Great Basin Naturalist

Volume 59 Number 1 Article 1

1-22-1999

Stoneflies () of the Black Hills of South Dakota and Wyoming, USA: distribution and zoogeographic affinities

Bret O. Huntsman Brigham Young University

Richard W. Baumann Brigham Young University

Boris C. Kondratieff Colorado State University, Fort Collins

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Recommended Citation Huntsman, Bret O.; Baumann, Richard W.; and Kondratieff, Boris C. (1999) "Stoneflies (Plecoptera) of the Black Hills of South Dakota and Wyoming, USA: distribution and zoogeographic affinities," Great Basin Naturalist: Vol. 59 : No. 1 , Article 1. Available at: https://scholarsarchive.byu.edu/gbn/vol59/iss1/1

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. The Great Basin Naturalist PuBUSHED AT PROVO, UTAH, BY M.L. BEAN LIFE SCIENCE MUSEUM BRIGHAM You C UNlVERSm'

ISS 0017-3614

VOLUME 59 30 JANUARY 1999 No.1

Great Basin Naturalist 59(l),ltll999. pp. 1-17

STONEFLIES (PLECOPTERA) OF THE BLACK HILLS OF SOUTH DAKOTA AND WYOMING, USA: DISTRIBUTION AND ZOOGEOGRAPHIC AFFINITIES

Bret O. Huntsman1, Richard ,v. Baumann1.2, and Boris C. Kondratieff3

ABSTRACT.-The Black Hills of South Dakota and Wyoming are an insular mountain range completely surrounded by the Great Plains. The stonefly (Plecoptera) fauna of the Black Hilts was surveyed and zoogeographic affinities exam­ ined. Twenty-seven species representing 22 genera and 6 families were found. Fifteen new state records for South Dakota and 2 for Wyoming are presented. Two species are removed from the South Dakota list. An analysis ofthe North American distribution of each species showed a strong relationship between the Black Hills and the Rocky Mountains, with much weaker relationships between the Black Hills and eastern and northern regions. Results ofa logistic regres­ sion analysis comparing factors contributing to long-distance dispersal ability against presence/absence in the Black Hills were inconclusive. However, other evidence suggests that the Black Hills fauna is a result ofexpansion and subse­ quent vicariance ofstonefly populations during Pleistocene climatic oscillations.

Key words: stonefl~, Plecoplera, zoogeography, B1llck Hills, South Dakot

The Black Hills are an island of mountains Melntosh 1931, Van Bmggen 1985). Mammal in a sea of grass (Tumer 1974). These isolated species of the Black Hills show similar mixed mountains, located in western South Dakota affinities. Approximately 14.3% are Rocky and eastern Wyoming, are completely sur­ Mountain, 8.6% are from eastern deciduous rounded by the Great Plains physiographic forests, 14.3% are from northern boreal forests, region. The Black Hills contain one ofthe most and the rest are a mix of widespread, steppe, unusual mixes of biotic affinities in orth and Sonoran species (Turner 1974). in America. For example, 30% ofthe plant species the Black Hills also appear to have mixed found in the Black Hills have Rocky Mountain affinities. For instance, the mayfly (Ephemer­ affinities, 9% have eastern deciduous forest optera) fauna of the Black Hills has western affinities, 6% have northern boreal forest affini­ species (36.8% of the fauna), eastern species ties, while the remainder are a mix: of steppe (21.1%), and widespread species (42.1%) and widespread species (Hayward 1928, (McCafferty 1990).

lDepartmeot ofZooIoP.', Brigbaro Young Un.ivecity; Provo. lIT 54600. :.\l,It/wr to whom reprint reque$l$ $bould be sent. 3Department of BioogrieultuOlI Scie.ooes and F'e$1 ;\laDagement. Color.ido Stile UnMmty. Fori

