. Lansbury, 1974b: 25, fig. 19; Polhemus, 1988: 26). It is most common in Florida. 530. Diagnosis. The penultimate antennal seg- Ranatra fusca Palisot de Beauvois ment has a lateral projection V2 to % the length of the terminal antennal segment (Fig. 14), Ranatra fusca Palisot de Beauvois, 1820: 235; which distinguishes it from R. fusca and R. Hungerford, 1922: 446; Lansbury, 1974b: 25, quadridentata, in each of which the projection figs. 22-23; Polhemus, 1988: 531. of the penultimate segment is longer. The Nepa 4-dentata [sic]: Uhler, 1871: 471 (mis- metaxyphus usually is long, raised, and keel-like, identification). almost reaching the base of the abdomen; Ranatra quadridentata: Uhler, 1876: 338 whereas R. fusca usually has a shorter, flatter (part) (misidentification). metaxyphus. The profemoral median spine usu- Ranatra americana Montandon, 1910b: 65 ally is situated midway between the trochanter (synonymized by Hungerford, 1922: 436). and apex (measured on the inner edge). The Ranatra americana edentula Montandon, profemoral apical notch usually is defined prox- 1910b: 66. New synonymy. imally by a rounded angle, without a tooth. The Ranatra fusca edentula: Hungerford, 1922: female subgenital plate is slender (in lateral 447; Polhemus, 1988: 531. view), and the ventral margin is rounded only in Diagnosis. The penultimate antennal segment the basal VA. has a lateral projection approximately Ys the Distribution. Baja, California Norte (Mexico) length of the terminal antennal segment (Fig. and southern California north to —40° latitude 15). The metasternal process usually is short, ex- (Fig. 26). This species apparently is adapted to a tending ~V2 the length of the metacoxal base Mediterranean climate, because it is not found (Fig. 17). The profemoral median spine usually east of the Sierra Nevada range or north of cen- is situated slightly closer to the apex than to the tral California. trochanter (measured on the inner edge). The 34 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 87, no. 1 profemoral apical notch usually is defined prox- cade, 23-IX-88, WSU. Kentucky: Nelson County: imally by a sharp angle (Fig. 11) or tooth. The near Bardstown, JTPC; Perry County: near Haz- female subgenital plate is broad (in lateral view), ard, JTPC. Montana: Flathead County: Thomp- with the ventral margin rounded in the basal V2. son River, 6.6 mi. [10.6 km] S Rt. 2, 25-IX-90, The paramere is long, slender, and curved, with WSU; Lincoln County: 1 9, Pipe Ck., 7 mi. [11.3 a short subapical tooth. km] N Libby, 24-IX-90, R. S. Zack, UMC; Mis- Discussion. Ranatra fusca is the most wide- soula County: Missoula, 21-V-55, WSU. Nebras- spread and common North American species. ka: Sioux County: 1 6, 1 9, Niobrara River, M. This species usually is an inhabitant of ponds, Harris, 16-IX-89, CSU, JTPC. Washington: Ad- where it sometimes is common and intermixed ams County: Othello, 10-VII-64, WSU; Asotin with dead grasses or weeds, although a few spec- County: Headgate Pk., 10 mi. [16.1 km] SW imens have been taken from rivers. It is morpho- Asotin, 26-V-86, WSU; Benton County: Benton, logically variable over its range, which has cre- Cedar River, CAS; Chelan County: 2 9 9, 17 mi. ated difficulty in establishing key characteristics. [27.4 km] N Leavenworth, T26N R17E sec. 3, Where the range overlaps that of a closely related 16-IX-88, R. S. Zack, UMC; Clark County: 6-IX- species (in this case R. brevicollis), the variabil- 39, WSU; King County: 1.3 mi. [2.1 km] E Red- ity is diminished, a phenomenon noticed by mond, 24-VIII-89, WSU; Columbia County: Bea- J.T.P. for taxa of distantly related groups (e.g., ver Lk., 16 mi. [25.7 km] SSW Pomeroy, T9N Microvelia pulchella Westwood). R41E sec. 2-3, 15-X-87, WSU; Snohomish Coun- The variety edentula was proposed by ty: Everett, 6-III-43, WSU; Stevens County: Montandon (1910b, as americana edentula) for Chewelah, Deer Lk., 6-X-72, WSU; Thurston those populations lacking the preapical tooth on County: Yelm, 22-VI-89, WSU; Whitman Coun- the profemur and