TAES 141: 405-429 RESEARCH ARTICLE ISSO 0002-8320 http://taes.entomology-aes.org/
Alderflies, Fishflies and Dobsonflies (Insecta: Megaloptera) of the Interior Highlands, U.S.A.
David E. Bowles1 and Robert W. Sites2
1 National Park Service, Heartland Inventory and Monitoring Program, c/o Department of Biology, Missouri State University, 901 South National Avenue, Springfield, MO 65897, USA. E-mail: [email protected]
2 Enns Entomology Museum, Division of Plant Sciences, University of Missouri, Columbia MO 65211, USA. E-mail: [email protected]
ABSTRACT
We report 15 species of Megaloptera from the Interior Highlands of Arkansas, Illinois, Missouri and Oklahoma including 2 genera and 9 species of Sialidae and 4 genera and 6 species of Corydalidae. New state distributional records are reported for Protosialis americana (Rambur) from Illinois, Sialis joppa Ross and S. vagans Ross from Missouri, and Chauliodes pectinicornis (L.) and Nigronia serricornis (Say) from Oklahoma. Distributional and biological data are summarized for each species where available. An illustrated key to the species occurring in the Interior Highlands is included. Key Words: Megaloptera, Protosialis, Sialis, Chauliodes, Corydalus, Neohermes, Nigronia, Interior Highlands.
INTRODUCTION Interior Highlands have poor soils, most agricultural activities involve small-scale livestock-rearing The Interior Highlands of the United States is operations. This region also contains numerous a geographically diverse area that includes portions national forests, wildlife refuges, national and state of Arkansas, Illinois, Missouri, and Oklahoma parks, and there is relatively little urbanization (Fenneman and Johnson 1946) (Fig. 1). This and industry. Due to a relatively low amount of 400,000 km2 area varies from 182 to 860 m above anthropogenic disturbance in the Interior Highlands, sea level and includes mountainous areas, plateaus, many streams in this region have relatively high and broad river valleys. The Interior Highlands have water quality and undisturbed hydrology (Petersen been variously divided into 15 subdivisions on the et al. 1998). basis of several factors such as topography, geology, Despite the geographical and biological vegetation communities, and climate (Thom and importance of the Interior Highlands (Allen Wilson 1980, Poulton and Stewart 1991, Foti 1983, 1990), few aquatic insect groups in this and Buckenhofer 1998). Named mountain ranges region have been fully inventoried (e.g., Schwardt in the Interior Highlands include the Arbuckle, 1936, McCafferty and Provonsha 1978, Poulton Ouachita, Ozark, and Wichita mountains of which and Stewart 1991, Bowles and Pinkovsky 1993, the Ozarks occupy the largest land area. The Moulton and Stewart 1996, Sarver and Kondratieff Ozarks are composed primarily of igneous rock 1997, Bowles 2006, Ferro and Sites 2007). The overlain by limestone and dolomite, which has insect order Megaloptera (Sialidae: alderflies, undergone substantial erosional processes to create Corydalidae: fishflies, dobonflies) is a relatively the current mountainous condition. By comparison, small group of physically striking aquatic insects the Arbuckle, Ouachita, and Wichita formations having aquatic larvae and terrestrial pupae and arose through orogeny with strong folding and adults. Megalopterans are important components faulting of sedimentary formations. Because the
Publication date: 29 December 2015 406 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Figure. 1. Interior Highlands geographic boundaries. ARB=Arbuckle Mountains, IOZ=Illinois Ozarks, OUA=Ouachita Mountains, OZR=Ozarks, and WIC=Wichita Mountains. in the structure and functioning of aquatic systems Tarter et al. 1978, Arnold and Drew 1987, Whiting through their predatory feeding habits, large body 1991, Penny et al. 1997, Engel 2004, Locklin sizes, and relatively rapid growth rates (Brown and 2008). The Interior Highlands is one region where Fitzpatrick 1978, Dolin and Tarter 1981, Epperson megalopteran assemblages are poorly documented and Short 1987, Short et al. 1987, Bowles 1990, and an exhaustive inventory of the species occurring Roell and Orth 1991, Phillips 1996, Contreras- there has not been previously conducted (Arnold Ramos 1998, Locklin et al. 2006). Although and Drew 1987, Bowles 1989, Bowles and Mathis megalopterans of eastern North America are well- 1992). A regional inventory of the Megaloptera described taxonomically (Davis 1903, Ross 1937, is an important step toward development of more Cuyler 1958, 1965, Flint 1964, 1965, Neunzig accurate water quality assessments, conducting 1966, Tarter et al. 1979, Stark and Lago 1980, other biodiversity and ecological studies, and for 1983, Bowles and Mathis 1992, Contreras-Ramos conservation planning. 1998, Rasmussen and Pescador 2002, Liu et al. Here we list the known species of Megaloptera 2015), species assemblages for most geographic occurring in the Interior Highlands and surrounding areas outside the eastern and southern United areas of Arkansas, Illinois, Missouri, and States are not well known (Smith 1925, 1934, Oklahoma, with an illustrated key to the species. We BOWLES and SITES 407 also briefly summarize the ecological information org/web/), also maintain copies of BugGuide.net relating to them in this region. taken at multiple instances in time. BugGuide.net shows several photographs MATERIALS AND METHODS of megalopterans that were taken at numerous locations in the U.S. The photographed specimens In addition to the distributional records we may not have been deposited in museums, may obtained from the published literature, we also no longer exist, and we did not physically exam examined specimens from the Illinois Natural them. However, the photographs are available for History Survey (INHS), Missouri State University future scrutiny, and they are of sufficient quality (MSU), Texas A&M University Insect Collection to allow identification of those specimens to the (TAMU), Oklahoma State University (OSU), species level. Several of the reported localities for Southern Illinois University (SIU), University of megalopterans listed on BugGuide.net represent Missouri (UMC), University of Arkansas Arthropod potential new state records for Interior Highlands. Museum (UAAM), and the collection of the senior Those records are included in this paper as author (DEB). Distributional records are presented supplemental records where appropriate. The as state and county data and include counties URL for each “page” with an image begins with bordering the Interior Highlands. Complete