1 2 GREAT BASIN NATURALIST [Volume 59

Another interesting aspect of Black Hills populations within the Black Hills. At the pre­ biogeography is the fact that many populations sent time no quantitative studies have been of plants and inhabiting the area are performed comparing vicariance and dispersal highly disjunct, being separated from the near­ models for the Black Hills fauna. est conspecific populations by as much as 600 Faunal affinities and distribution histories k-m (Turner 1974, Van Bruggen 1985, McCaf­ are most easily studied using organisms re­ ferty 1990). These disjunct populations also stricted to definable habitats (Sargent et al. often represent the extreme range limit for 1991, Houseman and Baumann 1997). One their species (Turner 1974, Van Bruggen 1985, outstanding example of this type of organism McCafferty 1990). is the Order Plecoptera (stonellies). The most common hypothesis explaining Most stonelly species require cold, clean these striking biogeographic patterns is that streams with rocky streambeds to complete these disjunct populations are vicariated relicts their life cycles (Surdick and Gaufin 1978, from the most recent period of glaciation Ward 1984, Stewart and Stark 1988). Adult (Ross 1965, Turner 1974, McCafferty 1990). stonellies are not powerful lliers and usually Palynological studies indicate that during the are not capable of long-distance migrations Full-glacial period of the Wisconsin glaciation (Ross 1965, Hynes 1988, Marden and Kramer (30,000-13,000 years BP) much of the central 1994, Griffith et aJ. 1998). Thus, dispersal of portion of the United States was covered with most stonelly species is thought to require boreal forest (Wright 1970), allowing typical adequate water connections (Surdick and boreal animals to inhabit this region. This Gaufin 1978, Baumann 1979, Flanagan and cooler, wetter climate also allowed deciduous Flanagan 1982). Many species have restricted gallery forests to form along stream banks, ranges, and distinct differences exist between prOViding avenues of expansion for eastern the stonelly faunas of western, eastern, and deciduous forest species into the Black Hills northern North America, primarily at the (Turner 1974). Later, during the Late-glacial generic and/or specific levels (Rickel' 1964, period (13,000-10,500 BP), the boreal forest Dosdall and Lemkuhl1979, Stark et aJ. 1986). followed the retreating ice sheet and was Few published records are available on the replaced by steppe habitat in the Great Plains stonellies of the Black Hills. Presently, only 14 (Holfman and Jones 1970). Eastern and north­ species have been reported from the Black Hills ern species that had expanded into the Black region (Stark and Gaufm 1976, Baumann et al. Hills during the FUll-glacial period were then 1977, Stark et al. 1986). Several families, such isolated in the refuge formed there by the as the , Pteronarcyidae, Taenioptery­ remnant deciduous and boreal forests (Turner gidae, and Leuctridae, have not been previ­ 1974). ously reported from the Black Hills. Unfortu­ Also during the.Full-glacial period of the nately, many of the streams within the Black Wisconsin the cordilleran-montane zone of Hills have been affected by pollution and the Rocky Mountains was depressed about other perturbations (Stewart and Thilenius 1200-1400 m. This displaced the biota of this 1964). It is poSSible that some populations of wne eastward, extending from the Laramie stonellies within the Black Hills have already and Big Horn Mountains ofWyoming into the been extirpated. In the face of the current Black Hills (Turner 1974). Again, when the ice challenge to document North American biodi­ sheet retreated, the montane species that had versity (Kosztarab and Schaefer 1990), inven­ expanded into the Black Hills were isolated in torying the stonelly fauna of the Black Hills the refuge formed there (Thrner 1974). should be a priority. . Although the hypothesis of Ross (1965), Our study had 3 main objectives: to docu­ Tnrner (1974), and McCafferty (1990) has great ment the stonelly fauna of the Black Hills, to explanatory power, it is not the only possibil­ detemune ifthe stonellies ofthis region rellect ity. It is also possible tbat tbe Black Hills pop­ the mixed affmities shown by other groups of ulations of plants and animals were established organisms, and to quantitatively test the vicari­ via long-distance dispersal. Sometime after the ance hypothesis of Ross (1965), Turner (1974), formation of the Great Plains, various species and McCafferty (1990) using these new stone­ may have dispersed across it and founded lIy distributional data. 1999] STONEFLIES OF THE BLACK HILLS 3