was reported to occur in Penn- ty: 1 9, Pullman, 21-V-84, R. W. Sites, UMC; sylvania and Texas. Although Pennsylvania is Yakima County: Dry Ck., 10 mi. [16.1 km] SW within the range of the nominate subspecies, Toppenish, 18-VIII-89, WSU. Oregon: Harney Texas is not, and it is likely that the specimen County: 1 9, 4-VIII-47, R. C. Erickson, JTPC. from the latter is R. australis Hungerford. A neo- Utah: Cache County: Green Canyon, USU; Utah type has been designated by Polhemus (1994) for County: 1 9, Provo, Bricky Pond, 6-XI-84, D. R. fusca edentula Montandon, and the type lo- McClain, NCSU. cality has been restricted to Pennsylvania to sta- bilize the nomenclature for the species R. fusca. Specimens with profemora of both forms (i.e., Ranatra kirkaldyi Torre Bueno with and without preapical tooth) and interme- Ranatra kirkaldyi Torre Bueno, 1905c: 187; diates are represented in a large sample from Hungerford, 1922: 440; Lansbury, 1974b: 28; Colorado in the JTPC. By definition, subspecies Polhemus, 1988: 531. must be spatially separated; thus edentula must Ranatra fusca: Montandon, 1910b: 63 (mis- fall as a species-group synonym of fusca. identification). Distribution. Ranatra fusca is found at lower Ranatra kirkaldyi hoffmanni Hungerford, elevations completely across the northern tier of 1922: 442. New synonymy. contiguous states in the United States and the Diagnosis. This species is relatively small and southern tier of Canadian provinces (Fig. 28). A robust for the genus, with short respiratory si- Florida record needs reconfirmation. Kuitert phons. The antennal segments are roughly cylin- (1947) discussed a specimen from Louisiana that drical, and the penultimate segment is without a was placed as this species, but it lacked the distal distolateral projection (Fig. 13), which distin- profemoral tooth and had a metaxyphus similar guishes it from all other United States and Cana- in form to that of R. australis; thus it is doubtful. dian species of Ranatra. The paramere is long, The range of R. fusca does not overlap that of the slender, and curved; the subapical tooth is closely related R. quadridentata (see Fig. 28). widely separated from the apical tooth, and the Material Examined. CANADA: British Colum- posterior diverticulum is slender, not expanded bia: Vancouver, Spencer, CAS. Prince Edward distally. The metaxyphus is truncate, and the lat- Island: Watervale, 21-VIII-71, P. Budd, CNC. eral margins are not membranous. Nova Scotia: Echo Hwy., 7-11-VIII-71, CNC. Discussion. Ranatra kirkaldyi is quite distinct UNITED STATES: Colorado: Boulder County: from all other North American species in several many 66, 9 9, Sawhill Ponds, Boulder, 9-X-84, respects and is most closely related to the South J. T. Polhemus, JTPC; Douglas County: 1 9, American fauna (S. L. Keffer, in litt). The strid- 20-111-73, J. T. Polhemus, JTPC; Jefferson Coun- ulatory mechanism is lacking, a characteristic ty: 1 6, 1 9, 21-111-86, J. T. Polhemus, JTPC; shared with only three other American species Yuma County: 1 6, Chief Creek, nr. Wray, CL (see Biology and Ecology section). 2690, 6-VI-92, J. T. Polhemus, JTPC. Idaho: La- The variety hoffmanni was proposed by Hun- tah County: 1 9, Moscow Mountain, 3500-4500 gerford (1922) on the basis of slight differences ft. [1,070-1,370 m], 21-XI-83, R. W. Sites, UMC; between populations from Minnesota and Kan- Valley County: Corral Ck., 3 mi. [4.8 km] S Cas- sas, particularly the presence or absence of a January 1994 SITES & POLHEMUS: NEPIDAE OF UNITED STATES AND CANADA 35 preapical tooth on the profemur. Both forms and mislabeled; therefore, it is not included in the intermediates are represented in each of two range presented here. widely separated samples (Kansas and Missis- Material Examined. Illinois: Jackson County: sippi) in the JTPC. By definition, subspecies 1