species of Sialis presented here, but these characters 6 (5). Middorsal abdominal line yellowish or light were based primarily on color and pigment patterns. brown ...... C. pectinicornis (L.) Because these characters fade or become distorted ─ Middorsal abdominal line black ...... in preserved specimens, and pigment patterns were ...... C. rastricornis Rambur highly variable among the specimens we examined, 7 (5). Spiracles on abdominal segment 8 sessile (Fig. we have elected not to separate larvae of Sialidae 8) ...... Neohermes concolor (Davis) to species in this key. The following key was ─ Spiracles on abdominal segment 8 set on short, developed and modified from those of Ross (1937), conical tubes (Figs. 9-10) . . . . Nigronia-8 Stark and Lago (1983), and Arnold and Drew (1987). 8 (7). Respiratory tubes of abdominal segment 8 about as long as 9th segment and nearly Key to the MEGALOPTERA OF THE contiguous at their bases (Fig. 9) ...... INTERIOR HIGHLANDS ...... N. fasciatus (Walker) ─ Respiratory tubes of abdominal segment 8 1. Wingless, aquatic ...... larvae-2 shorter than 9th segment and widely separated at ─ Winged, terrestrial/adults9 their bases (Fig. 10) . . .N. serricornis (Say) 2 (1). Abdomen with 7 pairs of segmented lateral filaments and a single long caudal filament; anal 9 (1). Ocelli absent, fourth tarsal segment dilated prolegs absent (Fig. 2); 4th tarsomere enlarged and bilobed; length 10-15 mm . . Sialidae-10 and wider than remaining segments; ocelli ─ Ocelli present; fourth tarsal segment cylindrical; absen ...... Sialidae-3 length over 25 mm . . . . . Coydalidae-26
─ Abdomen with 8 pairs of non-segmented lateral 10 (9). Apex of abdomen with an ovipositor filaments (Figs. 5-6); apex of abdomen with 2 (females) ...... 19 anal prolegs, each bearing a pair of claws (Figs. ─ Apex of abdomen without an ovipositor and 7-10); 4th tarsomere cylindrical and about the variously modified with sclerotized processes same width as other segments; three ocelli . . (males) ...... 11 ...... Corydalidae-4 3 (2). Mandibles with three subapical teeth; third or 11 (10) Legs entirely black or brown or with tibiae basal tooth usually about one-half the size of the slightly lighter in color than femora; head other two teeth (Fig. 3) ...... Protosialis primarily yellow with some black markings ─ Mandibles with two subapical teeth (Fig. 4) . . (Figs. 11, 12) ...... Protosialis-12 ...... Sialis ─ Legs with femora reddish brown and tibiae the same color or blackish; head primarily black but 4 (2). Abdominal segments 1-7 with ventral gill with yellow or orange markings (Figs. 13, 14) tufts at base of lateral filaments (Fig. 5 ...... Sialis-13 . . . .Corydalinae, Corydalus cornutus L. ─ Abdominal segments without ventral gill tufts at 12 (11) Head punctured and dull in appearance. base of lateral filaments (Fig. 6) ...... Eleventh gonocoxite with processes ...... Chauliodinae-5 approximately as long as length of ninth gonocoxite in ventral view and roughly 5 (4). Respiratory tubes of abdominal segment 8 at parallel-sided for most of their length (Fig. 15); least twice as long as segment and unequal in processes of eleventh gonocoxite appearing length (Fig. 7) ...... Chauliodes-6 straight in lateral view (Fig. 16); ectoproct ─ Respiratory tubes of abdominal segment 8 nearly round in caudal view and with a small slightly longer than or shorter than segment but distal notch (Fig. 17) . P. americana (Rambur) equal in length, or sessile (Figs. 8-10) . . . 7 BOWLES and SITES 409
─ Head smooth and polished in apperance. approximately one-third its length (Figs. 36); Eleventh gonocoxite with processes ectoproct egg-shaped in caudal view (Fig. 37 . approximately as long as length of ninth ...... S. infumata Newman gonocoxite, and having the apical one-half ─ Ninth gonocoxite not produced into prominent markedly narrowed; processes of eleventh apicolateral knobs; basal lobe of median gonocoxite appearing gently arched in lateral processes of eleventh gonocoxite at least one- view (Fig. 19) ...... P. glabella (Ross) half its length ...... 18
13 (11). Ninth sternite produced posteriorly 18 (17) Basal lobe of eleventh gonocoxite markedly covering most of genitalia (Figs. 20); processes robust and generally thicker in comparison with of eleventh gonocoxite hook-shaped and distal lobe (Fig. 39-40); ectoproct shorter than approximately same length as tenth tergite in parasagittal processes of eleventh gonocoxite lateral view (Fig. 21); tenth tergite pear-shaped are high in lateral view (Fig. 40) and heart- in caudal view and with a small distal notch shaped in caudal view (Fig. 41) ...... (Fig. 22) ...... S. vagans Ross ...... S. itasca Ross ─ Ninth sternite not produced posteriorly and not ─ Basal lobe of parasagittal arm of eleventh covering most of genitalia ...... 14 gonocoxite small and about the same thickness as distal lobe (Fig. 42-43); ectoproct as long as 14 (13). Eleventh gonocoxite short and horn-like, median processes of eleventh gonocoxite are triangular in shape and distinctly shorter than high (Fig. 44) and spade–shaped in caudal view length of ninth gonocoxite in ventral view (Figs. (Fig. 45) ...... S. velata Ross 24); processes of eleventh gonocoxite about one-third as long as ectoproct in lateral view 19 (10). Legs entirely black or brown or with (Fig. 25); ectoproct broadly pentagonal in shape tibiae slightly lighter in color than femora; head (Fig. 26) ...... S. joppa Ross primarily yellow (Figs. 11-12) ...... ─ Eleventh gonocoxite produced into a pair of ...... Protosialis …20 long median processes or lobes (Figs. 29, 36) . ─ Legs with femora reddish brown and tibiae the ...... 15 same color or blackish; head primarily black but with yellow or orange markings (Figs. 13-14) . 15 (14). Ectoproct produced into prominent apical ...... Sialis…21 processes (Figs. 29, 32) ...... 16 ─ Ectoproct not produced into prominent apical 20 (19). Head punctured and dull in appearance. processes (Figs. 36, 40, 43) ...... 17 Female terminalia with gonocoxite 8 rectangular, weakly sclerotized; eighth 16 (15). Lobes of eleventh gonocoxite shorter than gonapophyses diamond-shaped and weakly apical arms of ectoproct and with the basal lobe scelerotized (Fig. 18) ...... distinct (Figs. 28-29); apical arms of ectoproct ...... P. americana (Rambur) separated at their bases by a distance almost ─ Head smooth and polished in appearance. equal to their length (Fig. 30) . . . S. hasta Ross Terminalia of female indistinguishable from that ─ Lobes of eleventh gonocoxite as long or longer of . . . S. americana . . . P. glabella (Ross) than apical arms of ectoproct and without a distinct basal lobe (Figs. 31-32); apical arms of 21 (19). Eighth gonapophyses large, triangular- to ectoproct close together at base and converging diamond-shaped, distinctly sclerotized (Fig. 34) at tip (Fig. 33) ...... S. mohri Ross ...... S. mohri Ross ─ Eighth gonapophyses not triangular- or 17 (15). Ninth gonocoxite produced into prominent diamond-shaped, appearing membranous and apicolateral knobs (Fig. 35); basal lobe of difficult to distinguish in uncleared specimens median process of eleventh gonocoxite ...... 22 410 MEGALOPTERA OF THE INTERIOR HIGHLANDS