STUDY AREA Adult stoncflies were collected nsing beat­ ing sheets or aerial nets, or by hand. Nymphal The Black HilIs region consists of the mack stondlies were collected with aquatic dip Hills proper and the Bear Lodge Mountains in nets. At some sites nymphs of selected species \vestern South Dakota and northeastern were set aside and reared to adults to assist in Wyoming (approximately 43'10'-44'50' N lat., specific identification. All adults (and the 103'20'-104'50' W long.; Tnrner 1974). The nymphs of certain species) were identified Black Hills were created by intermittent domal using keys in Baumann et al. (1977) and Stew­ uplifts during the Cretaceous, Miocene, and art and Stark (1988). Pleistocene (Turner 1974). Elevations in tbe In addition to field-collected specimens, we Black Hills region range from 1000 m on the examined specimens from the following insti­ western plains to 2228 m at Harney Peak. The tutions: Monte L. Bean Life Science Museum, area is completely surrounded by the North­ Brigham Young University, Provo, Utah (BYUC); ern Great Plains physiographic province, which C.P. Gillette Museum of Diversity, consists of arid to semiarid short grass prairie Colorado State University. Fort Collins (CSUC); (Fig. 1). The Black Hills are located approxi­ Illinois Natural History Survey, Champaign mately 240 km east and northeast of the Big (INHS); North Dakota State University Research Horn and Laramie ranges of the Rocky Moun­ Collection, Fargo (NDSU); Purdue Entomo­ tains, respectively. The nearest component of logical Research Collection, Purdue Univer­ northern boreal forest is in Saskatchewan, sity, West Lafayette, Indiana (PURC); Sonth approximately 600 km to the north. The near­ Dakota State University Research Collection, est component of eastern deciduous forest is Brookings (SDSU); United States National Mu­ in Minnesota, approximately 600 km to the seum, Smithsonian Institution, "'ashington, east, althongh gallery forests exist along the D.C. (USNM); and University of Nebraska Re­ Missouri River, approximately 200 km to the search Collection, Lincoln (UNSM). Specimens east. The Black Hills are drained by the collected during the course of this study are Cheyenne and Belle Fourche rivers, which deposited in the Monte L. Bean Life Science circle around the southern and northern ends Museum and C.P. Gillette Museum of Arthro­ of the mack Hills (Fig. 1). Since these rivers pod Diversity. originate in the plains of eastern Wyoming, The North American distribution of each there are no direct water connections between species collected in the Black I-Iills was exam­ the Rocky Monntains and the Black Hills. The ineel to determine its region of faunal affinity Belle Fourche drains into the Cheyenne (Baumann et al. 1977, Stark et a1. 1986, Stark approximately 80 km east of the Black Hills 1998). This allowcd us to determine probable (Fig. 2), which in turn drains into the Missouri source pools for species inhabiting the Black River. Hills. The stonefly faunas of Colorado (Stark et al. 1973, Baumann et al. 1977, Stark et al. METHODS 1986), Minnesota (Harden and Mickel 1952, Stark et al. 1986), and Saskatchewan (Dosdall We made collections at 112 sites within the and Lemkuhl1979, Stark et al. 1986) were ini­ Black Hills region. These collection sites were tially selected to represent western, eastern, chosen to provide coverage of all major drain­ and northern source pools for purposes of a ages and most major creeks and streams with­ statistical test of the long-distance dispersal in the Black Hills. Although not considered model. These areas were chosen hoth for their part ofthe Black Hills proper, the Belle Fourche proximity to the Black Hills and availability of and Cheyenne rivers were included in our col­ published records (Baumann et al. 1977, Dos­ lections because stonefly species have direct dall and Lemkuhl 1979, Harden and Mickel water connections to the Black Hills via these 1952, Stark et al. 1986). drainage systems. Collections were made in If the Black Hills were colonized by ran­ February and June 1995, April and August dom, long-distance dispersal flights across the 1996, and July 1997 to include species emerg­ Great Plains, one would expect those stonefly ing in the winter, spring, and summer. All speci­ species with greater dispersal abilities tu be mens collected were preserved in 70% ethyl over-represented in the Black Hills fauna alcohol. (Houseman and Baumann 1997). We used a 4 GREAT BASIN NATURALIST [Volume 59 logistic regression procedure (PROC LOGIS­ TIC; SAS 1990) to dete.mine ifdispersal abil­ MONTANA GREAT PLAINS ity would predict the occurrence of stonelly WYOMING species in the Black Hills muna. A significant positive relationship between dispersal ability and presence in the Black Hills would provide Belle Fourche River evidence that long-distance dispersal is a viable alternative to vicariance as a mode of colonization. Flying insects can disperse long distances SLACK Spemf"h either by active, powered flight (adaptive dis­ HILLS Creek persal) or by being accidentally caught up in convective upcurrents, which deliver them to the stronger winds present at higher altitudes (inadvertent dispersal; Johnson 1969, Drake and Farrow 1988). For purposes ofthis test we assumed that either type of dispersal could occur. The dispersal ability of all stonelly species occurring in the Black Hills and the hypothesized source pools were quantified /'-t'~,­ based on the follOwing factors: season ofemer­ CheyenneRjve'~''''''''/ gence, length of emergence, ecological toler­ ance, and overall distribution in the source range (adapted from Houseman and Baumann SOUTH DAKOTA 1997). However, due to the low number of NEBRASKA eastern and northern species inhabiting the Black Hills (see Results section below), only western species (represented by the Colorado Fig. 1. The Black Hills of South Dakota and Wyoming. fauna) were tested to determine if dispersal ability predicted presence in the Black Hills. Each species was given a score for each factor longer emergences for 2 reasons. First, adults based on data published in Baumann et aJ. are more likely to be present during extended (1977) and Baumann (1979). These scores periods ofcalm weather, which is conducive to were determined using the follOwing criteria. adaptive dispersal (Johnson 1969). Second, a SEASON OF EMERGENCE.-Muscle power longer period of emergence would increase output of insects is severely resbicted at low the probability of inadvertent dispersal via temperatures (Josephson 1981, Marden and convective upcurrent events. Species with Kramer 1994). Stoneflies are no exception. adults present for 3 months or less in the Even winter-emerging species are sluggish source pool were scored 1, those with adults and rarely lIy at temperatures near or below present 4-5 months were scored 2, and those freezing (Frison 1929, Marden and Kramer with adults present for more than 5 months 1994). In addition, during periods of extreme were scored 3. cold, winter stonellies seek shelter under rocks, ECOLOGICAL TOLERANCE.-Baumann (1979) bark, and leaves (Frison 1929) and are unlikely identified 3 major stonelly environments (cold to be engaged in dispersal activities. Therefore, lotic, warm lotic, and cold lentic) and the gen­ we assumed species that emerge in colder era which inhabit them. All species in this months are less likely to successfully disperse. analysis primarily inhabit cold lotic habitats. We scored species with September to March However, those that can tolerate warm lotic emergence as 1, species that emerge from habitals would be more likely to survive in April to June as 2, and those that emerge from intervening areas of the Great Plains during July to August as 3. dispersal, since cold habitats are rare in the LENGTH OF EMERGENCE.-We assumed prairies. In addition, warm-tolerant species that dispersal is more likely for species with could more easily colonize the Black Hills upon 1999] STONEFLIES OF THE BLACK HILLS 5

MONTANA

WYOMING SOUTH DAKOTA

Butte Co. BELLE FOURCHE RIVER Meade Co.

SPEAR- FISH • Crook Co. CREEK Sturgis

- - ~~~~!...!:;.i=::;EL~K~C~RE~E:K::::=:Lawrence Co. BEAVER CREEK "",-.r,,_~-:.~Rapid City RAPID CREEK Pennington Co.

Custer Weston Co. •

Cuter eO.

--...... _Hot Springs• Shannon Co. CHEYENNE RIVER

Niobrara Co. Fall River Co.