Figs. 18-23. Ranatra texana: (18) Lateral view of left paramere. (19) Lateral view of terminal abdominal segment of male. (20) Left antenna. (21) Digitate metaxyphus of metasternum. (22) Median ventral spine of profemur. (23) Lateral view of terminal abdominal segment of female. em California (Fig. 28). In Mexico, it is common species by the elongated, sickle-shaped median and widespread, known from the states of Chia- tooth of the profemur. Superficially, because of pas, Chihuahua, Durango, Guanajuato, Michoa- its slender habitus and lack of profemoral distal c£n, Oaxaca, Sinaloa, Sonora, and Veracruz. It tooth, R. texana most closely resembles R. nigra, ranges southward through Guatemala to Hondu- with which it is parapatric, but differs in the ras and Nicaragua. longer penultimate antennal segment and ex- Material Examined. Arizona: Maricopa Coun- pression of the profemoral median tooth. ty: 2 6 6, 5 mi. [8 km] SE Wickenburg, 10-IX-80, Description. Male: (Male described by Hun- USU; Yavapai County: 2 6 6, 1 9, In, 5 mi. [8 gerford [1930]; only additional notes given here.) km] NE Castle Hot Springs, CL 312, 7-X-64, J. T. Paramere long, slender, curved, with short sub- Polhemus, JTPC. New Mexico: Dona Ana Coun- apical tooth (Fig. 18); posterior diverticulum ex- ty: 1 9, Las Cruces, Jornado Exptl. Range, To- panded; connexiva of abdominal segment VII bosa Playa, 30-VII-70, C. R. Ward & L. G. Rich- embracing operculum distally (Fig. 19). ardson, UMC. Texas: Cherokee County: 5 6 6, 4 Female: Body length, 35.30 ± 0.31 mm (mean 9 9, 10 mi. [16.1 km] N Jacksonville, 15-X-67, T. ± SEM; n = 21); respiratory siphon, 18.43 ± B. Tennison, TAM; Houston County: 1 9, Crock- 0.45; head width, 3.02 ± 0.03. Body slender, cy- ett, 24-X-65, TAM; San Patricio County: 1 6, lindrical. Overall coloration medium brown to Lake Corpus Christie St. Pk., 23-V-72, UMC; straw-yellow dorsally, medium to dark brown Smith County: 3 6 6, Bullard, 28-X-67, T. B. ventrally. Head, pronotum slightly lighter than Tennison, TAM; Waller County: 2 9 9, 3 mi. [4.8 hemelytra and respiratory siphons. km] W Hempstead, 21-IX-70, R. K. LaVal, UMC. Vertex of head elevated above level of eyes; GUATEMALA: 1 9, 10 mi. [16.1 km] N Ascun- tylus longer, slightly more prominent (dorsally cion Mita, CL 1313, 10-1-70, J. T. Polhemus, elevated) than juga. Antenna 3-segmented; pen- JTPC. HONDURAS: 1 9, Dept. Comayagua, ultimate segment with lateral projection ~V% pond, Quebrada Montanuelos, 1,080 m, 21-111- length of terminal segment (Fig. 20). 84, A. R. Gillogly, JTPC. NICARAGUA: 4

Figs. 24-30. Distribution maps of United States and Canadian Nepidae.

R. fusca from invading this range, as the climate to those of R. texana, although the median is no more severe than in many other places it profemoral tooth is much shorter. occurs. The anomalous presence of R. fusca near Denver, CO, at —1,700 m, is therefore confound- ing. Acknowledgments Ranatra operculata Kuitert is known from sev- eral localities in Chihuahua and Sonora, Mexico, We thank Thomas J. Henry (U.S. National Museum), as close as 200 km to the Arizona border; thus, Sharon Jasper (Texas A&M University), Steven L. Kef- fer (103 Dinkel Ave., Ridgewater, VA 22812), Frank W. we expect that this species will eventually be Mead (University of Florida), and Michael Schwartz found in the United States. The females are very (Canadian National Collection) for sharing unpub- easy to recognize by the extremely long, slender lished information; and Becky J. Nichols (University of subgenital plate (operculum) that far surpasses Missouri) for assistance with preparation of distribu- the base of the respiratory tube; males are similar tion maps. We are indebted to the following for the January 1994 SITES & POLHEMUS: NEPIDAE OF UNITED STATES AND CANADA 39 loan or gift of material: Dean W. Blinn and Clay Runck Bobb, M. L. 1974. The aquatic and semi-aquatic (Northern Arizona University), Robert Blinn (North Hemiptera of Virginia. Va. Polytech. Inst. Res. Div. Carolina State University), Wilford J. Hanson (Utah Bull. 87. State University), Boris C. Kondratieff (Colorado State Brooks, A. R. & L. A. Kelton. 