ILLINOIS NATURAL HISTORY SURVEY The Winter Stoneflies of Illinois (Insecta: Plecoptera): 100 Years of Change Donald W. Webb Illinois Natural History Survey Bulletin Volume 36, Article 5 December 2002 [linois Natural History Survey, David L. Thomas, Chief A Division of the Illinois Department of Natural Resources, Brent Manning, Director A catalog of the publications of the Illinois Natural History Survey is available without charge from the address below. A price list and an order blank are included with the catalog. This catalog may also be accessed on the World Wide Web at http://www.inhs.uiuc.edu/chf/pub/ pub-catalog/spring00/index00.html Illinois Natural History Survey Distribution Office 607 E. Peabody Dr. Champaign, IL 61820 Editors: Thomas Rice and Charles Warwick Citation: Webb, Donald, W. 2002. The winter stoneflies of Illinois (Insecta: Plecoptera): 100 years of change. Illinois Natural History Survey Bulletin 36(5): 195-274. US ISSN 0073-4918 Printed by authority of the State of Illinois (MJ128056—1.5M—12-02) Equal opportunity to participate in programs of the Illinois Department of Natural Resources (IDNR) and those funded by the U.S. Fish and Wildlife Service and other agencies is available to all individuals regardless of race, sex, national origin, disability, age, religion, or other non-merit factors. If you believe you have been discriminated against, contact the funding source's civil rights office and/or the Equal Employment Opportunity Officer, IDNR, One Natural Resources Way; Springfield, IL. 62702-1274; 5° 217/785-0067; TTY 217/782-9175. Printed with soy ink on recycled and recyclable paper. ILLINOIS NATURAL CSTE O Ray: SURVEY The Winter Stoneflies of Illinois (Insecta: Plecoptera): 100 Years of Change Donald W. Webb * pg ae Photo by Mark J. Wetzel, INHS. Illinois Natural History Survey Bulletin Volume 36, Article 5 December 2002 Acknowledgments Support for this study was provided by the IIli- nois Natural History Survey and grants from the Illinois Nature Preserves Commission; the Natu- ral Heritage Division, Illinois Department of Conservation; the Illinois Groundwater Consor- tium; and The Environmental Protection Trust Fund Commission. I would also like to thank the curators of en- tomology at Eastern Illinois University, South- ern Illinois University, Western Illinois Univer- sity, and the Field Museum of Natural History for the loan of specimens relative to this study. My thanks also go out to Mitch Harris and Ed DeWalt for their assistance in collecting winter stoneflies and to Frank Hutto for his assistance in preparing the distribution maps for this study. Contents Acknowledgements 1i Abstract 195 Introduction 195 Historical Background 195 Environmental Change in Illinois 196 Present Habitat in Illinois 197 Biology of Stoneflies 198 Methods 202 Results 202 Discussion 202 Natural Divisions of Illinois 204 River Drainages 208 : Galena, Apple, and Plum River systems 208 Rock River System (Rock, Pecatonica, Kishwaukee, and Green River drainages) 209 Middle Mississippi River tributaries (Edwards River, Henderson Creek, Bear Creek, Sny Creek, Bay Creek) 209 Des Plaines River and Lake Michigan tributaries (Des Plaines, DuPage rivers) 210 Fox River system 211 Little Vermilion River, Big Bureau and Kickapoo Creek systems 211 Kankakee and Iroquois River systems 211 Vermilion and Mazon River systems 212 Spoon River system 212 LaMoine River system 213 Mackinaw River system 213 Sangamon River system 213 Lower Illinois River tributaries and American Bottoms 214 Kaskaskia River system 214 Big Muddy system 214 Cache River system 215 Massac, Bay, Lusk, Big Grand Pierre, and Big Creek systems 215 Saline River system 215 Little Wabash River and Bonpas Creek systems 216 Embarras River and Wabash River tributaries 216 Vermilion and Little Vermilion River systems 217 Illinois River 217 Mississippi River 217 Ohio River 218 Wabash River 218 Summary 218 Keys 220 Key to Families of Illinois Winter Stoneflies 220 Capniidae 221 Key to Adults of Illinois Capniidae 221 Key to Mature Nymphs of Illinois Capniidae 222 Allocapnia Claassen 222 Key to Males 222 Key to Females 223 Key to Mature Nymphs 224 Allocapnia forbesi Frison 228 Allocapnia granulata (Claassen) 230 Allocapnia illinoensis Frison 232 Allocapnia mystica Frison 234 Allocapnia nivicola (Fitch) 236 Allocapnia recta (Claassen) 238 Allocapnia rickeri Frison 240 Allocapnia smithi Ross and Ricker 242 Allocapnia vivipara (Claassen) 243 Nemocapnia Banks 245 Nemocapnia carolina Banks 245 Paracapnia Hanson 246 Paracapnia angulata Hanson 246 Leuctridae 248 Zealeuctra Ricker 248 Key to Adults of Illinois Zealeuctra 248 Zealeuctra claasseni (Frison) 249 Zealeuctra fraxina Ricker and Ross 251 Zealeuctra narfi Ricker and Ricker 252 Nemouridae 253 Prostoia Ricker 253 Prostoia completa (Walker) 253 Taeniopterygidae 254 Key to Genera of Taeniopterygidae in Illinois (Stewart 2000). 254 Strophopteryx Frison 255 Strophopteryx fasciata (Burmeister) 255 Taeniopteryx Pictet 258 Key to Adults of Illinois Taeniopteryx (Ricker and Ross 1968) 258 Taeniopteryx burksi Ricker and Ross 260 Taeniopteryx lita Frison 262 Taeniopteryx metequi Ricker and Ross 264 Taeniopteryx nivalis (Fitch) 265 Taeniopteryx parvula Banks 267 Literature Cited 268 December 2002 Winter Stoneflies of Illinois 195 ABSTRACT Winter stoneflies (Insecta: Plecoptera) are an aquatic group of insects whose adults emerge in IIli- nois from late November to early April. Twenty-one species have been reported from Illinois. Ex- tensive collections of winter stoneflies were made in Illinois during the 1920s and 1930s by Frison, the 1960s by Ross and “the winter stonefly club’, and the 1990s by Webb. These specimens are housed in the Insect Collection of the Illinois Natural History Survey and allowed for an evaluation of the current status of these species following a century of environmental change. Over the past century (1900-2000), the species diversity of winter stoneflies averaged 2.5 spe- cies per county with species reported from every county but 3 (Carroll, DuPage, Ford) and with 10 counties recording 5 or more species. Pope County (13 species) reported the greatest species diver- sity. During the recent resurvey (1976-2000), species diversity average 1.9 species per county with specimens not collected in 11 counties, and only 3 counties (Hardin, Pope, and Saline) exhibited 5 or more species. Four species are considered extirpated from Illinois: Allocapnia illinoensis, Nemocapnia carolina, Paracapnia angulata, and Taeniopteryx parvula. Seven species were found to be common (known from more than 15 localities): Allocapnia forbesi, A. granulata, A. mystica, A. rickeri, A. vivipara, Taeniopteryx burksi, and T: nivalis. Four species are considered uncommon (known from 4—15 localities): Allocapnia recta, Strophopteryx fasciata, Taeniopteryx metequi and Zealeuctra claasseni. Six species are considered rare (known from 1-3 localities): Allocapnia nivicola, A. smithi, Prostoia completa, Taeniopteryx lita, Zealeuctra fraxina, and Z. narfi. Significantly reduced patterns of dis- tribution were noted in three species: A. granulata, A. mystica, and Strophopteryx fasciata. Only Taeniopteryx nivalis, previously rare, is now spreading its distribution across northern [linois. There has been a modest decline during the past century in the diversity of stonefly species within various counties. Also, a trend was observed toward an increase in pollution-tolerant, “generalist” species with a decrease in pollution-sensitive, habitat “specialists.” INTRODUCTION Historical Background Since its inception as the Natural History Soci- ety of Illinois in 1858, the Illinois Natural His- tory Survey (INHS) has established a strong commitment to identifying and preserving the fauna and flora of Illinois (Mills 1958). This has been particularly true with regard to insects. At present, the INHS Insect Collection houses over 6 million specimens from throughout the world, but is most strongly focused on the cen- tral United States. Early INHS entomologists were encouraged to undertake faunal studies (mayflies, Burks 1953; leafhoppers, DeLong 1948; stoneflies, Frison 1929, 1935; damsel-flies, Garman 1917; pentatomids, Hart 1919; earwigs, grasshoppers and crickets, Hebard 1934; plant- lice, Hottes and Frison 1931; mirids, Knight 1941; midges, Malloch 1915; deerflies and horseflies, Pechuman et al. 1983; sialids, Ross 1937; caddis flies, Ross 1944; mosquitoes, Ross 1947; scorpionflies, Webb et al. 1975) which focused on Illinois, but were considerably broader in their geographic scope. Such was the case for the stoneflies (Plecoptera) of Illinois. Early in his career, Theodore H. Frison loved to collect on the hillsides along the Salt Fork of the Vermilion River south of Oakwood. It was here that Frison observed that “...in some of the very small streams, the smallest of the stonefly nymphs kept increasing in size as winter ap- proached.” From this observation arose his **,..abiding interest in and love of stoneflies which continued through the rest of his life” (Ross 1958). Through the 1920s, Frison collected and reared a diverse array of stoneflies, culminating in his study of the “Fall and Winter Stoneflies, or Plecoptera, of Illinois” (Frison 1929). With the assistance of H. H. Ross, collecting contin- ued through the 1930s and was broadened be- yond the borders of Illinois. Frison completed a faunal study of “The Stoneflies, or Plecoptera, of Illinois” in 1935; a “Study of Nearctic Insects. II. Descriptions of Plecoptera, with special ref- erence to the Illinois Species” in 1937; and cul- minated his interest in stoneflies in his “Studies of North American Plecoptera, with special ref- erence to the fauna of Illinois” in 1942. All of the material collected by Frison and Ross dur- ing those years was deposited in the collections 196 Illinois Natural History Survey Bulletin of the Illinois Natural History Survey. In the 1960s, H. H. Ross and W. E. Ricker established the “Winter Stonefly Club” which enlisted the assistance of over 200 enthusiasts to collect win- ter stoneflies throughout eastern North America. The dedication of these individuals to brave the winter environs in search of small, black, creepy- crawly things that ventured over the snow and ice and seemed to seek pleasure in running across the railings of bridges or crawling out to the tips of branches in search of the warm sunlight, again added a multitude of specimens to the INHS collections. Through the diligence of the club members, Ross and Ricker were able to revise three genera of the winter stoneflies, Taeniopteryx (Ricker and Ross 1968), Zealeuctra (Ricker and Ross 1969), and Allocapnia (Ross and Ricker 1971). Environmental Change in Illinois With the arrival of European settlers into Illi- nois during the 1820’s, significant changes be- gan to take place in the landscape and water- ways of Illinois. At that time, 61.2% of Illinois was dominated by prairie, 38.2% by forest, and 0.6% water (Iverson 1991). In the following 100 years the forests of Illinois were reduced to 22% of what had existed in 1820. There has been some recovery in the reforestation of Illinois, but all of this is secondary growth (Iverson 1991). The desire for tillable land drastically altered all of the prairies of Illinois (Anderson 1991). For many years broad hedgerows of multiflora rose (Rosa multiflora) separated farm boundaries, providing abundant habitat for small game, but also acting as a significant barrier to soil runoff. Over the past 30 years this practice has changed significantly with hedgerows being removed, allowing farmers to cultivate row crops from property line to property line. The buffer strips of vegetation along IJlinois’ streams were greatly reduced, adding significantly to the sediment load borne by our waterways. Streamside ripar- ian buffer strips are an integral part of rivers and streams. They stabilize stream banks, reduce erosional input, filter agricultural runoff of sedi- ments, and protect streams from excessive nu- trient loading (Illinois Department of Energy and Natural Resources, IDENR 1994). Only now are we attempting to modify our views on how close we can cultivate along our waterways. Vol. 36 Art. 5 Illinois possesses over 42,000 km of flow- ing water (IDENR 1994). These waterways have been drastically altered since presettlement times. Dams have been constructed to control water levels for navigation along the Mississippi, Illinois, and Ohio rivers. Many of Illinois’ smaller rivers have been dammed for flood con- trol and water supplies and the added attraction of water recreation. In our efforts to improve and stabilize agricultural production, most fields in Illinois are criss-crossed with drainage tiles to lower groundwater levels and increase the avail- ability of tillable land. Increased runoff from agricultural land with its inherent sediment load has required the need to channelize many streams and ditches of Illinois. This channelization has drastically altered the flow and bottom substrates of many streams as we attempt to move water more rapidly downstream. Stream channelization drastically reduces the habitat diversity for aquatic macroinvertebrates and fish, while in- creasing stream bank erosion and sediment trans- portation (IDENR 1994). Generally, following channelization there has been a reduction in the biodiversity and abundance of aquatic macroinvertebrates and fish (Henegar and Harmon 1973, Hortle and Lake 1982), and many of these effects may persist for some time (Arner et al. 1976). Currently, less than 1% of the origi- nal landscape of Illinois remains in its natural state, as defined by criteria established for an inventory of the natural areas of Illinois (White 1978). Because of this manipulation, streams in the agricultural areas of Illinois are perhaps now more homogeneous than presettlement times (IDENR 1994, Ross 1944). Stoneflies present an excellent group of aquatic macroinvertebrates for examining envi- ronmental change over time because many spe- cies are intolerant to environmental perturba- tions. Hynes (1993) warned that stoneflies are probably the insect order most threatened by human activity. The diversity and abundance of stoneflies reflects the physical, chemical, and biological nature of a stream (Surdick and Gaufin 1978). Because of the intensive agricultural practices, deforestation, and industrial and sub- urban development over the past 100 years, - stream quality in Illinois has undergone signifi- cant change (Page and Jeffords 1991). Increased public awareness of the environ- ment and the use of stoneflies as important indi- December 2002 cators of stream quality (Harper 1984, Hynes 1972; Jones et al. 1981; Resh and Unzicker 1975, Stewart and Harper 1996) stress the need for up- to-date knowledge of Illinois stoneflies. Major refinements in North American stonefly system- atics, in conjunction with the scope and histori- cal value of the INHS collections, provide us with a unique opportunity to re-examine the system- atics and distribution of the Illinois fauna in light of environmental change within stream systems. In Illinois, species of winter stoneflies are found in the families Capniidae, Leuctridae (in part), Nemouridae (in part), and Taeniopterygidae, and collected from Novem- ber through early April (Table 1). When Frison (1929) published research on the winter stoneflies of Illinois, he included five genera and nine species. By 1935, 11 species were listed, and 14 in 1942. Overall, his work covered 137. sites in 54 counties. Ricker and Ross (1968, 1969, 1975) and Ross and Ricker (1971) and the collections of “the winter stonefly club” further contributed to our knowledge of Illinois winter stoneflies with specimens collected at 121 sites in 46 counties from 1946 to 1975. By this time, seven genera and 21 species of winter stoneflies were known from Illinois. During the past 10 years, winter stoneflies were sought at 1,259 sites in all 102 counties of Illinois. These recent collections allow us to as- sess the historical changes in species diversity and distribution during the “Frison” era (1900- 1945); the “Ross and Ricker” era (1946-1975) and the “Present” (1976-2000). These records are especially important today with government mandates to protect aquatic habitats and their biota, and state agencies actively working to identify high- quality streams and populations of stream-dependent species at risk of extirpa- tion (Page et al. 1992). Present Habitat in Illinois Although the general perception of Illinois may be field after field of corn and soybean extend- ing south from the metropolitan area of Chicago, its north-south alignment offers a varied physi- ography. Large rivers (Wabash, Ohio, and Mis- sissippi) surround two-thirds of Illinois with an- other broad river (Illinois River) running diago- nally across the state from Chicago southwest to Alton. Within its boundaries lies a northern Winter Stoneflies of Illinois 197 boreal region of natural lakes and tamarack bog (Lake and McHenry counties); finger-like rem- nants of the eastern deciduous forest still project into its eastern border (Vermilion County); a broad Appalachian-Ozark corridor spans the southern fourth of the state composed of lime- stone and sandstone hills with numerous small, pristine, gravel streams (Fig. 1|A&B); a coastal plain with lowland marshes and tupelo-bald cy- press swamps (Alexander, Massac, Pulaski coun- ties); and a driftless (unglaciated) area in north- western Illinois (Carroll and Jo Daviess coun- ties). Schwegmann (1973) has divided Illinois into 14 natural divisions (Fig. 2). Within Illinois, 25 major river drainages have been outlined (Page et al. 1992) (Fig. 3). For these drainages Hite and Bertrand (1989) devel- oped an index from “A” to “E” to identify bio- logically significant streams in Illinois based on fish populations, water quality, and aquatic macroinvertebrates. A stream segment catego- rized as “A” was a high-quality stream. Page et al. (1992) expanded this list of biologically sig- nificant streams by considering additional infor- mation on biodiversity, in particular, informa- tion on endangered and threatened species, and Mollusca and crustacean diversity. Within these drainages, the Illinois Water Quality Report (IEPA 1990, 1996) describes water quality con- ditions in terms of the degree waters attain des- ignated uses such as: Full Support: The water quality meets the needs of all designated uses protected by appli- cable water-quality standards. Full Support/Threatened: Water quality is presently adequate to maintain designated uses, but if a declining trend continues, only partial support may be attained in the future. Partial Support/Minor Impairment: Water quality has been impaired, but only to a minor degree. There may be minor exceedences in applicable water- quality standards or criteria for assessing the designated use attainment. Partial Support/Moderate Impairment: Wa- ter-quality conditions are impaired to a greater degree inhibiting the waterbody from meeting all the needs for that designated use. Nonsupport: Water quality is severely im- paired and not capable of supporting the desig- nated use to any degree. These conditions give the most recent assess- ment of the general health of drainage basin. 198 [linois Natural History Survey Bulletin Vol. 36 Art. 5 Figure 1. Winter stonefly habitats. (A). Ephemeral stream. (B). Gravel riffles. (C). Tributary of Sangamon River. (D). Mississippi River. Photos by Donald W. Webb, INHS. Biology of Stoneflies Winter stoneflies (Table 1, Fig. 4) play a vital role in the energy and production of these streams (Stewart and Stark 1988). They undergo univoltine (one year) or semivoltine (two or more years) life cycles. These cycles may be heterodynamic (Wigglesworth 1974, Butler 1984) in having an egg or nymphal diapause, or homodynamic, and having no regular period of dormancy (Stewart and Stark 1988). Species with heterodynamic development cycles are re- ferred to as fast (short-term development) and those with homodynamic cycles as slow (devel- opment over most of the required generation time) (Stewart and Stark 1988). Stewart and Stark (1988) and Ernst and Stewart (1985) found that Allocapnia rickeri exhibited a univoltine, fast cycle in Oklahoma, whereas it exhibited a univoltine, slow cycle in southern Canada (Harper 1973). Krueger and Cook (1981) noted a longer developmental time for A. rickeri in Minnesota with small nymphs appearing in July, growing rapidly until December, slowing growth in December-January, then resuming growth until a March-April emergence. Snellen and Stewart (1979) reported an interesting example of indeterminate voltinism for Zealeuctra spe- cies in intermittent Texas streams. Their con- tinuous field and laboratory study during the period November 1974 to April 1977 showed that given cohorts of both Z. claasseni and Z. hitei were able to undergo both univoltine cycles from nondiapausing and short-term diapausing eggs in wet and normal rainfall years, and semivoltine cycles from diapausing eggs follow- ing at least two years of drought. Both nondia- pausing and diapause eggs from given adult co- horts were confirmed with laboratory incubation experiments. Small numbers of adults were reared from the nondiapausing eggs in the labo- ratory to corroborate the univoltine aspect of the cycle. This is perhaps a good example in stoneflies of the “relict seed phenomenon” (Giesel 1976), in which adults effectively pro- long their reproductive effort by producing eggs with delayed and asynchronous development ~ (Stewart and Stark 1988). Eggs: In some stoneflies several matings may occur and several batches of eggs laid (Hitchcock 1974). Capniids apparently lay only a single December 2002 Figure 2. The Natural Division of Illinois (Schwegman 1973). See Table 3 for division names. batch of 100-700 eggs (Coleman and Hynes 1970, Khoo 1964). Egg development: There appear to be four basic patterns of embryonic nymphal develop- ment (Stewart and Stark 1988): short term, syn- chronous (3—8 weeks) in homodynamic species; optional short- to long-term in homodynamic species; long-term, synchronous, or asynchro- nous (5>24 months) in diapausing eggs of heterodynamic species; and a combination of short-term nondiapausing eggs and long-term, synchronous or asynchronous diapausing eggs. In the long-term pattern, development to the ad- vanced eyespot protonymph stage may be (a) short-term, within about four weeks or (b) long- term, requiring most of the diapause period. Nymphal growth and development: First instar nymphs open the chorion by pushing with an egg tooth located on, or consisting of, the frons (DeGrange 1957; Khoo 1968a, 1968b; Hynes 1976). The shell splits into two halves (Komatsu 1971, Brittain 1973, Hynes 1976) Winter Stoneflies of Illinois 199 one | A Y 3 nS ‘ 4 y A ’ s aan \ ‘ | : t (BP OE iy a Ra ef" tesla. 