NEBRASKA

Fig. 2. Counties, major cities, and major drainages in the Black Hills region of South Dakota and Wyoming.

arrival, since more habitats would be available RESULTS for them. Species limited to cold lotic environ­ ments were scored 1, while those known from We collected 27 species of stoneflies repre­ both cold and warm lotic habitats were scored senting 22 genera and 6 families in the Black 2 (we omitted cold lentic habitats from our Hills region. They are listed in Table 1 by analysis since few species are known from state and county in which they were collected. them and few of these habitats exist in the Fifteen of these species represent new state Great Plains). records for South Dakota, while 2 represent DISTRIBUTION WITHIN THE SOURCE POOL.­ new state records for Wyoming (Table 1). We assumed that species with wider distribu­ The winter stonefly genus Isocapnia was tions within the source pool possess higher represented in our study by a single female, vagility and thus have a higher probability of designated Isocapnia sp. A, collected in 1982. performing adaptive dispersal flights. Also, This brachypterous female may represent an those species with wider dishibutions are more undescribed species. likely to be picked up by local, random con­ The zoogeographic affinities of the stonefly vective currents and dispersed by high­ species in the Black Hills fall into 4 major altitude winds. The number ofColorado coun­ categories: western (21 species), eastern (2 ties in which each species occurs was used to species), northern (1 species), and widespread represent overall distribution (range 1-22). (3 species; Table 2). Western species are defined 6 GREAT BASIN NATURAlJST [Volume 59

TABLE 1. Stonefly species ofthe Blacl: Hills by slate and county.

South Dakota Wyoming

Species Butte Custer Fall RiveT Lawrence Meade Pennington Crook Weston

Amphine1mlra b(J/lksi X X X XX Malellka colomdensis X X X X Nelrwum trispinosut:,t X X X X ProsioUJ hesametsa* X X X X X Zapada cinctipes X X X X XX Capnia confusa* X X X Capnia gracilariaol< X X X Eucapnopsis bl'evicauda* X X X lsocapnia sp. A'" X ParcJ.capnia angulata '" XXX X Utaculmia lemonianu* X X X Para1euctra L"'ersrunu'" X X XX isogenoides elongatus· X X Isoperla lonnt'liselo X X X {sorlcrla phalerato X X X X X X X ISoperlll quinquepunctata X X X XX X X Isoper-Itl tmmmat'ino"',t X X Skwa](/ umericUfW· X X X X Acruneuria abrwnnis' X X Cwussenia subulosu X f1esperoper/a pacific" X X X XX Perlesta decipiens* X X Pamperla frontalis X X X Suwalua linco.m* X Sweltsa. borealis X X Swellsa colnradensis X X X TrizHaka pintada X X X XX

~inditat(l~ oow South Dakota ~t;lt(l record tiIlJkc-lt'" ,,"W \\'y",nin~ .tate l'ecoo-d as those whose region of greatest abundam..-c Rocky Mountains, Baumann et a1. (1977) treated falls west or northwest of the Great Plains, the Black Hills as an exteusion of tl,e Rockies. while ea.fiitem species are those most abundant They listed 1I spe<.'ies ofstoneflies in the Black east of the Great Plains. Northern species are Hills. However, they neglected C. sabulos". those occurring primarily in the previously Stark et a1. (1986) listed 14 species from South glaciated areas of Canada and the Great Lakes Dakota. In addition to tl,e 12 species listed by region. Finally, widespread species are those previous authors, Stark et a1. added Isoperla with a largely east-west transcontinental dis­ kmgiseta Banks and Triznaka pintada (Ricker). tribution. It is interesting to note that no Botll Baumann et a1. (1977) and Stark et al. southern species or endemic species were col­ (1986) list Nemoum arctica Esben-Peterson as lected in the Black Hills. occurring in the Black Hills. These records Logistic regression analysis of factors were based on a single female collected in affecting dispersal ability found 2 factors, 1953 and 2 females collected in 1965. Our length of emergence (X21 = 4.28, P = 0.038) subsequent efforts failed to locate any speci­ and distribution in the source pool (x2, = mens of N. arctica, although a congener, N. 6.71, P = 0.010), to be significant predictors of trispinosa Claassen, proved to be common. The l'resence in the Black Hills. 3 females previously identified as N. arctic" are actually N. trispinosa. There appears to he DISCUSSION no evidence that N. arcUca occurs within the Black Hilfs. In addition, both Baumann et al. In his catalog ofthc world Plecoptera fauna, (1977) and Stark et a1. (1986) list T signttta IlIies (1966) listed 3 species from the Black (Banks) from the Black Hills region. Several Hills region. Stark and Gaufin (1976) reported vials of specimens from the Illinois Natural sabulosa (Banks) from Lawrence History Survey containing T pintada from the County. In their review of the stone£lies of the Black Hills were inmrrectly identi6ed as T 1999] STONEFLIES OF THE BLACK HILLS 7

TABLE 2. Geographic affinities ofthe stonefly species ofthe Black Hills.