1967. Aquatic and University), Carl Olson (University of Arizona), Ed- semiaquatic Heteroptera of Alberta, Saskatchewan, ward G. Riley and Sharon Jasper (Texas A&M Univer- and Manitoba (Hemiptera). Mem. Entomol. Soc. sity), and Richard S. Zack, Jr. (Washington State Uni- Can. 51: 1-92. versity). Reviews of the manuscript were graciously Chagnon, G. & O. Fournier. 1948. Contribution a provided by Carl W. Schaefer (University of Connect- l'e'tude des He"mipteres aquatiques du Quebec. icut) and J. E. McPherson (Southern Illinois Universi- Contrib. Inst. Biol. Univ. Montreal 21: 1-66. ty). The Wilbur R. Enns Entomology Museum pro- Champion, G. C. 1901. Insecta: Rhynchota vided word processing and graphics resources. (Hemiptera-Heteroptera), vol. II, pp. i-xvi + 345- Funding for R.W.S. was provided in part by project no. 416. In F. D. Godman & O. Salvin [eds.], Biologia PSSL0232. This is Missouri Agricultural Experiment Centrali Americana. London. Station journal series paper no. 11,927. Chockalingam, S. & P. Somasundaram. 1987. Effi- ciency of mosquito predation by an aquatic insect, maculatus (Hemiptera: Nepidae). Arch. Hydrobiol. Beih. Ergeb. Limnol. 28: 525-528. References Cited Clark, L. B. 1926. Aquatic Hemiptera from Mani- Abbott, C. E. 1940. Some reflex responses of Rana- toba. Can. Entomol. 58: 203-205. tra fusca to contact stimuli. Bull. Brooklyn Ento- Cloarec, A. 1976. Interactions between different re- mol. Soc. 35: 133-134. ceptors involved in prey capture in Aiken, R. B. 1985. Sound production by aquatic in- (Insect, Heteroptera). Biol. Behav. 1: 251-266. sects. Biol. Rev. 65: 163-211. 1986. Distance and size discrimination in a water Aldrovandi, U. 1602. De animalibus insectis libri stick insect Ranatra linearis (Heteroptera). J. Exp. septem, cum singulorum iconibus ad vivum expres- Biol. 120:59-77. sis. Denuo Impresi: Bonon. Apud Clementem Fer- Cochran, P. A., A. P. Gripentrog & K. M. Stack. 1992. ronium. (Reprinted in 1638 with same title page but Distribution of the water Nepa apiculata different date.) (Hemiptera: Nepidae) in Wisconsin. Gr. Lakes Anonymous. 1878. Ranatra linearis attacking carp Entomol. 25: 83-84. eggs. The Entomologist 11: 95. Cross, J. L. 1972. New state records of aquatic in- Baunacke, W. 1912. Statische Sinnesorgane bei den sects from Virginia. Proc. Entomol. Soc. Wash. 74: Nepiden. Zool. Jahrb. Abt. Anat. Ontog. 34: 179- 476. 342. Davis, C. C. 1961. A study of the hatching process Bennett, D. V., III. 1987. Evolution of a waterscor- in aquatic invertebrates. II. Hatching in Ranatra pion species complex in North America (Insecta: fusca P. Beauvois (Hemiptera, Nepidae). Trans. Hemiptera: Nepidae). Ph.D. dissertation, Univer- Am. Microsc. Soc. 80: 230-234. sity of Connecticut, Storrs. De Carlo, J. A. 1967. Diferencias entre Ranatridae y Bisht, R. S. & S. M. Das. 1981. Observations on Nepidae. Division de la familia Ranatridae en Rana- aquatic insects as food of fishes and the predatory trinae y Curictinae. Rev. Soc. Entomol. Argent. 29: action of some aquatic insects on fish and fish food. 21-29. J. Inl. Fish. Soc. India 13: 80-86. 1968. Redescripcion y aparato genital masculino de Blatchley, W. S. 1926. Heteroptera or true bugs of Austronepa angusta (Hale) (Hemiptera, Rana- eastern North America with especial reference to tridae). Physis (B. Aires) 27: 343-347. the faunas of Indiana and Florida. Nature Publ. Co., 1972. Clave de las especies Americanas descriptas Indianapolis. del genero Ranatra Fabricius (Hemiptera, Rana- Blinn, D. W. & C. Runck. 1989. Substratum require- tridae, Ranatrinae). Rev. Soc. Entomol. Argent. 34: ments for oviposition, seasonal egg densities, and 177-185. conditions for egg eclosion in Ranatra montezuma Drake, C. J. & J. A. De Carlo. 1953. American spe- (Heteroptera: Nepidae). Ann. Entomol. Soc. Am. cies of Ranatra annulipes Stal group (Hemiptera: 82: 707-711. Ranatridae). J. Wash. Acad. Sci. 43: 109-117. Blinn, D. W. & M. W. Sanderson. 