22 (21). Eighth gonocoxite divided, each side 55). Female ovipositor largely membranous roughly oval in shape; eighth gonapophyses and with a conspicuous lateral sclerite on ninth small and located between the two lobes of the gonocoxite (Fig. 56) ...... eighth goncoxite (Fig. 23) . . S. vagans Ross . . . . Corydalinae, Corydalus cornutus L. ─ Eighth gonocoxite not divided, or if divided, ─ M vein of forewing with 2 branches reaching then they are not oval-shaped; eighth the wing margin; wing maculations absent or gonapophyses prominent ...... 23 covering large areas of wings (Figs. 47-51); males with mandibles always shorter than 23 (22). Eighth gonocoxite flat, not tilted so that length of head ...... Chauliodinae-27 the apical margin is deflexed; bursa copulatrix prominent and with a terminal fingerlike 27 (26). Wings black, patterned with white (Figs. process (Fig. 45) ...... 50-51) ...... Nigronia-28 ...... S. itasca Ross . . ..S. velata Ross ─ Wings gray or brown, without white (Figs. 47- ─ Eighth gonocoxite with apex rounded, or 49) ...... 29 with distal portion of apical margin without conspicuous lobes; bursa copulartrix without a 28 (27). Anal area of hind wing white (Fig. 50). terminal fingerlike process ...... 24 Male with gonoxoxite of sternite10 rounded distally; apex of gonocoxite of tergite 9 truncate 24 (23). Eighth gonocoxite with a small round and forming a complex with the ectoproct (Fig. depression apically on meson; bursa copulatrix 70) ...... N. fasciatus (Walker) prominent, broadly triangular in shape and with ─ Anal area of hind wing black (Fig. 51). Male the tip entirely membranous (Fig. 27) . . . . with gonocoxite 9 truncate distally; apex of ...... S. joppa Ross gonocoxite 10 rounded (Fig. 71)...... ─ Eighth gonocoxite with a narrow V-shaped ...... N serricornis (Say) depression extending along the meson of the Anal area of hind wing black (Fig. 51). Male entire segment and with the distal portion of with gonocoxite of sternite 10 truncate distally; the apical margin conspicuously swollen and apex of gonocoxite of tergite 9 rounded and appearing as lobes; bursa copulatrix not broadly forming complex with ectoproct (Fig. 71) . . . . . triangular in shape ...... 25 ...... N serricornis (Say)
25 (24). Seventh sternite entirely sclerotized, apical 29 (27). Antennae pectinate or serrate. Male margin entire; eighth gonocoxites divided and with apex of sternum 9 with spinules on broadly rectangular (Fig. 38) ...... inner margins (Figs. 59, 61). Apex of female ...... S. infumata Newman gonocoxite 9 smoothly rounded in lateral view ─ Seventh sternite with only apical half (Fig. 63) ...... Chauliodes-30 sclerotized and with apical margin shallowly ─ Antennae moniliform. Male aedeagus and emarginated ...... S. hasta Ross ectoproct highly variable in shape, sometimes with grossly expanded dorsal membrane (Figs. 26 (9). M vein of forewing with 3 branches 64-68). Apex of female gonocoxite 9 with reaching the wing margin and with evenly slender finger-like processes (Fig. 69) . . . . maculated small white spots surrounded by ...... Neohermes concolor (Davis) uniform gray of the wing membrane (Fig. 46); males usually with prominent mandibles 30 (29). Elongate spots on vertex of head posterior approximately one-half to two-thirds body to ocelli yellow to light brown (Fig. 57); length (Fig. 52), those of female short but antennae generally pectinate in both sexes. prominently toothed (Fig. 53). Male terminalia Ectoproct of male with small spicules on inner with ninth gonostyli elongate and approximately margins (Fig. 59); lateral aspect cylindrical (Fig. three-fourths as long as ectoproct (Figs. 54- 60) ...... C. pectinicornis (L.) BOWLES and SITES 411
─ Elongate spots on vertex of head posterior to Protosialis glabella (Ross) ocelli dark brown to black (Fig. 58); antennae Fig. 19 pectinate (male) or serrate (female). Ectoproct of males with prominent tooth and small This species previously was placed in the spicules on inner margins (Fig. 61); lateral genus Sialis, but it was reassigned to Protosialis aspect triangular (Figs. 62) ...... by Liu et al. (2015). Ross (1937) described the ...... C. rastricornis Rambur species from specimens collected in the Wabash River system in southern Illinois. The holotype Family Sialidae and allotype were collected at Mt. Carmel, Wabash County, Illinois located 214 km south of Muncie, Protosialis americana (Rambur) Vermillion County, Illinois where the paratypes Figs. 3, 11-12, 15-18 were collected. In addition to the type specimens, Canterbury (1978) reported a population of P. Liu et al. (2015) reassigned Sialis americana to glabella from a woodland pond in Nelson County, Protosialis americana. This uncommonly collected Kentucky, and Stark and Lago (1980) reported a species is distributed primarily in the eastern U.S. single male at Natchez, Adams County, Mississippi. (Ross 1937, Whiting 1991) and has been collected These few widely-scattered distributional records only from Illinois and Missouri in the Interior suggest this rare species may be broadly distributed Highlands. The species previously had not been in the Mississippi River drainage. It is most closely reported from Illinois, but its occurrence there was related to P. americana with which it can be expected. Missouri may represent the westernmost confused. However, adults of P. glabella have the distributional boundary for P. americana. This head smooth and polished in appearance rather than species is most closely related to S. glabella, but punctured and dull as in P. americana, and on the adults of P. americana can be distinguished from it basis of male genitalia. Adults have been collected in having the head punctured and dull in appearance in May and June. rather than smooth and polished, and on the basis of male genitalia. Adults have been collected during Illinois: [Vermillion, Wabash counties] (Ross April through June. 1937, Whiting 1991).
Illinois [new state record]: Union Co., Alto Sialis hasta Ross Pass, 14.iv.1963, T. F. Hartline, 1 female (SIU). Figs. 28-30 Missouri: Non-specific (Whiting 1991); Barton Co., Shawnee Trail C.A., 7.vi.1999, Whiteman and This species has been collected only from a few Landwer, blacklight, 1 male (UMC); Greene Co., scattered locations in Arkansas and Missouri in the 4.6 km SW Strafford, Lovett Pinetum, McKerral Interior Highlands. It can be distinguished from Spring, 37o 14’ 15.83” N, 93o 09’ 7.19” W, congeners presented here in having the apical arms 9.vi.2012, UV-light, D. E. Bowles, 1 male; Henry of the ectoproct greatly elongated and separated Co., Ferro farm, 10.vi.2001, M. Ferro, blacklight, 1 basally by a distance roughly equal to their length. female (UMC); Johnson Co., Knob Noster St. Pk., Adults have been collected in April and May. 7.vi.1979, E. G. Riley, blacklight; 1 female (UMC); Shelby Co., Crooked Creek, T56N, R11W, Sec. 6, Arkansas: Garland Co. (Flint 1964, Bowles 4.iv. 2001, E. Kosnicki, 1 female (UMC); Taney 1989), Montgomery Co. (Flint 1964); non-specific Co., Bull Shoals Field Station, small spring/pond, (Tarter et al. 1978); Washington, Yell counties 36o 33′ 59.46″ N, 93o 05′ 41.71″ W, 14.vi.2009, D. (Bowles 1989). Missouri: no specific (Tarter et E. Bowles, UV-light (DEB); same, but Malaise trap, al. 1978); Greene Co. (Stark and Lago 1980); 24-31.v.2012, 1 female (DEB). [Callaway Co.] (UMC). 