WESTERN SPECIES Amphinemura barzksi Isocapnia sp. A ClaMsenia sabulosa Ma1enka coloradensis Utacapnia lemoniana Hesperoperla pacifica Prostoia bcsametsa Paraleuctra vershina Paraperla frontalis Zapada cinctipcs Isogenoides elongatus Suwallia Uneasa Capnia con~a Isoperla phalerata Sweltsa borealis Capnia gracilaria Isoperla quinquepunctata Sweltsa coloradensis Eueapnopsis brevicauda Skwala americana Triznaka pintada

WIDESPREAD SPECIES Isoperlalongiseta Acroneuria ahnormis Perlesta decipiens

EASTERN SPECIES angulata Isoperla transmarina

NORTHERN SPECIES Nemoura trispinosa

signata. It is probable that these samples would be expected in a vicariance model ifthe represented the errant records reported by habitat connections before the vicariance event these autbors. were of short duration or poor quality, since Analysis offactors affecting dispersal ability only the most vagile species would then be showed 2 factors, length of emergence and expected to reach the Black Hills. Thus, the distribution in the source pool, to be signifi­ question of mode of origin for the Black Hills cantly related to presence/absence in the Black fauna was not resolved by this analysis. Hills. Upon further analysis, however, it appears A vicariarice explanation for the origin of the significant result for length of emergence the Black Hills stonefly fauna is supported by is due entirely to low numbers of capniid the similarity of its distributional pattern (Table species in the Black Hills. Only 6 species of 2) to that of mammals, plants, and mayflies capniids have been found in the Black Hills, (Turner 1974, Van Bruggen 1985, McCafferty compared to 20 species in Colorado. In addi­ 1990). It is unlikely that 4 separate groups, tion, one of the Black Hills species, Isocapnia each with very different dispersal modes and sp. A, could not be included in the original abilities, would have similar distributional pat­ analysis due to lack of specific identification. terns originating independently via long­ Therefore, only 5 capniid species from the distance dispersals. The most parsimonious Black Hills were included in the original explanation is that the entire biota was iso­ analysis. Since most species in the family Cap­ lated at the same time by a vicariance event, niidae have short emergences (Baumann et aI. most likely by the formation ofthe Great Plains 1977), a paucity of capniids in the Black Hills at the end ofthe Pleistocene. could bias the results. To determine if this Although the Black Hills stonefly fauna ex­ happened, we repeated the analysis with the hibits a pattern of mixed zoogeographic affini­ species in the family Capniidae removed from ties similar to other groups, there are some the data set. No significant relationship be­ differences. The percentage of eastern species tween emergence length and presence in the of stoneflies (7.4%) agrees well with those of Black Hills was found among the species of plants, mammals, and mayflies (range 8.6­ the 5 remaining families (x2[ = 1.42, P = 0.23). 21.1%), while the percentage of northern stone­ There was a significant relationship between flies (3.7%) resembles that of other groups distribution in the source pool and presence (range 0-14.3%; Turner 1974, Van Bruggen in the Black Hills even after capniids were 1985, McCafferty 1990). However, the percent­ removed from the analysis (X2, = 5.36, P = age of western species of stoneflies (77.8%) is 0.021). This positive result provides some evi­ far higher than the percentage of western dence that long-distance dispersal may have species of other groups (range 14.3-31.6%; been a factor in the origin of tbe Black Hills Turner 1974, Van Bruggen 1985, McCafferty fauna, although this evidence is far from con­ 1990). Also, only 14.8% of the Black Hills clusive. It is possible to argue that this result stonefly fauna consists of either prairie or 8 GREAT BASIN NATURALIST [Volume 59 widespread species, compared to 42.1~2.8% ranges within the region. Some species, such as for other groups (Turner 1974, Van Bruggen lsoperla trans-marina ewman, Isogenoi.des 1985, McCafferty 1990). There are few species elonga/us (Hagen), lsucapnia sp. A, and Suwal­ of stoneflies able to tolerate streams typical of lia lineosa (Banks), were found in only 1 or 2 the prairies (Ricker 1946, 1964). This explains streams, suggesting relatively limited popula­ the paucity of prairie or widespread stonefly tions in the Black Hills. This makes them sus­ species in the Black Hills. The large number ceptible to stochastic extinction events. of western species, however, is more difficult Of the groups examined in the Black Hills, to interpret. There are roughly 7 times more only 1, the plants, contains an endemic species western species than eastern and northern (Van Bruggen 1985). This single endemic species in the Black Hills. This suggests that species of plant is a hybrid, meaning it could direct water connections between the Rocky have arisen very recently. This lack ofendemics Mountains and the Black Hills during the is unusual for an area as large and isolated as Wisconsin glaciation were mOTe extensive than the Black Hills. In comparison, the Ozarks and mnnections to the east. This agrees with Pleis­ Ouachita Mountains of the south nentral U.S. tocene glacial drainage patterns. During the eontain 25 endemic species of stonellies, fully Pleistocene, the general drainage pattern of 28% of the stonefly fauna of that region (Poul­ the Great Plains consisted of parallel rivers ton and Stewart 1991). Lack of endemics in running from the Rud,,, Mountains to near the the Black Hills indicates a young fauna, which glacial boundary (Cmss et al. 1986). Here these lends credence to the assumption that the rivers were deflected to the south by the lau­ Black Hills have been isolated ecologically rentide ice sheet, converging to form a large only since the latest period ofglaciation. river which flowed southeast to Kansas and Due to the isolated location of the Black Oklahoma (Cross et al. 1986). While Rocky Hills, most populations ofstoneflies in the area Mountain species had a relatively direct path represent the extreme range limit for their across the Great Plains to the Black Hills, respective species (15 of 21 western species, 1 eastern species had to travel the longer route of 2 eastern species, 1 of 1 northern species). around the southern tip of the ice lobe to Also, because of the Black Hills location, reach the Black Hills. However, this is not the almost all stonefly populations in the area are only interpretation. The disparity in the ratio highly disjunct, in some cases being separated ofeastern and western species may also reflect by over 600 km from their nearest known con­ the fact that the prevailing winds in the Black specillc population. This same pattern is exhi­ Hills are from the west (Orr 1975), and thus bited by mammals, plants, and mayflies ill this adaptive and inadvertent dispersaillights from area (Tumer 1974, Van Bruggen 1985, McCaf­ the west are more probable. ferty 1990). The Black Hills is one of the very A total of 27 species of stoneflies is rela­ few places in which the eastern, westenl, and lively low for an area as large as the Black northern biota overlapped during the Wiscon­ Hills. In compmison, 37 species of stoneOies sin glaciation, and which also contained the have been reported from the Raft River Moun­ elevation necessary to provide a refuge from tains of Utah, which cover an area onJy one~ the encroaching steppe for cold-adapted organ­ tenth as large (Houseman and Baumann 1997). isms like stoneflies. McCafferty (1990) found the diversity of may­ One of the most interesting distributional flies (Ephemeroptera) in the Black Hills to be patterns in the Black Hills was exhibited by low as well. This may reflect a low rate ofcolo­ the 3 species with widespread distributions: nization of the Black Hills by aquatic insects. Acroneuria abnorrnis (Newman), Perlesta dedp­ However, activities hy man also may have re­ iens (Walsh), and lsoperla longiseta. These 3 duced the diversity of the area. As previously species were found only in the Belle Fourche mentioned, most streams in the Black Hills and Cheyenne rivers. As was stated earlier, have been heavily inlpacted by reduned 1I0ws, these rivers are not in the Black Hills proper, mine drainage, grazing, and agriculture (Stew­ but rather surround and drain the area. These art and Thilenius 1964). Thus, it is possible 3 species are not montane species and seemed that some populations ofaquatic insects native unable to invade the streams of the Black Hills to the area have been extirpated. Many stonelly and, other than a single collection of Isoperla species in the Black Hills have very restricted quinquepuru;tal