1989. Aquatic in- DuBois, M. B. 1978. A recent record of Curicta pro- sects in Montezuma Well, Arizona, USA: a traver- notata Kuitert (Hemiptera: Nepidae) from its type tine spring mound with high alkalinity and dis- locality. Entomol. News 89: 207. solved carbon dioxide. Great Basin Nat. 49: 85-88. Blinn, D. W., C. Pinney & M. W. Sanderson. 1982. Dufour, L. 1821. Recherches anatomiques sur la Nocturnal planktonic behavior of Ranatra monte- Ranatre liniaire et la Nepe cendre"e. Ann. Gen. Sci. zuma Polhemus (Nepidae: Hemiptera) in Monte- Phys. Brux. 7: 194-213, 3 taf. [1820]. zuma Well, Arizona. J. Kans. Entomol. Soc. 55: 481- 1833. Recherches anatomiques et physiologiques 484. sur les H6mipteres accompagne'es de consider- Blinn, D. W., C. Runck & R. W. Davies. 1993. The ations relatives a l'histoire naturelle et a la classifi- impact of prey behaviour and prey density on the cation de ces insectes. Mem. Savans Etrang. Acad. foraging ecology of Ranatra montezuma (Het- Sci. 4: 129-462, pis. 1-19. eroptera): a serological examination. Can. J. Zool. Ellis, L. L. 1952. The aquatic Hemiptera of south- 71: 387-391. eastern Louisiana (exclusive of the ). Am. Blois, C. & A. Cloarec. 1983. Density-dependent Midi. Nat. 48: 302-329. prey selection in the water stick insect, Ranatra Farris, J. L. & G. L. Harp. 1982. Aquatic macroin- linearis (Heteroptera). J. Anim. Ecol. 52: 849-866. of three acid bogs on Crowley's Ridge 40 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 87, no. 1

in northeast Arkansas. Arkansas Acad. Sci. Proc. 36: 1930. Two new water bugs from the western U.S.A. 23-27. (Nepidae and ). Can. Entomol. 62: Ferrari, E. 1888. Die Hemipteren-Gattung Nepa 216-218. Latr. (sens, natur.). Ann. Naturhist. Mus. Wien 3: 1932. Concerning a fossil water bug from the Flo- 161-194, 2 pis. rissant. Univ. Kans. Sci. Bull. 20: 327-331. Frisch, J. L. 1728. Beschreibung von allerley In- Keffer, S. L. 1991. Taxonomic revision of the neo- secten in Teutschland. Nicolai, Berlin. Teil 7, tropical genus Curicta Stal (Insecta: Heteroptera: 1+7+31 pp., 22 taf. Nepidae). Ph.D. dissertation, Southern Illinois Uni- Froeschner, R. C. 1962. Contributions to a synopsis versity, Carbondale. of the Hemiptera of Missouri, Part V. Hy- 1993. Synonymy of Curicta howardi Montandon drometridae, , , , with Curicta scorpio Stal (Heteroptera: Nepidae). J. , , , Belosto- N.Y. Entomol. Soc. vol. 101:434-435. matidae, Nepidae, Notonectidae, , Corixi- Keffer, S. L. & J. E. McPherson. 1993. Curictan cop- dae. Am. Midi. Nat. 67: 208-240. ulation and waterscorpion higher classification Gonsoulin, G. J. 1975. Seven families of aquatic and (Heteroptera: Nepidae). Proc. Entomol. Soc. Wash. semiaquatic Hemiptera in Louisiana. Part V. Fam- 95: 74-78. ily Nepidae Latreille, 1802. "Water ." Kirkaldy, G. W. & J. R. de la Torre Bueno. 1909. A Entomol. News 86: 23-32. catalogue of American aquatic and semi-aquatic Hamilton, M. A. 1931. The morphology of the wa- Hemiptera. Proc. Entomol. Soc. Wash. 10: 173-215. ter-scorpion, Nepa cinerea Linn. (Rhynchota, Het- Kuitert, L. C. 1947. Nepidae of the Western Hemi- eroptera). Proc. Zool. Soc. Lond. 1931: 1067-1136. sphere (Hemiptera). Ph.D. dissertation, University Harris, H. M. 1937. Contributions to the South Da- of Kansas, Lawrence. kota list of Hemiptera. Iowa State Coll. J. Sci. 11: 1949a. Some new Ranatra from the Americas. J. 169-176. Kans. Entomol. Soc. 22: 24-34. Harris, H. M. & W. E. Shull. 1944. A preliminary 1949b. Some new species of Nepidae (Hemiptera). list of Hemiptera of Idaho. Iowa State Coll. J. Sci. J. Kans. Entomol. Soc. 22: 60-68. 18: 199-208. Lago, P. K. & S. Testa III. 1989. 6. The aquatic and Henry, T. J. & R. C. Froeschner [eds.]. 