5 NB ' FESS Sarai, when z. A, — via i nm uo Figure 3. The drainage basins of Illinois (Page et al. 1992). See Table 4 for drainage names. The first instars are consistent in being un- pigmented, without ocelli, and having antenna:cercal segment formulae of 8-11:3-5; compound eyes of 2-4 ommatidia; gills absent, reduced, or represented only by knobs or stubs, generally few hairs compared with later instars; and three tarsal segments, the first two short and together subequal to the longer third (Snellen and Stewart 1979, for Zealeuctra claasseni ). Harper (1979) observed that first instars of Ontario euholognathan species began feeding immedi- ately after eclosion. Nymphal diapause has been reported in a variety of winter stoneflies: Capniidae: Allocapnia granulata (Harper and Hynes 1970, Pugsley and Hynes 1985), A. pygmaea (Coleman and Hynes 1970, Harper and Hynes 1972, Pugsley and Hynes 1985) and A. vivipara (Coleman and Hynes 1970, Pugsley and Hynes 1985); Nemouridae: Prostoia completa (Ernst and Stewart 1985); and Taeniopterygidae: Strophopteryx fasciata (Harper and Hynes 1970, 200 Illinois Natural History Survey Bulletin Vol. 36 Art. 5 Table 1. Winter Stoneflies of Illinois (Harris and Webb 1995) Capniidae Leuctridae Allocapnia Zealeuctra forbesi Frison claasseni (Frison) granulata (Claassen) fraxina Ricker and Ross illinoensis Frison narfi Ricker and Ross mystica Frison Nemouridae nivicola (Fitch) Prostoia recta (Claassen) completa (Walker) rickeri Frison Taeniopterygidae smithi Ross and Ricker Strophopteryx vivipara (Claassen) fasciata (Burmeister) Nemocapnia Taeniopteryx carolina Banks burksi Ricker and Ross Paracapnia lita Frison angulata Hanson metequi Ricker and Ross nivalis (Fitch) parvula Banks * ea Re urksi, male (C), female (D). Photos by Figure 4. Allocapnia vivipara, male (A), female (B). Taeniopteryx b Michael Jeffords, INHS. December 2002 1972), Taeniopteryx burksi and T. nivalis (Harper and Hynes 1970, 1972, Pugsley and Hynes 1985). Instar number varies within many species (Hitchcock 1974, Hynes 1976, Butler 1984, Jop and Szczytko 1984) and is relatively large, 10— 22 plus (Butler 1984, Sephon and Hynes 1982). Winter- and most early-spring-emerging euholognathan families grow fastest in cold wa- ter (Stewart and Stark 1988). Nymphs crawl out of the water on objects, plant their claws, and adults emerge from the nymphal skin, often over a surprisingly short period of time (i.e., 5 min- utes, Stewart and Stark 1988) to several hours. Winter species, particularly capniids, emerge under surface ice (Harper and Hynes 1972). Food habits: Stoneflies have diversified their food habits so that the different species fill about every conceivable major food niche in streams (Stewart and Harper 1996, Stewart and Stark 1988). They generally fall into three feeding groups: phytophagous, feeding strictly on algae and organic detritus (Brinck 1949, Frison 1929, Hitchcock 1974, Hynes 1941, Jones 1950, Wu 1923); polyphagous or omnivorous, feeding on plant and animal material; and carnivorous, feed- ing strictly on aquatic invertebrates, predomi- nately chironomids (Fuller and Stewart 1977, Hitchcock 1974, Richardson and Gaufin 1971, Short and Ward 1980). Natural enemies: Hitchcock (1974) reported hunting spiders attacking winter stoneflies and found an immature of Pityohyphantes phrygianus feeding on Taeniopteryx burksi. He also reported on a few records of birds, odonates, frogs and bats feeding on adult stoneflies and Hamilton (1932) reported bluejays feeding on stoneflies. In contrast, Hitchcock (1974) cited numerous references where stonefly nymphs were fed upon by fish, water shrews, turtles, water dippers, Cinclus mexicanus, salamanders, crayfish, and other aquatic insects. He also cites a few references of parasites inhabiting stoneflies. Drumming: Hitchcock (1974), Ziegler and Stewart (1977, 1985, 1988), Snellen and Stewart (1979), Szcezytko and Stewart (1979), Stewart et al. (1982a, 1982b), Maketon and Stewart (1984, 1988), Stewart and Maketon 1991), Stewart (1997) cite a variety of references for drumming activity within the Plecoptera. Ziegler and Winter Stoneflies of Illinois 201 Stewart (1977) and Snellen and Stewart (1979) described the drumming behavior of Zealeuctra claasseni. Mating: Hitchcock (1974) reported that the male mounts the back of the female, placing his abdomen to her side and recurving the apex of his abdomen to position his genitalia in line with the female terminalia on the ventral surface Drift: Most Plecoptera have a low propen- sity to drift (Brusven 1970; Elliot 1967a, 1967b; Stewart and Szczytko 1983) but when it occurs it is generally in pre-emergent instars, just after sunset. For Allocapnia rickeri (Ernst and Stewart 1985) drift was correlated to their standing stock. Ernst and Stewart (1985: Fig. 3) reported that nymphs of Prostoia completa drifted during Feb- ruary and March and that the size of the drifting nymphs was always greater than the average size of nymphs concurrently found in benthic samples. Ernst and Stewart (1985: Fig. 4) re- ported that a few nymphs of A. rickeri displayed pre-sunset drifting at the beginning of emer- gence. In contrast to drifting, adults and nymphs of Plecoptera can disperse upstream (Hitchcock 1974). Both sexes fly with the wind (Elliot 1967b, Hitchcock 1974) and may disperse sig- nificant distances. Adults also have been re- ported to walk upstream once they reach the shoreline (Hitchcock 1974, Thomas 1966). Nymphs of Allocapnia pygmaea have been ob- served moving upstream by Hultin et al. (1969) and Bishop and Hynes (1969). Economic importance: In general, adult stoneflies are of little economic importance. They have been reported as a minor pest of or- chards and ornamental plants (Hitchcock 1974; Kawai 1967; Newcomer 1918, 1950; Schuh and Mote 1948). Distribution and seasonal activity: 21 spe- cies of winter stoneflies have been reported for Illinois, although 4 species now appear to be extirpated from the state (Allocapnia illinoensis, Nemocapnia carolina, Paracapnia angulata, Taeniopteryx parvula). Eight of the extant win- ter stoneflies are restricted to the Shawnee Hills of southern Illinois (Allocapnia forbesi, A. mystica, A. smithi, Prostoia completa, Taeniopteryx metequi, Zealeuctra claasseni, Z. fraxina, and Z. narfi). Six species have a gen- eral distribution over much of Illinois (Allocapnia recta, A. rickeri, A. vivipara, 202 Illinois Natural History Survey Bulletin Strophopteryx fasciata, Taeniopteryx burksi, T: lita), two species are found in the northern third of the state (Allocapnia granulata, Taeniopteryx nivalis), and one species was found in the rem- nants of the eastern deciduous forest in Cole and Vermilion counties of eastern Illinois (A. nivicola). METHODS To re-evaluate the biodiversity and distribution of winter stoneflies in Illinois, attempts were made to collect adults from every county in the state. Extensive efforts were made to collect stoneflies in areas where specimens had been previously reported by Frison, Ross, and Ricker and for those species that were rare in Illinois. Specimens were collected as they walked on bridge railings, bridge abutments, ice, and snow; by beating or sweeping low hanging brush, weedy vegetation and grass along the stream margins; by separating leaf packs and woody debris in a large tray; and by collecting speci- mens sitting on or under rocks or logs that pro- jected from a stream. Specimens were preserved in 80% ethyl alcohol, identified, and entered into our database. The entire Plecoptera collection of the Illi- nois Natural History survey has been recurated and the species reorganized to conform to the current nomenclature (Stark et al. 1986). Col- lection information on all stonefly species in the INHS collections has been integrated into a Filemaker Pro database, utilizing field param- eters similar to those used for existing INHS databases (fishes, molluscs, crayfishes, amphi- pods). Keys to the families, genera, and species of Illinois winter stoneflies are provided. For each species, their nomenclatoral history is cited, along with a diagnostic description of the male, female, and nymph (when known) and informa- tion on their biology, habitat preference, distri- bution, seasonal activity, and current status in Illinois. Changes in the distribution pattern of each species for the historical period 1900-1945, from 1946-1975, and from 1976-2000 are mapped (Figs. 5—7) for each species. For those species with limited distributions in Illinois (10 localities or less), specific locality information is provided. Specific locality information on II- linois stoneflies may be obtained through the II- Vol. 36 Art. 5 linois Natural History Survey Web site at www.inhs.uiuc.edu/cbd.EPT/index.html. RESULTS Table 2 outlines the species diversity of winter stoneflies in Illinois from 1900-1945, 1946—- 1975, 1976-2000, and for the past century 1900- 2000. Table 3 outlines the species diversity of win- ter stoneflies within the Natural Divisions of Il- linois for the past century 1900-2000. Table 4 outlines the species diversity of win- ter stoneflies within the drainage basins of IIli- nois for the past century 1900-2000. Historical changes within a species’ distri- bution pattern in Illinois from 1900-1945, 1946— 1975, and 1976—2000 are presented within each species account. DISCUSSION The collections of winter stoneflies in Illinois made by Frison and colleagues from 1900-1945, by Ross and his “winter stonefly club” from 1946 to 1975, and my own extensive collecting dur- ing the 1990s as well as other specimens col- lected by colleagues for the period 1976—2000 provide us with a unique opportunity to exam- ine the changing patterns of species diversity and distribution within three distinct time frames, as well as to look at the total overall picture for the past 100 years. Frison (1900-1945) collected specimens from 54 counties at an average of 2.1 species per county. Ross and his “winter stonefly club” (1946-1975) collected in 46 counties, with an average of 1.9 species per county. From 1976 to 2000, attempts were made to collect winter stoneflies from every county in the state (Fig. 5), with an average of 1.9 species per county collected. When viewed over the past 100 years, the species diversity of winter stoneflies in IIli- nois averaged 2.5 species per county, with win- ter stoneflies reported in every county but 3- (Carroll, DuPage, Ford) and with 10 counties recording five or more species. In comparison to the century picture, recent collections (1976— 2000) indicate that the overall species diversity December 2002 Winter Stoneflies of Illinois 203 1m o eF e m4 3 2 tJ eN b e 1 > YOO ‘ i) ie) 4 e * Figure 6. Collection sites for Illinois winter stoneflies Figure 5. Recent (1976-2000) collecting sites for win- 1900 through 1945. ter stoneflies in Illinois. Closed circles=specimens col- lected. Open circles=negative results. Figure 7. Collection sites for Illinois winter stoneflies during 1946-1975. 204 Illinois Natural History Survey Bulletin within the various counties has dwindled with winter stoneflies not found in 11 counties, and only 3 counties (Hardin, Pope, and Saline) ex- hibiting 5 or more species. Over the past century (1900-2000), Pope County, with 13 species, has been the most spe- cies-rich county for winter stoneflies, although Taeniopteryx lita and T. burksi have not been collected there since the 1930s. Saline County with nine species reported, is interesting in that six of these species were collected only during the recent (1976-2000) resurvey. Clark County with eight species, has lost five of these species since 1975. Four of these species were from Rocky Branch, a favorite collecting site of Frison and Ross. Hardin County with seven species, has lost two of these species since 1946. In gen- eral, there has been a decline in species diver- sity per county. Of the 21 species of winter stoneflies discov- ered during the past century (1900-2000), 7 spe- cies were found to be common (currently known from more than 15 localities): Allocapnia forbesi, restricted to the Shawnee Hills; A. granulata, its previous distribution reduced and now restricted to the Rock River drainage; A. mystica, its pre- vious distribution reduced and now restricted to the Shawnee Hills; A. rickeri, abundant in the Vol. 36 Art. 5 Shawnee Hills, with extensions up the eastern and western borders of Illinois; A. vivipara and Taeniopteryx burksi, widespread throughout state; T: nivalis, previously rare but now spread- ing its distribution across northern Illinois. Four species are considered uncommon (currently known from 4—15 localities): Allocapnia recta, scattered in eastern Illinois; Strophopteryx fasciata, widespread but disappearing from the state; Taeniopteryx metequi and Zealeuctra claasseni, restricted to the Shawnee Hills. Six species are considered rare (currently known from 1-3 localities): Allocapnia nivicola, known only from a single locality in Vermilion County; A. smithi, found in the eastern Shawnee Hills; Prostoia completa, known only from Hutchins Creek in the western Shawnee Hills; Taeniopteryx lita, scattered in east central Ili- nois; Zealeuctra fraxina, eastern Shawnee Hills; and Z. narfi, found at single localities in Saline, Union, and Vermilion counties. Four species are considered extirpated from Illinois: Allocapnia illinoensis, Nemocapnia carolina, Paracapnia angulata, and Taeniopteryx parvula. Natural Divisions of Illinois Schwegman (1973) outlined 14 Natural Divi- sions in Illinois (Fig. 2), and the species diver- Table 2. The species diversity of winter stoneflies in Illinois by county. Species per County 1900-2000 1900-1945 1946-1975 1976-2000 0 3 11 1 31 26 16 38 2 #7 13 15 28 3 18 I ie 16 4 13 3 2 6 = 4 2 1 l 6 1 Zz 0 0 7 2 0 0 0 8 1 1 1 0 7 1 1 10 0 1 1] 0 iB 1 Number of counties 102 54 46 102 Species average/county 2D mal 1.9 Lo 205 Winter Stoneflies of Illinois December 2002 Ta bl e 3. Th e hi st or ic al di st ri bu ti on (1 90 0- 20 00 ) of wi nt er st on ef li es wi th in th e Na tu ra l Di vi si on s of Il li no is Spe cie s/D ivi sio ns [| WD D [R RH CD ] NE MD | GP D_ [U M/ RB T] IM RS A] WF PD _[ MM BD | ST PD | WB D | OD [L MR BD |_ SH D | CP D | Occ urr enc e per Div isi on r ili noe nsi C o N T H D R I R E S E l O , Aii vic ala a r e a t a r a F a n ac e a a S Ce ) le a e E M a | A S | [ c r e | [ T = [n ec es | ca t] a c | ee e ee e | e e e | g o o e ae E P a B d EE A O N P e a e e e (y n 1 Ea ee e d E d a Sd E L a S e a e e SS L d E R S A T A R E EE E T C E S p e e r ee s | Ba e) e e e | | e e | e e s | e e e a s [ R E p e e | Se | e e S y a l a E M O T T R E W D D ( 1 * ) : Wi sc on si n Dr if tl es s Di vi si on . R R H D ( 2 ) : R o c k Ri ve r Hi ll Co un tr y Di vi si on . N E M D ( 3 ) : No rt he as t Mo ra in al Di vi si on . GP D( 4) : Gr an d Pr ai ri e Di vi si on . U M / I R B D ( 5 ) : Up pe r Mi ss is si pp i/ II li no is Ri ve r Bo tt om la nd s Di vi si on . I M R S A ( 6 ) : Il li no is /M is si ss ip pi Ri ve r Sa nd Ar ea s Di vi si on . W F P D ( 7 ) : We st er n Fo re st -P ra ir ie Di vi si on . M M B D ( 8 ) : Mi dd le Mi ss is si pp i B o t t o m l a n d s Di vi si on . S T P D ( 9 ) : So ut he rn Ti ll Pl ai n Di vi si on . W B D ( 1 0 ) : W a b a s h Bo rd er Di vi si on . OD (1 1) : Oz ar k Di vi si on . L M R B D ( 1 2 ) : L o w e r Mi ss is si pp i Ri ve r Bo tt om la nd s Di vi si on . S H D ( 1 3 ) * : S h a w n e e Hi ll s Di vi si on . CP D( 14 ): Co as ta l Pl ai n Di vi si on . *T he nu mb er in pa re nt he si s re fe rs to th e Na tu ra l Di vi si on s nu mb er ed on th e ma p in Fi gu re 2 ( S c h w e g m a n 19 73 ). 206 Illinois Natural History Survey Bulletin Vol. 36 Art. 5 Table 4. The distribution of winter stoneflies within the drainage basins in Illinois. LVBBKCS VMRS | LVBBKCS | aot BP Ya eye | te) ao peter eetereas| Bed apleraes | aes Elaine! ey ta) aa! Boel aes Se] 4 45 3 | 42) 4 ee ee [Species/Drainage | GAPRS | RRS | MMRT| DPRLMT| FRS | LVBBKCS| KIRS | VMRS| SPRS|LMRS| MRS | SRS | LIRABS| KRS | BMRS | A. forbesi 6 es Pe er | ee ae | ae | es OT ed al OP. Gl Ce EN LN CG I Lee A. illinoensis 8 Ye ep |r| a WOT ie PCG we A. nivicola 3) ee ee erie ee TST, re Ne Nd A OS ea RCD Pe PE 77 es i Se aoe FC i ee Cee) el Re ee Re iia Ail oe IN. caroling «250 3a) et] 988 7 | ht fonts eon] eae] See | | ee ey | |. ee IP. angulata | © fs Cea ts Se ee ge | gee De ee ee |e TT es i el el oe Te S.fasciata ee ee es es | eee Fe burkei SO. A] een 1) OC | PKS [ae aie: ee Xs ewe X eae ae Xe ee | | A Emir ri Tita 35,2 PS WIE] aS | Boats. ee) ee Seeger, tats |e ee ee ot | eae] Ly i ee Roe Rs Ree Eke ead ea Et CoS Se) eee) Be, es es oe Ee es a teat] [Te eee Red Bee ed et a lt | Eas Z. claasseni Oa! | ess] “he we] semen | Semen Fae Sane Soa Za Beco PA eee os ee) eed Meee ee Es Eel Znarp ate Teal CAPE RN) ae a Re OE SES | _Spectes Richness [ot | 4 | 37 8 GAPRS(1): Galena, Apple, Plum Rivers Systems. RRS(2): Rock River System. MMRT(3): Middle Mississippi River Tributaries. DPRLMT(4): Des Plaines River & Lake Michigan Tributaries. FRS(5): Fox River System. LVRBBKCS(6): Little Vermilion River, Big Bureau & Kickapoo Creek Systems. KIRS(7): Kankakee-Iroquois River Systems. VMRS(8): Vermilion-Mazon River Systems. SPRS(9): Spoon River System. LMRS(10): LaMoine River System. MRS(11): Mackinaw River System. SRS(12): Sangamon River System. LIRT-ABS(13): Lower Illinois River Tributaries-American Bottoms Systems. KRS(14): Kaskaskia River System. BMRS(15): Big Muddy River System. * The numbers in parentheses ( ) refer to the drainage basins on the map in Figure 3. December 2002 Winter Stoneflies of Illinois 207 Table 4 continued. 1900-2000 pecies/Drainages MBLBGPBCS A. illinoensis A. mystica |A. rickeri .vivipara N. carolina z | Tn ed Ee Eas P. completa . fasciata T. burksi T. lita T. metequi T. nivalis . parvula pears | Ex Z claasseni rae ee pear: | Z. fraxina an = “lelolel> > I> |> a pecies Richness 5 pecies/Drainages SalRS | LWRB a . forbesi . granulata . illinoensis . mystica . nivicola _recta eee hs | . rickeri F223 . smithi Simm .vivipara eae N. carolina ees Ella ee D> [> > [> [> | > | > [> | > [Mm — \©o — ON 3 P. angulata P. completa oe ee ae Bee ed Ea 2 ce a ER) a HL oo EE SM Pe ee Me Base coin te we ea SC Ee Z. claasseni SE AERA TR YE KEM) eSNG) | Ew: | Teves aR] dees PPS, boy oe ee VA OS ST a ee a SS EE ES ES a aa ed a es ee cease SSS) TR Bd ed ee CRS(16): Cache River System. MBLBGPBCS(17): Massac, Bay, Lusk, Big Grand Pierre, Big Creek Systems. SalRS(18): Saline River System. LWRBCS(19): Little Wabash River-Bonpas Creek Systems. ER-WRTS(20): Embarras River-Wabash River Tributaries. VLVRS(21): Vermilion-Little Vermilion River Systems. IR(22): Illinois River System.MR(23): Mississippi River System. OR(24): Ohio River System. WB(25): Wabash River System * The numbers in parentheses ( ) refer to the drainage basins on the map in Figure 3. 208 Illinois Natural History Survey Bulletin sity of winter stoneflies within these divisions over the past 100 years (Table 3) has undergone a significant decline. Winter stoneflies have been collected in 13 of these divisions with only the Illinois/Mississippi River Sand Areas lacking winter stoneflies. In the last century (1900-2000), winter stoneflies had been reported on 73 occasions in the natural divisions of Illinois with an average of 5.2 species per division. Fourteen species and 12 species were reported, respectively, from the Shawnee Hills and Wabash Border Divisions. The remaining natural divisions recorded 0-8 species at an average of 3.4 species per division. Early on, Frison and his colleagues (1900- 1945) recorded winter stoneflies on 58 occasions within the natural divisions with an average of 4.1 species per division. Twelve species were recorded from the Shawnee Hills Division and 10 species within the Wabash Border. The re- maining 12 divisions recorded 0-6 species. From 1946 to 1975, winter stoneflies were re- corded on only 40 occasions within the natural divisions with an average of 2.9 species per di- vision, with only 8 species found within the Shawnee Hills Division and 6 species within the Wabash Border Division. The remaining 12 di- visions recorded 0-5 species. During the recent resurvey (1976-2000), winter stoneflies were recorded on 52 occasions in the natural divisions with an average of 3 spe- cies per division. The Shawnee Hills Division (11 species) was still the most species- rich area although Allocapnia granulata, and Taeniopteryx lita have disappeared from this division. Six spe- cies—Allocapnia illinoensis, A. granulata, A. mystica, Nemocapnia carolina, Strophopteryx fasciata, and Zealeuctra claasseni—have disap- peared from the Wabash Border Division, al- though Allocapnia nivicola and Zealeuctra narfi have been added to this area recently. This divi- sion displayed the greatest loss of winter stoneflies. The remaining 12 divisions recorded 0-5 species with the Western Forest-Prairie and Ozark divisions reporting an increase in species richness. Overall, the species diversity of win- ter stoneflies within the Natural Divisions of II- linois has declined by an average of over two species per division. This decline appears to be a pattern over the entire state with the exception of the Western Forest Prairie and the Ozark di- visions. Vol. 36 Art. 5 Allocapnia vivipara (collected in 13 divi- sions) and Taeniopteryx burksi (collected in 10 divisions) displayed the greatest geographic range as measured by natural divisions. Allocapnia forbesi, A. illinoensis, A. smithi, Nemocapnia carolina, Paracapnia angulata, Prostoia completa, Taeniopteryx metequi, T. parvula, and Zealeuctra fraxina were restricted to one natural division. River Drainages Page et al. (1992) outlined the 25 river drain- ages within Illinois (Fig. 3) and the following summary evaluates the occurrence of winter stoneflies within these basins (Table 4) over the past century. The number in parentheses follow- ing the name of each drainage refers to Figure 3. Galena, Apple, and Plum River systems (1) These rivers and their tributaries lie within the Wisconsin Driftless Area of Illinois and drain 2,214 sq. km (855 sq. mi.) (Page et al. 1992). Streams in this area are highly variable in their substrates, fanging from gravel and silt to cobble and bedrock. Row crops and pasture form the major land use and silt run-off and barnyard pol- lution appear to be the major pollutants of the streams (Page et al. 1992). Smith (1971) rated the Apple River as good, with an excellent rating in the upper reaches. The Galena River was rated as good and the Plum River as fair. The [Illinois Water Quality Report (IEPA 1996) rated most of the Apple and Ga- lena Rivers as Full Support and 14 km of the Galena River, Coon Creek, Lilly Branch, Wolf Creek and the South Fork Apple River as Par- tial Support/Minor Impairment. The Illinois Water Quality Report (IEPA 1990) rated the Plum River and its tributary Carroll Creek as Partial Support/Minor Impairment. Phosphorus re- leased in municipal wastewater discharges and agricultural runoff led to the lower ratings. AlI- though this area is one of the most attractive ar- eas of Illinois, there are no “A” streams in this region (Hite and Bertrand 1989, Page et al. 1992). The rivers and tributaries in this area were rated from “B” to “C.” Only two species of winter stoneflies have - been collected from this area. Allocapnia rickeri was collected prior to 1946 in the Apple River drainage in and around Apple River State Park. Although several recent trips were made to this December 2002 area, this species was not collected. Much of the natural areas around the Apple River and it tributaries have been cleared for pastureland. Here, the trees and vegetation have been removed up to the stream edge. Allocapnia vivipara has been collected at several localities within Jo Daviess County, but no winter stoneflies have been collected in Carroll County through which much of the Plum River and its tributaries run. The absence of winter stoneflies from Carroll County is somewhat perplexing, as the county is not heavily impacted by agriculture and nu- merous pristine gravel streams run off the lime- stone bluffs along its western border. Rock River System (Rock, Pecatonica, Kishwaukee, and Green River drainages) (2) The Rock River (Page et al. 1992) drains an area of 13,838 sq. km and runs from the Wisconsin border north of Rockford to the Mississippi River at Rock Island. The river is predominately a gravel-bottomed stream interspersed with sand, rubble, and silt. The creation of channel dams has significantly altered the current and depth of this river. Channelization has been extensive in the southern and eastern areas of this basin. The principal land use is row crops and pasture leading to extensive siltation. This river has also had a history of domestic and industrial pollu- tion, especially below Rockford and Sterling. Phosphorus, siltation, and channel modifications have adversely affected this river and its tribu- taries (IEPA 1996). The Pecatonica River (Page et al. 1992) drains an area of 2,085 sq. km with the primary stream substrate being sand and silt. Phospho- rus, siltation, and municipal wastewater dis- charges have impacted this river and its tributar- ies (IEPA 1996). The Kishwaukee River (Page et al. 1992) drains an area of 3,173 sq. km and is separated into a North and South Branch. The upper reaches of the North