280 17~ 30N, 9-VII-I997, 30 6~. Lawrence Co.: 23-IV-1996, 130 7~, 13-VII-1997, 19; trihutaryof Iron Creek, 7-VI-199S, 10, 13-VII-1997, H; Little Bear Butte Creek, 22-IV-1996, 9N; Wbitetail Spearfish Creek, 21-V-19S1, 30; South Fork Rapid Creek, IO-I1I-19S1, 2N; Whitewood Creek, IS-I1I­ Creek, 12-VII-1997, 10; Spearfish Creek, 20-V­ 19S1, 3N, 20-V-19S1, 10 149, 25-1II-19S2, 4N. Pen­ 19S1, 7N, 9-XII-19S1, 4N, 7-VI-199S, 160 3~, 13­ nington Co.: Battle Creek, S-II-1995, ION; Beaver VII-1997, 1 ~ ; tributary of Bear Butte Creek, 22-iv­ Creek, 16-VII-1997, 1 ~; Grizzly Bear Creek, 20­ 1996,4N, 12-VII-1997, 30 7~; tributary of White­ IV-1996, ION; creek in Sunday Gulch, S-VI-199S, wood Creek, 12-VII-1997, 10 2 ~; Whitewood 10 6~, 9-VII-1997, 10 19. Creek, 7-VI-199S, So S~. Pennington Co.: Elkhorn WYOMING DISTRIBUTlON.-Crook Co.: Beaver Spring, S-VI-199S, 140 23 ~ , 22-VIII-1996, 3 ~ . Creek, 24-IV-1996, 100 1 ~; North Redwater Creek, WYOMING DISTRIBUTION,-Crook Co.: Blacktail 24-IV-1996, 3N; Whitelaw Creek, 16-VII-1997, 3 ~ . Creek, 16-VII-1997, 10 1 ~. Weston Co.: Beaver Creek, 23-IV-1996, 6N. DISCUSSION.-This species is most common in DISCUSSION.-This species occurs throughout the southern portion of the Rocky Mountains (Bau­ the Rocky, Cascade, Coast, and Sierra Nevada mann et aI. 1977). The Black Hills population rep­ Mountains (Baumann et al. 1977). The Black Hills resents the northeast range limit for this species. In population represents the easternmost range limit the Black Hills M. coloradensis was collected in for this species. habitats ranging from small springs to larger creeks. Genus Zapada Genus Nenwura Zapaaa cinctipes (Banks) Nemoura trispinosa Claassen SOUTH DAKOTA DISTRIBUTION.-Custer Co.: Iron 220 19~; SOUTH DAKOTA DISTRIBUTION.-Custer Co.: Syl­ Creek, 20-IV-I996, stream entering Syl­ ~, van Lake, IS-VI-19S3, 1 ~ (USNM). Lawrence Co.: van Lake, S-VI-199S, 1 9-VII-1997, H; stream Englewood (probably Whitewood Creek), 17-VI­ 10 mi S of Sylvan Lake on Needles Hwy, 2S-IlI­ 1936, I ~ (SDSU); Spearfish Creek, 28-VI-196S, 2 ~ 1968, 10 I? (INHS); Willow Creek, 19-IV-1996, 4~, 1~. (USNM), 27-VI-I99S, S20 21 ~; stream in Black 40 4-VI-199S, Lawrence Co.: Bear Butte Hills Experimental Forest, 14-VI to 26-VIII-199S, Creek, 22-IV-1996, 40 39; Boxelder Creek, 24-1II­ 1965, 10 (INHS), 6-11-199S, 2N, 22-IV-1996, 30 2So IS ~ (SDSU); tributary of Bear Butte Creek, 14~ 12-VII-1997, 10 19; tributary ofWhitewood Creek, 39; East Spearfish Creek, 20-V-19S1, ISo IN, 7-II-I09S, 12N, 7-VI-190S, 100 14?, 23-IV-I996, 12-VII-1997, 19; Whitewood Creek, 17-VI-1936, 130 13 ~ , 25-VIII-I096, SN; Elk Creek, 25-I1I-1965, 19, 7-VI-199S, 10 1 ~. Pennington Co.: Burnt Fork, 19 (INHS), 22-IV-I996, 90 39, 2S-VIII-1996, 4N; 6-VI-199S, 130 269, 11-VII-I997, 40 99. Estes Creek, 24-111-196S, 240 9~ (INHS); False WYOMING DlSTRlBUTION.