1988. Cat- semiaquatic Hemiptera and Coleoptera of Point alog of the Heteroptera, or true bugs, of Canada and Clear Island, Hancock County, Mississippi. J. Miss. the continental United States. Brill, Leiden. Acad. Sci. 34: 33-^38. Herrich-Schaeffer, G.A.W. 1849-1853. Die Wanze- Lansbury, I. 1972. A review of the Oriental species nartigen Insecten. Getreu nach der Natur abgebil- of Ranatra Fabricius (Hemiptera-Heteroptera: Ne- det und beschrieben. Vol. 9: 1-44 (1849); 45-256 pidae). Trans. R. Entomol. Soc. Lond. 124: 287-341. (1850); 257-358 (1851); ["historischer ubersicht" 1974a. A new genus of Nepidae from Australia with und "index"] 1-210 (1853). J. L. Lotzebeck, Nurn- a revised classification of the family (Hemiptera: berg. Heteroptera). J. Aust. Entomol. Soc. 13: 219-227. Herring, J. L. 1951a. The aquatic and semiaquatic 1974b. Notes on Ranatra (Amphischizops) com- Hemiptera of northern Florida. Part 3: Nepidae, pressicollis Montandon with a review of its system- , Notonectidae, Pleidae and Corixi- atic position within the American Ranatra dae. Fla. Entomol. 34: 17-29. (Hemiptera-Heteroptera, Nepidae). Zool. Scr. 3: 1951b. The aquatic and semiaquatic Hemiptera of 23-30. northern Florida. Part 4: Classification of habitats Larimore, R. W., W. F. Childers & C. Heckrotte. and keys to the species. Fla. Entomol. 34: 146-161. 1959. Destruction and reestablishment of stream Hilsenhoff, W. L. 1984. Aquatic Hemiptera of Wis- fish and invertebrates affected by drought. Trans. consin. Gr. Lakes Entomol. 17: 29-50. Am. Fish. Soc. 88: 261-285. Hinton, H. E. 1961. The structure and function of Larsen, O. 1949a. Uber die kataleptische Akinese the egg-shell in the Nepidae (Hemiptera). J. Insect der Nepiden. Opusc. Entomol. 14: 5-25. Physiol. 7: 224-257. 1949b. Die Ortsbewegungen von Ranatra linearis 1962. A key to the eggs of the Nepidae (Hemiptera). L. Ein Beitrag zur vergleichenden Physiologie der Proc. R. Entomol. Soc. Lond. Ser. A Gen. Entomol. Lokomotionsorgane der Insekten. Lunds Univ. Ar- 37: 65-68. sskrift N. F. Avd. 2, 45: 1-82. 1970. Insect eggshells. Sci. Am. 223: 84-91. Lauck, D. R. 1959. The taxonomy and bionomics of Hoffmann, W. E. 1925. Some aquatic Hemiptera the aquatic Hemiptera of Illinois. M.S. thesis, Uni- having only four nymphal stages. Bull. Brooklyn versity of Illinois, Urbana. Entomol. Soc. 20: 93-94. Lee, C. E. 1991. Morphological and phylogenetic 1930. Notes on the life history of Ranatra chinensis studies on the true water bugs (Hemiptera: Het- Mayr. Proc. Nat. Hist. Soc. Fukien Christian Univ. eroptera). Nature and Life 21: 1—183. 3: 31-37. 1992. A new name for the subgenus Miyamotoa of Holmes, S. J. 1906. Death-feigning in Ranatra. J. Nepidae (Heteroptera, Hemiptera). Nature and Life Comp. Neurol. Psychol. 16: 200-216. 22: 62. 1907. Observations on the young of Ranatra Lindeman, R. L. 1941. Seasonal food-cycle dynam- quadridentata Stal. Biol. Bull. 12: 158-164. ics in a senescent lake. Am. Midi. Nat. 26: 636-673. Hungerford, H. B. 1920. The biology and ecology of Locy, W. A. 1884. Anatomy and physiology of the aquatic and semiaquatic Hemiptera. Univ. Kans. family Nepidae. Am. Nat. 18: 250-255, 353-367, Sci. Bull. 11: 1-328 [1919]. pis. 9-12. 1922. The Nepidae of North America (Further stud- Marshall, W. S. & H.H.P. Severin. 1904. Some ies in aquatic Hemiptera). Univ. Kans. Sci. Bull. 14: points in the anatomy of Ranatra fusca P. Beauv. 425-467. Trans. Wis. Acad. Sci. Arts Lett. 14: 487-508, 3 pis. January 1994 SITES & POLHEMUS: NEPIDAE OF UNITED STATES AND CANADA 41

Martin, J. 1898. Descriptions d'especes nouvelles Penn, G. H. & L. L. Ellis. 1949. Notes on aquatic de Nepidae (Hem.). Bull. Soc. Entomol. Fr. 4: 66- Hemiptera from Lafayette and Marshall counties, 68. Mississippi. Fla. Entomol. 32: 158-159. McPherson, J. E. & R. J. Packauskas. 1987. Life his- Pierce, W. D. 1948. Fossil of California. tory and laboratory rearing of Nepa apiculata (Het- 15. Some Hemiptera from the McKittrick asphalt eroptera: Nepidae), with descriptions of immature field. Bull. South. Calif. Acad. Sci. 47: 21-33. stages. Ann. Entomol. Soc. Am. 80: 680-685. Polhemus, D. A. 1988. Family Nepidae Latreille, Menke, A. S. 1964. Lectotype designation for Rana- 1802: the water scorpions, pp. 528-532. In T. J. tra quadridentata Stal. Bull. Brooklyn Entomol. Henry & R. C. Froeschner [eds.], Catalog of the Soc. 58: 112-113. Heteroptera, or true bugs, of Canada and the conti- 1979. Family Nepidae/water scorpions, pp. 70-75. nental United States. Brill, Leiden. In A. S. Menke [ed.], The semiaquatic and aquatic Polhemus, J. T. 1976. Notes on North American Ne- Hemiptera of California (Heteroptera: Hemiptera). pidae (Hemiptera: Heteroptera). Pan-Pac. Entomol. Bull. Calif. Insect Surv. 21: 1-166. 