Branch have a substrate of gravel, which changes to silt and sand down- stream. The South Branch consists primarily of rocks with a mixture of gravel and sand. Phos- phorus, organic enrichment, agricultural runoff, and municipal wastewater discharges have im- pacted this river and its tributaries (IEPA 1996). The Green River and its tributaries (Page et al. 1992) drain an area of 2,930 sq. km over a Winter Stoneflies of Illinois 209 lake plain of sand and gravel outwash from the Wisconsinan Glacier. Most of this river has been channelized. Siltation, agricultural runoff, channelization, and animal, domestic, and indus- trial wastes have vastly altered this stream (IEPA 1996). Smith (1971) rated the Rock River as good to excellent, except where it ran through exten- sive urban or industrial areas. Hite and Bertrand (1989) and Page et al. (1992) rated the Kishwaukee River, upstream from South Branch, Piscasaw Creek, upstream from West Branch, and the entire Rush Creek as “A” streams along with the Sugar River, and upstream from Otter Creek in the Pecatonica River drainage. Over the past century (1900-2000), seven species, Allocapnia granulata, A. mystica, A. vivipara, Strophopteryx fasciata, Taeniopteryx burksi, T. nivalis, and T: parvula, have been col- lected from this drainage area. Allocapnia mystica, S. fasciata, and T: parvula have not been collected here since the 1930s. Allocapnia vivipara, although it is the most common spe- cies of winter stonefly in Illinois, has not been collected often within the Rock, Pecatonica, and Green River drainages and is still scarce within these drainages. Allocapnia granulata is well established along the Rock River and its tribu- taries, as well as on the Green River and the Kishwaukee. Taeniopteryx burksi is the most widespread winter stonefly in this area, being found all along the Rock River, with populations still present on the Green River, the Pecatonica, and the Kishwaukee. Taeniopteryx nivalis is a northern species of Taeniopteryx that was first collected in Illinois along the Fox River and it tributaries in the 1960s. During the recent re- survey (1976-2000), it was collected widely along the upper Rock River, the Green River, the Pecatonica, and the Kishwaukee River. Middle Mississippi River tributaries (Edwards River, Henderson Creek, Bear Creek, Sny Creek, Bay Creek) (3) This area (Page et al. 1992) drains numerous small streams along the eastern Mississippi and western Illinois River bottomlands, which are primarily covered by the Middle Mississippi Border (Schwegman 1973). This is an area of mixed forest and upland prairie, although row crops and pasture are the primary land uses. 210 Illinois Natural History Survey Bulletin Agricultural siltation, herbicides, fertilizers, and livestock sewage contributes much of the pollu- tion to the streams of this area. The Edwards River (Page et al. 1992) drains an area of 1,129 sq. km, although it has been channelized for much of its length. The stream substrate is principally sand and silt. Henderson Creek (Page et al. 1992) and its tributaries drain an area of 1,572 sq. km with a bottom substrate of silt and sand with scattered gravel riffles. The lower extremities of this creek have been channelized. Bear Creek and its tributaries (Page et al. 1992) drain an area of 2,595 sq. km with a bot- tom substrate of sand and gravel. The lower extremities of this creek have been channelized as well as various tributaries. Sny Creek and its tributaries (Page et al. 1992) originally drained an area of 1,961 sq. km, but now after diversion drain 787 sq. km, witha bottom substrate of sand and gravel. Bay Creek (Page et al. 1992) drains an area of 456 sq. km with a bottom substrate of sand and gravel. The lower extremities of this creek are confined within a levee for 24 km before entering the Mississippi River. Smith (1971) categorized the streams of this area as poor to fair, with agricultural pollution, siltation, and desiccation from drought as the major environmental problems in this area. The [linois Water Quality Report (IEPA 1996) rated half of the Edwards River, much of Henderson River and Bear Creek as Full Support, with the upper portion of Cedar Creek and Middle Henderson Creek as Partial Support/Moderate Impairment and Bay Creek, half of the Edwards River and 126.5 stream km of Bear Creek as Partial Support/Minor Impairment. Nonsupport was limited to 2.1 km of Cedar Creek due to organic enrichment and elevated levels of am- monia. Hite and Bertrand (1989) and Page et al. (1992) listed no “A” streams in this area, with some streams in this area listed as “D” streams (streams with limited aquatic resource). Over the last century (1900-2000), four spe- cies of winter stoneflies, Allocapnia rickeri, A. vivipara, Strophopteryx fasciata, and Taeniopteryx burksi have been collected in the various streams of this drainage area. Allocapnia rickeri is rare in this area and recently has been collected at a single site in each of Calhoun and Vol. 36 Art. 5 Pike counties. Strophopteryx fasciata has not been collected in these streams since the 1920s. Currently, A. vivipara is widespread throughout this area. Taeniopteryx burksi was collected at two sites in Calhoun County during the 1920s but recently (1976-2000) has been collected at several sites in each of Calhoun and Pike coun- ties, as well as being widespread within the Edwards River and Henderson Creek drainages. Des Plaines River and Lake Michigan tribu- taries (Des Plaines, DuPage rivers) (4) This area (Page et al. 1992) encompasses prin- cipally Lake, Cook, DuPage, and Will counties and drains an area of 3,188 sq. km, which falls within the Northeastern Morainal and Grand Prairie Natural Divisions (Schwegman 1973). Over the past 40 years, urbanization and indus- trial use have heavily impacted it. Eutrophica- tion from treated and untreated sewage is a ma- jor pollution problem for streams in this drain- ACC Aes The Des Plaines River historically contained large areas of rapids, riffles, and rocky shallows, although today little remains of what was con- sidered a dangerous river (Vierling 1977). The river bottom is bedrock covered with sand and gravel (Page et al. 1992) although the sand and gravel is often imbedded with organic muck. Considerable dredging and channelization has altered this stream, in addition to the construc- tion of two major lock and dam structures. This stream has been heavily impacted by pollution. The DuPage River (Page et al. 1992) has two major branches with the West Branch having numerous riffles and gravel flats, and the East Branch having a substrate of silt and muck. The DuPage River proper contains numerous riffles and small rapids. Forty-one km of this stream have been channelized and several small dams impound its flow. Smith (1971) rated the Des Plaines River as poor with domestic and industrial sewage being the major pollution problem. The Illinois Water Quality Report (IEPA 1996) rated 80% of the Des Plaines River as Partial Support/Minor Im- pairment and 17% as Partial Support/Moderate | Impairment with nearly 90% of the DuPage River rated as Full Support. Hite and Bertrand (1989) and Page et al. (1992) rated Manhattan Creek as an “A” stream. December 2002 Over the past century (1900-2000), only Allocapnia vivipara and Taeniopteryx burksi have been collected in this drainage. Allocapnia vivipara can still be collected in northern Will County, but Taeniopteryx burksi has not been collected in this drainage since the 1920s. Fox River system (5) The Fox River and its tributaries (Page et al. 1992) drain an area of 4,455 sq. km and fall within the Northeastern Morainal and Grand Prairie Natural Divisions (Schwegman 1973). The upper portion of this drainage contains nu- merous natural lakes, while the lower portion of the Fox River and its tributaries have a bottom substrate of gravel, cobble, and sand. Smith (1971) rated the Fox River as good to excellent with domestic and industrial pollution its dominant pollution problems. The Illinois Water Quality Report (EPA 1996) rated 80.4% of the Fox River drainage as Full Support with 15.6 % as Partial Support/Minor Impairment. Hite and Bertrand (1989) and Page et al. (1992) rated Buck Creek as the only “A” stream in this drainage. Over the past century (1900-2000), six spe- cies of winter stoneflies, Allocapnia granulata, A. vivipara, Paracapnia angulata, Strophopteryx fasciata, Taeniopteryx burksi, and T: nivalis have been collected within this drainage. Paracapnia angulata and Strophopteryx fasciata have not been collected here since the 1920s. Paracapnia angulata was restricted in Illinois to several seep springs in the Elgin Botanical Garden (now Trout Park) and was not collected recently although several specific trips were made to this site in search of this species. Allocapnia granulata has not been collected within this drainage since the 1960s. Allocapnia vivipara and Taeniopteryx burksi are still found abundantly within this drainage. Taeniopteryx nivalis began being col- lected within this drainage in the 1960s and re- cently (1976-2000) has expanded its range throughout this drainage. Little Vermilion River, Big Bureau and Kickapoo Creek systems (6) The Little Vermilion River, Big Bureau Creek, Kickapoo Creek, and their tributaries are located in north-central Illinois on the west side of the Illinois River (Page et al. 1992) and fall within the Grand Prairie, Upper Mississippi River and Winter Stoneflies of Illinois Zhi [Illinois Bottomlands, and Western Forest-Prai- rie Natural Divisions (Schwegman 1973). Row crops and pasture are the principal land use within this drainage. The Little Vermilion River (Page et al. 1992) drains an area of 326 sq. km with a bottom sub- strate of sand and gravel. Smith (1971) rated this drainage as fair. The Illinois Water Quality Report (IEPA 1990) rated this drainage as Par- tial Support/Moderate Impairment. Big Bureau Creek (Page et al. 1992) drains an area of 1,259 sq. km with a bottom substrate of sand and gravel. Smith (1971) rated this drain- age as good to excellent. The Illinois Water Quality Report (IEPA 1996) rated 76.2% of this drainage as Full Support and 23.6% as Partial Support/Minor Impairment. Kickapoo Creek (Page et al. 1992) drains an area of 793 sq. km, principally in Peoria County. Much of this area is utilized for row crops, which has created a problem with siltation. Local dis- charge of domestic sewage is also a pollution problem. Smith (1971) rated this drainage as good. The Illinois Water Quality Report (IEPA 1990) rated this drainage as Partial Support/ Minor Impairment. Smith (1971) rated Big Bureau Creek as good to excellent, Kickapoo Creek as good, and the Little Vermilion River as fair. Hite and Bertrand (1989) and Page et al. (1992) rated no streams in this system as “A.” Over the past century (1900-2000), four spe- cies of winter stoneflies, Allocapnia granulata, A. vivipara, Strophopteryx fasciata, and Taeniopteryx burksi have been collected within this drainage. Both Allocapnia granulata and Strophopteryx fasciata were collected in this system at single localities during the 1960s but were not collected recently (1976-2000). Allocapnia vivipara and Taeniopteryx burksi were collected at scattered localities by both Frison (1900-1945) and Ross (1946-1975) but after the recent resurvey (1976-2000) are con- sidered widespread through these drainages. Kankakee and Iroquois River systems (7) The Kankakee River System (Page et al. 1992) drains an area of 5,618 sq. km principally in the Grand Prairie Natural Division (Schwegman L973): 212 [llinois Natural History Survey Bulletin The Kankakee River flows through the flat terrain on an old glacial lake (Page et al. 1992) over a substrate of bedrock covered with a thin layer of sand and gravel upstream of Kankakee. Downstream from Kankakee the river runs through silted pools and long stretches of bed- rock. The Iroquois River (Page et al. 1992) drains an area of 3,212 sq. km, principally in [Iroquois County, with primarily a silt substrate. Smith (1971) rated the Kankakee and Iroquois rivers as excellent. The [linois Water Quality Report (IEPA 1996) rated 89.3 % of this drainage as Full Support and 2.8% as Full Sup- port/Threatened. Hite and Bertrand (1989) rated no streams in this system as “A.” Over the past century (1900-2000), five spe- cies of winter stoneflies, Allocapnia granulata, A. vivipara, Strophopteryx fasciata, Taeniopteryx burksi, and T: nivalis have been collected within this drainage area. No winter stoneflies were collected in this drainage basin by Ross and his colleagues (1946-1975). Allocapnia granulata has not been collected here since the 1920s. Recently (1976-2000), Taeniopteryx nivalis has moved into this area and has been collected in both the Kankakee and Iroquois River drainages. Strophopteryx fasciata is somewhat rare in this system but is still present at two localities along the Iroquois River. Both Allocapnia vivipara and Taeniopteryx burksi are widespread throughout this area. Vermilion and Mazon River systems (8) The Vermilion and Mazon rivers (Page et al. 1992) drain an area of 5,180 sq. km in central [llinois south of the Illinois River, predominately in the Grand Prairie Natural Division (Schwegman 1973). Row crops are the domi- nant land use and siltation and agricultural run- off are the major pollution problems. The Vermilion River (Page et al. 1992) drains an area of 3,447 sq. km. Upstream of Streator, the substrate is sand, silt, and gravel. Down- stream of Streator, the substrate becomes gravel and cobble. Smith (1971) rated this river as fair with domestic and agricultural pollution a prob- lem. The Illinois Water Quality Report (IEPA 1996) rated 85.4 % of this drainage as Full Sup- port and 14.6% as Partial Support/Minor Im- pairment. Vol. 36 Art. 5 The Mazon River (Page et al. 1992) drains an area of 1,419 sq. km with a bottom substrate of gravel and rock. Smith (1971) rated this river as good. The Illinois Water Quality Report (EPA 1996) rated this drainage as Full Support. Hite and Bertrand (1989) rated no “A” streams in this area. Over the past century (1900-2000), four spe- cies of winter stoneflies, Allocapnia granulata, A. vivipara, Strophopteryx fasciata, and Taeniopteryx burksi, have been collected in this area. Strophopteryx fasciata has not been col- lected here since the 1920s, and Allocapnia granulata has not been taken here since the 1960s. Most recently (1976-2000), only the two commonest species of winter stoneflies in Illi- nois, Allocapnia vivipara and Taeniopteryx burksi, have been collected in this drainage sys- tem. Spoon River system (9) The Spoon River (Page et al. 1992) drains an area of 4,804 sq. km on the west side of the IIli- nois River, predominately in the Western For- est-Prairie Division with the upper reaches in the Grand Prairie Natural Division (Schwegman 1973). This drainage system flows through flat prairie land that is now dominated by row crops and has a bottom substrate of gravel and sand. The major pollution problems are siltation, pes- ticides, stripmine and industrial waste, and do- mestic and animal waste. Smith (1971) rated this drainage area as fair, with excessive siltation and agricultural pollution as problems. The Illi- nois Water Quality Report (IEPA 1996) rated portions of the Spoon River and most of the tribu- taries as Full Support and some portions of the Spoon River and Big Creek as Partial Support/ Minor Impairment. Hite and Bertrand (1989) and Page et al. (1992) rated no “A” streams in this area. Over the past century (1900-2000), four spe- cies of winter stoneflies, Allocapnia vivipara, Strophopteryx fasciata, Taeniopteryx burksi, and T. nivalis, have been collected within this drain- age. Strophopteryx fasciata has not been col- lected here since the 1920s and T. nivalis, a win- ter stonefly that is expanding its distribution in northern Illinois, has recently (1976-2000) been — collected here at a single site. Allocapnia vivipara and Taeniopteryx burksi are both wide- spread within this drainage. December 2002 LaMoine River system (10) The LaMoine River drainage (Page et al. 1992) consists primarily of the LaMoine River and McKee Creek. This system encompasses an area of 3,497 sq. km principally in the Western For- est-Prairie Natural Division (Schwegman 1973), with bottom substrates principally of sand with some gravel. Smith (1971) rated the LaMoine River as fair with siltation and agricultural pollution affect- ing the quality of the LaMoine River and its tribu- taries. The Illinois Water Quality Report (IEPA 1996) rated 79.2% of this drainage as Full Sup- port and 20% as Partial Support/Minor Impair- ment. An 11-km stretch of the South Branch of the LaMoine River was rated as Partial Support/ Moderate Impairment. Hite and Bertrand (1989) and Page et al. (1992) rated the LaMoine River from Flour Creek to Cedar Creek as an “A” stream. Over the past century (1900-2000), three species of winter stoneflies, Allocapnia rickeri, A. vivipara, and Taeniopteryx burksi have been collected in this system. Allocapnia vivipara has been recorded from this drainage since the 1920s. Recently (1976-2000), Taeniopteryx burksi has been collected at several localities and A. rickeri at one site along the LaMoine River. This is one of the few instances when the overall species diversity has increased within a drainage sys- tem. Mackinaw River system (11) The Mackinaw River (Page et al. 1992) and its three main tributaries, Panther, Walnut, and Money creeks, drain an area of 2,942 sq. km pri- marily in the Grand Prairie Natural Division (Schwegman 1973) of central Illinois. The bot- tom substrate of these streams is sand and gravel with scattered silt and cobble. The Mackinaw River has no dams on it, but the upper 17.7 km have been channelized. Ninety percent of the drainage area has been converted to row crops, and siltation and agricultural pollution are the major environmental problems. Smith (1971) rated this drainage as good to excellent and the Illinois Water Quality Report (IEPA 1996) rated this drainage as Full Support. Hite and Bertrand (1989) and Page et al. (1992) rated several tributaries and parts of the Macki- naw River as “A” streams. As a percentage of Winter Stoneflies of Illinois 213 the total drainage area, the Mackinaw River sys- tem has the highest number of “A” streams in Illinois. Over the past century (1900-2000), four spe- cies of winter stoneflies, Allocapnia granulata, A. vivipara, Strophopteryx fasciata, and Taeniopteryx burksi, have been collected within this system. Strophopteryx fasciata has not been collected here since the 1920s and Allocapnia granulata has not been reported here since the 1960s. Allocapnia vivipara and Taeniopteryx burksi are still well established within this sys- tem. Sangamon River system (12) The Sangamon River system (Page et al. 1992) drains an area of 14,035 sq. km primarily in the Grand Prairie Natural Division (Schwegman 1973) of central Illinois. The topography of the basin is rolling with flat valleys along the rivers. Five major impoundments impede the flow of this stream. Two areas, between Decatur and Springfield and between Petersburg and Beardstown, have been channelized. Row crops cover much of the topography and coal mining occurs within the southern half of this basin. Siltation, agricultural pesticides and fertilizer, domestic sewage, and acid mine waste are the dominant pollutants of the streams. The bottom substrate of the streams is sand, with gravel, silt, and cobble. Smith (1971) rated the Sangamon from good to fair. The Illinois Water Quality Report (IEPA 1990) rated most of this river as Partial Sup- port/Minor Impairment with 6 km rated as Par- tial Support/Moderate Impairment. Hite and Bertrand (1989) and Page et al. (1992) rated three streams, Drummer, Goose, and Ten Mile creeks, as “A.” Most tributaries were rated “B,” with the main stream of the Sangamon as “C.”” Asmall segment of the upper Sangamon was rate “D.” Over the past century (1900-2000), four spe- cies of winter stoneflies, Allocapnia granulata, A. vivipara, Strophopteryx fasciata, and Taeniopteryx burksi, have been collected within this system. Strophopteryx fasciata has not been collected here since the 1920s and Allocapnia granulata has not been reported here since the 1960s. Allocapnia vivipara and Taeniopteryx burksi are still well established within this sys- tem. 214 Illinois Natural History Survey Bulletin Lower Illinois River tributaries and Ameri- can Bottoms (13) The American Bottoms and the tributaries of the lower Illinois River (Page et al. 1992) drain an area of approximately 5,856 sq. km, primarily in the Western Forest-Prairie, Lower Mississippi River Bottomlands, and the Ozark Natural Di- vision (Schwegman 1973) in southwestern [li- nois. The topography is rolling uplands and bot- tomlands along the Illinois and Mississippi riv- ers. Row crops form the major land use, with urban and industrial development prevalent in the southern bottomland around East St. Louis. Siltation, pesticides, and fertilizers from the up- lands and domestic and industrial waste in the lowlands form the major pollution problems to streams in this system. Sand forms the predomi- nate bottom substrate with some gravel and cobble with considerable silt in the lower reaches. Smith (1971) and Page et al. (1992) rated sev- eral streams in this area as variable, with Wood River and Cahokia Creek as poor. The Illinois Water Quality Report (IEPA 1990) rated the up- per reaches of Cahokia Creek, Sandy Creek, and part of Macoupin Creek as Full Support with most streams rated as Partial Support/Minor Use. The lower 4.8 km of Wood River were rated as Non Support. Hite and Bertrand (1989) and Page et al. (1992) rated streams in this region as “B” and “C.” Siltation, pesticides and fertilizers from agricultural practices, as well as pollution from coalmines and oil fields affect the quality of streams in this area. Over the past century (1900-2000), only Allocapnia vivipara has been reported from this area and it has been consistently collected in this drainage since the 1920s. Kaskaskia River system (14) The Kaskaskia River drainage (Page et al. 1992) extends from Champaign County in east-central [llinois to the Mississippi River and drains an area of 15,022 sq. km within the Grand Prairie, Southern Till Plain, and Ozark Natural Divisions (Schwegman 1973). Sand and gravel is the pri- mary substrate for most streams with some silt. Shelbyville and Carlyle Reservoirs form two major impoundments on this stream and the lower reaches have been channelized. Row crops form the major land use but coal underlies much Vol. 36 Art. 5 of this basin and several oilfields are active. Silt- ation, pesticides and fertilizers from agricultural practices, as well as pollution from coalmines and oil fields affect the quality of streams in this area. Smith (1971) rated the Kaskaskia River as variable and the Illinois Water Quality Report (IEPA 1996) rated 2,407 km as Full Support, 96 stream km as Full Support/Threatened, 1,841 stream km as Partial Support/Minor Impairment, and 397 stream km as Partial Support/Minor Impairment. Hite and Bertrand (1989) and Page et al. (1992) rated parts of Ramsey Creek and the West Okaw River as “A.” The majority of the basin was rated as “B” with the lower reaches rated as “C” or “D.” Over the past century (1900-2000), six spe- cies, Allocapnia granulata, A. mystica, A. rickeri, A. vivipara, Taeniopteryx burksi, and T: lita, have been reported from this system. Allocapnia granulata, A. mystica, and A. rickeri have not been collected in this drainage since the 1960s. Taeniopteryx lita has always been rare in this drainage, and recent collecting (1976-2000) found it at only one locality. Allocapnia vivipara and Taeniopteryx burksi have always been wide- spread over this system. Big Muddy system (15) The Big Muddy system (Page et al. 1992) drains 6,112 sq. km in southern Illinois within the Southern Till Plain, Shawnee Hills, and Lower Mississippi River Bottomlands Natural Divisions (Schwegman 1973). Half of this drainage area supports row crops and coal underlies the