-Crook Co.: Blacktail Bottom Creek, 24-I1I-196S, 20 (INHS); Hay Creek, 16-VII-1997, 40 7~; Whitelaw Creek, 16­ Creek, 22-IV-I996, 10 IN, 25-VIII-1996, SN; Iron VII-1997,Io. Creek, 7-11-199S, SO* 7~* 20N, 7-VI-199S, 30 DISCUSSION.-This species is known from Labra­ 1 ~, 23-IV-1996, 140 9~, 26-VIII-1996, 2N; Jim dor west to Manitoba and south to Illinois (Stark et Creek, 6-11-199S, 20* 1 ~ *; Little Elk Creek, 22­ al. 19S6). The Black Hills represent the western­ IV-1996, 29; Little Spearfisb Creek, 17-I1I-19S1, most range limit for this species, In the Black Hills 150 21 ~ 20N, 21-V-19S1, 7~ (INHS), IS-X-1994, N. trispinosa was associated with spring-influenced IN (SDSU), 27-IV-199S, 30 S?, 7-VI-199S, 2 ~, systems. 23-IV-1996, 100 II ~, 26-VIII-1996, 2N, 13-VII­ 1997,19; North Boxelder Creek, 22-IV-1996, 20 Genus Prostoia I ~ ; creek SE of Pluma [probably West Strawberry Prostoia besametsa (Ricker) Creek], 24-III-196S, 1 ~ IN (INHS); South Fork Boxelder Creek, 22-IV-I996, 100 10~; South Fork SOUTH DAKOTA DISTRlBUTlON,-Custer Co.: Rapid Creek, 22-V-19S1, 10 1 ~; Spearfish Creek, Custer State Park, 17-VI-19SS, 10 (SDSU); Iron 27-VII-1940, 2N (INHS), 2S-I1I-196S, 270 36~ Creek, 20-IV-1996, 100 8 ~ 12N, 10-VII-1997, 19. (INHS), 19-I1I-19S1, 70 19 70N, 20-V-19S1, So Lawrence Co.: Boxelder Creek, 22-IV-1996, 1 ~ 2N; 169, 6-X-1981, 20N, 9-XII-19S1, 20N, 25-I1I-19S2, East Spearfisb Creek, 20-V-19S1, 1 ~, 7-VI-199S, S30 40~, 7-11-199S, 25N, 7-VI-199S, 140 179,23­ 4~, 23-IV-1996, 30 6N; Hay Creek, 22-IV-1996, IV-1996, S40 4S ~, 26-VIII-1996, 13N, 13-VII-1997, loIN; Iron Creek, 23-IV-1996, ISo 6?; Little 1 ~; Strawberry Creek [probably West Strawberry Spearfish Creek, 17-I1I-19S1, 19N, 21-V-19S1, 40 Creek], 10-XII-19S1, 20N, 23-I1I-19S2, 1 ~ 3N; 69, 27-IV-199S, 3~ (SDSU), 23-IV-1996, 30 I?; tributary of Bear Butte Creek, 22-IV-1996, 90119, South Fork Boxelder Creek, 22-IV-I996, 30 6N; 24-VIII-1996, 4N; Whitetail Creek, 19-111-19S1, South Fork Rapid Creek, 22-V-19S1, 2 ~: Spearfish 30 S~ 20N, 20-V-19S1, 10 12~, 16-VII-19S1, IN, Creek, 2S-I1I-1968, IN (INHS), 10-I1I-19S1, 47N, S-X-19S1, SN, 9-XII-19S1, 3N; Whitewood Creek, 20-V-19SI, 10 4 ~, 25-I1I-19S2, IN, 7-1I-199S, 2N, IS-I1I-19S1, 140 79 26N,20-V-19S1,So 102~, 15- 1999] STONEFLIES OF THE BLACK HILLS 11