412 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Sialis infumata Newman features of both species. Figs. 13-14, 35-38 Arkansas: Craighead Co. (Tarter et al. 1978). Sialis infumata is widely distributed throughout Missouri: [Miller Co.] (Ross 1937); Boone, Cass the Interior Highlands. The knob-like projections of counties (UMC). Illinois: [Coles, Kankakee, Ogle, the male ninth gonocoxites readily distinguish this Piatt counties] (Ross 1937); Kankakee Co. (Whiting species from other sialids occurring in the region. 1991); Iroquois Co. (INHS). Oklahoma: non- We have records of adults from April through specific(Tarter et al. 1978); Murray, Noble, Pawnee June and September, and from both streams and counties (Arnold and Drew 1987), and Payne Co. impounded waters. We have collected this species (Arnold and Drew 1987, Whiting 1991). syntopically with S. velata in Missouri and the April to May flight periods of the two species are similar. Sialis joppa Ross Figs. 24-27 Arkansas: Washington Co. (Tarter et al. 1978, Bowles 1989, UAAM); Garland, Pike counties Although widely distributed in the Interior (Bowles 1989, UAAM). Ilinois: [Champaign, Highlands, this species is seldom collected. Jo Daviess, Lake, Mason, McHenry, Vermillion, The collection of S. joppa in Missouri is a new Will counties] (Ross 1937); Champaign, Cook, Jo distributional record for the state. This species Davies, Kankakee, McHenry counties (Whiting is distinct from all other species addressed in 1991). Missouri: non-specific (Tarter et al. 1978); this study; it may be diagnosed by the small, Barry, Boone, Johnson, Macon, Maries, Miller, triangular shaped eleventh gonoxite, which is Pike, Pulaski, Reynolds, Texas, Vernon counties distinctly shorter than the ninth gonocoxite in (UMC); Christian, Greene, Stone counties (DEB, ventral view. Adults of S. joppa have been collected MSU); Newton Co. (DEB, UMC).Oklahoma: during April and May and primarily from small Pittsburg Co. (Stark and Lago 1980); Payne Co. woodland streams and spring systems. Bowles (Arnold and Drew 1987). (1989) collected adults of S. joppa from a small woodland spring in Arkansas where the annual Sialis itasca Ross water temperature was 15oC (range of 10.0-21.0oC), Figs. 39-41 the substrate was covered with a deep layer of detritus, and there was little observable current This species is widely distributed throughout velocity (Bowles 1998). This spring also had copius the Interior Highlands. It has the broadest amounts of ferric hydroxide precipitate that colored flight period of any species ofSialis occurring the water orangish red. Mather (1981) and Pittman in the Interior Highlands, and adults of have et al. (1996) reported the occurrence of larval S. been collected every month from April through joppa in carnivorous pitcher plants in New Jersey September. Sialis itasca primarily occurs in lotic and West Virginia, respectively. It is not known if habitats but also can be found in association with the larvae actively use these plants as habitat, or if impounded waters. Sialis itasca can be easily the association was adventitious. confused with S. velata. The basal lobes of the eleventh gonocoxite in males of S. itasca vary Arkansas: Crawford, Montgomery counties considerably in thickness within and among (Bowles 1988); Benton (DEB), Fulton Co. (UMC). populations we have examined. However, this lobe Missouri: [new state record]- Carter Co., Ozark is never as thin as that observed for S. velata, which National Scenic Riverways, Big Spring, 20.iv.2011, usually is about the same thickness as the distal D. E. Bowles, at rest, 1 male, 3 females (DEB); lobe or thinner. In S. itasca, the basal lobe is always Crawford Co. 9.v.1982, R. Morris, 1 female, same thicker than the distal lobe. Locklin et al. (2006) data, but Wurdack Farms, 25.v.2001, E. Kosnicki, indicated that where their ranges overlap, these two 1 male (UMC); Phelps Co., Meramec Pk. nr. St.; species may produce hybrids with some genitalic James, 8.vi.1961, Wilbur R. Enns, 1 male (UMC); BOWLES and SITES 413 same data, but Meramec Springs, St. James, Sialis vagans Ross 8.vi.1961, Clark, Neal, Enns, 1 male (UMC); Pike Figs. 20-23 Co., 16.v.1963, R. Morris, 1 male (UMC); Taney Co., Bull Shoals Field Station, small spring/pond, This species has been collected from only a few 36o 33′ 59.46″ N, 93o 05′ 41’.71″ W, 3-10.v.2011, locations in the Interior Highlands. Our collection D. E. Bowles, Malaise trap, 1 female (DEB). from Texas County, Missouri, represents a new Illinois: non-specific (Ross 1937,Whiting 1991). state record for this species. Adults have been Oklahoma: Latimer Co. (Bowles 1989). collected in March, May and June. The prominent flap-like ninth sternite of the male distinguishes Sialis mohri Ross this species from all other sialids occurring in the Figs. 31-34 Interior Highlands. We have collected S. vagans from a variety of aquatic habitats, but they seem to Sialis mohri is the most common and favor small streams and backwaters. widespread sialid occurring in the Interior Highlands. The male genitalia of this species are Arkansas: Greene Co. (Tarter et al. 1978, distinct from those of all other species occurring Bowles 1989); Johnson Co. (Bowles 1989). Illinois: in the Interior Highlands. The apical arms of the [Cook, Logan, Mason counties] (Ross 1937); ectoproct are long, close together at their base, and Cook, Cumberland, DuPage, Franklin, Kankakee, converging at their apex. The lobes of the eleventh McHenry, Pope counties (Whiting 1991); Jackson gonocoxite also are greatly elongated and as long as Co. (SIU). Missouri [new state record]: Texas Co., the apical arms of the ectoproct. We have collection Ozark National Scenic Riverways, oxbow of Jacks data for this species from a wide variety of habitats Fork, 0.3 km upstream of Buck Hollow Bridge, including small streams, large rivers, farm ponds, 37o 03′ 13.30″ N, 91o 40′ 09.98″ W, elev. 282 m, and large lakes. Adults have been collected from 9.v.2014, D. E. Bowles, at rest, 1 male, 2 females March through June. (DEB). Oklahoma: Latimer Co. (Bowles 1989).