south- western part of this basin. The bottom substrate of most streams is clay and three impoundments Rend, Crab Orchard, and Kinkaid reservoirs impede the flow of this system. Siltation, pesti- cides and fertilizers from agricultural practices, as well as pollution from coalmines and oil fields affect the quality of streams in this area. Smith (1971) rated the lower reaches as good and the upper reaches above Murphysboro as very poor. The Illinois Water Quality Report (IEPA 1996) rated 15.3% of this drainage as Full Sup- port, 81% as Partial Support/Minor Impairment, and 3.6% as Partial Support/Minor Impairment. Hite and Hebrand (1989) and Page et al. (1992) rated Miller creek as an “A” stream, but the re- mainder of the system varied from “B” to “D.” December 2002 Over the past century (1900-2000), six spe- cies, Allocapnia mystica, A. rickeri, A. vivipara, Prostoia completa, Zealeuctra claasseni, and Z. narfi, have been collected within this system. Zealeuctra narfi has not been collected here since the 1920s, but Z. claaseni was recently (1976- 2000) collected in a tributary of the Big Muddy River. Prostoia completa is a rare species in Il- linois, collected at a single site by Frison (1900- 1945) from Hutchins Creek and has been recol- lected there recently (1976-2000). Allocapnia mystica, A. rickeri, and A. vivipara are still well established within this system. For some un- known reason, no species of Taeniopteryx have been collected within this drainage basin. Cache River system (16) The Cache River system (Page et al. 1992) drains an area of 2,717 sq. km in southern Illinois pri- marily within the Coastal Plain Natural Division (Schwegman 1973), with a small portion of its upper reaches extending into the Ozark and Shawnee Hills divisions. The upper reaches drain through rolling hills and the stream sub- strate is primarily gravel riffles, but once out of the Shawnee Hills, the basin is low-lying, and has been severely impacted by dredging and channelization with the substrate becoming silt. Row crops and the Shawnee National Forest cover most of this basin, and siltation, pesticides, and fertilizers from agricultural practices affect the quality of streams in the lower reaches. Smith (1971) rated the Cache River as good, and the Illinois Water Quality Report (IEPA 1990) rated this drainage as Partial Support/ Minor Impairment, with Lick Creek rated as Full Support. Hite and Bertrand (1989) and Page et al. (1992) rated the streams within this basin as “C” (moderate aquatic resource). Over the past century (1900-2000), three species, Allocapnia mystica, A. rickeri, and A. vivipara, have been collected from this drain- age. Allocapnia mystica and A. rickeri have not been collected in this drainage since the 1960s. Allocapnia vivipara has remained well estab- lished within this system. Massac, Bay, Lusk, Big Grand Pierre, and Big Creek systems (17) These systems of streams (Page et al. 1992) drain a small area of 1,331 sq. km, principally in the Winter Stoneflies of Illinois mis Shawnee Hills Natural Division (Schwegman 1973) with Massac Creek and Mud Creek fall- ing within the Coastal Plain Division. The to- pography here is primarily rolling hills in the Shawnee Hills and flat lowland in eastern Massac County. The Shawnee National Forest covers much of this area, with row crops being culti- vated in the lowlands. Stream substrates are pri- marily gravel with patches of bedrock, cobble, and some sand. Streams in the lowlands have bottom substrates of silt. Occasional siltation is a problem in the upper reaches but heavier silt- ation, pesticides, and fertilizers from agricultural practices affect the quality of lowland streams in eastern Massac County. Smith (1971) rated Big Creek and Lusk Creek as outstanding streams in this area, with the majority of streams being rated as excellent, with only the lowland streams in eastern Massac County being rated as good. The Illinois Water Quality Report (EPA 1990, 1996) rated the streams in the Shawnee Hills area as Full Sup- port with Massac and Mud Creeks being con- sidered Partial Support/Minor Impairment. Hite and Bertrand (1989) and Page et al. (1992) rated upper Lusk and Big Creeks as “A” streams with the remaining streams in Hardin and Pope coun- ties as “B” except for their lowest extremities. Over the past century (1900-2000), 13 spe- cies of winter stoneflies, Allocapnia forbesi, A. granulata, A. mystica, A. rickeri, A. smithi, A. vivipara, Strophopteryx fasciata, Taeniopteryx burksi, T. lita, T: metequi, Zealeuctra claasseni, Z. fraxina, and Z. narfi, have been collected from this region. Only Allocapnia granulata and Taeniopteryx lita have not been collected here since the 1920s. All of the other species are still well established within this region. Saline River system (18) The Saline River System (Page et al. 1992) drains an area of 3,048 sq. km primarily within the Southern Till Plain and the Wabash Border Natural Divisions (Schwegman 1973) of south- eastern Illinois, with a small amount of the Sa- line River and the South Fork of the Saline River extending into the Shawnee Hills Division. Much of this basin is flat with the bottom sub- strate of the slow moving streams being covered with silt. Row crops form the dominant land 216 Illinois Natural History Survey Bulletin use, but coal mining and oil wells are present. Siltation, pesticides, and fertilizers from agricul- tural practices as well as leaching of acid water from abandoned coal mines, strip mine waste, and leakage of brine from oil wells affect the quality of streams in this basin (Allen and Wayne 1973, Page et al. 19972). Smith (1971) rated the Saline River as poor and the Illinois Water Quality Report (IEPA 1996) rated a 28-km reach of Bear Creek in the North Fork and Sugar Creek (except for the low- ermost 46.8 km) in the South Fork as Full Sup- port; the North Fork, Middle Fork, and the lower 2.3 km of the Saline River as Partial Support/ Minor Impairment; a 22.4-km reach of the Sa- line River below the confluence of the North and South Forks as Partial Support/Moderate Im- pairment, and 3.4 km of the South Fork and 6.8 km of Sugar Creek as Nonsupport. Hite and Bertrand (1989) and Page et al. (1992) rated no “A” streams within this drainage, with the streams varying in characterization from “B” (highly valued aquatic resource) to “D” (limited aquatic resource). Over the past century (1900-2000), six spe- cies of winter stoneflies, Allocapnia forbesi, A. mystica, A. rickeri, A. smithi, A. vivipara, and Zealeuctra claasseni, have been collected from this area. Allocapnia rickeri and A. smithi were not collected by Frison (1900-1945) or Ross (1946-1975) from this drainage, and for some unknown reason species of Taeniopteryx have not been collected in this drainage. Still, overall there has been a increase in the diversity of spe- cies collected within this system. Little Wabash River and Bonpas Creek sys- tems (19) This drainage system (Page et al. 1992) encom- passes an area of 8,936 sq. km primarily in the Southern Till Plain and Wabash Border Natural Divisions (Schwegman 1973) of southeastern Illinois. This area is characterized by broad flat uplands and U-shaped valleys with the substrate of the streams consisting of silt and sand. Land use 18 primarily row crops, with all of the area underlain by coal (Barker et al. 1967), in addi- tion to one-third of the oil produced in Illinois coming from this area (Page et al. 1992). Silt- ation, pesticides, fertilizers from agricultural practices, and oil field pollution affect the qual- ity of streams in this basin. Vol. 36 Art. 5 Smith (1971) rated the lower reaches of the Little Wabash River as poor and the upper reaches as good. He rated Bonpas Creek as fair. The Illinois Water Quality Report (IEPA 1990) has generally rated the Little Wabash River as Partial Support/Minor Impairment. Hite and Bertrand (1989) and Page et al. (1992) rated the streams in this system as “B” (highly valued aquatic resource) and “C” (moderate aquatic re- source). Over the past century (1900-2000), five spe- cies of winter stoneflies, Allocapnia rickeri, A. vivipara, Strophopteryx fasciata, Taeniopteryx burksi, and T: lita, have been collected in this system. Allocapnia rickeri and Taeniopteryx lita were not collected by either Frison (1900-1945) or Ross (1946-1975) in this system, but recently (1976-2000), each species has been collected at single localities. TYaeniopteryx burksi has not been collected here since they 192Z0seand Strophopteryx fasciata not since the 1960s. Only Allocapnia vivipara is well established in this basin. Embarras River and Wabash River tributar- ies (20) The Embarras River and Wabash River tributar- ies (Page et al. 1992) encompass an area of 7,646 sq. km in southeastern Illinois within the Grand Prairie, Southern Till Plain, and Wabash Border Natural Divisions (Schwegman 1973). This ba- sin has a topography of rolling hills and low flat land in the Wabash bottoms. The substrate of streams is gravel and sand in the upper reaches and silt in the lowlands. Row crops are the prin- cipal land use, with scattered oil fields. Siltation, pesticides, fertilizers from agricultural practices, and oil field pollution affect the quality of streams 1n this basin. Smith (1971) rated the Embarras River as variable, with the middle reaches of the Embarras designated as one of Illinois’ outstanding streams (Evers and Page 1977, Smith 1971). The Ilh- nois Water Quality Report (IEPA 1996) rated 51.9% of the Embarras River as Full Support, 43.3% was rated as Partial Support/Minor Im- pairment, and 4.8% was rated as Full Support/ - Threatened. Hite and Bertrand (1989) and Page et al. (1992) rated Riley Creek as an “A” stream. Most of this basin was rated “B” with the lower reaches rated “C.” December 2002 Over the past century (1900-2000), eight species of winter stoneflies, Allocapnia granulata, A. illinoensis, A. mystica, A. recta, A. vivipara, Strophopteryx fasciata, Taeniopteryx burksi, and T: lita, have been collected within this basin. Allocapnia mystica has not been col- lected here since the 1920s and A. granulata, A. illinoensis, and S. fasciata have not been col- lected here since the 1960s. Taeniopteryx lita was not collected in this drainage by either Frison (1900-1945) or Ross (1946-1975), but recently (1976-2000) was collected at a single locality. Allocapnia recta has been uncommon in this drainage basin but still can be found at two lo- calities. Allocapnia vivipara and Taeniopteryx burksi remain widespread throughout this sys- tem. Vermilion and Little Vermilion River systems (21). The Vermilion and Little Vermilion systems (Page et al. 1992) drain an area of 4,268 sq. km in east-central Illinois within the Grand Prairie and Wabash Border Natural Divisions (Schwegman 1973). Much of the topography here is gently rolling with stream substrates be- ing sand and gravel with some cobble and silt. Row crops are the major land use but much of this area is also underlain by coal. Siltation, pesticides and fertilizers from agricultural prac- tices, and domestic sewage affect the quality of streams in this basin. Smith (1971) rated the streams in this basin as variable, with the Middle Fork of the Vermil- ion River as one of the outstanding streams in the state. The Illinois Water Quality Report (IEPA 1996) rated all of the Vermilion River and Little Vermilion River as Full Support. Hite and Bertrand (1989) and Page et al. (1992) rated Jor- dan Creek, Spoon River, and the Middle Fork from Knights Branch to its mouth as “A.” The remainder of the basin was rated “B” with the Little Vermilion River not being rated. Over the past century (1900-2000), six spe- cies of winter stoneflies, Allocapnia granulata, A. mystica, A. nivicola, A. recta, A. vivipara, and Taeniopteryx burksi, have been collected in this system. Allocapnia mystica has not been col- lected since the 1920s, and A. granulata has not been collected since the 1960s. Allocapnia nivicola is rare in Illinois and a single new lo- cality was found at Forest Glen Forest Preserve. Winter Stoneflies of Illinois 217 Allocapnia recta, A. vivipara, and Taeniopteryx burksi remained well established within this drainage basin. Illinois River (22) The Illinois River proper (Page et al. 1992) ex- tends 439 km from the confluences of the Kankakee and Des Plaines rivers to the Missis- sippi. Much of it length has undergone human disturbances from the dumping of untreated sew- age and industrial waste to the development of six dams. Much of the original flow of this river has now been impeded and the substrate is pri- marily sand and silt. Row crops are the major land use along this river and siltation, pesticides, and fertilizers from agricultural practices se- verely affect the quality of this stream. The Ulinois Water Quality Report (IEPA 1996) rated all of the mainstem of the Illinois River as Full Support/Minor Impairment. Over the past century (1900—2000), four spe- cies of winter stoneflies, Allocapnia granulata, A. vivipara, Taeniopteryx burksi, and T. nivalis have been collected from this river. Recently (1976-2000), Allocapnia granulata and Taeniopteryx nivalis each have been collected at one locality along the river. Allocapnia vivipara and Taeniopteryx burksi remain well established in the Illinois River. Mississippi River (23) The Mississippi River proper (Page et al. 1992) forms a 1,094-km border along the western side of Illinois. Because of the need for navigation in this river, much of it original character has been altered. Twenty-six locks and dams from Minneapolis, Minnesota, to Alton, Illinois, have seriously impounded the flow of this river. The substrate is principally sand. Siltation has been a major form of pollution as well as domestic and industrial pollution and run off of agricul- tural pesticides and fertilizers into the streams that feed this river. The Illinois Water Quality Report (IEPA 1990) rated the Mississippi River from East Dubuque to Quincy as Full Support, from Quincy to the mouth of Chain of Rocks Canal as Partial Support/Minor Impairment, from there to the Meramec River as Non Support, and from the Meramec River to the Ohio River as Partial Support/Moderate Impairment. 218 Illinois Natural History Survey Bulletin Over the past century (1900—2000), two spe- cies, Allocapnia vivipara and Taeniopteryx burksi, have been collected along this river. No winter stoneflies were collected by Frison (1900— 1945) or Ross (1946-1975) from the Mississippi River. Both records were made during recent collecting (1976-2000). Attempts to collect winter stoneflies along this river were made from several of the bridges that cross the river, but little collecting along the shoreline proper was conducted. Ohio River (24) The Ohio River proper (Page et al. 1992) ex- tends from its confluence with the Wabash River across the southern tip of Illinois to the Missis- sippi River. Its flow has been impeded over it entire length by 20 locks and dams, creating a substrate of silt. Over the past century (1900—2000), four spe- cies, Allocapnia vivipara, Strophopteryx fasciata, Taeniopteryx burksi, and T: lita, have been collected along this river in Illinois. Strophopteryx fasciata, and T: lita have not been collected from the river since the 1920s, but A. vivipara and T: burksi are still established. Wabash River (25) The Wabash River proper (Page et al. 1992) runs for 319 km along the southeast border of Illi- nois. The substrate is sand, gravel, and rock with pools and riffles. No locks or dams impede the flow of this river. The Illinois Water Quality Report (IEPA 1996) rated the Wabash River from Terre Haute, Indiana, to the Ohio River as Partial Support/ Minor Impairment. Industrial, municipal, and agricultural pollution impinges upon the quality of this river. Over the past century (1900-2000), four spe- cies of winter stoneflies, Allocapnia vivipara, Nemocapnia carolina, Strophopteryx fasciata, and Taeniopteryx burksi, have been collected from the Illinois portion of this river. Nemocapnia carolina has not been collected along the Illinois portion of the river since the 1920s, and Allocapnia vivipara, Strophopteryx fasciata, and Taeniopteryx burksi have not been collected along here since the 1960s. Although numerous trips were made recently (1976-2000) to collect winter stoneflies along the Wabash Vol. 36 Art. 5 River, not a single specimen was collected. This is a rather strange situation as easy access to the shoreline of the river is available at several sites. Over the past 100 years, the species richness of winter stoneflies within the 25 drainage ba- sins of Illinois ranged from | in the lower Illi- nois River-American Bottoms to 13 within the Massac, Bay, Lusk, Big Grand Pierre, and Big Creek System. Following the recent resurvey (1976-2000), the species richness ranged from 0 in the Wabash River (where 4 species had been collected previously) to 11 within the Massac, Bay, Lusk, Big Grand Pierre, and Big Creek System. Twenty-two of the 25 drainage basins displayed a reduction in species richness. Four species (Allocapnia illinoensis, Nemocapnia carolina, Paracapnia angulata, and Taeniopteryx parvula) are considered extirpated from the state and eight other species displayed a reduction in the diversity of drainage basins in which they had previously been collected. Strophopteryx fasciata and Allocapnia granulata which historically had been collected in 12 and 11 drainage basins respectively, had their distri- butions reduced to 2 drainage basins. SUMMARY Over the past century (1900-2000), the species diversity of winter stoneflies averaged 2.5 spe- cies per county with species reported from ev- ery county but 3 (Carroll, DuPage, Ford) and with 10 counties recording 5 or more species. Pope County (14 species) reported the greatest species diversity of winter stoneflies. Frison (1900-1945), collected in 54 counties finding an average of 2.1 species per county. Ross and his “winter stonefly club” (1946-1975) col- lected in 46 counties, reporting an average of 1.9 species per county. During the recent resurvey of the state (1976— 2000), attempts were made to collect winter stoneflies from every county, with an average of 1.9 species per county collected. Compared to the overall picture of winter stoneflies in Illinois (1900-2000), species diversity within the various counties has dwindled and now winter stoneflies are not found in 11 counties, and only 3 counties — (Hardin, Pope, and Saline) exhibited 5 or more species. Allocapnia granulata and Taeniopteryx lita have disappeared from species-rich Pope December 2002 County. Saline County with nine species, is in- teresting in that six of these species were only collected recently. Clark County with eight spe- cies, has lost five species since 1975; four of these species were from Rocky Branch, a favor- ite collecting site of Frison and Ross. Hardin County with seven species, has lost two species since 1946. Seven species were found to be com- mon (currently known from more than 15 lo- calities): Allocapnia forbesi, restricted to the Shawnee Hills; A. granulata, its previous distri- bution reduced and now restricted to the Rock River drainage; A. mystica, its previous distri- bution reduced and now restricted to the Shawnee Hills; A. rickeri, abundant in the Shawnee Hills, with extensions up the eastern and western borders of Illinois; A. vivipara and Taeniopteryx burksi, widespread throughout state; T nivalis, previously rare but now spread- ing its distribution across northern Illinois. Four species are considered uncommon (currently known from 4—15 localities): Allocapnia recta, scattered in eastern Illinois; Strophopteryx fasciata, widespread but disappearing from the state; Taeniopteryx metequi and Zealeuctra claasseni, restricted to the Shawnee Hills. Six species are considered rare (currently known from 1-3 localities): Allocapnia nivicola, known only from a single locality in Vermilion County; A. smithi, found in the eastern Shawnee Hills; Prostoia completa, known only from Hutchins Creek in the western Shawnee Hills; Taeniopteryx lita, scattered in east central I]li- nois; Zealeuctra fraxina, eastern Shawnee Hills; and Z. narfi, found at single localities in Saline, Union, and Vermilion counties. Four species are considered extirpated from Illinois: Allocapnia illinoensis, Nemocapnia carolina, Paracapnia angulata, and Taeniopteryx parvula. Over the past century (1900-2000), the vari- ous species of winter stoneflies had been reported on 73 occasions within the 14 Natural Divisions of Illinois on an average of 5.2 species per divi- sion. The remaining natural divisions recorded 0-8 species. Only the Illinois/Mississippi River Sand Areas Division lacked winter stoneflies. The Shawnee Hills (14 species) and Wabash Bor- der (12 species) divisions were the most speciose. Allocapnia vivipara (13 divisions) and Taeniopteryx burksi (10 divisions) displayed the greatest diversity. Allocapnia forbesi, A. Winter Stoneflies of Illinois 219 illinoensis, A. smithi, Nemocapnia carolina, Paracapnia angulata, Prostoia completa, Taeniopteryx metequi, T: parvula, and Zealeuctra fraxina were restricted to one natural division. From 1900-1945, they were collected on 58 occasions at an average of 4.1 species per divi- sion. Twelve and 10 species were recorded, re- spectively, from the Shawnee Hills and Wabash Border divisions, with the remaining 12 divisions recording 0-6 species. From 1946-1975, they were recorded on 40 occasions at an average of 2.9 species per divi- sion. Eight and six species were found, respec- tively, within the Shawnee Hills and Wabash Border divisions with the remaining 12 Divisions recording 0-4 species. During the recent resurvey (1976-2000), winter stoneflies were recorded on 52 occasions at an average of 3 species per division. Overall, the species diversity within the Natural Division of Illinois has declined by an average of over two species per Division. The Shawnee Hills Division (11 species) was still the most species- rich area although Allocapnia granulata, and Taeniopteryx lita have disappeared from this di- vision. Six species Allocapnia illinoensis, A. granulata, A. mystica, Nemocapnia carolina, Strophopteryx fasciata, and Zealeuctra claasseni have disappeared from the Wabash Border Di- vision, although Allocapnia nivicola and Zealeuctra narfi have been added to this area recently. This division displayed the greatest loss of winter stoneflies. The remaining 12 divisions recorded 0-5 species with the Western Forest- Prairie and Ozark Divisions reporting an increase in species richness. Over the past 100 years the species richness of winter stoneflies within the 25 drainage ba- sins of Illinois ranged from 1 in the lower Illi- nois River-American Bottoms to 13 within the Massac, Bay, Lusk, Big Grand Pierre, and Big Creek System. Following the recent resurvey (1976-2000), the species richness ranged from 0 in the Wabash River (where 4 species had been collected previously) to 11 within the Massac, Bay, Lusk, Big Grand Pierre, and Big Creek System. The absence of winter stoneflies from the Wabash River is a strange situation because this river is considered to have good water qual- ity. Several attempts were made to collect adults during the 1990s along the river’s banks and on 220 Illinois Natural History Survey Bulletin Vol. 36 Art. 5 bridges spanning it, however no adults were in 12 and 11 drainage basins respectively, had found. Strophopteryx fasciata and Allocapnia their distributions reduced to 2 drainage basins. granulata which historically had been collected KEYS Twenty-one species of winter stoneflies have been reported for Illinois. Keys to the various fami- lies, genera, and species are given. Within each species I have tried to pull together all of the pertinent related information; the nomenclatoral changes; the diagnostic characteristics of the males, females, and nymphs (when possible); information of their biology, habitat preference, distribution, and seasonal activity; and the current status of each species based on the recent resurvey of the state (1976-2000). Maps outlining their distribution patterns within Illinois for the Frison period (1900- 1945), Ross period (1946-1975) and the recent resurvey of the state (1976-2000) are provided. When distribution records are limited, no distribution map is provided, rather the individual locality data are given. Key to Families of Illinois Winter Stoneflies Adults (from Poulton and Stewart 1991, Stewart and Harper 1996, Stark and Nelson 2000) 1. First and second tarsal segments subequal in length (as in Fig. 8.................... Taenioptery gidae First tarsal segment much longer than sécond (as in Fig’ 9) re neeceeseeeteee ee 2 2. Cerci multisegmented; forewing vein A, simple, unforked (Fig 12), intercubital crossveins few, usuallyil-25., 2257p ete eee. c scenes ences Capniidae Cerci one-segmented; forewing vein A, forked (Fig. 13), intercubital crossveins numerous, USUALLY 5 OF MOTE.........0.ccvensealeceéescees copnanonedecannetdsnecteasaseeen: noe cette ee ea s 3. Wings lying flat over back; costal space with crossvein just beyond cord, forming X-pattem with agjoming Veins C19, 13) sce vee sctece ries ee eee cere ee Nemouridae Wings rolled, covering dorsum and sides of abdomen; costal space with no CFOSSVEINS DEVON ‘COLD kee eee eee ncn css ee NET Ort eee oe ieee Leuctridae Nymphs (from Stewart and Stark 1988, Poulton and Stewart 1991) 1. First and second tarsal segments subequal in length (Fig. 8); coxae with single, telescoping gills or abdomen with large triangular ventroapical plate............. Taenioptery gidae Second tarsal segment wedge-shaped, shorter than first (Fig. 9); coxal gills and apical abdominal plates ADSENE ...4.....s010:s01crsesecosereteas¥ecseseageudavesssrecesséuusteay sa4 leleetne ent tata 2 2. Metathoracic wingpads strongly diverging from body axis (as in Fig. 17); cervical gills sometimes present; body short and robust; extended hindlegs reaching approximately 10 apex Of aDdOMen yout eteere ere Nemouridae (only Prostoia completa) Metathoracic wingpads essentially parallel to body axis (Fig. 16); cervical gills absent; body slender and elongate; extended hindlegs reaching far short of apex of abdomen............. 