52: 204-208. Menke, A. S. & L. A. Stange. 1964. A new genus of 1994. Neotype designation for Ranatra americana Nepidae from Australia with notes on the higher edentula Montandon (Heteroptera: Nepidae). J. classification of the family. Proc. R. Soc. Queensl. N.Y. Entomol. Soc. (in press). 75: 67-72. Radinovsky, S. 1964. Cannibal of the pond. Nat. Miller, S. E. 1983. Late Quaternary insects of Ranch Hist. 73 (Nov.): 16-25. La Brea and McKittrick, California. Quat. Res. Rao, T.K.R. 1976. Bioecological studies on some (N.Y.) 20: 90-104. aquatic Hemiptera-Nepidae. Entomon 1: 123-132. Millspaugh, D. D. 1939. Bionomics of the aquatic Roemhild, G. 1976. The aquatic Heteroptera (true and semi-aquatic Hemiptera of Dallas County, bugs) of Montana. Mont. Agric. Exp. Stn. Res. Rep. Texas. Field and Lab. 7: 67-87. 102. Montandon, A. L. 1910a. Hydrocorises de l'Ame- Runck, C. & D. W. Blinn. 1990. Population dynam- rique du Nord, notes et descriptions d'especes nou- ics and secondary production by Ranatra monte- velles. Bull. Soc. Roum. Sci. 18: 180-191, 1909. zuma (Heteroptera: Nepidae). J. North Am. 1910b. Notes sur quelques formes Nord-Americains Benthol. Soc. 9: 262-270. du genre Ranatra (Hem.). Bull. Soc. Roum. Sci. 19: 1992. The foraging ecology of Ranatra montezuma 62-67. (Heteroptera): an optimal forager in Montezuma Moufet, T. 1634. Insectorum sive minimorum ani- Well, Arizona. J. Ariz. Nev. Acad. Sci. 26: 119-130. malium theatrum. T. Cotes, London. 9 + 326 + 2 Sanderson, M. W. 1982. Aquatic and semiaquatic pp. Heteroptera, pp. 6.1-6.94. In A. R. Brigham, W. U. Neiswander, C. R. 1925. On the anatomy of the Brigham & A. Gnilka [eds.], Aquatic insects and head and thorax in Ranatra (Heteroptera). Trans. oligochaetes of North and South Carolina. Midwest Am. Entomol. Soc. (Phila.) 51: 311-320. Aquatic, Mahomet, IL. Oldham, T. W. 1978. New records of aquatic and Sarbu, S. M. & R. Popa. 1992. A unique chemoau- semi-aquatic Hemiptera in Kansas. Tech. Publ. totrophically based cave ecosystem, pp. 640-666. State Biol. Surv. Kans. 6: 59-69. In A. L. Camacho [ed.], The natural history of bio- Ormerod, E. A. 1878. Ranatra linearis. Entomolo- speleology. Museo Nacional de Ciencias Natural, gist 11: 119-120. Madrid. Ouda, N. A., Abdul-Hussain, M. Al-Faisal & H. H. Schaefer, K. F. 1966. The aquatic and semiaquatic Zayia. 1986. Laboratory observations on the effi- Hemiptera of Oklahoma. Ph.D. dissertation, Okla- ciency of seven mosquito larvae predators. J. Biol. homa State University, Stillwater. Sci. Res. 17: 245-252. Schaefer, K. F. & W. A. Drew. 1964. Check list of Packauskas, R. J. & J. E. McPherson. 1986. Life his- aquatic and semiaquatic Hemiptera (Insecta) of tory and laboratory rearing of Ranatra fusca Oklahoma. Southwest. Nat. 9: 99-101. (Hemiptera: Nepidae) with descriptions of imma- 1968. The aquatic and semiaquatic Hemiptera of ture stages. Ann. Entomol. Soc. Am. 79: 566-571. Oklahoma, pp. 125-134. In Proceedings, 55th Okla- Palisot de Beauvois, A.M.F.J. 1820-1821. Insectes homa Academy of Science Meetings, 2—3 Decem- recueillis en Afrique et in Amerique, dans les ber 1966, Norman, OK. Oklahoma Academy of Sci- Royaumes d'Oware et de Benin, a Saint Dominique ence, Weatherford, OK. et dans les Etats Unis, pendant les annees 1786- Scudder, G.G.E. 1977. An annotated checklist of 1797. Levrault, Paris, xvi + 276 pp. the aquatic and semiaquatic Hemiptera (Insecta) of Parshley, H. M. 1922. Report on a collection of British Columbia. Syesis 10: 31-38. Hemiptera-Heteroptera from South Dakota. S.D. Severin, H.H.P. & H. C. Severin. 1911a. Habits of State Coll. Tech. Bull. 2. Belostoma (=Zaitha) flumineum Say and Nepa api- Parsons, M. C. 1972. Respiratory significance of the culata Uhler, with observations on other closely thoracic and abdominal morphology of Belostoma related aquatic Hemiptera. J. N.Y. Entomol. Soc. and Ranatra. Z. Morphol. Tiere 73: 163-194. 