Arkansas: Montgomery (Flint 1964, Bowles Sialis velata Ross 1989); Pike, Washington counties (Bowles 1989, Figs. 42-45 UAAM); Clark, Conway counties (UAAM). Illinois: [Brown, Cook, DuPage, Kankakee, This species has a sporadic distribution in Lake, Mason, Marshall, McHenry, Morgan, Rock the Interior Highlands and is not known from Island, Washington, Will, Winnebago counties] Arkansas. This species can be easily confused with (Ross 1937); Champaign, Cook, Coles, DuPage, S. itasca. In males, the basal lobe of the eleventh Edgar, Jackson, Kankakee, Lake, Marshall, Mason, gonocoxite is about the same thickness of the distal Massac, McHenry, Montgomery, Morgan, Rock lobe or thinner, whereas in S. itasca this lobe is Island, Washington, Williamson, Winnebago always thicker than the distal lobe. Adults of this counties (Whiting 1991); Williamson Co. (Whiting uncommon species have been collected during 1991, SIU); Jackson, Marion, Union counties April and May, and similar to S. itasca, it seems (SIU). Missouri: non-specific (Tarter et al. 1978); to prefer lotic systems although it does well in Boone, Callaway, Howard, Jefferson, Macon, impoundments. Randolph, counties (UMC); Taney Co. (DEB); Franklin Co. (SIU). Oklahoma: [Payne Co.] (Flint Illinois: [Champaign, Coles, Gallatin, Johnson, 1964); non-specific (Tarter et al. 1978); Pawnee, Lee, Logan, Massac, Mason, Rock Island, Wabash, Payne, Pittsburg, and Tulsa counties (Arnold and Washington, White counties] (Ross 1937); Drew 1987). Champaign, Coles, Gallatin, Jersey, Johnson, Madison, Pope, Rock Island, Wabash, Washington, White counties (Whiting 1991); Jackson, Williamson counties (SIU). Missouri: [Boone 414 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Co.] (Ross 1937); Boone Co. (UMC); Greene Co. Contreras-Ramos 1998; UAAM); Arkansas Co. (DEB). Oklahoma: Garfield Co. (Whiting 1991). (Chordas et al. 1996); Sharp Co. (Moss and Harp 1993); Fulton, Lawrence, Sharp counties (Harp Sialis spp. and Robison 2006); Stone Co. (Contreras-Ramos 1998); Garland, Marion, Washington counties We examined additional larval Sialis, but these (Contreras-Ramos 1998, UAAM); Garland Co. specimens could not be identified to species. These (DEB); Arkansas, Drew, Hempstead, Hot Springs, records are indicated here for the purpose of aiding Lincoln, Logan, Madison, Monroe, Newton, Sevier further distributional studies of this group in the counties (UAAM). Illinois: Champaign, Mason, region. McHenry, Moultrie, Pope, Rock Island, Will, Winnebago counties (Contreras-Ramos 1998); Ogle Arkansas: Crawford Co. (DEB). Illinois: Co. (Contreras-Ramos 1998, INHS); Cook, Kane, Jackson, Union counties (SIU). Missouri: Bates, Knox, Lee, Mason, Vermillion counties (INHS). Barry, Benton, Boone, Callaway, Camden, Carter, Missouri: non-specific (Davis 1903, Sullivan Cass, Cedar, Christian, Cole, Crawford, Dade, 1929), Niangua River [Webster, Dallas, Laclede, Dallas, Dent, Douglas, Dunklin, Gasconade, Camden counties]; Howell, Taney, Barry, Ozark, Franklin, Harrison, Henry, Holt, Iron, Jasper, McDonald, Polk, Shannon, Reynolds, Franklin, Jefferson, Laclede, Lawrence, Livingston, Macon, Wayne, Madison, Texas, Douglas, Laclede, Phelps, Madison, Maries, Morgan, McDonald, New Gasonade, St. Francois counties (Harp et al. 2008); Madrid, Newton, Oregon, Osage, Phelps, Polk, Gasconade, Newton, Ozark, Pulaski counties Pulaski, Ray, Reynolds, Ripley, Shannon, St. Clair, (Contreras-Ramos 1998); Reynolds, Shannon, St. St. Francois, St. Genevieve, St. Louis, Stone, Louis, Washington counties (Contreras-Ramos Taney, Texas, Vernon, Warren, Washington counties 1989, UMC), Boone, Callaway, Camden, Carter, (UMC), Green Co. (DEB, UMC). Chariton, Christian, Cole, Cooper, Crawford, Dallas, Dunklin, Howard, Jefferson, Mississippi, Family Corydalidae Pike, Putnam, Taney counties (UMC). Oklahoma: Subfamily Corydalinae Bryan, Carter, Delaware, Johnston, Kay, LeFlore, Mayes, McClain, Nowatta, Ottawa, Pontotoc, Corydalus cornutus L. Pushmataha, and Washita counties (Arnold and Figs. 5, 46, 52-56 Drew 1987); Canadian, [Carter], [Comanche], Latimer, Oklahoma, Osage counties (Contreras- Corydalus cornutus is common and widespread Ramos 1998); Adair, Cherokee, Choctaw, throughout the Interior Highlands. Contreras- McCurtain, Murray, Payne counties (Arnold and Ramos (1998) reported this species from numerous Drew 1987, Contreras-Ramos 1998). counties in Arkansas, Missouri and Oklahoma. Adults have been collected from May through Corydalus sp. October. We examined two male specimens collected in Missouri during early October, and a We have collected larval Corydalus from single female specimen collected during September numerous locations throughout the Interior in Arkansas. Larvae of C. cornutus are commonly Highlands. Although these specimens are probably found under rocks in clear, flowing streams, but C. cornutus, we were unable to make a specific we have also collected specimens of this species determination. Because additional species of on woody snag habitat in stream pools. Phillips Corydalus occur in North America (Contreras- (1996) reported that C. cornutus occurred in benthic Ramos 1998), the possibility exists that some of habitats at densities roughly four times greater than these larvae may not be C. cornutus. those on coarse woody debris. Arkansas: Benton, Clark, Columbia, Arkansas: Johnson Co. (Bowles 1989, Montgomery, Pike, Sebastian (UAAM). Illinois: BOWLES and SITES 415
McHenry, Pope, Pulaski, Union, Williamson Montgomery, Oregon, Reynolds, Ripley, St. Louis, counties. Missouri: Barry, Bates, Benton, Bollinger, Vernon, Warren counties (UMC); Christian Co. Boone, Callaway, Camden, Cape Girardeau, (DEB). Oklahoma: [new state record] Logan Co., Carter, Cass, Cedar, Christian, Cole, Crawford, 4 mi S Guthrie, 18.viii.1998, blacklight trap, R. Dade, Dallas, Douglas, Franklin, Gasconade, Grantham, 1 female (OSU); Mayes Co., Locust Greene, Henry, Hickory, Holt, Illinois, Iron, Grove, 30.viii.1989, D. C. Arnold, blacklight, 1 Jasper, Jefferson, Laclede, Lawrence, Livingston, male (OSU). Madison, Maries, McDonald, Miller, Monroe, Morgan, Newton, Oregon, Osage, Perry, Phelps, Chauliodes rastricornis Rambur Polk, Pulaski, Ralls, Reynolds, Ripley, Shannon, St. Figs. 49, 58, 61-63 Charles, St. Clair, St. Francois, St. Genevieve, St. Louis, Stone, Texas, Vernon, Washington, Wayne This species has a widespread distribution in counties (UMC). North America including the Interior Highlands and it is commonly collected from a variety of Subfamily Chauliodinae aquatic habitats. The black elongate spots on the vertex of adults and the black middorsal abdominal Chauliodes pectinicornis (L.) stripe of the immatures distinguish this species Figs. 48, 57, 59-60 from its congener, C. pectinicornis. Adults of C. rastricornis have been collected from March This species is widely distributed in the Interior through September. Tarter et al. (1976) reported Highlands, but it had not been previously reported that the larvae of C. rastricornis are found in lotic from Oklahoma. Adults have been collected from and lentic habitats including ponds, swamps, and May through September throughout the range of marshes. We have also collected larvae of this this species. The light colored elongate spots on species from small springs and seeps. the vertex of adults and the yellowish middorsal abdominal stripe of the immatures distinguish this Arkansas: non-specific (Tarter et al. 1976); species. Cuyler (1958) and Tarter et al. (1976) Arkansas Co. (Chordas et al. 1996); Desha, reported that the larvae of C. pectinicornis primarily Franklin, Pulaski, Washington counties (UAAM); occur in lentic habitats, including ponds, marshes, Ashley, Logan counties (UAAM, DEB); Drew, and swamps, and our collections were made from Montgomery, St. Francis counties (DEB); Johnson similar habitats. Co. (DEB, OSU). Supplemental Record: Faulkner Co., Conway, Arkansas, 24.iii.2007, 1 male Arkansas: non-specific (Tarter et al. 1976); [BugGuide 99375].Illinois: non-specific (Tarter et Fulton, Sharp counties (Harp and Robison 2006); al. 