5 December 2002 Winter Stoneflies of Illinois 221 3. Abdominal terga widest posteriorly (Figs. 18 and 19), with posterior setal fringe; membranous pleural fold on abdominal sterna 1-9; metathoracic wingpads about as wide as long, reduced or absent ................ Te wn ea ee eee Capniidae Abdominal terga essentially parallel-sided, often without posterior fringe; membranous pleural fold not extending beyond sternum 7; metathoracic wingpads usually longer than LUCE bette 0 A oS A a Capniidae This family contains 10 North American gen- era and 152 species (Bill Stark 2001 http:// www.mce.edu/campus/users/stark). Adult capniids (Harper 1984, Stewart and Harper 1996, Stark and Nelson 2000) are char- acterized by having the glossae and paraglossae subequal in length and size (Fig. 15); the absence of gill remnant on the sides of the thorax; the wings at rest folded flat over the back and the forewing with only 1-2 median crossveins and one cubital crossvein; vein A» of forewing simple and unforked (Fig. 12); few intercubital crossveins; the first tarsal segment long, much longer than second (as in Fig. 9); the cerci multisegmented; and females with small subgenital plates. Hanson (1946) discussed the adult taxonomy of the Capniidae and gives a comparison of the morphology of the various genera within the Capniidae. Bebe AVN are ene ara hel th Sa A a ee Leuctridae Nymphs (Harper and Hynes 1971a, Harper 1984, Stewart and Harper 1996) are small and dark with the glossae and paraglossae subequal in length (Fig. 15); thoracic sterna not overlap- ping posteriorly; the first tarsal segment long, much longer than second (Fig. 9); abdominal segments |—2 lacking ventral gill tufts; abdomi- nal terga and sterna on segments 1-9 separated by membranous pleural fold; abdominal terga widest posteriorly, with posterior fringe of setae (Fig. 18, 19). Nymphs of the Capniidae remain the most poorly known of all North American families (Stewart and Stark 1988) although Harper and Hynes (1971a) produced a key to various capniid species. Biology: Capniids are principally sprawler- clingers in lotic situations where they feed as shredder-detritivores (Hynes 2000, Stewart and Harper 1996). Key to Adults of Illinois Capniidae (Harper 1984, Poulton and Stewart 1991, Stewart and Harper 1996) PSL ALISO Ue eetezne Sec eae eo cake cose ae dene hoe tasas o> PARTON DRCSCTILICnL Forres tek, | outs aes erst weeeeercl with less'tnan | l*segments.../..2..2-5...... Cerci with more than 11 segments.................. 3. Anal lobe of hindwing enlarged, subequal in size to rest of wing (Fig. 20)............... Allocapnia Anal lobe noticeably smaller than rest of wing (Fig. 21).............seeeeeeeeeeeees Paracapnia angulata Illinois Natural History Survey Bulletin Vol. 36 Art. 5 Key to Mature Nymphs of Illinois Capniidae (Harper and Hynes 1971a, Harper 1984, Poulton and Stewart 1991, Stewart and Stark 1988) 1. Basal or apical cercal segments or both with multiple, long, fine hairs forming a prominent vertical fringe (Fig. 11)............... Site ceirevsethericeti commer ee ees eet ee Nemocapnia carolina Long cercal hairs restricted to apical segmental whorls, no prominent vertical fringe: (B19 910) see ees 2. Body and appendages densely clothed with long, stout bristles (Fig. 19), some abdominal bristles one-half or more as long as their segment.................... Paracapnia angulata Bristles on body and appendage few or short (Fig. 18); abdominal hairs or bristles less than one-third length of segment ............ Allocapnia Claassen Allocapnia Claassen (1928:667, 1931:111): Hanson™ ClL94672 1 e235 lee Harper (1984:105), Harper and Hynes (1971a:924), Iilies (1966:122), Poulton and Stewart (1991:16), Stewart and Harper (1996:228, 246), Stewart and Stark (1988:98), Zwick (1973:360). New name for Capnella. Genotype: Allocapnia granulata (Claassen). Capnella Claassen (1924:43). Name preoccu- pied. Capnellula Strand (1935:304). New name for Capnella Claassen. This genus contain 43 North American spe- cies (Bill Stark 2001 http://www.mc.edu/cam- pus/users/stark). Adults (Hanson 1946, Stewart and Harper 1996) of Allocapnia are characterized by hav- ing Ry straight beyond origin of Rs; anal lobe of Dede ncce dhercatiore scent (cette ates ee Allocapnia hindwing subequal in size to remainder of wing (Fig. 20), wings occasionally absent (A. vivipara males); cerci with more than 11 segments. Color dark brown. Length about 5 mm in males, 6 mm in females. Wings slightly smoky with brown veins; female wings extend beyond apex of abdomen; male wings highly variable within and among species. Hanson (1946:211) dis- cussed the comparative morphology of the Allocapnia relative to other genera of capniids. Nymphs (Stewart and Harper 1996) of Allocapnia are characterized by having few bristles on body and appendages; wingpads brac- hypterous in males (absent in A. vivipara) and variable in females; the medial margin of the hindwing pad unnotched or notched close to apex; abdominal terga with posterior fringe of short setae, and scattered intercalary setae (Fig. 18); long hairs on cerci restricted to apical seg- mental whorls (as in Fig. 10). Key to Males (from Ross and Ricker 1971) Wings vestigial (Fig. 4A); dorsal process on tergite 8 with low, sharp profile (Fig. 47) avuvsdodassdaddissedecasiacye bodeweedeadectibenceesterat cuit eeeeerer deer iStT aC a ee UT TCE TTT ROR net tt aes are ne vivipara Wings about half as long as abdomen, or greater::,...:....c..--.2.-c-ee.+--censestssectesrescsieceyse en 2 Apical segment of upper limb of epiproct long, flat to tip, very thin and regular in profale, with short dorsal retrorse spines (Hig, 39) j..ceecccserstes-seeteoeete tans ee ee recta Apical segment of upper limb of epiproct either short, narrow, or tip expanded...................00+ 3 December 2002 Winter Stoneflies of Illinois 223 3. Dorsal process of tergite 8 forming pair of oblique rugose areas, their dorsal aspect (SUS SITET GUD 2oea Oe Sin tee nae a ees Fog ete pe Soe Se eM ee ene granulata Dorsal process forming a transverse bar or oval having 2 or 3 projecting apical points RoE et setae rey sete. WAU ACE SS) cerca eo ies cet te ey ie Feat oiet ate Ba re as 4 4. Tergite 7 with sclerotized process as large as sclerotized portion of tergite 8 (Figs. 24 area te ere See er, Seen Reais ERPs oe eM Cea. TAGs davies, Senin cae Lh ak howe, Sie ee oe. 5 Tergite 7 lacking sclerotized process or with small sclerotized ridge (Figs. 34, 37, 42, 45) (on Suh pb ange nea MERC cee 9 Sot Mie ge a ean Dan aco epee bor te Bee ee REE es Dee ee ee 6 5. Sclerotized projection of tergite 8 triangular in lateral view, apex pointed posteriorly ON tM ead sauce oc nya Coto toca cctons\s coer saondses saensoveetesasecursececenisenaenesndeses illinoensis Sclerotized projection of tergite 8 quadrate in lateral view, apex angulate (Fig. 24)........ forbesi 6. Dorsal process of tergite 8 with mesal point at least half height of lateral lobe (Fig. 45) ......... 4 Dorsal process of tergite 8 with only two apical lobes (Figs. 34 and 42) ooo... ee eeeeestreeeeees 8 7. Dorsal process of tergite 8 as high as wide, without a posterodorsal projecting corner; lower limb of epiproct wider and extending laterally considerably beyond upper limb, nN Tt Le Ce A110 WAAC, ( 1779.1, Neeng sn een the ck be foie tacks aa da Sloe dG Saal ya Lecce el ancatadee nivicola Dorsal process wider, posterodorsal corner markedly projecting; lower limb of epiproct extending only slightly beyond sides of upper limb (Fig. 45)... eee eeeeeeeeeee smithi 8. Apical segment of upper limb of epiproct ovoid, less than one-half length of base RN soe var ata see ne scceca cn cavaydoath debadas sos tentcvs baeasclds tedgeehovanbesvessdunsinese rickeri Apical segment more slender and more than one-half length of base (Fig. 34) .............. mystica Key to Females (from Ross and Ricker 1971) 1. Tergite 8 with sclerotization covering more that half of tergite (as in Fig. 22)... eeeeeeeeees ,3 Tergite 8 with mesal area membranous (as in Fig. 23) over length of tergite........... eee 3 2. Sternites 7 and 8 fused, mesal area forming solid transverse area, intersegmental SEEM TILCT ALCL 1a any Views Sane enna ie chee deret fe. wat sane ek tt oblate ak feces oy ly teccek ccsasucesas. vivipara Sternites 7 and 8 with distinct intersegmental suture (Fig. 25) ...........ceesesssessseeeseeseeeeeeees forbesi Sra sternites7 and S:solidly fused medially (Figs.32; 38; 43; 40). .cc..ccccstecccssti ede odsectebesentelsorweseoes 4 Sternites 7 and 8 completely separated by membranous strip (Figs. 28, 35, 40).........:ccceeseeee 7 Illinois Natural History Survey Bulletin Vol. 36 Art. 5 Sternites 7 and.8 with medial suture not evident (Fig.43) sive. circ ceee ee eee rickeri Sternites 7 and 8 with medial suture indicated by a dark line (Figs. 32, 38, 46) ........ eee 5 Apex of sternite 8 with wide sagittate medial portion contrasting sharply with less sclerotized remainder of mesal’area (Fig, 32) te-c-ercecteeesees eoeeereceee see illinoensis Apex of sternite 8 with mesal area not contrasting noticeably with more basal area of SCIOLICE 05 jheskecssfoscelsceeed aden abe Meese cecebuhasec cetirenerc ce eeeadunmac Oc eeeeccteen ce tat ek tent nn 6 Sternite 8 with apical flap moderately wide; lateral humps of sternite chiefly membranous; the combination of the two producing an hourglass-shaped mesal dark area (Higa 4G), 57 ei PR cee eee en ve eee smithi Sternite 8 with apical flap narrower; lateral humps more extensively sclerotized (FIG 38) gcd, coe ees SE DRE Pa eee, ee ce cen nivicola Sternite 8 with apicomedial area differentiated into a sagittate or emarginate fluted area set off laterally by pale membranous areas (Fig. 28) 2.02.02... eee granulata Sternite 8 with apicomedial area not markedly differentiated from lateral areas (Figs35, 40) oo cacc. 2h, Cd recdicrelsetesecdsesanet lesdoiser ebb Solus. tuassecs tate tie cageneR Re chet an 8 Sternite $ with apical margin concave (Fig. 40) .....222a.crepcee recente nee ee recta Sternite 8 with apical margin convex (Fieg235) see ccn eee eee eres ee mystica Key to Mature Nymphs (from Harper and Hynes 1971a) Abdominal tergites with erect hairs throughout (Harper and Hynes 197 1a: Fig. 64) Abdominal tergites with erect hairs not evenly distributed, more abundant near posterior margin and sometimes at midlength of tergites (Harper and Hynes 1971a: Figs. 59-63) cnt AAR eee eee teow d ones Meedase ene eee einer ee ane ne) ee 2 Abdominal tergites, in addition to prostrate clothing hairs, bearing only a terminal row of erect hairs (lateral view) (Harper and Hynes 197 1a: Fig? 60) 2.700002.) \ce creer 3 Abdominal tergites, in addition to prostrate clothing hairs, bearing terminal and intermediate rows of erect hairs (lateral view) (Harper and Hynes 1971a: Figs. 61-63) .......... 4 Terminal abdominal tergites with intermediate erect hairs only on lateral margins; supra-anal lobe of male long; abdominal tergites, in addition to prostrate clothing hairs, bearing only a terminal row of erect hairs (lateral view) (Harper and Hynes 197 1a: as In Pig. 59) piss ceeeeece ess pates te celcpeg setae seen tet et nivicola Terminal abdominal tergites with intermediate erect hairs on lateral third of tergite; supra-anal lobe of male short (Harper and Hynes 197 1a: Fig. 60) .........cecceessccesseeeseeeeeeees recta December 2002 Winter Stoneflies of Illinois fi be 4. Abdominal tergites with erect bristles on margin long, about half as long as mid-dorsal tergite length; intermediate bristles few (Harper and Hynes 197 1a: Fig. 61); nymphs SBME TOL SEZ Cots: LTIIT) ee sachin er eee ee ic See se seks tne rocker Cee ete TT aE TEER een eta dosbeses rickeri Abdominal tergites with erect bristles on margin short, about one-third as long as mid-dorsal tergite length; intermediate bristles numerous (Harper and Hynes 1971a: Figs. 62-63); PAU StC ALCO SIZ OL MT ierreeeetss corse crepe cass crateaper canes? couche ct eerie, teats erent bres 7haastessefaetacns 5 5. Head and pronotum hairy (lateral view); male without wing pads; supra-anal lobe of male twice as long as abdominal segment 10 (Harper and Hynes 197 1a: Fig. 63) ........ vivipara Head and pronotum with fewer hairs (lateral view); male with short wing pads; supra-anal lobe of male short, about as long as abdominal segment 10 (Harper and mM tot eR) 2) ree meee tere taree ttre etre craters tatereeheat cies wechenetsctetetisigeie raven eos granulata 226 Illinois Natural History Survey Bulletin Vol. 36 Art. 5 Z Yl ; j pp fi 2 Ba on an ee coe ees coms oe aes LLL Mh, LZ \ \\ 10 11 Figures 8-15. 8. Nymphal leg, Zaeniopteryx burksi. 9. Nymphal leg, Prostoia completa.10. Nymphal cerci, Paracapnia angulata. 11. Nymphal cerci, Nemocapnia carolina. 12. Forewing, Nemocapnia carolina. 13. Fore- wing, Prostoia completa. 14. Nymphal labial palps.15. Nymphal labial palps. a=costal space crossvein, g=glossae, p=paraglossae. (Fig. 8-15 from Poulton & Stewart 1991). December 2002 Winter Stoneflies of Illinois 227 Figures 16-23. 16. Nymphal wingpads, Allocapnia granulata. 17. Nymphal wingpads, Taeniopteryx burksi: 18. Nymphal abdominal terga, Allocapnia granulata. 19. Nymphal abdominal terga, Paracapnia angulata. 20. Hindwing, Allocapnia granulata. 21. Hindwing, Paracapnia angulata. 22. Female terminalia, Allocapnia tennessa. 23. Female terminalia, Allocapnia brooksi. (Figs. 16-21 from Poulton & Stewart 1991; 22-23 from Ross & Ricker 1971). 228 Illinois Natural History Survey Bulletin Allocapnia forbesi Frison Allocapnia forbesi Frison (1929:397): Claassen (1940:89), Harris and Webb (1995:342); Hitchcock (1974:50), Illies (1966:123), Poulton and Stewart (1991:17), Ross and Ricker (1971:35), Zwick (1973:362). Type locality: Illinois, Herod, tributary North Branch, Hicks Creek. Allocapnia forbesi cornuta Frison (1935:363): Claassen (1940:89), Harris and Webb (1995:342), lies (1966:123), Ross and Ricker (1971:35). Type locality: Illinois, Pope County, Dixon Springs, Hills Branch Creek. Adult: (Ross and Ricker 1971). Male (Fig. 24 A-D): Wings highly variable, ranging from re- duced vestiges to reaching abdominal tergites 5 or 6. Tergite 6 occasionally with dorsal process similar to that on tergite 7. Tergite 7 with dorsal process on posterior half of segment, conical, distinctly raised in lateral view; conical in pos- terior view. Tergite 8 with dorsal process dis- tinctly raised, quadrate, apex truncate; emargin- ate medially producing two short, well-separated, rounded lobes in posterior view. Upper limb of epiproct elongate, apical segment short, dis- tinctly shorter than basal half; lower limb with apical segment narrow, fingerlike. Female (Fig. 25): Wings varying from vestigial to reaching apex of abdomen. Tergite 8 generally sclero- tized, occasionally with narrow, medial, mem- branous area. Sternite 7 more sclerotized than sternite 6, distinctly separated from sternite 8, posterior margin produced into broadly rounded lobe. Sternite 8 with distinct lateral lobes, me- dial area wedge-shaped, rounded posteriorly. Nymph: unknown. Biology: Hitchcock (1974) reported that Frison (1929) found the adults feeding on algae from stones and posts. This reference is in error. Frison (1929:360) did report observing several species of winter stoneflies feeding “...upon blue- green algae growing on tree trunks, stones, old logs, etc....” but Allocapnia forbesi was not one of the species. Habitat: Ross and Ricker (1971) reported A. forbesi from clear, cool, rocky, spring-fed streams that had a tendency to become dry dur- ing the summer. Surdick and Gaufin (1978) re- Vol. 36 Art. 5 ported A. forbesi as being oligothermal, eulichtophilous, and found in rivers or perma- nent and intermittent streams. Adults collected from 1976-2000 were taken on the railings and abutments of bridges, on and under moderate sized rocks, and from leaf packs collected near the water surface. Adults were collected near small streams with a slow to moderate flow, 30 cm—1 m deep and 60 cm-7 m wide with a sub- strate of gravel, sand, cobblestone and bedrock. At two localities they were collected from streams with moderate flow, 35 m wide, 30 cm— 1 m deep, with a substrate dominated by gravel. Distribution and Seasonal Activity: Allocapnia forbesi ranges from western Virginia to Illinois (Ross and Ricker 1971, Fig. 98; Stark et al. 1986). In Illinois, adults were collected from 30 November to 24 March. Figure 26 displays the distribution of Allocapnia forbesi from 1900- 1945, 1946-1975, and 1976-2000. Current Status: In Illinois, Allocapnia forbesi is an uncommon species that historically (1900- 2000) was collected across the Shawnee Hills Natural Division of southern Illinois. Recent collections (1976-2000) indicate that A. forbesi is now restricted to the eastern Shawnee Hills in the Bay, Lusk, Big Grand Pierre, Big Creeks drainage system and the Saline River drainage system (Table 4). December 2002 Winter Stoneflies of Illinois 229 phe 26 1976-2000 @ 1946-1975 1900-1945 4m Figures 24-26. Allocapbnia forbesi. 24. Male terminalia, A=lateral view, B=dorsal view, C=posterior view of tergite 8, D=posterior view of tergite 7. 25. Female terminalia, 26. Distribution in Illinois. (Figs. 24-25 from Ross & Ricker 1971). 230 Illinois Natural History Survey Bulletin Allocapnia granulata (Claassen) Allocapnia granulata (Claassen): Claassen (1931:114, 1940:89), Finni and Chandler (1977:243), Frison (1929:394, 1935:364); Hanson (1946:211); Harper and Hynes (1971a); Harris and Webb (1995:342); Illies (1966:123), Poulton and Stewart (1991:17), Ross and Ricker (1971:28), Zwick (1973:362). Type locality: Johnstown, New York. Capnella granulata Claassen (1924:44): Illes (1966:123), Needham and Claassen (1925 212): Adult: (Ross and Ricker 1971). Male (Fig. 27 A-C): Wings vary from extremely short to mod- erately long, reaching tergite 7. Hindwing (Harper 1984, Fig. 13.168; Poulton and Stewart 1991, Fig. 13; Stewart and Harper 1996, Fig. 14.172). Tergite 6 without dorsal process. Terg- ite 7 with slightly raised, nonsclerotized bulge on posterior half. Tergite 8 with dorsal process distinctly raised, sloping posteriorly from base, apex rounded with V-shaped medial emargin- ation and small lateral tubercle bearing a few hairs in posterior view. Upper limb of epiproct with apical segment subtriangular, equal in length to basal segment, apical segment bulbous, wider apically than basal half in dorsal view; lower limb narrow, elongate, apex rounded, of- ten with short, blunt projection. Other figures: Frison (1929, Fig. 51 male habitus; 1935, Fig. 227 male terminalia), Needham and Claassen (1925, pl. 47, Fig. 3 male wings, 5 female wings, pl. 50, Figs. 6 and 6a male terminalia, 7 female terminalia), Harper and Hynes (1971a, Fig.12 terminalia), Stewart and Harper (1996, Fig. 14.172 hindwing). Female (Fig. 28), Tergite 8 membranous dorsally. Sternite 7 separated from sternite 8. Sternite 8 more heavily sclerotized than sternite 7, posterior margin triangular me- dially, although this area varies considerably. Other figures: Frison (1929, Fig. 12 female habi- tus; Figs. 44-45 female habitus; Fig. 71, female abdominal sterna 7-8; 1935, Fig. 220 female ab- dominal sterna 7-8; Figs. 294—295, female habi- tus); Harper and Hynes (1971a, Fig. 33 subgenital plate); Needham and Claassen (1925, pl. 47, Fig. 5 female wings, pl. 50, Fig. 7 female terminalia). Nymph (Fig. 29) (Harper and Hynes 1971a): Length 7-9 mm. Medium to dark yellowish Vol. 36 Art. 5 brown. Head and pronotum with sparse pubes- cence, pronotum fringe very conspicuous; terg- ites with well-developed intermedial and termi- nal bristles, about one-third as long as mid-dor- sal length of tergite (Harper and Hynes 197 1a, Fig. 62, male setal pattern). Other descriptions of the nymphs are found in Claassen (1931:114); Frison (1929:395, Fig. 35, female habitus; Fig. 40 female terminalia; 1935, Fig. 289, female terminalia); Harden and Mickel (1952:29, plate Il, Fig. 1 habitus, pl. [X, Fig. 1 maxilla, Fig. 2 mandible); Harper (1984, Fig. 13.12, abdominal ventrum; Fig. 13.37, male wing pads), and Stewart and Harper (1996, Fig. 14.11, abdomi- nal venter; Fig. 14.12, abdominal terminalia; Fig. 14.37, male wing pads). Biology: Harper and Hynes (1970:925) deter- mined Allocapnia granulata to have a univoltine life cycle with eggs deposited in May. Nymphs hatched within a month and underwent a diapause during the summer, with development proceed- ing rapidly through the fall and winter. Finni and Chandler (1977) reported five post-diapausal instars for A. granulata. Habitat: Ross and Ricker (1971) reported Allocapnia granulata from larger, slower, and more turbid streams but also collected this spe- cies from smaller tributaries of large rivers. Surdick and Gaufin (1978) reported A. granulata as being a herbivore or scavenger that was alkaliphilous, mesotrophic, saprophobic, eu- and mesoxyphilous, meso-, meta-, and oligothermal, eu- and mesolichtophilous, rheophilous and rheobiontic, epilithic and epiphytic, and found in rivers or permanent and intermittent streams. Gaufin (1958) reported A. granulata to be intol- erant of pollution, preferring water with high lev- els of dissolved oxygen (70-115% saturation). Poulton and Stewart (1991) reported A. granulata being collected up to 1 km away from moving water. Adults from 1976—2000 in Illinois, were collected on the railings and abutments of bridges and from leaf packs collected near the water sur- face. They were collected from small- to mod- erate-sized streams, 6-25 m wide, 1-2 m deep with a gravel and sand substrate, although col- lections from the Leaf River were from a stream that has a bottom substrate of 70% sand and 30% silt. Allocapnia granulata was also collected from two localities on the Rock River, a deep river, Over a quarter of a km wide, with a sub- strate of sand and silt. December 2002 Distribution and Seasonal Activity: Allocapnia granulata is widely distributed in eastern North America from Alabama to Quebec west to Min- nesota and Texas (Ross and Ricker 1971, Fig. 96, Stark et al. 1986). In Illinois, adults were collected from 19 January to 3 April, with a single collection made on 3 December. Figure 30 displays the distribution of A. granulata from 1900-1945, 1946-1975, and 1976-2000. Current Status: Allocapnia granulata is con- sidered an environmentally tolerant species and for many years in Illinois it ranged from Hardin Winter Stoneflies of Illinois 231 County in southern IIlinois to the Wisconsin bor- der in 5 of the Natural Divisions of Illinois (Table 3) and 11 of the drainage systems (Table 4). Over the past 25 years, its distribution pattern has been reduced to 3 natural divisions (Table 3) and the Rock River Hill Country drainage system, with a single disjunct collection found at Morris, on the Illinois River (Table 4). Although broadly distributed within the Rock River Hill Country drainage, A. granulata appears to be disappear- ing from the remainder of the state. Est 30A. 1900-1945 30B. 1946-1975 30C. 1976-2000 Figures 27-30. Allocapnia granulata. 