19: 99-108. 1974. Anterior displacement of the metathoracic 1911b. An experimental study on the death-feign- spiracle and lateral intersegmental boundary in the ing of Belostoma (=Zaitha Aucct.) flumineum Say pterothorax of Hydrocorisae (aquatic Heteroptera). and Nepa apiculata Uhler. Behav. Monogr. 1: 1-44. Z. Morphol. Tiere 79: 165-198. Slater, A. 1981. Aquatic and semiaquatic Het- Penn, G. H. 1951. Additional records of aquatic eroptera in the collection of the State Biological Hemiptera in Louisiana I. Nepidae, Survey of Kansas. Tech. Publ. State Biol. Surv. and Naucoridae. Proc. La. Acad. Sci. 14: 67-71. Kans. 10: 71-88. 42 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 87, no. 1

Stal, C. 1862. Nova methodus familias quasdam during the expedition of 1869, pp. All-All. In F. V. Hemipterorum disponendi. Ofv. Kongl. Svensk Vet. Hayden [ed.], Preliminary report of the United Akad. Forh. 18: 195-212 [1861]. States Geological Survey of Wyoming and portions Stys, P. & A. Jansson. 1988. Check-list of recent of the contiguous territories [1870]. Government family-group and genus-group names of Nepomor- Printing Office, Washington, D.C. pha (Heteroptera) of the world. Acta Entomol. 1876. List of Hemiptera of the region west of the Fenn. 50: 1-44. Mississippi River, including those collected during Swammerdam, J. 1737-1738. Biblia naturae; sive the Hayden explorations of 1873. Bull. U.S. Geol. Historia Insectorum, in classes certas redacta, nee Geog. Surv. Terr. 1: 267-361, pis. 19-21. non exemplis, et anatomico variorum animaculorum 1884. Order VI.—Hemiptera, pp. 204-293. In J. S. examine, aeneisque tabulis illustrata. Severin etc., Kingsley [ed.], The standard natural history. Vol. II. Leydae. 2 vols, 910 + 56 + 36 + 124 pp., 53 pis. Crustacea and insects. S. E. Cassino, Boston. Thorpe, W. H. & D. J. Crisp. 1947. Studies on plas- tron respiration. III. The orientation responses of Usinger, R. L. 1974. Aquatic insects of California Aphelocheirus [Hemiptera, (Nau- with keys to North American genera and California coridae)] in relation to plastron respiration; together species. University of California Press, Berkeley. with an account of specialized pressure receptors in Van Duzee, E. P. 1917. Catalogue of the Hemiptera aquatic insects. J. Exp. Biol. 24: 310-328. of America north of Mexico excepting the - Torre Bueno, J. R. de la. 1905a. The tonal apparatus idae, and Aleurodidae. Univ. Calif. Publ. of Ranatra quadridentata, Stal. Can. Entomol. 37: Tech. Bull. Entomol. 2: 1-902. 85-87. Weissmann, M. J. 1986. Biology of the water scor- 1905b. Practical and popular entomology.—No. 4. pion Ranatra fusca Palisot de Beauvois in Colo- Notes on collecting, preserving and rearing aquatic rado, with notes on mosquito control capability. Hemiptera. Can. Entomol. 37: 137-142. M.S. thesis, University of Colorado, Boulder. 1905c. The three Ranatras of the north-eastern Wiley, G. O. 1922. Some notes on the biology of United States. Can. Entomol. 37: 187-188. Curicta from Texas. Univ. Kans. Sci. Bull. 14: 507- 1906. Life histories of North American water-bugs. 511. II. Life-history otRanatra quadridentata. Stal. Can. 1924. On the biology of Curicta drakei Hungerford Entomol. 38: 242-252. (Heteroptera, Nepidae). Entomol. News 35: 324- 1923. Family Nepidae, pp. 399-402. In W. E. Brit- 331. ton [ed.], Guide to the insects of Connecticut. Part Wilson, C. A. 1958. Aquatic and semiaquatic IV. The Hemiptera or sucking insects of Connecti- Hemiptera of Mississippi. Tulane Stud. Zool. 6: cut. Conn. Geol. Nat. Hist. Surv., Bull. No. 34. 115-170. Uhler, P. R. 1862. Hemiptera, pp. 11-12, 192-256, Wood, T. 1884. Note on Nepa cinerea. Entomol. pi. I. In T. W. Harris, A treatise on some of the Mon. Mag. 21: 20. insects injurious to vegetation. W. White, Boston, xi + 640 pp. 1871. A list of Hemiptera collected in eastern Col- Received for publication 24 May 1993; accepted 19 orado and northeastern New Mexico, by C. Thomas, July 1993.