1976); Massac, Franklin, Jackson, Perry, Pope, Douglas, Reynolds counties (Harp et al. 2008); Union, Williamson counties (SIU). Missouri: non- Benton, Clark, Crawford, Desha, Washington specific (Davis 1903); Boone, Clinton, Douglas, counties (UAAM); Johnson Co. (OSU); Franklin, Dunklin, Howard, Jackson, Johnson, Lincoln, Linn, Montgomery counties (DEB). Supplemental Macon, Montgomery, Randolph, Vernon counties Record: Searcy Co., Ozark Mountains, 21.v.2008, (UMC); Cedar, Jefferson, Stoddard counties 1 male [BugGuide 184222-184223]. Illinois: (MSU); Newton Co. (DEB); Wayne Co. (SIU). Carroll, Champaign, DeWitt, Hardin, Marshall, Supplemental Records: Franklin Co., Shaw Nature Piatt, Vermillion counties (Tarter et al. 1976); Cook, Reserve, 4.ix.2011, 1 male [BugGuide 572446]; Madison counties (INHS); Alexandria, Jackson, Jackson Co., Blue Springs, September 13.ix.2008, 1 Pope, Union, Wayne, Williamson counties (SIU). male [BugGuide 228164]. Oklahoma: non-specific Missouri: non-specific (Davis 1903,Tarter et al. (Tarter et al. 1976); Alfalfa, Carter, Cherokee, 1976); Audrain, Benton, Boone, Callaway, Camden, Jackson, Ottawa, Payne, Rogers, and Tulsa counties Carter, Cole, Crawford, Dallas, Dent, Douglas, (Arnold and Drew 1987); Latimer Co. (UAAM); Dunklin, Henry, Lincoln, Miller, Mississippi, Logan, Mayes, McCurtain, Oklahoma, Pushmataha 416 MEGALOPTERA OF THE INTERIOR HIGHLANDS counties (OSU). Supplemental Records: Okmulgee Washington counties (Bowles and Mathis 1992); Co., Henryetta , 10.v.2013; found while collecting Montgomery, Newton, Yell counties (UAAM). an arrowhead plant, 1 female. [BugGuide 768787]; Illinois: Pope, Union counties (Tarter et al. Johnston Co., Tishomingo National Wildlife 1976, INHS, SIU); Jackson, Wayne, Williamson Refuge, 28.iv.2012, 1 female [BugGuide 645268]; counties (SIU). Missouri: Greene Co. (Flint Osage Co., Sand Springs, 19.iv.2009, 1 male 1965); no data (Tarter et al. 1976); Pulaski, [BugGuide 267215]; same, but 27.iv.2008, 1 male Randolph counties (Bowles and Mathis 1992); [BugGuide 179282]; Okmulgee Co., Deep Fork Boone, Butler, Callaway, Christian, Cole, Dent, River, 22.iv.2006, 1 male, 1 female [BugGuide Douglas, Henry, Jefferson, Lincoln, Randolph, 48973, 48972]; Oklahoma Co., Nicoma Park, Ozark counties (UMC); Taney Co. (DEB, UMC); 24.ix.2003, 1 male [BugGuide 229717- 229719, Dallas Co. (DEB). Supplemental Record: Franklin labeled as C. pectinicornis]. Co., 12.viii.2012, 1 male [BugGuide 729109]. Oklahoma: Latimer Co. (Tarter et al. 1976, Bowles Chauliodes spp. and Mathis 1992); Choctaw, Johnston, LeFlore, McCurtain, Osage, Payne counties (Arnold and We collected several specimens of Chauliodes Drew 1987). from Missouri that could not be identified to species because they were poorly preserved or damaged. Nigronia fasciatus (Walker) Those distributional records are presented here to Figs. 9, 50, 70 illustrate the potential distribution of the respective species. This species seldom has been collected in the Interior Highlands. Although we have no collection Missouri: Christian, Crawford, Iron, Jefferson, records of N. fasciatus from Oklahoma, it may Lawrence, St. Francis, Warren, Washington occur in the eastern portion of the state. Larvae of counties (UMC). N. fasciatus are readily distinguished from those of N. serricornis in having the abdominal respiratory Neohermes concolor (Davis) tubes close together basally and as long as the 9th Figs. 6, 47, 64-69 segment compared to widely separated and shorter than the 9th segment in the latter species. The This species is widely distributed in the Interior wings of this species have considerably more white Highlands. Bowles and Mathis (1992) reported a markings than do those of N. serricornis. Adults wide range of variation in the male terminalia of have been collected only during June from the this species from throughout its range, particularly Interior Highlands. The larvae of N. fasicatus prefer in the size and shape of the ectoprocts, but this small, low-gradient woodland streams often with variation was not geographically coordinated. rocky substrates (Cuyler 1965, Neunzig 1966). Roughly one-half of the specimens collected by Bowles and Mathis (1992) had grossly expanded Arkansas: non-specific (Davis 1903, Tarter et genital membranes. Adults of this species have al. 1976). Illinois: non-specific (Tarter et al. 1976); been collected from May through August. Bowles Jackson, Pope, Union counties (SIU). Missouri: and Mathis (1992) reported N. concolor larvae non-specific (Davis 1903, Tarter et al. 1976); from a variety of small springs, spring-run and Carter, Jefferson, Phelps, Shannon, Wright counties seep habitats, including a small spring located (UMC). well within a wet cave. This species also occurs in temporary, spring-fed streams that become dry Nigronia serricornis (Say) during periods of drought. Figs. 10, 51, 71-72
Arkansas: Hot Springs, Lawrence counties This species is widely distributed and common (Flint 1965); Crawford, Garland, Pulaski, in Interior Highland streams. A report of N. BOWLES and SITES 417 fasciatus on BugGuide (2014) from McCurtain ACKNOWLEDGEMENTS County, Oklahoma actually shows a representative of N. serricornis, which represents a new We thank the numerous people who provided distributional record for this species. Although we us with material or assisted us during this study did not physically examine this specimen, we could including Jay McPherson (Southern Illinois easily determine its identity from the photograph University), Chris Carlton (Louisiana State provided. Larvae are easily distinguished in having University), Mike Ferro (Clemson University), the abdominal tubes widely spaced basally and Bob Lovett (Lovett Pinetum), Bob Kipfer they are shorter than the 9th segment. Unlike N. (Springfield, Missouri), Steve Jensen (Missouri fasciatus, the wings of this species are largely State University), Ely Kosnicki (College of black with small, white maculations. This species Brockport, New York), Karl Stephan (late, Latimer probably occurs throughout eastern Oklahoma. Co., Oklahoma), Mike Mathis (late, University Adults have been collected from May through July. of Central Arkansas), Ed Riley (Texas A&M We have found the larvae of N. serricornis only in University), Jeff Barnes (University of Arkansas), lotic habitats where they occur almost exclusively Richard Leschen (New Zealand Landcare Research, in riffles and glides with coarse substrates. Cara Rowbotham (University of the Ozarks), Donald Arnold (Oklahoma State University), and Arkansas: non-specific (Hagen 1861, Tarter Robert T. Allen (Mississippi State University). We et al. 1976); Lawrence Co. (Harp and Robison also thank the American Entomological Society, 2006); Barry, Taney, Madison, Texas counties (Harp the Entomological Society of America, the Central et al. 2008); Crawford, Garland, Johnson, Polk States Entomological Society, and the Illinois counties (DEB), Washington Co. (DEB, UAAM); Natural History Survey for giving us permission to Montgomery Co. (UAAM). Supplemental Record: reproduce