27. Male terminalia, A=lateral view, B=dorsal view, C=posterior view of tergite 8. 28. Female terminalia, 29. Nymph. 30. Distribution in Illinois. (Figs. 27-28 from Ross & Ricker 1971; 29 Stewart & Stark 1988). 232 Illinois Natural History Survey Bulletin Allocapnia illinoensis Frison Allocapnia illinoensis Frison (1935:365): Claassen (1940:89), Harris and Webb (1995:342); Hanson (1946:238), Harper and Hynes (1971a:924, 934), Illies (1966:123), Ross and Ricker (1971:33), Zwick (1973:363). Type locality: Illinois, Clark County, Dolson (Clarksville), on fence post near Rocky Branch. Adult: (Ross and Ricker 1971). Male (Fig. 31 A-D): Wings reaching abdominal tergites 6 or 7. Tergite 6 without dorsal process. Tergite 7 with dorsal process on posterior half moderately produced into a conical tubercle; with slight medial emargination in posterior view. Tergite 8 with dorsal process not very high, sloping no- ticeably posteriorly, with distinct, widely sepa- rated medial emargination in posterior view. Upper limb of epiproct with apical segment short, spatulate, distinctly shorter than basal segment; lower limb with apical segment shallow. Other figures: Frison (1935: Fig. 212 male genitalia), Harper and Hynes (1971a, Fig. 17, terminalia. Female (Fig. 32): Tergite 7 with posterior mar- gin slightly arcuate, joined to tergite 8 on meson by faint sclerotized area. Tergite 8 slightly more sclerotized than tergite 7, lateral lobes distinct, posterior margin produced into wide sagittate process, dark along posterior extremities and rap- idly becoming less sclerotized anteriorly. Other figures: Frison (1935, Fig. 225, female genita- lia), Harper and Hynes (1971a, Fig. 34, subgenital plate). Nymph (Harper and Hynes 1971a): Length 8- 10 mm. Yellowish brown with venter and ap- pendages paler. Body covered with short, dense pubescence, with a few bristles that are mark- edly longer than the others. Tergites covered with erect bristles in addition to prostrate clothing bristles (Harper and Hynes 1971a, Fig. 64). Habitat: Ross and Ricker (1971) reported Allocapnia illinoensis from clear, rocky, spring- fed streams. Surdick and Gaufin (1978) reported A. illinoensis as being neutral, eury- and oligothermal, eulichtophilous, and found in streams. Distribution and Seasonal Activity: Allocapnia illinoensis ranges from New York to Quebec west to Minnesota and Illinois (Ross and Ricker 1971, Vol. 36 Art. 5 Fig. 96; Stark et al. 1986). In Illinois adults were collected from 5 January to 23 March. Figure 33 displays the distribution of A. i/linoensis from 1900-1945. This species was collected only once from 1946-1975 (Clark County, Rocky Branch, 4.2 km NE Dolson (Clarksville)). Current Status. Allocapnia illinoensis has al- ways been rare in Illinois, being restricted to the Wabash Border Natural Division and the Wabash tributaries drainage. Ross et al. (1967) and Ross and Ricker (1971) discussed the postglacial dis- persal of A. illinoensis from the Cumberland Pla- teau into the northeastern United States and southern Canada. Prior to 1946, Frison collected 12 males and | female at 3 sites on Rocky Branch and the East Fork of Big Creek west of Oliver. From 1946-1975, only 22 specimens (1male 1958, 21 females 1965) were collected from Rocky Branch. Since 1965, no specimens of A. illinoensis were collected in Clark County al- though extensive collecting (Fig. 5) has been done on several occasions. Historically (1900- 2000), four winter stoneflies were collected at Rocky Branch. Frison recorded A. granulata, A. illinoensis, A. recta, and A. vivipara; Ross and Ricker recorded only A. illinoensis; and currently only A. vivipara has been collected from this lo- cality. Rocky Branch is currently protected as an Illinois Nature Preserve yet three species of winter stoneflies have disappeared from this lo- cality, with A. illinoensis appearing to be extir- pated from Illinois. December 2002 Winter Stoneflies of Illinois 233 Figures 31-33. Allocapnia illinoensis. 31. Male terminalia. A=lateral view, B=dorsal view, C=posterior view of tergite 8, D=posterior view of tergite 7. 32. Female terminalia. 33. Distribution in Illinois. (Figs. 31-32 from Ross & Ricker 1971). 234 Illinois Natural History Survey Bulletin Allocapnia mystica Frison Allocapnia mystica Frison (1929:399): Claassen (1931:116, 1940:89), Illies (1966:125), Har- ris and Webb (1995:342), Hitchcock (1974:57), Poulton and Stewart (1991:18), Ross and Ricker (1971:31), Zwick (1973:365). Type locality: Illinois, Vermil- ion County, Oakwood, a small tributary of the Salt Fork of Vermilion River. Adult: (Ross and Ricker 1971). Male (Fig. 34 A-C): Wings reaching abdominal tergites 5 or 6. Tergite 6 and 7 without dorsal process. Terg- ite 8 with dorsal process large, sloping posteri- orly and ending in small apical point, truncate apically with small medial emargination in pos- terior view. Upper limb of epiproct elongate, apical segment shorter than basal segment; lower limb with apical segment short, with flat apical point. Other figures: Frison 1929, Figs. 57, 67, (PR IB GRRL EeE UISY AGM OAM OOS), PAN. 3 Ire male (Fig. 35): Tergite 8 membranous dorsally. Sternite 7 separated from sternite 8. Sternite 8 more heavily sclerotized than sternite 7, poste- rior margin produced medially into a highly pol- ished bulge. Nymph: The male and female nymphs were de- scribed by Frison (1929:402—403) and re- described by Claassen (1931:116). Habitat: Ross and Ricker (1971) reported Allocapnia mystica from cool, rapid, clear, gravel or rocky streams, usually spring-fed and small. Surdick and Gaufin (1978) reported A. mystica as being a herbivore or scavenger that was oligothermal, rheobiontic, epilithic, and found in streams. From 1976-2000, adults of A. mystica were collected from bridge railings and abutments, in leaf pack near the water surface, and from tree trunks. It has been collected from small moderately flowing streams 1-7 m wide, 30 cm—1 m deep, with a substrate of gravel, sand, cobblestone, and bedrock. It has also been col- lected from moderate sized streams, 35 m wide, 30 cm-l m deep with a bottom substrate of gravel. Distribution and Seasonal Activity: Allocapnia mystica ranges from Georgia and Virginia west to Missouri and Arkansas (Ross and Ricker 1971: Fig. 96, Stark et al. 1986). In Illinois, adults were Vol. 36 Art. 5 collected from 18 September to 16 March. Fig- ure 36 displays the distribution of A. mystica from 1900-1945, 1946-1975, and 1976-2000. Current Status. Prior to 1946, Allocapnia mystica had been collected across the Shawnee Hills Natural Division of southern Illinois, the Wabash Border Division, and the Rock River Hill Country Division (Table 3). From 1946-1975 it was still well established in the Shawnee Hill Natural Division, with a single record from the Southern Till Plain Division. Current records indicate that this species is still well established across the Shawnee Hills Natural Division, but has disappeared from the northern three-fourths of the state. Historically (1900—2000), A. mystica was collected from seven drainage systems (Table 4), but now is only found in the Big Muddy, the Bay, Lusk, Big Grand Pierre, Big Creek, and the Saline River drainage systems. December 2002 Winter Stoneflies of Illinois 235 36A.1900-1945 36B. 1946-1975 36C. 1976-2000 Figures 34-36. Allocapnia mystica, 34. Male terminalia. A=lateral view, B=dorsal view, C=posterior view of tergite 8. 35. Female terminalia. 36. Distribution in Illinois. (Figs. 35-36 from Ross & Ricker 1971). 236 Illinois Natural History Survey Bulletin Allocapnia nivicola (Fitch) Allocapnia nivicola (Fitch): Hanson (1942:83), Harper and Hynes (1971a:925, 935), Illies (1966:125), Harris and Webb (1995:342), Hitchcock (1974:58), Ricker (1952:164), Ross and Ricker (1971:49), Zwick (1973:365). Type locality: New York. Perla nivicola Fitch (1847:278): Ilies (1966:125). Adult: (Ross and Ricker 1971). Male (Fig. 37 A-D): Tergite 6 without dorsal process. Tergite 7 with low medial dorsal process only slightly raised above tergite, truncate apically; broad, shallow truncate apically in posterior view. Terg- ite 8 with dorsal process higher than long, di- rected vertically with three apical teeth, mesal tooth usually slightly lower than lateral ones in posterior view. Upper limb of epiproct with api- cal segment triangular in dorsal view, tapered apically, shorter than basal segment; lower limb in dorsal view much wider than upper limb. Other figures: Hanson (1942, Fig. 2, male terminalia; Fig. 4 female terminalia), Harper and Hynes (197 1a, Fig. 16). Female (Fig. 38): Terg- ite 8 membranous dorsally. Sternite 7 lightly sclerotized, separated from sternite 8. Sternite 8 with posterior margin concave with narrow, medial, truncate flap. Other figures: Hanson (1942, Fig. 4, female terminalia), Harper and Hynes (197 1a, Fig. 39). Nymph (Harper and Hynes 1971a:935) found the nymphs of A. nivicola to be similar to A. pygmaea in the abdominal setal bristles (Harper and Hynes 1971a, Fig. 59), each species being separated by the setal pattern on the outer sur- face of the galea. Length 6—7.5 mm. The tuft of setae on the outer surface of the galea is poorly developed in A. nivicola (rarely more the 10 se- tae) (Harper and Hynes 1971a, Fig. 45). Nymphs move from the gravel (where they were re- cruited) to leaf habitats for active growth (Mackay 1969, Stewart and Stark 1988). Habitat: Ross and Ricker (1971) reported col- lecting Allocapnia nivicola from small stony brooks. Surdick and Gaufin (1978) reported A. nivicola as being neutral or alkaliphilous, saprophobic, euoxyphilous, eury-, meso-, meta-, and oligothermal, epiphytic, and found in streams. Gaufin (1958) reported A. nivicola to Vol. 36 Art. 5 be intolerant of pollution, preferring water with high levels of dissolved oxygen (70—115% satu- ration). From 1976—2000, adults of A. nivicola in Illinois were collected only from a small slow- flowing stream, 30 cm-1 m wide, 15 cm deep, with a gravel and sand substrate. Adults were collected on and under moderate size stones, from leaf packs, and from woody debris. Distribution and Seasonal Activity: Allocapnia nivicola ranges from northern Alabama to Nova Scotia west to eastern Illinois (Ross and Ricker 1971: Fig. 96, Stark et al. 1986). In Illinois, adults were collected from 28 January to 25 Feb- ruary. Illinois Record: 1900-1945. Coles County: Fox Ridge State Park. 1976-2000. Vermilion County: Forest Glen Forest Preserve, 8.5 km SE Westville. Current Status. Allocapnia nivicola is a rare species in Illinois originally found at a single locality in the Grand Prairie Natural Division and currently known from a single locality in the Wabash Border Natural Division (Table 3). Frison collected three males in a small, shallow creek in Fox Ridge State Park in conjunction with A. granulata, A. recta, and A. vivipara. No speci- mens were collected there during 1946-1975, although a collecting trip to this park had been made in the winter of 1964. During 1993-1995, five collecting trips were made to this park with only A. recta and A. vivipara being collected. In 1993 and 1995, a population was collected in Forest Glen Forest Preserve in Vermilion County in the Vermilion-Little Vermilion River drain- age system. Efforts were made to collect winter stoneflies from this type of habitat in other seep areas of Vermilion County with no success. December 2002 Winter Stoneflies of Illinois 237 D hivicola 37 38 Figures 37-38. Allocapnia nivicola. 37. Male terminalia. A=lateral view, B=dorsal view, C=posterior view of tergite 8, D=posterior view of tergite 7. 38. Female terminalia. (Figs. 37-38 from Ross & Ricker 1971). 238 Illinois Natural History Survey Bulletin Allocapnia recta (Claassen) Allocapnia recta (Claassen): Claassen (1931:113, 1940:90), Frison (1929:395), Harper and Hynes (197 1a: 925, 936), Har- ris and Webb (1995:342), Hanson (1946:238), [lies (1966:127), Ross and Ricker (1971:23), Zwick (1973:367). Type locality: Ithaca, New York. Capnella recta Claassen (1924:44): [lies (1966:127). Adult: (Ross and Ricker 1971). Male (Fig. 39 A-C): Tergite 6 and 7 lack a dorsal process. Terg- ite 8 with dorsal process quadrate, apical mar- gin highly variable, in posterior view broad ba- sally with narrow, quadrate, apical process, apex truncate. Upper limb of epiproct flat, elongate, apical segment distinctly longer than basal seg- ment, apical segment clavate in dorsal view; lower limb bulbous apically, wider apically than apical segment of upper limb in dorsal view. Otherhirgures; = hrisone (9307 bipw 221, terminalia); Hanson (1946, Fig. 60); Harper and Hynes (1971a, Fig. 13); Needham and Claassen (1925, pl. 50, Fig. 8, terminalia); Kondratieff and Kirchner (2000, Figs. 21-26, male terminalia). Female (Fig. 40): Tergite 8 membranous dor- sally. Sternite 7 distinctly separated from stern- ite 8. Sternite 8 markedly darker than sternite 7, posterior margin concave. Other figures: Harper and Hynes (1971a, Fig. 35); Needham and Claassen (1925, pl. 50, Fig. 9, terminalia); Frison (1935, Fig. 213, sternites 7 and 8). Nymph (Harper and Hynes 1971a:936): Length 6-—7.5 mm. Light yellowish brown. Femora cov- ered with short bristles, anterior bristles equal in length to posterior bristles. Abdomen with pros- trate clothing bristles and erect bristles on pos- terior margin (about one-third as long as mid- dorsal length), intermediate erect setae on lat- eral third of caudal 4—5 tergites, not on middle of tergite (Harper and Hynes 1971a, Fig. 60). Other figures: Frison (1929:396, Fig. 36, female habitus; Fig. 37, male habitus; Fig. 39, female habitus; Fig. 55, female habitus; 1935:367, Fig. 290, female abdomen; Fig. 291 male habitus), and Claassen (1931:113). Habitat: Ross and Ricker (1971) reported A. recta from fairly rapid, clear, cool streams with gravel or rocky bottoms and very common in Vol. 36 Art. 5 small spring-fed brooks. Harper and Hynes (1971a) found A. recta in small spring-fed streams that did not freeze. Surdick and Gaufin (1978) reported A. recta as being a herbivore or scavenger that was alkaliphilous, saprophobic, epiphytic, and found in permanent and intermit- tent streams. Gaufin (1958) reported A. recta to be intolerant of pollution, preferring water with high levels of dissolved oxygen (70-115% satu- ration). From 1976—2000, adults of A. recta in Illinois, were collected from bridge railings and abutments, on and under cobblestones, and in leaf packs from small, slow- to moderate- flow- ing streams, 1-15 m wide, less than 30 cm-1 m deep, with a gravel, sand, and cobblestone sub- strate. Distribution and Seasonal Activity: Allocapnia recta ranges from Alabama to Nova Scotia west to Illinois (Ross and Ricker 1971: Fig. 90, Stark et al. 1986). In Illinois, adults were collected from 17 November to 25 March. Figure 41 dis- plays the distribution of Allocapnia recta from 1900-1945, 1946-1975, and 1976-2000. Current Status. Prior to 1946, Allocapnia recta had been collected abundantly in Clark, Coles, and Vermilion counties in the Grand Prairie and Wabash Border Natural Divisions of eastern Il- linois (Table 3). Only one collection of this spe- cies was obtained during 1946-1975. From 1976-2000, abundant populations were collected in Coles and Vermilion counties (Fig. 41). East- ern Illinois appears to be the western tip of its distribution, but it is still well established in the Embarras River-Wabash Tributaries drainage system and the Vermilion-Little Vermilion River drainage systems (Table 4). December 2002 Winter Stoneflies of Illinois 239 41A. 1900-1945 41B. 1946-1975 41C. 1976-2000 Figures 39-41. Allocapnia recta. 39. Male terminalia. A=lateral view, B=dorsal view, C=posterior view of tergite 8. 40. Female terminalia. 41. Distribution in Illinois. (Figs. 39-40 from Ross & Ricker 1971). 240 Illinois Natural History Survey Bulletin Allocapnia rickeri Frison Allocapnia rickeri Frison (1942:269): Hanson (1946:238), Harper and Hynes (197 1a: 925, 936), Harris and Webb (1995:342), Illies (1966:127), Poulton and Stewart (1991:20), Ricker (1945:227), Zwick (1973:365). Type locality: Illinois, Pope County, Golconda, and Big Grand Pierre Creek. Allocapnia pygmaea (Burmeister 1839:874): Frison (1929:396, 1935:367). Misidentification. Adult: (Ross and Ricker 1971). Male (Fig. 42 A-C): Wings reaching tergites 4 or 5. Tergites 6 and 7 lacking dorsal process. Tergite 8 with dor- sal process raised vertically above tergite, apex tapered posteriorly, in posterior view apex trun- cate with wide medial emargination. Upper limb of epiproct with apical segment bulbous, dis- tinctly shorter than basal segment; lower limb narrow, tapered apically. Other figures: Frison (1942: Fig. 36), Harper and Hynes (1971a: Fig. 14), Stark and Nelson (2000: Fig. 2.1, triocellate head; Fig 2.8, mouthparts). Female (Fig. 43): Tergite 8 membranous dorsally. Sternites 7 and 8 broadly fused over three-fourths of width. Ster- nite 8 with lateral lobes usually lighter than me- sal portion, posterior margin truncate. Other fig- ures: Harper and Hynes (197 1a: Fig. 37). Nymph (Harper and Hynes 1971:936): Length 6-7.5 mm. Light to medium yellowish brown. Pronotum with well-developed anterior and pos- terior fringe; few setae on disc and not much shorter than on fringe. Abdomen covered with short prostrate setae, a few long erect bristles on posterior margin (about half as long as mid-dor- sal length), a few erect intermediate setae on middle of each segment (three-fourths length of segment) (Harper and Hynes 1971a, Fig. 61). Biology: Allocapnia rickeri has a univoltine fast cycle of development with a nine-month dia- pause in Oklahoma (Ernst and Stewart 1985, Stewart and Stark 1988) and a univoltine slow cycle of development in southern Canada (Harper 1973, Ernst and Stewart 1985). Krueger and Cook (1981) noted a longer developmental time in Minnesota with small nymphs appear- ing in July, growing rapidly until December, slowing growth in December—January, then re- suming growth until a March-April emergence. Vol. 36 Art. 5 A few nymphs of A. rickeri displayed pre- sunset drifting at the beginning of emergence (Ernst and Stewart 1985, Fig. 4) and drift was related to their standing stock (Krueger and Cook 1981). Krueger and Waters (1983) and Jop and Stewart (1987) give annual production rates of this herbivore-detritivore. Habitat: Ross and Ricker (1971) reported Allocapnia rickeri from clear cool rocky or grav- elly streams. Surdick and Gaufin (1978) reported A. rickeri as being neutral or alkaliphilous, euphilous, eury-, meta-, and oligothermal, eulichtophilous, rheobiontic, and found in streams. From 1976—2000, adults of A. rickeri were collected in Illinois on bridge railings and abutments, on and under cobblestones, in leaf packs, generally from small, slow-moving streams, 1-10 m wide, less than 30 cm—1 m deep, with a gravel substrate. At one locality, it was collected abundantly from a channelized creek (Max Creek) with a clay/silt substrate. Distribution and Seasonal Activity: Allocapnia rickeri 1s widespread, ranging from Alabama to New York west to Minnesota and Oklahoma (Stark et al. 1986). Ross and Ricker (1971) and Stewart and Stark (1988) did not list this species from Illinois, although the type locality is Big Grand Pierre Creek, Golconda, Illinois. In Ili- nois, adults were collected from 1 January to 22 June. Figure 44 displays the distribution of A. rickeri from 1900-1945, 1946-1975, and 1976— 2000. Current Status. Allocapnia rickeri was de- scribed by Frison (1942) from Big Grand Pierre Creek, Pope County, Illinois. Ross and Ricker (1971) neglected to list Illinois in their distribu- tion of this species (p. 41), although their map (p. 102) correctly indicates its Illinois distribu- tion. Consequently, subsequent authors have omitted this species from the Illinois fauna. Allocapnia rickeri remains an abundant mem- ber of the winter stoneflies of Illinois. Frison reported it abundantly from across the Shawnee Hills, with a disjunct record from Jo Daviess County. Ross and Ricker (1971) reported nu- merous records from across the Shawnee Hills, with additional records from Calhoun, Fayette, Fulton, and Pike counties in central and western Illinois. From 1976—2000, A. rickeri was one of the most abundant species in the Shawnee Hills with a continuing distribution up the west December 2002 side of Illinois into Calhoun, Pike, and Schuyler counties. Two populations were also found in eastern Illinois in Vermilion and Wayne coun- ties. On two separate occasions, efforts were made to collect this species in northeastern Jo Daviess County near Warren and Apple River Canyon State Park with no success. Much of this area has been transformed into pastureland with trees being removed up to the edges of the rickert Winter Stoneflies of Illinois 241 streams. Historically (1900-2000), this species was collected from within eight Natural Divi- sions of Illinois (Table 3) and nine drainage ba- sins (Table 4). Recent records (1976-2000) in- dicate that the distribution of this species has been reduced to five natural divisions (Table 3) and six drainage basins (Table 4) yet this spe- cies 1S well established in Illinois. 44A. 1900-1945 44B. 1946-1975 44C. 1976-2000 Figures 42-44. Allocapnia rickeri. 42. Male terminalia. A=lateral view, B=dorsal view, C=posterior view of tergite 8. 43. Female terminalia. 44. Distribution in Illinois. (Figs. 42-43 from Ross & Ricker 1971). 242 Illinois Natural History Survey Bulletin Allocapnia smithi Ross and Ricker Allocapnia smithi Ross and Ricker (1971:48): Harris and Webb (1995:342), Hitchcock (1974:64), Poulton and Stewart (1991:20), Zwick (1973:368). Type locality: Kentucky, Butler County, 5.5 km west of South Hill. Adult: (Ross and Ricker 1971). Male (Fig. 45 A-D): Wings long, reaching tergite 7. Tergite 6 lacking dorsal process. Tergite 7 with dorsal process slightly raised above tergite, apex trun- cate, in posterior view apex truncate, broad. Tergite 8 with dorsal process raised vertically above tergite, quadrate, apical margin angulate; in posterior view dorsal process with three points, mesal one slightly higher than laterals. Upper arm of epiproct with apical segment subequal to basal segment, in dorsal view apical segment narrow, slightly tapered apically; lower limb in lateral view narrow, slightly swollen medially, tapered apically and recurved ventrally, in dor- sal view about as wide as upper limb. Female (Fig. 46): Wings reaching to or beyond apex of abdomen. Tergite 8 membranous dorsally. Ster- nites 7 and 8 fused for most of their width, line of fusion indicated by sharp crease. Sternite 8 with lateral areas semi-membranous; posterior margin produced into wide, sclerotized reflexible flap. Habitat: Ross and Ricker (1971) reported Allocapnia smithi from small spring fed creeks. From 1976-2000, adults of A. smithi in Illinois, were collected from leaf packs in small slow flowing streams, 5-7 m wide, 30 cm—1 m deep, with a gravel, cobblestone, and sand substrate. Distribution and Seasonal Activity: Allocapnia smithi is a southern species ranging from north- ern Alabama to western Kentucky, southern IIli- nois, and southern Ohio (Ross and Ricker 1971, Fig. 198; Stark et al. 1986). In Illinois, adults were collected from 20 January to 5 March. Illinois Record 1946-1975. Pope County: Lusk Creek, 5 km SE Eddyville. 