previously published illustrations. Newton Co., Buffalo National River, 19.v.2007, 1 female [Bugguide 111408, 111410]. Illinois: non- REFERENCES specific (Tarter et al. 1976); Monroe, Vermillion, Winnebago counties (INHS); Pope, Union counties Allen RT (1983) Distributional patterns of among (INHS, SIU); Jackson, Williamson counties (SIU). arthropods of the North Temperate Deciduous Missouri: non-specific (Hagen 1861,Tarter et al. Forest biota. Annals of the Missouri Botanical 1976); Barry, Bates, Bollinger, Camden, Carter, Garden70: 616-628. Cole, Crawford, Dade, Dallas, Dent, Douglas, Allen RT (1990) Insect endemism in the Interior Franklin, Gasconade, Greene, Iron, Jasper, Laclede, Highlands of North America. Florida Lawrence, Madison, Maries, McDonald, Miller, Entomologist 73: 539–569. Morgan, Phelps, Polk, Pulaski, Reynolds, Ripley, Arnold DC, Drew WA (1987) A preliminary survey St. Clair, St. Francois, Texas, Warren, Washington, of the Megaloptera of Oklahoma. Proceedings Wayne, Wright counties (UMC); Benton Co. of the Oklahoma Academy of Science 67: 23- (INHS, UMC); Christian, Stone counties (UMC, 26. MSU); Ozark Co. (MSU); Oregon, Shannon Aspöck U, Aspöck H (2008) Phylogenetic relevance counties (DEB, INHS, SIU, UMC); St. Genevieve of the genital sclerites of Neuropterida (Insecta: (INHS, SIU, UMC). Supplemental Record: Holometabola). Systematic Entomology 33: McCurtain Co., Ouachita National Forest, 6.v.2011, 97-127. D. Arbour, flying around a spring seep in a pine/ Baker JR, Neunzig HH (1968) The egg masses, hardwood forest, 1 male [514563, labeled as eggs, and first instar larvae of eastern North Nigronia fasciatus]. American Corydalidae. Annals of the Entomological Society of America 61: 1181- 1187. 418 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Bowles DE (1989) New records of Sialis larvae of sialid Megaloptera of east central (Megaloptera: Sialidae) from Arkansas and North Carolina with keys to and descriptions Oklahoma. Entomological News 100: 27-28. of larvae of genera known to occur in the Bowles DE. (1990) Life history and variability of United States. MS thesis, North Carolina State secondary production estimates for Corydalus College, Raleigh, North Carolina 150 pp. cornutus (Megaloptera: Corydalidae) in Cuyler RD (1958) The larvae of Chauliodes an Ozark stream. Journal of Agricultural Latreille (Megaloptera: Corydalidae). Annals Entomology 7: 61-70. of the Entomological Society of America 51: Bowles DE (1998) Life history of Bittacomorpha 582-586. clavipes (Fabricius) (Diptera: Ptychopteridae) Cuyler RD (1965) The larva of Nigronia fasciatus in an Ozark spring, U.S.A. Aquatic Insects 20: Walker (Megaloptera: Corydalidae). 29-34. Entomological News 76: 192-195. Bowles DE (2006) Spongillaflies (Neuroptera: Davis KC (1903) Sialidae of North and South Sisyridae) of North America with a key to the America. Bulletin 68. New York State Museum larvae and adults. Zootaxa 1357: 1-19. of Entomology 18: 442-486. Bowles DE, Mathis ML (1992) Variation in the Dolin PS, Tarter DC (1981) Life history and terminalia of Neohermes concolor with a ecology of Chauliodes rastricornis Rambur key to the males in eastern North America and C. pectinicornis (Linnaeus) (Megaloptera: (Megaloptera: Corydalidae: Chauliodinae). Corydalidae) in Greenbottom Swamp, Cabell Insecta Mundi 6: 145-149. County, West Virginia. Brimleyana 7: 111-120. Bowles DE, Pinkovsky DD (1993) Occurrence of Engel MS (2004) The alderflies of Kansas larval black flies and horse flies in an Ozark (Megaloptera: Sialidae). Transactions of the headwater stream. Southwestern Naturalist 38: Kansas Academy of Science 107: 119-125. 86-87. Epperson CR, Short RA (1987) Annual production Brown AV, Fitzpatrick LC (1978) Life history of Corydalus counutus (Megaloptera) in the and population energetics of the dobson fly, Guadalupe River, Texas. American Midland Corydalus cornutus. Ecology 59: 1091-1108. Naturalist 118: 433-438. BugGuide (2014) Available from: http://www. Fenneman NM, Johnson DW (1946) Physical bugguide.net (accessed 1 May 2014). divisions of the United States. US Geological Canterbury LE (1978) Studies on the genus Sialis Survey (USGS), map. Washington, DC. (Sialidae: Megaloptera) in eastern North Ferro ML, Sites RW (2007) The Ephemeroptera, America. Ph D dissertation, University of Plecoptera, and Trichoptera of Missouri Louisville, Louisville, Kentucky 186 pp. state parks, with notes on biomonitoring, Canterbury LE, Neff SE (1980) Eggs of Sialis mesohabitat associations, and distribution. (Sialidae: Megaloptera) in eastern North Journal of the Kansas Entomological Society America. Canadian Entomologist 112: 409- 80: 105-129. 419. Flint OS, Jr (1964) New species and new state Chordas SW, III, Harp GL, Wolfe GW (1996) records of Sialis (Neuroptera: Sialidae). The aquatic macroinvertebrates of the White Entomological News 75: 9-13. River National Wildlife Refuge, Arkansas. Flint OS, Jr (1965) The genus Neohermes. Psyche Proceedings Arkansas Academy of Science 50: 72: 255-263. 42-51. Foti TL, Bukenhofer GA (1998) A description of Contreras-Ramos A (1998) Systematics of the the sections and subsections of the Interior dobsonfly genus Corydalus (Megaloptera: Highlands of Arkansas and Oklahoma. Journal Corydalidae). Thomas Say Publications of the Arkansas Academy of Science 52: 54-62. in Entomology. Entomological Society of Hagen HA (1861) Synopsis of the Neuroptera of America, Lanham, Maryland 360 pp. North America. Smithsonian Miscellaneous Cuyler RD (1956) Taxonomy and ecology of Collections 4: 1-347. BOWLES and SITES 419
Harp G, Harp P, McCord S (2008) Aquatic Petersen JC, Adamski JC, Bell RW, Davis JV, macroinvertebrates collected from thirty- Femmer SR, Freiwald DA, Joseph RL (1998) two Missouri Ozark streams. Journal of the Water Quality in the Ozark Plateaus, Arkansas, Arkansas Academy of Science 62: 61-74. Kansas, Missouri, and Oklahoma, 1992–95. Harp GL, Robison HW (2006) Aquatic USGS circular 1158, US Geological Survey, macroinvertebrates of the Strawberry River Little Rock, Arkansas 33 pp. system in North-central Arkansas. Journal of Phillips EC (1996) Habitat preference of large the Arkansas Academy of Science 60: 46-61. predatory aquatic insects (Megaloptera and Liu X, Hayashi F, Yang D (2015) Phylogeny of the Odonata) in Ozark streams of Arkansas. Texas family Sialidae (Insecta: Megaloptera) inferred Journal of Science 48: 255-260. from morphological data, with implications Pittman JL, Turner TS, Frederick L, Petersen RL, for generic classification and historical Poston ME, Mackenzie M, Duffield RM (1996) biogeography. Cladistics 31: 18-49. Occurrence of alderfly larvae (Megaloptera) in Locklin JL, Arsuffi TL, Bowles DE (2006) Life a West Virginia population of the pitcher plant, history of Sialis (Megaloptera: Sialidae) in a Sarracenia purpurea L. (Sarraceniaceae). lentic and lotic ecosystem in Central Texas. Entomological News 107: 137-140. American Midland Naturalist 155: 50-62. Poulton BC, Stewart KW (1991) The Stoneflies Locklin JL (2008) Texas alderflies (Megaloptera: (Plecoptera) of the Ozark and Ouachita Sialidae): new records, distributions, and a key Mountains. Memoirs of the American to adults in Texas. Southwestern Naturalist 53: Entomological Society No. 38. The American 524-528. Entomological Society, Philadelphia, Mather B (1981) Larvae of alderfly (Megaloptera: Pennsylvania 116 pp. Sialidae) from pitcher plant. Entomological Rasmussen AK, Pescador ML (2002) A guide to the News 92: 32. Megaloptera and aquatic Neuroptera of Florida. McCafferty WP, Provonsha AV (1978) The Department of Environmental Protection, State Ephemeroptera of mountainous Arkansas. of Florida, Tallahassee, Florida 78 pp. Journal of the Kansas Entomological Society Roell MJ, Orth DJ (1991) Production of dobsonfly 51: 360-379. (Corydalus cornutus) larvae in the New Moss LE, Harp GL (1993) Effects of downstream River of West Virginia. Journal of Freshwater wastewater effluent on the water quality and Ecology 6: 1-9. aquatic macroinvertebrates in a Sharp County, Ross HH (1937) Studies of Nearctic aquatic insects, Arkansas, stream. Proceedings of the Arkansas I. Nearctic alder flies of the genusSialis Academy of Science 47: 82-85. (Megaloptera, Sialidae). Bulletin of the Illinois Moulton SR, III, Stewart KW (1996) Caddisflies Natural History Survey 21: 57-78. (Trichoptera) of the Interior Highlands of Sarver R, Kondratieff BC (1997) Survey of North America. Memoirs of the American Missouri mayflies with the first description Entomological Institute. The American of adults of Stenonema bednariki Entomological Institute, Gainesville, Florida (Ephemeroptera: Heptageniidae). Journal of the 313 pp. Kansas Entomological Society 70: 132-140. Neunzig HH (1966) Larvae of the genus Nigronia Schwardt HH (1936) Horseflies of Arkansas. Banks. Proceedings of the Entomological Bulletin of the Arkansas Agricultural Society of Washington 68: 11-16. Experiment Station 332: 1-66. Penny ND, Adams PA, Stange LA (1997) Species Short RA, Stanley EH, Harrison JW, Epperson catalog of the Neuroptera, Megaloptera, and CH (1987) Production of Corydalus cornutus Raphidioptera of America north of Mexico. (Megaloptera) in four streams differing in size, Proceedings of the California Academy of flow, and temperature. Journal of the North Sciences 50: 39-114. American Benthological Society 6: 105-114. 420 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Smith RC (1925) The Neuroptera and Mecoptera Tarter DC, Watkins WD, Ashley DL, Goodwin of Kansas. Bulletin of the Brooklyn JT (1978) New state records and seasonal Entomological Society 20: 165-171. emergence patterns of alderflies east of the Smith RC (1934) The Neuroptera and Mecoptera Rocky Mountains (Megaloptera: Sialidae). of Kansas, with keys for the identification of Entomological News 89: 231-234. species. Journal of the Kansas Entomological Tarter DC, Watkins WD, Etnier DA (1979) Larval Society 7: 120-145. description and habitat notes of the fishfly Stark BP, Lago PK (1980) New records of Nearctic Neohermes concolor (Davis) (Megaloptera: Sialis (Megaloptera, Corydalidae). Entomological News 90: 29-32. Sialidae), with emphasis on Mississippi fauna. Thom RH, and Wilson JH (1980) The natural Entomological News 91: 117–121. divisions of Missouri. Transactions of the Stark BP, Lago PK (1983) Studies of Mississippi Missouri Academy of Science 14: 9-23. fishflies (Megaloptera: Corydalidae: Whiting MF (1991) A distributional study of Sialis Chauliodinae). Journal of the Kansas (Megaloptera: Sialidae) in North America. Entomological Society 56: 356-364. Entomological News 102: 50-56. Sullivan KC (1929) Notes on the aquatic life of the Niangua River, Missouri, with special reference to insects. Ecology 10: 322-325. BOWLES and SITES 421
Figures 2-6. Larval Megaloptera: 2, Sialis sp., dorsal (from Ross 1937); 3, Protosialis americana, mandible, dorsal; 4, Sialis sp., mandible, dorsal; 5, Corydalus cornutus, dorsal (from Rasmussen and Pescador 2002); 6, Neohermes concolor, dorsal (from Tarter et al. 1979). 422 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Figures 7-10. Larval Chauliodinae, terminalia, dorsal: 7, Chauliodes sp. (redrawn from Cuyler 1958); 8, Neohermes sp. (from Cuyler 1958); 9, Nigronia fasciatus; 10, Terminalia of Nigronia serricornis. RT = respiratory tube. BOWLES and SITES 423
Figures 11-12. Protosialis americana: 11, lateral; 12, head, dorsal. Figures 13-14. Sialis infumata: 13, lateral; 14, head, dorsal. 424 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Figures. 15-18. Protosialis americana, terminalia (from Ross 1937): 15, male, ventral; 16, male, lateral; 17, male, ectoproct, caudal; 18, female, ventral. Figure 19. Protosialis glabella, terminalia, male, lateral. Figure 20-23. Sialis vagans, terminalia (from Ross 1937): 20, male, ventral, with right side removed to reveal genitalia; 21, male, lateral; 22, male, ectoproct, caudal; 23, female, ventral. Figures 24-27. Sialis joppa, terminalia (from Ross 1937): 24, male, ventral; 25, male, lateral; 26, male, ectoproct, caudal; 27, female. BOWLES and SITES 425
Figures 28-30. Sialis hasta, terminalia (from Ross 1937): 28, male, ventral; 29, male, lateral; 30, male, ectoproct, caudal. Figures 31-34. Sialis mohri, terminalia (from Ross 1937): 31, male, ventral; 32, male, lateral; 33, male ectoproct, caudal; 34, female, ventral. Figures 35-38. Sialis infumata, terminalia (from Ross 1937): 35, male, ventral; 36, male, lateral; 37, male, ectoproct, caudal; 38, female, ventral. Figures 39-41. Silais itasca, terminalia (from Ross 1937): 39, male, ventral; 40, male, lateral; 41, male, ectoproct, caudal. Figures 42-45. Sialis velata, terminalia (from Ross 1937): 42, male, ventral; 43, male, lateral; 44, male, ectoproct, caudal; 45, female, ventral. 426 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Figures. 46-51. Wings of Corydalidae: 46, Corydalus cornutus; 47, Neohermes concolor; 48, Chauliodes pectinicornis; 49, Chauliodes rastricornis; 50, Nigronia fasciatus; 51, Nigronia serricornis. BOWLES and SITES 427
Figures 52-56. Corydalus cornutus, adults: 52, male head, dorsal; 53, female head, dorsal; 54, male terminalia, dorsal; 55, male terminalia, ventral; 56, female terminalia, lateral (Figs. 54-56 from Contreras- Ramos 1998).
Figures 57-58. Chauliodes, adult heads, dorsal: 57, Chauliodes pectinicornis; 58, Chauliodes rastricornis. 428 MEGALOPTERA OF THE INTERIOR HIGHLANDS
Figures 59-60. Chauliodes pectinicornis, terminalia (from Stark and Lago 1983): 59, male, dorsal; 60, male, lateral.
Figures 61-63. Chauliodes rastricornis, terminalia (from Stark and Lago 1983): 61, male, dorsal; 62, male, lateral; 63, female, lateral.
Figures 64-69. Neohermes concolor, terminalia (from Bowles and Mathis 1992): 64, male, dorsal; 65, male, dorsal; 66, male, lateral; 67, male, lateral with expanded dorsal membrane; 68, male, ventral; 69, female, lateral.
Figure 70. Nigronia fasciatus, male, terminalia, ventral (from Stark and Lago 1983). Figures 71-72. Nigronia serricornis, terminalia: 71, male, ventral; 72, female, ventral (from Stark and Lago 1983). BOWLES and SITES 429