1976-2000. Pope County: East Fork, Little Lusk Creek, 7.4 km SW Herod. Saline County: tribu- tary Battle Ford Creek, 3.7 km NE Delwood. Vol. 36 Art. 5 Current Status. Allocapnia smithi is rare in II- linois, even with the extensive collecting that has been done recently across the Shawnee Hills (Fig. 5). It is restricted to the Shawnee Hills Natural Division (Table 3) and the Lusk Creek and Saline River drainages (Table 4). Ross and Ricker (1971) reported A. smithi from Lusk Creek, southeast of Eddyville, Pope County. Recent collections have added an additional lo- cality in both Pope and Saline counties, but it was not collected at Lusk Creek, which is a fa- vored collecting site in the Shawnee Hills. 46 Figures. 45-46. Allocapnia smithi. 45. Male terminalia. A=lateral view, B=dorsal view, C=posterior view of tergite 8, D=posterior view of tergite 7. 46. Female terminalia. (Figs. 45-46 from Ross & Ricker, 1971). December 2002 Allocapnia vivipara (Claassen) Allocapnia vivipara (Claassen): Claassen (1931:114, 1940:90), Frison (1929:392, 1935:370, 1942:265), Gaufin (1958), Hanson (1946:238), Harper and Hynes (1971a:925, 937), Harris and Webb (1995:342), Hitchcock (1974:65), Harden and Mickel (1952:32), Illes (1966:128), Poulton and Stewart (1991:20), Ross (1962:105), Ross and Ricker (1971:21), Zwick (1973:369). Type locality: Lake For- est, Illinois. Capnella vivipara Claassen (1924:46): Ilies (1966:128). Capnia minima Walsh (1862) from Rock Island probably is A. vivipara (Frison 1935, Ross and Ricker 1971). Adult: (Ross and Ricker 1971). Male (Fig. 47 A-D): Wings apterous. Tergite 6 lacking dorsal process. Tergite 7 slightly raised posteriorly. Tergite 8 with dorsal process moderately raised above tergite, apical margin slanted anteriorly, broad, rounded in posterior view. Upper limb of epiproct elongate, apical segment bulbous, distinctly shorter than basal segment, triangular, tapered apically in dorsal view; lower limb with apical half bulbous with apical steplike projec- tion in lateral view, lower limb wider than api- cal segment of upper limb in dorsal view. Other figures: Frison (1929, Fig. 49, habitus; 1935, Fig. 228, terminalia, Fig. 293, habitus); Hanson (1946, Fig. 38a, metatergum); Harper and Hynes (197 1a, Fig. 10); Needham and Claassen (1925, pl. 6 Fig. 9, egg, pl. 50 Figs. 4—5, terminalia). Female (Fig. 48): Wing length variable (Ross 1962, Fig. 44), reaching from tergite 1 to apex of abdomen. Tergite 7 often with much of terg- ite sclerotized. Tergite 8 sclerotized completely. Sternites 7 and 8 with broad, medial, cone- shaped fusion. Sternite 8 with inconspicuous lateral bulges, posterior margin truncate. Other figures: Frison (1929, Fig. 46 habitus, Figs. 52— 53 habitus, Fig. 74, sterna 7-8; 1935, Fig. 218, sterna 7-8, Fig. 292 habitus); Harper and Hynes (197 1a, Fig. 36); Needham and Claassen (1925, pl. 50, Fig. 5 terminalia). Egg. Needham and Claassen (1925, pl. 6, Fig. 9). Nymph (Harper and Hynes 197 1a: 937): Length 7-10 mm. Setation pattern almost identical to that of A. granulata (Harper and Hynes 197 1a, Winter Stoneflies of Illinois 243 Figs. 62 and 63). Mature male nymphs lack wingpads. Other figures: Frison (1929:395, Fig. 15 male labium, 19 male maxilla, 23 male man- dibles, 27 male hind leg, 33 male habitus, 34 female habitus; 1935:370, Fig. 57 female man- dibles, 87 female maxilla, 117 female labium, 144 female mentum and submentum, 257 male hind leg, 297 male habitus, 298 female habitus), and Claassen (1931:114). Harper and Hynes (1970:925, Fig. 1) illustrated the diapausing nymph. Biology: Allocapnia vivipara has a univoltine life cycle (Harper and Hynes 1970) with eggs deposited in May, hatching within a month, un- dergoing a diapause during summer (Coleman and Hynes 1970, Harper and Hynes 1970, Pugsley and Hynes 1985), and developing rap- idly through the fall and winter. - Habitat: Frison (1929) reported Allocapnia vivipara from small and medium streams in Illi- nois, and it has been reported from small streams in Minnesota (Harden and Mickel 1952:32). Ross and Ricker (1971) found A. vivipara in streams of various sizes and abundant in streams heavily loaded with organic materials, as it is especially tolerant of barnyard pollution. Gaufin (1958) reported A. vivipara to be intolerant of pollution, preferring water with high levels of dissolved oxygen (70—-115% saturation). Surdick and Gaufin (1978) reported A. vivipara as being a herbivore or scavenger that was neutral or alkaliphilous, saproxenous or saprophobic, euoxyphilous, eury-, meso-, meta-, and oligothermal, rheophilous and rheobiontic, epilithic and epiphytic, and found in rivers or permanent and intermittent streams. From 1976— 2000, adults of A. vivipara were the most com- monly collected winter stonefly in Illinois. Adults were collected from bridge railings and abutments, on and under cobblestone and cement blocks, in leaf packs and woody vegetation, and as they walked across snow and ice. This spe- cies displayed a wide diversity in habitat selec- tion, being collected from small slow flowing streams, 1-10 m wide, less thanl—3 m deep, with a gravel, sand, and cobblestone substrate. In the central part of the state, A. vivipara was collected predominately from clay/silt or sand bottom streams. It was also collected from large streams over 35 m wide and greater than 2 m deep with clay/silt and sandy substrates to big rivers over 244 Illinois Natural History Survey Bulletin a quarter of a kilometer wide with clay/silt and sandy bottoms. Distribution and Seasonal Activity: Allocapnia vivipara is widely distributed from Virginia to southern Quebec west to eastern Wisconsin, east- ern Kansas and northeastern Oklahoma (Ross and Ricker 1971, Fig. 88; Stark et al. 1986), with a single locality from northeastern Arkansas (Poulton and Stewart 1991). In Illinois, adults were collected from 10 December to 30 April. Figure 49 displays the distribution of A. vivipara from 1900-1945, 1946-1975, and 1976-2000. Current Status. Allocapnia vivipara is the most = dOrsdll process Siem) Vie \ cae \ | D\ IC 47 Vol. 36 Art. 5 widespread and commonest species of winter stonefly in Illinois. It also is the most environ- mentally tolerant species; being collected from pristine streams to those heavily impacted with livestock waste. During the resurvey of Illinois, it was collected in all natural divisions, except the Illinois/Mississippi River Sand Areas (Table 3). Historically (1900-2000), it has been col- lected from all drainage basins within the state (Table 4). Recently (1976-2000), it was col- lected in all drainage basins except the Missis- sippi River and Wabash Rivers proper (Table 4). 48 49A. 1900-1945 49B. 1946-1975 49C. 1976-2000 Figures 47-49. Allocapnia vivipara. 47. Male terminalia. A=lateral view, B=dorsal view. 48. Female terminalia. 49. Distribution in Illinois. (Figs. 47-48 from Ross & Ricker 1971). December 2002 Nemocapnia Banks Nemocapnia Banks (1938:74): Claassen (1940:96), Frison (1942:262), Hanson (1946:223, 236), Harper and Hynes (1971a:938), Harper (1984), Hitchcock (1974:70), Illies (1966:160), Stewart and Harper (1996). Genotype Nemocapnia carolina Banks. This genus contain only one North American species (Bill Stark 2001 http://www.mc.edu/ campus/users/stark). Nemocapnia (Hanson 1946, Hitchcock 1974) is separated from other capniid genera by hav- ing the presternum united to the basisternum on each thoracic sternite; its mesothoracic postfurcasternum is large and united to the spinasternum and furcasternum (Hitchcock 1974, Fig. 13); vein A; of forewing is slightly bent just beyond the origin (Fig. 50) and there is no costal crossvein beyond Sc in the forewing. The epiproct is bluntly pointed, with no lower limb separate this genus from Allocapnia (Poulton and Stewart 1991). Hanson (1946:236) also defined this genus and discussed the com- parative morphology of Nemocapnia within the Capniidae. There is only one Nearctic species. Nemocapnia carolina Banks Nemocapnia carolina Banks (1938:74): Claassen (1940:96), Frison (1942:262), Harper and Hynes (1971a:926, 939), Harris and Webb (1995:342), Illies (1966:160), Poulton and Stewart (1991:21), Ricker (1945:227), Stark and Gaufin (1979:428). Type locality: Morgantown, North Carolina. Adult: (Poulton and Stewart 1991). Male (Figs. 50 and 51): Length 7 mm. Forewing (Harper 1984, Fig. 13.162; Poulton and Stewart 1991, Fig. 15; Stewart and Harper 1996, Fig. 14.166). Epiproct unipartite, sternite 9 lacking ventral lobe (vesicle) Frison (1942, Fig. 28, terminalia), Harper and Hynes (1971a, Fig. 9, terminalia), Hitchcock (1974, Fig. 68 terminalia), Stark and Gaufin (1979, Figs. 61-62, terminalia), Stewart and Stark (1988, Fig. 7.14 F—H), Poulton and Stewart (1991, Fig. 55, terminalia). Female (Fig. 52): Subgenital plate slightly rounded, but set Winter Stoneflies of Illinois 245 before the hind margin of the sternite (Hitchcock 1974), Frison (1942, Fig. 30), Hanson (1946, Fig. 51), Stark and Gaufin (1979, Fig. 63), Poulton and Stewart (1991, Fig. 54). Nymph (Fig. 53) (Harper and Hynes 197 1a: 939, Stewart and Stark 1988:122). Nemocapnia dif- fers from /socapnia in having the fringe less dense, the enclosure of the anterior mesosternal Y ridge is narrowly rectangular, and the body has fewer erect setae. Length 6-7 mm. Yellow- ish brown. Most of body with well-developed prostrate clothing hairs, no erect setae (Harper and Hynes 1971a, Fig. 52). Antennae short, about one-half length of body; mandibles with- out patches of hairs near ventral socket; legs very short, usually with fringe of setae on tibiae and tarsi. Abdomen with prostrate clothing setae, no erect setae (Harper and Hynes 1971la: Fig. _ 52). Cerci short, about 15 segments, apical seg- ments with vertical fringe of long setae which are absent of the basal 5 or 6 segments (Fig. 11). Other figures: Frison (1942, Fig. 29 mandibles, maxilla, labium, Fig. 31, habitus); Harper and Hynes (1971a, Fig. 52, abdominal setation); Harper (1984, Fig. 13.33, cercus); Poulton and Stewart (1991, Fig. 49, cercus); Stewart and Stark 1988:122, Fig. 7.13, habitus, Fig. 7.14, A head-pronotum, B mandible, C lacinia, D front leg, E mesosternum, F—H male terminalia lat- eral, dorsal ventral, I female terminalia, J cer- cus), Stewart and Harper (1996, Fig. 14.33, cer- cus). Distribution and Seasonal Activity: Nemocapnia carolina ranges from Florida to Quebec, west to Illinois and Arkansas (Stark et al. 1986). In Illinois, adults were collected 8 March. Illinois Records 1900-1945. White County: Wabash River, Grayville. Current Status: Nemocapnia carolina has been collected only once in IIlinois in 1928 from the Wabash River at Grayville. Several attempts were made to collect this species at Grayville and other localities along the Wabash River but were unsuccessful. This species must be con- sidered as extirpated from the state. 246 Illinois Natural History Survey Bulletin Vol. 36 Art. 5 ok, Figures 50-53. Nemocapnia carolina. 50. Forewing. 51. Male terminalia. 52. Female terminalia. 53. Nymph. (Figs. 50-52 from Poulton & Stewart 1991; 53 Stewart & Stark 1988). Paracapnia Hanson Paracapnia Hanson (1946:225, 236): Hanson (1961:25), Harper (1984:117), Illies (1966:161), Stewart and Harper (1996:228, 246, 264), Zwick (1973:389). This genus contains five North American spe- cies (Bill Stark 2001 http://www.mc.edu/cam- pus/users/stark). Paracapnia (Hanson 1946, Hitchcock 1974, Stewart and Harper 1996) is separated from other capniid genera by having the mesothoracic postfurcasternal plate united with the furcasterna and the spinasternus; the mesofurcasternum and metafurcasternum transverse and almost rectan- gular; forewing with 0-3 costal crossveins; Ry; of forewing slightly curved cephalad then more broadly curved distad beyond origin of Rs; and the apical portion of Cu, of the hindwing gener- ally missing; anal lobe of hindwing smaller than remainder of wing; cerci with more than 11 seg- ments. Hanson (1946:236) discusses the compara- tive morphology of Paracapnia within the Capniidae. Paracapnia angulata is separated from P. opis (Newman) in the shape of the male epiproct (Hanson 1961). Paracapnia angulata Hanson Paracapnia angulata Hanson (1961:29): Harper and Hynes (1971a:926, 939), Harris and Webb (1995:342), Hitchcock (1974:69), Illies (1966:161), Poulton and Stewart (1991:21), Zwick (1973:389). Type local- ity: Pelham, Massachussetts. Capnia opis (Newman): Frison (1942:264, in part), Ricker (1938:134, in part). Capnia vernalis Needham and Claassen (1925:356) (male and female figures ques- tionable): Frison (1942:264), Hanson (1961:29). Adult (Hanson 1961): Male (Figs. 54-55): Epiproct thick, broadened once then tapered to apex. Other figures: Frison (1942, Fig. 32, terminalia, as P. opis); Hanson (1946, Fig. 52, as P. opis); Hanson (1961, Fig. 1-4 epiproct), Harper and Hynes (1971a: Fig. 2 terminalia), Hitchcock (1974, Fig. 65, epiproct); Poulton and Stewart (1991, Figs. 51-52, terminalia). Female (Fig. 56): indistinguishable from P. opis. Other figures: Frison (1942, Fig. 32, terminalia, as P opis); Poulton and Stewart (1991, Fig. 53, terminalia). Nymph (Figs. 10, 19, 57) (Harper and Hynes 1971a:939): Length 6-8 mm. Reddish brown with distinct purplish markings on head; append- ages and wingpads yellowish. Body generally hairy. Head with long setae, especially on antero- December 2002 lateral margins of frons and near compound eyes. Antennae long, about three-fourths the length of body. Pronotum with numerous long setae (Harper and Hynes 1971a, Fig. 56); meso- and metanota with tufts of long bristles on anterior angles. Wingpads bordered with long bristles. Outer margins of legs with long bristles; bristles on inner surface of tibiae short, half as long as width of tibia; middle and distal segments of cerci with long intermediate bristles at ventral base of segment. Abdomen with clothing setae yellow, many erect purplish setae on margins of tergites. Cerci about as long as abdomen, each segment with whorl of short bristles. Other fig- ures: Harper (1984, Fig. 13.34, abdominal terga); Stewart and Harper (1996, Fig. 14.34, abdomi- nal terga); Stewart and Stark (1988, Fig. 7.15, habitus, Fig. 7.16 A—J, A head-pronotum, B man- dible, C lacinia, D front leg, E mesosternum, F— H male terminalia lateral, dorsal, ventral, I fe- male terminalia, J-L cercus, basal, middle, and apical sections). First instar nymph: Harper (1979, Fig. 8, first- instar habitus). Biology: In Illinois, adults emerged over a short period in March. Eggs presumably hatch within a few weeks (Harper and Hynes 1972) and the nymphs grow rapidly throughout the summer and autumn. The last instar occurring sometime in December with a slowing of growth until emer- gence in early April. No nymphal diapause has Winter Stoneflies of Illinois 247 been observed (Harper and Hynes 1970, 1972). Surdick and Gaufin (1978) reported Paracapnia angulata as being neutral or alkaliphilous, eury- meta- and oligothermal, and found in streams. Habitat: In Illinois, Paracapnia angulata has been collected from small, spring-fed streams, 30-60 cm wide and less than 30 cm deep with a gravel and sand substrate. Distribution and Seasonal Activity: Paracapnia angulata ranges from North Caro- lina to Maine west to Washington and Oregon (Stark et al. 1986, Stewart and Stark 1988). In Illinois, adults were collected from 7—20 March. Illinois Records 1900-1945. Kane County: Elgin, Trout Park (Elgin Botanical Garden). - Current Status: Paracapnia angulata in Mlinois has been collected on two separate dates from an area of springs and spring-fed streams in Trout Park (Elgin Botanical Garden) in 1940. These springs and shallow streams support a rich vari- ety of aquatic organisms, with several species of caddisflies and one species of mayfly found nowhere else in the state (Ross 1944). Several attempts were made to collect P. angulata dur- ing the past five years but with no success. At this time, this species must be considered as ex- tirpated from the state. a0 Figures 54-57. Paracapnia angulata. 54-55. Male terminalia. 56. Female terminalia. 57. Nymph. (Figs. 54-56 from Poulton & Stewart 1991; 57 Stewart & Stark 1988). 248 Illinois Natural History Survey Bulletin Leuctridae This family contains 9 North American genera andiisauspecies: (Bill@StarkeZOOImnttip. 77 www.mce.edu/campus/users/stark). Adults (Stewart and Harper 1996, Stark and Nelson 2000) have the glossae and paraglossae subequal in length and size, set at same level on labium, apical segment elongate, but not much larger than preceding segment; the thorax lack- ing gill remnants; first tarsal segment much longer than second; wings at rest rolled over the back and sides of the abdomen, giving leuctrids a cylindrical, stick-like appearance; no crossvein in the costal space beyond the cord; vein A> in forewing forked; intercubital crossveins numer- ous, usually five or more; cerci one-segmented. Nymphs (Stewart and Harper 1996) have the glossae and paraglossae subequal in length and size; lack cervical gills; have the extended hindlegs reaching far short of the abdominal apex; first tarsal segment longer than second; the hindwing pads usually longer than wide; abdomi- nal terga 1-2 or 1-3 lacking gill tufts; at most abdominal segments 1—7 separated by a mem- branous pleural fold; a posterior fringe of setae on abdominal terga (lacking in Zealeuctra). Biology: Leuctrids are principally sprawler- clingers in lotic situations where they feed as shredder-detritivores (Stewart and Harper 1996). Vol. 36 Art. 5 Zealeuctra Ricker Zealeuctra Ricker (1952:173): Harper (1984:116), Illies (1966:120), Stewart and Harper (1996: 218-219, 228, 249, 263), Zwick (1973:412). Type species Leuctra claasseni (Frison 1929). This genus contains eight North American spe- cies (Bill Stark 2001 http://www.mc.edu/cam- pus/users/stark). Adults (Poulton and Stewart 1991, Stewart and Harper 1996) have the glossae and paraglossae subequal in length and size (Fig. 15); gill remnants absent on sides of thorax and ab- dominal segments |—2; at rest, wings rolled, cov- ering both back and sides of abdomen; rN of forewing forked (as in Fig. 13); intercubital crossveins numerous; crossveins absent beyond cord in costal space; first tarsal segment longer than second (as in Fig. 9); cerci one-segmented (Fig: S8C)- Nymphs (Poulton and Stewart 1991, Stewart and Harper 1996, Stewart and Stark 1988) have the wing pads nearly parallel to axis of body (as in Fig. 16); abdominal segments cylindrical, seg- ments 1-6 separated by a pleural fold, posterior margin of terga without setal fringe (Fig. 60). Key to adults of Illinois Zealeuctra (Hitchcock 1974, Ricker and Ross 1969) 1. Male epiproct with secondary cusp near tip of main cusp (Fig. 62B); female sternite 7 with hind margin emarginate medially to contain median lobe (Fig. 63)............ccesseecceeeeeseeeeees raxina Male epiproct ending in simple tapered cusp (Figs. 58B and 64B); female sternite 7 with hind Marcin (roncale (E1vs OU and OD) eer ars 2. Male tergite 9 with medial margin of central depression minutely and irregularly serrated ante- riorly (Fig. 58A); base of epiproct rounded, as long as wide (Fig. 58B); female sternite 7 with median lobe thick, white, as broad as long (Fig. 59); sternite 7 with posterior marginal hairs NUMETOUSAIOUS see eee eee eee ollie ee ee Ol ee eee claasseni Male tergite 9 with medial margin of central depression smooth except for rounded terminal and subterminal projections (Fig. 64A); base of epiproct half as long as wide (Fig. 64B); female sternite 7 with median lobe brown, about twice as broad as long (Fig. 65); sternite 7 with poste- Hormareinabhairs tewsshortqess ee eee Of the three species of Illinois Zealeuctra, only the nymphs of Z. claasseni are known. December 2002 Zealeuctra claasseni (Frison) Zealeuctra claasseni (Frison): Harris and Webb (1995:342); Hitchcock (1974:87), Illies (1966:120), Poulton and Stewart (1991:23), Ricker and Ross (1969), Snellen and Stewart (1979:65), Stewart et al. (1974), Ziegler and Stewart (1977), Zwick (1973:41). The type local- ity was given as Bushy Fork, Herod, Illi- nois. On the 1916 Equality Quadrangle topographic map this fork is listed as Brushy Fork and on current topographic maps is cited as Pinhook Creek. Leuctra classeni Frison (1929:404, in part): Claassen (1931:100, 1940:77), Frison (1935:354), Ricker (1945:227), Zwick (1973:412). Leuctra (Paraleuctra) claasseni Frison: (Frison 1942:256). Leuctra (Zealeuctra) claasseni: Jewett (1956:168), Ricker (1952:173). Paraleuctra claasseni (Frison): Hanson (1941:58). Adult: (Hitchcock 1974, Poulton and Stewart 1991, Ricker and Ross 1969). Length 8-10 mm. Male (Fig. 58): Lateral margins of medial de- pression strongly convergent anteriorly, almost V-shaped (Fig. 58A), terminating posteriorly in a sharp medially directed point and smaller sub- terminal projection. Epiproct with single sharp tooth on broadly rounded base (Fig. 58B). Cerci with rounded hump on upper side (Fig. 58C). Other figures: Frison (1929, Figs. 68 and 69, terminalia; 1935, Figs. 178, 194, 230, 285; 1942, Fig. 18). Female (Fig. 59): Sternite 7 with nu- merous long hairs, especially near hind margin; posterior margin straight; median lobe white, about as broad as long. Sternite 8 with anterior pair of sclerotized knobs bearing long hairs, and lying on either side of the median lobe of stern- ite 7. Nymph: (Fig. 60). Frison (1929, Fig. 16, labium; Fig. 20, maxillae; Fig. 24 mandibles; Fig. 28, legs; Fig. 38, male nymph; 1935, Figs. 59, 88, ise tS3. 298, 253; 284) and Claassen (1931:100) have described the nymph. In 1935, Frison described anal gills on the nymph but Hitchcock (1974:8) concluded that these were probably fungal growths. Other figures: Harper Winter Stoneflies of Illinois 249 (1984, Figs. 13.13 and 13.48, abdominal ventrum; Fig. 13.49, male abdominal dorsum); Stewart and Stark (1988, Fig. 8.14 a—l, a head- pronotum, b mandible, c lacinia, d front leg, e mesosternum, f—h male terminalia lateral dorsal ventral, 1 female terminalia, j—l basal, middle, and apical sections of cercus) and Stewart and Harper (1996, Fig. 14.13, abdominal venter; Fig. 14.48, abdominal venter; fig. 14.49, abdominal dorsum). The first instars (Snellen and Stewart 1979, Fig. 9 and Fig. 11 A mandible, B maxilla, C hind leg) are consistent in being unpigmented, with- out ocelli, and having antenna:cercal segment formulae of 8-11:3-5; compound eyes with 2-4 ommatidia; gills absent, reduced, or represented only by knobs or stubs, generally few hairs com- pared with later instars; and three tarsal seg- _ ments, the first two short and together subequal to the longer third. Biology: Snellen and Stewart (1979) found Zealeluctra claasseni to have a flexible life cycle. Both diapausing and nondiapausing eggs are pro- duced with extended periods of egg diapause in dry years and a very fast nymphal growth. Adults displayed a peak in emergence when the daily water temperature was about 12°C. Emergence was diurnal and teneral adults and mating pairs were found from dawn to late evening. Ziegler and Stewart (1977) described the drumming of the male to be a multi-beat signal with more than 20 beats per signal and an interbeat interval that shortened as the signal progressed. Snellen and Stewart (1979) also reported on the drumming behavior. Laboratory reared nymphs fed on de- tritus and algae (Snellen and Stewart 1979) and Frison (1929) found decaying vegetation in the gut of one nymph. Habitat: Zealeuctra claasseni emerges early in the spring (Frison 1935) and can be found even in streams that dry up in midsummer (Ricker 1952) or in intermittent limestone-base streams (Stewart and Stark 1988). Surdick and Gaufin (1978) reported Z. claasseni as being a herbi- vore, alkaliphilous, saprophobic, euoxyphilous, meso- and metathermal, rheobiontic, epilithic and epiphytic, and found in permanent and in- termittent streams. From 1976-2000, Z. claasseni was collected from bridge railings and leaf packs near small, permanently flowing streams, 1—2 m wide, less than 60 cm deep, with a gravel, sand, and cobble substrate. 250 Illinois Natural History Survey Bulletin Distribution and Seasonal Activity: Zealeuctra claasseni ranges from West Virginia to Okla- homa and eastern Texas, with isolated popula- tions in the Wichita and Arbuckle mountains of southwestern and south-central Oklahoma (Stewart et al. 1974, Stark et al. 1986). In Tli- nois, adults were collected from 27 February to | May. Figure 61 displays the distribution of Z. claasseni from 1900-1945, 1946-1975, and 1976-2000. 58A Vol. 36 Art. 5 Current Status: Zealeuctra claasseni is uncom- mon in Illinois but it remains established across the Shawnee Hills. Historically (1900-2000), it was collected in the Shawnee Hills and South- ern Till Plain Natural Divisions of Illinois (Table 3) and four river drainages in the Shawnee Hills (Table 4). Recently (1976-2000), it was collected only in the Shawnee Hills Division (Table 3). | he 4 Pos ey ry bS ey 60 rT 59 Y< a F hae +t . wf => ] pom 61 1976-2000 @ 1946-1975 1900-1945 4m On Figures 58-61. Zealeuctra claasseni. 58. Male terminalia (A). B=epiproct, C=cercus. 59. Female terminalia. 60. Nymph. 61. Distribution in Illinois. (Figs. 58-59 from Poulton & Stewart 1991; 60 Stewart & Stark 1988). December 2002 Zealeuctra fraxina Ricker and Ross Zealeuctra fraxina Ricker and Ross (1969:1117): Harris and Webb (1995:342), Poulton and Stewart (1991:23), Zwick (1973:413). Type locality Hardinsburg, Breckenridge County, Kentucky. Leuctra claasseni Frison (1929, in part, Fig. 68 and 69 are of the female paratype of Zealuctra fraxina. Leuctra claasseni Frison (1935, in part, Fig. 194 and 230 are Zealuctra fraxina. Adult: (Hitchcock 1974, Poulton and Stewart 1991, Ricker and Ross 1969). Length 8-10 mm. Male: Tergite 9 with medial depression U- shaped, lateral margins heavily sclerotized (Fig. 62A), terminating posteriorly in a sharp medi- ally directed point and smaller subterminal pro- jection. Epiproct with secondary subapical cusp (Fig. 62B). Cerci with a blunt, fingerlike, lat- eral terminal lobe and a dorsal lobe near middle of cercum (as in Fig. 64C). Female (Fig. 63): Sternite 7 with median portion well sclerotized and with numerous hairs; posterior margin strongly notched medially; median lobe white. Habitat: Adults of Zealeuctra fraxina were 62B Winter Stoneflies of Illinois 251 collected from leaf packs along a small, slow flowing stream, 3 m wide, 30 cm deep, with a cobble, gravel, and sand substrate. Distribution and Seasonal Activity: Zealeuctra fraxina ranges from Tennessee to West Virginia, west to southern Illinois (Stewart et al. 1974, Stark et al. 1986). In Illinois, adults were col- lected from 20 January to 6 March. Illinois Records. 1900-1945. Pope County: Gibbons Creek, Herod. 1976-2000. Pope County: tributary Hart Creek, 2.5 km SW Herod. Saline County: tributary Battle Ford Creek, 3.7 km NE Delwood. Current Status: Zealeuctra fraxina is rare in Illinois but is still established at two sites in the eastern Shawnee Hills. Historically (1900- 2000), it has been collected only in the Shawnee Hills Natural Division (Table 3). Prior to 1946, it had been collected only in the Big Grand Pierre Creek drainage (Table 4), but recently (1975-— 2000) it was also collected in the Saline River Drainage (Table 4). Figures 62-63. Zealeuctra fraxina. 62. Male terminalia (A), B=epiproct. 63. Female terminalia. (Figs. 62-63 from Ricker & Ross 1969). 252 Illinois Natural History Survey Bulletin Zealeuctra narfi Ricker and Ricker Zealeuctra narfi Ricker and Ross (1969:1118): Harris and Webb (1995:342), Hitchcock (1974:89), Poulton and Stewart (1991:24), Zwick (1973:413). Type locality Otter Creek, Sauk County, Wisconsin. Leuctra claasseni Frison (1929, in part) the fe- male specimen from Fountain Bluff, Illinois is Z. narfi. Adults: (Hitchcock 1974, Poulton and Stewart 1991, Ricker and Ross 1969). Length 6.0—7.5 mm. Male: Tergite 9 with lateral margins of medial depression terminating in a rounded point and rounded subterminal tooth (Fig. 64A). Epiproct with single tooth on a subrectangular base that is wider than long (Fig. 64B). Cerci with a blunt, fingerlike, lateral terminal lobe and a dorsal lobe near middle of cercum (Fig. 64C). Female (Fig. 65): Sternite 7 with scattered short hairs, not forming a tuft posteromedially, poste- rior margin straight, median lobe brown, about twice as wide as long. Habitat: Surdick and Gaufin (1978) reported Zealeuctra narfi as being neutral with regards to pH, eury- and oligothermal, eu- and mesolichtophilous, and rheophilous. In Illinois, adults of Z. narfi were collected from leaf packs along a small, slow flowing stream, 3 m wide, 64A Vol. 36 Art. 5 30 cm deep, with a cobble, gravel, and sand sub- strate. Distribution and Seasonal Activity: Zealeuctra narfi ranges from Oklahoma to Wisconsin (Stark et al. 1986). In Illinois, adults were collected from 6 February to 5 May. Illinois Records 1900-1945. Jackson County: Fountain Bluff. 1976-2000. Saline County: tributary Battle Ford Creek, 3.7 km NE Delwood. Union County: Alto Pass, Cave Spring Cave. Vermilion County: Kickapoo State Park. Current Status: Zealeuctra narfi was originally known only from Fountain Bluff, Jackson County, Illinois. Several attempts were made to collect this species at Fountain Bluff with no success. From 1976—2000, specimens were col- lected at three new localities in Saline, Union, and Vermilion counties in eastern Illinois. AlI- though rare in Illinois, this species is still estab- lished. Prior to 1946, this species was known only from the Shawnee Hills Natural Division and within the Big Muddy Drainage System. Recently (1975-2000), it has been collected within the Shawnee Hills and Wabash Border Natural Divisions and the Big Muddy, Saline, and Vermilion River drainage basins (Table 4). 65 Figures 64-65. Zealeuctra narfi. 64. Male terminalia (A), B=epiproct, C=cercus. 65. Female terminalia. (Figs. 64-65 from Poulton & Stewart 1991). December 2002 Nemouridae This family contains 12 North American genera and./1 species (Bill Stark 2001 http:// www.imce.edu/campus/users/stark). Adults (Poulton and Stewart 1991:28, Stewart and Harper 1996:239, Stark and Nelson 2000) have the glossae and paraglossae subequal in length and size; apical segment of labial palpus circular, larger than penultimate segment; gill remnants lacking on sides of thorax; at rest, wings folded flat over dorsum; an extra crossvein in the costal space beyond the cord of the fore- wing (Fig. 13); first tarsal segment longer than second (as in Fig. 9); and cerci one-segmented (Fig. 66); female subgenital plate usually small. Nymphs have divergent wing pads similar to Taeniopterygidae. Only one species Prostoia completa is considered a winter stonefly in [li- nois. Biology: Nemourids are principally sprawler- clingers in lotic situations where they feed as shredder-facultative detritivores or collector- gatherers (Stewart and Harper 1996). Prostoia Ricker Prostoia (Ricker): Harper (1984:116), Illies (1966:220), Poulton and Stewart (1991:29), Stewart and Harper (1996:29). Type spe- cies Prostoia completa (Walker), Zwick (1973:345). Nemoura (Prostoia) Ricker (1952:47): Ilhes (1966:220). This genus contains four North American spe- cies (Bill Stark 2001 http://www.mc.edu/cam- pus/users/stark). Adults (Harper 1984, Stewart and Harper 1996) have gill remnants absent on the submentum or cervix; forewing with terminal crossvein join- ing R (Fig. 13), veins Aj and A2 separate; male sternum 9 with vesicle, epiproct long and simple, cerci membranous; female wings distinctly banded, sternum 8 unsclerotized except at mar- gin, and not extending over sternum 9. Nymphs (Harper 1984, Stewart and Stark 1988) have the submentum and cervix lacking gills; width of the pronotum usually subequal to dis- tance between eyes, with rounded corners, with- out a lateral notch and with tiny inconspicuous Winter Stoneflies of Illinois 253 bristles; fore tibial fringe of outer hairs complete, without distinct rows of robust bristles; tibia with short, stout bristles along outer margins (Fig. 9); middle and apical cercal segments with dorsal or ventral intersegmental hairs or both. Biology: Nymphs of Prostoia are herbivore- detritivores (Krueger and Waters 1983) or shredder-detritivores (Stewart and Harper 1996). Prostoia completa (Walker) Prostoia completa (Walker): Illies (1966:221), Poulton and Stewart (1991:29), Zwick (1973:346). Type locality Nova Scotia, Canada. Nemoura (Prostoia) completa (Walker): Poulton and Stewart (1991:29), Ricker (1952:49) - Nemoura completa Walker (1852:191): Claassen (1940:53), Harden and Mickel (1952:16), Poulton and Stewart (1991:29), Zwick (1973:346). Nemoura glabra Claassen (1923:281): [lies (1966:221). Type locality Nova Scotia, Canada. Adult: (Poulton and Stewart 1991). Male (Fig. 66): Epiproct widened with a short basal pro- cess. Other figures: Needham and Claassen (1925: pl. 34, Figs. 1-4), Ricker (1952, Fig 23, epiproct). Female (Fig. 67): Posterior margin of sternite 8 not extending over sternite 9. Nymph: (Fig. 68). Harden and Mickel (1952, pl. 1, Fig. 2, pl. 8, Figs. 1-2). First instar nymph: Harper (1979, Fig. 4, first instar habitus). Biology: In Oklahoma, Ernst and Stewart (1985) reported Prostoia completa to have a univoltine fast life cycle with a probable egg diapause of 6—7 months during spring and summer. Harper (1973) reported a four-month diapause for eggs from southern Ontario reared in the laboratory. Ernst and Stewart (1985, Fig. 3) reported nymphs during February and March and that the size of the drifting nymphs was always greater than the average size of nymphs concurrently found in benthic samples. Bishop and Hynes (1969) re- ported on the upstream migration of P. completa (as Nemoura) in the Speed River, Ontario. Habitat: Surdick and Gaufin (1978) reported Prostoia completa as being alkaliphilous, eury-, 254 Illinois Natural History Survey Bulletin meso-, and oligothermal, and found in rivers and streams. Stewart and Poulton (1991) collected P. completa from permanently flowing streams. From 1976-2000, adults of P. completa were col- lected from leaf packs along a permanent, mod- erately flowing stream, 3-4 m wide, less than 30 cm deep with a gravel and sand substrate. Distribution and Seasonal Activity: Prostoia completa ranges from Alabama to Quebec west to Minnesota and Oklahoma (Stark et al. 1986, Stewart and Stark 1988). In Illinois, adults were collected from 13 March to 19 April. Figures 66—68. Prostoia completa. 66. Male terminalia. 67. Female terminalia. 68. Nymph. Poulton & Stewart 1991; 68 Stewart & Stark 1988). Taeniopterygidae This family contains 6 North American genera and 34 species (Bill Stark 2001 http:// www.me.edu/campus/users/stark). Adults (Harper 1984, Poulton and Stewart 1991, Stewart and Harper 1996, Stark and Nelson 2000) have the glossae and paraglossae subequal in length and size (as in Fig. 15); cervical gill remnants lacking; the first tarsal segment long, although subequal in length to the second (Fig. 8); cerci multisegmented or one-segmented. Vol. 36 Art. 5 Illinois Records 1946-1975. Union County: Hutchins Creek. 1976-2000. Union County: Hutchins Creek, 5.4 km E Wolf Lake. Current Status. Prostoia completa is rare in [llinois, having been collected at only one site in the Big Muddy Drainage System in the Shawnee Hills Natural Division. It still can be collected at that site. Figs. 66-67 from Nymphs (Harper 1984, Poulton and Stewart 1991, Stewart and Harper 1996) have the glossa and paraglossae subequal in length (as in Fig. 15); wing pads strongly divergent from the body: axis (Fig. 17); tarsal segments 1 and 2 subequal in length (Fig. 8), and ventral gill tufts absent on abdominal segments | and 2. Biology: Taeniopterygids are principally sprawler-clingers in lotic situations where they feed as shredder-detritivores or facultative col- lector-gatherers (Krueger and Waters 1983, Stewart and Harper 1996). Key to Genera of Taeniopterygidae in Illinois (Stewart 2000). 1. Gill scar present, appearing as a membranous circle in coxa; sternum 9 extended little, if any, distad to tergite 10 in males, and not reaching base of subanal lobes in:stemales"3.-3 = oe FE PPP Pet AREY td PRS Sead RK sen Taeniopteryx Gill scar absent on coxa; sternum 9 extended distinctly distad to tergite 10 in males, and distad to base of subanal lobes 'in fermales......................sccccssccsssseeceeseeceeees Strophopteryx December 2002 Strophopteryx Frison Strophopteryx Frison (1929:374): Claassen (1940:41), Harper (1984:205), Illies (1966:67), Stewart (2000:59), Stewart and Harper (1996: 224,242), Zwick (1973:315). Type species Strophoteryx (Semblis) fasciata (Burmeister). Taeniopteryx (Rhabdiopteryx) Banks (1907:14); Illies (1966:67). This genus contains 5 North American species (Bill Stark 2001, http://www.mc.edu/campus/us- ers/stark). Adults (Harper 1984, Ricker and Ross 1975, Stewart and Harper 1996) have 1 or 2 costal crossveins; Rs with two branches, Cu, with 2-3 branches; coxa lacking a gill scar; posterior mar- gin of the male sternum 9 symmetrical, elevated medially, corners low; apex of the epiproct sym- metrical with one prong; female sternum 9 subtriangular, tapered posteriorly (Fig. 71). Nymphs (Harper 1984, Stewart and Harper 1996) have the coxae without gills; tibiae with dorsal and ventral hair fringe (Strophopteryx fasciata) or only a dorsal hair fringe; some basal cercal segments or entire cercus with dorsal hair fringe. Biology: Nymphs have a univoltine life cycle and within the functional feeding groups they are scrappers (Krueger and Waters 1983). Frison (1929) reported that the nymphs of Taeniopterygidae fed on decaying leaves and seven species of diatoms. Brinck (1949), Hynes (1941), and Richardson and Gaufin (1971) re- ported taeniopterygids feeding primarily on de- tritus, with appreciable amounts of plant mate- rial, diatoms, and algae. Strophopteryx fasciata (Burmeister) Strophopteryx fasciata (Burmeister): Claassen (1940:41), Frison (1929:384, 1935:347), Funk and Sweeney (1990), Harper and Hynes (1971b:1942), Harris and Webb (1995:342), Illies (1966:67), Poulton and Stewart (1991:27), Stewart (2000:61), Ricker and Ross (1975:143), Zwick (1973:315). Type locality: Pennsylvania. Semblis fasciata Burmeister (1839:875): Illies (1966:67), Ricker and Ross (1975:143). Winter Stoneflies of Illinois 255 Brachyptera fasciata (Burmeister): Frison (1942:250), Harper and Hynes (1972:302), Harden and Mickel (1952:14), Hitchcock (1974:127). Brachyptera (Strophopteryx) fasciata (Burmeister): Harper and Hynes (1971b:942), Hitchcock (1974:127). Nemoura fasciata (Burmeister): Pictet (1841:359). Rhabdiopteryx fasciata (Burmeister): Banks (1907:14), Ilies (1966:67). Taeniopteryx (Strophopteryx) fasciata (Burmeister): Frison (1942:250), Ricker (1945:226). Taeniopteryx fasciata (Burmeister): Claassen (1931:106), Hagen (1861:34), Needham and Claassen (1925:243). Taeniopteryx frigida Hagen (1861:35): Illies (1966:67). Adults: (Hitchcock 1974, Poulton and Stewart 1991:26). Length 10-15 mm. Male (Figs. 69 and 70): Tergite 9 with two conspicuous poste- rior setose lobes extending posteriorly. Sternite 9 broad, elliptical anteriorly, posterior half nar- rowed, recurved, bifurcated apically. Epiproct connected to tergite 10 by three sclerotized bands. Other figures: Frison (1935, Figs. 199 226 280, terminalia; 1942, Figs. 9 and 10, terminalia); Needham and Claassen (1925, pl. 33, Fig. 2, pl. 45 Figs. 3 and 4, pl. 46, Fig. 2); Harper (1984, Fig. 13.127, terminalia); Stewart and Harper (1996, Fig. 14.130 terminalia); Stark and Nelson (2000, Figs. 2.28 and 2.29 epiproct); Stewart (2000, Figs. 5.43—5.46, terminalia). Female (Fig. 71): Subgenital plate with narrow tonguelike extension, tapered posteriorly, trun- cate apex. Other figures: Frison (1929, Fig. 30, coxae; 1935, Fig. 2b, coxae, Fig. 172 forewing, Fig. 280 terminalia); Stewart (2000, Fig. 5.47, genitalia). Nymph: (Fig. 72) (Harper and Hynes 1971b:942, Fig. 1, Poulton and Stewart 1991:27, Stewart and Stark 1988:207). Mature nymphs 8-9 mm. Overall color yellow with distinctive dark patterns on head and thorax; antenna subequal in length to body length; yellow with basal three segments dark, mouthparts yellow, labrum with indistinct median marking. Wings pads with dark markings. Legs yellow, femora becoming darker distally but with terminal yel- 256 Illinois Natural History Survey Bulletin low band, tibiae with proximal dark band and often a longitudinal median line; legs with dor- sal and ventral fringe of long hairs. Abdomen yellow, basal half and posterior margin of terg- ites dark; sternites 1—7 unsclerotized; dorsum of tergites with short bristles, posterior margin with row of short bristles and a few long hairs; cerci longer than body, yellow with light apical band, each segment with terminal whorl of short api- cal bristles. Other figures: Claassen (1931:106, pl. 29 Fig. 215 male terminalia, 216 female terminalia), Frison (1929, Fig. 11, female exuvium, Fig. 14, male labium, Fig. 18, male maxillae, Fig. 22, male mandibles, Fig. 26, male hind leg, Fig. 31, female habitus, Fig. 42, female nymph, Fig. 43, male and female nymphs); Frison (1935, Fig. 56, female mandible, Fig. 86, female maxillae, Fig. 116, female labium, Fig. 148, female mentum and submentum, Fig. 255, male hind leg, Fig. 278, female habitus, Fig. 279, female terminalia); Harper (1984, Fig. 13.4c); Stewart and Harper (1996, Fig. 14.4d, habitus); Stewart and Stark (1988, Fig. 10.7, habitus, Fig. 10.8 a—h, a lacinia, b mandible, c front leg, d, mesosternum, e, f, g, male terminalia, lateral, dorsal, ventral, h, female terminalia). First instar nymph: Harper (1979, Fig. 2, habitus); Stewart and Harper (1996, Fig. 14.4c, habitus). Harper and Hynes (1970, Fig. 2) de- scribed the diapausing nymph. Biology: Strophoteryx fasciata has a fast univoltine life cycle, anymphal diapause (Frison 1929, 1935; Harper and Hynes 1970, 1972) be- ginning in the fourth instar (Harper and Hynes 1970, Puglsley and Hynes 1985) and a peak emergence during the last half of March and the first half of April (Frison 1929, 1935). Eggs develop directly following oviposition and nymphs in the laboratory underwent diapause in their fourth instar (Harper and Hynes 1972). The nymphs prefer the gravel areas of the stream and are herbivorous, feeding on leaf fragments and diatoms (Frison 1929, 1935). Frison (1929) re- ported the adults feeding on the blossoms of elm trees. Habitat: Surdick and Gaufin (1978) reported Strophopteryx fasciata as being a herbivore that was neutral or alkaliphilous, saprophobic, euoxyphilous, eury-, meso-, and metathermal, eulichtophilous, rheophilous and rheobiontic, epilithic, and found in rivers and streams. Harper Vol. 36 Art. 5 and Hynes (1971b) reported S. fasciata from large streams and rivers and in 1972 they re- ported it to be a warm river species. Poulton and Stewart (1991) reported S. fasciata from permanently flowing streams in mountainous regions. From 1976-2000, S. fasciata in Illi- nois has been collected from leaf packs along broad, moderate flowing streams 35 m wide and 30 cm—1 m deep with a gravel substrate. Distribution and Activity: Strophopteryx fasciata ranges from Alabama to Quebec, west to Manitoba and Oklahoma (Stark et al. 1986, Stewart and Stark 1988). In Illinois, adults were collected from 23 November to 23 April. Fig- ure 73 displays the distribution of S. fasciata from 1900-1945, 1946-1975, and 1976-2000. Current Status: Through 1945, Strophopteryx fasciata was recorded from 39 localities through- out Illinois. It was collected at five localities from 1946-1975 and four localities from 1976— 2000. The distribution of this species has been severely reduced in the state, currently being found only in the Kankakee River in northern [linois and Lusk Creek in the Shawnee Hills. December 2002 Winter Stoneflies of Illinois 257 73A. 1900-1945 73B. 1946-1975 73C. 1976-2000 Figures 69-73. Strophopteryx fasciata. 69. Male terminalia, lateral view. 70. Male terminalia, dorsal view. 71. Female terminalia. 72. Nymph. 73. Distribution in Illinois. (Figs. 69-71 from Poulton & Stewart 1991; 72 Stewart & Stark 1988). 258 Illinois Natural History Survey Bulletin Taeniopteryx Pictet Taeniopteryx 1963: Official list of generic names in Zoology. Opinion Number 652, ICZN: Harper (1984:205), Stewart (2000:63), Stewart and Harper (1996: 219, 242, 262), Zwick (1973:304). Nemoura (Taeniopteryx) Pictet (1841:345): Banks (1907:14), Illies (1966:71), Needham and Claassen (1925:236). Type species: Phryganea nebulosa Linnaeus. Neophopteryx Klapalek (1902:179): Illes (1966:71). (Preoccupied). Type species: Phryganea nebulosa Linnaeus. Nephelopteryx Klapalek (1903:42). (new name for Neophopteryx): Ilies (1966:71). Vol. 36 Art. 5 This genus contains 11 North American species (Bill Stark 2001 http://www.mc.edu/campus/us- ers/stark). Adults (Harper 1984, Stewart and Harper 1996) of Taeniopteryx are characterized by having the glossae and paraglossae subequal in length (as in Fig. 15); gill remnants lacking from sides of the thorax; and the first tarsal segment subequal in length to the second (as in Fig. 8). Nymphs (Harper 1984, Stewart and Harper 1996) of Taeniopteryx have the glossae and paraglossae subequal in length (Fig. 15); thoracic sterna not overlapping on succeeding sterna; wing pads divergent posteriorly (Fig. 17); coxae with unmodified gills; gills absent on abdomi- nal segments 1-2; first and second tarsal seg- ments equal in length (Fig. 8); cerci with com- plete vertical fringe. Key to Adults of Illinois Taeniopteryx (Ricker and Ross 1968) Males 1:» Sternite 9 with ventral lobe (vesicle) (Fig. 76).2..........2s..c...cegece-scdeesucese s.0e ie ee Z Sternite 9 lacking Ventral LODE .....o 6s .fcsescanteecsdeosseeabencedecssecceve-tetenesucusde ttagtseie: (e seett eaten 3 2. Sternite 9 with hairs on posterior margin pointing ventrally and anteriorly, usually much shorter than more anterior hairs and with ventral lobe 2—3 times longer than wide; extruded aedeagus lacking brown band between lateral lobes .................cceeseseeeeees nivalis Sternite 9 with hairs on posterior margin directed posteriorly, much longer than anterior hairs and with ventral lobe 3—5 times longer than wide (Fig. 76); extruded aedeagus with brown band between lateral lobes (Fig. 74)... eee eeeceeeeessneeceeeeeesnenees burksi 3. Apical half of subanal lobes broad, flat, apical margin broadly rounded (Fig. 80) ..................4+ lita Apical half of subanal lobes reduced, clavate (Figs. 84; 91)..,....2c...--t--ccceccsscesestesecceseeeeeae teen 4 4. -Apex of subanal lobes bent outward; subacute (Fig. 91) <0... c.ccscc-<.ccsceesetesssuencseeeseueeeaanane parvula Apex of subanal lobes straight, somewhat swollen, narrowly rounded at apex (Fig. 84).............. EE LT Te ere me EAM Pere re esis aie rote en RN TEEPE ri PRE erericcgorce cis metequi Females 1. Sternite 8 with posterior emargination deep, broadly U-shaped, heavily sclerotized laterally (Fig:81) sis ee pe ee lita Sternite 8 with posterior emargination shallow, V-shaped, normally sclerotized (Pigs.77,'85, 89,92) 055 Be Pe ie relack coun ger sos ace ee 2 December 2002 Winter Stoneflies of Illinois 259 2. Sternite 8 with medial half lightly sclerotized, pale, contrasting strongly with central plate and narrow dark margin of emargination (Fig. 92)...............sscccccssserccecsscecessssnceees parvula Sternite 8 with medial half normally sclerotized, occasionally with pale anterior area PMT CANT R IOS RI OOO ieee seek tapas. ccepe eae eee eae RTA ee vada 3 3. Sternite 8 with margins of posterior emargination meeting anteriorly at obtuse angle Coie! TN ce niindl ee nde ee eh oi iy Ae De GAA a SRE er eae ce SP Ewe Sn Rete aaenna ted ee ake eh tales burksi Sternite 8 with margins of posterior emargination meeting anteriorly at acute angle (Figs. 85, 89), usually strongly sclerotized near hind margin.................... metequi, nivalis Nymphs: (Poulton and Stewart 1991). i eciamoorsal stripe absent or incomplete (Fig 593) ici 0h cc. esicessdeescetoonstnsesenccasebecsensddentésceecenses 2 meter esc ati Pe eCOMIIC(C (PIGS. O.t0ms. 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