VII-1981, 9N, 7-X-1981, 37N, 9-XIl-1981, 3IN, 25­ Capnia gracilaria Claassen 111-1982, 180 23~ 35N, 7-II-1995, 4N. Meade Co..' Elk Creek, 22-IV-1996, 2 ~; Meadow Creek, 22-IV­ SOUTH DAKOTA DISTRIBUTION.-Cusfer Co.: Iron 1996, 150 7~. Pennington Co.: Battle Creek, 8-11­ Creek, 20-IV-1996, 20 6~; Willow Creek, 19-IV­ 1995, 10* 5N; Beaver Creek, 23-Vlll-1996, 7N; 1996, 30 1 ~. Lawrence Co.: East Spearfish Creek, Burnt Fork, 6-VI-1995, 4~, 21-1V-1996, 110 7~; 20-V-1981, 1 ~, 7-VI-1995, 1 ~,23-IV-1996, 20 1 ~; Castle Creek, 9-II-1995, 19N, 21-IV-1996, 240 13~; Elk Creek, 25-111-1968, 1 ~ (INHS); Iron Creek, 7­ Deer Creek, 21-IV-1996, 10 1 ~; Ditch Creek, 23­ II-1995, 20*, 23-1V-1996, 200 1O~; Little Spear­ Vlll-1996, IN; Grizzly Bear Creek, 20-IV-1996, 2 ~ ; fish Creek, 17-IIl-1981, 1~, 23-IV-1996, 70 1~; Middle Fork Boxelder Creek, 6-VI-1995, 2 ~. 22-IV­ Spearfish Creek, 25-UI-1968, 300 20~ (INHS), 1996, 150 10~; Newton Fork, 9-II-1995, 6N, 6-VI­ 19-IIl-1981, 70 1~, 20-V-1981, 8~, 25-IIl-1982, 1995, 1 ~, 21-1V-1996, 50 3 ~; North Fork Castle 400 9N, 7-VI-1995, 4~, 23-IV-1996, 190 6~; trib­ Creek, 21-1V-1996, 20, 24-Vlll-1996, IN; Palmer utary of Bear Butte Creek, 22-1V-1996, 220 21 ~. Creek, 8-II-1995, 7N, 20-1V-1996, 10 8~; Pine Pennington Co.: Burnt Fork, 21-IV-1996, 220 26 ~: Creek, 20-IV-1996, 20 1 ~; Rapid Creek, 24-111­ Deer Creek, 21-IV-1996, 3 ~; Elkhorn Spring, 5-VI­ 1968, 240 15~, 6-II-1995, 20* 13N, 6-VI-1995, 1995, 10 12~, 20-1V-1996, 440 35~; Pine Creek, 10 2~, 21-IV-1996, 60 6~: Slate Creek, 21-IV­ 5-VI-1995, 1 ~, 20-1V-1996, 60 7~; Spokane Creek, 1996, 90 1O~; South Fork Castle Creek, 21-1V­ 20-IV-1996, 20 1 ~; creek in Sunday Gulch, 19-IV­ ~: ~, 1996,140 n; Spokane Creek, 20-IV-1996, 10 6~; 1996, 3 Willow Creek, 5-VI-1995, 1 20-IV­ 5~. Spring Creek, 19-111-1988, 10 2~, 19-1V-1996, 10 1996, 90 2~; creek in Sunday Gulch, 8-II-1995, 4N, 5-Vl­ DISCUSSION.-This widespread species is very 1995, 5 ~, 19-1V-1996, 110 13 ~; Willow Creek, 20­ common throughout western North America, occur­ IV-1996, 3~. ring from Alaska south to Baja California and east to Manitoba (Nelson and Baumann 1989). The \VYOMING mSTRIBUTIOK.-Crook Co.: Beaver Black Hills represent the northeastern limit for C. Creek, 24-IV-1996, 40 6~, 27-VIII-1996, IN; Lytle gracilaria. Creek, 27-VIII-1996, 5N; North Redwater Creek, 24-1V-1996, 2~, 27-Vlll-1996, 7N. Weston Co.: Genus Eucapnopsis Beaver Creek, 23-IV-1996, 70 H; Cold Creek, 23­ Eucapnopsis brevicauda Claassen IV-1996,100 14~. DlSCUSSION.-This species is perhaps the most SOUTH DAKOTA DISTRIBUTION.-Custer Co.: Iron common and vagile species of the Euholognatha in Creek, 5-VI-1995, 2 ~, 20-IV-1996, 170 9 ~. Law­ It western North America. is found throughout the rence Co.: Iron Creek, 23-IV-1996, 1Ii; Spearfish Rocky, Cascade, Coast, and Sierra Nevada Moun­ Creek, 23-IV-1996, 220 17~: Whitewood Creek, tains (Baumann et al. 1977). It is also one of the 20-V-1981, I? Pennington Co.: Pine Creek, 5-VI~ most common stonefly species in the Black Hills, 1995, 2~; Rapid Creek, 21-IV-1996, 20 6~; creek occurring in both small and large streams. in Sunday Gulch, 5-VI-1995, 10 1~, 9-VIl-1997, 2 ~; Willow Creek, 5-VI-1995, 1 ~. Family Capniidae DISCUSSIOK.-This late-emerging winter stone~ fly is found throughout the Rocky, Cascade, Coast, Genus Capnia and Sierra Nevada Mountains. Capnia confusa Claassen Genus Isocapnia SOUTH DAKOTA DISTRIBUTION.-Custer Co.: Grace Isocaprtia sp. A Coolidge Creek, 5-VI-I995, 2 ~; Iron Creek, 5-Vl­ ~ ~ 1995, 2 , 20-1V-1996, II0 4 . Lawrence Co.: East SOUTH DAKOTA DISTRlBUTION.-Lawrence Co.: Spearfish Creek, 7-VI-1995, 4~, 23-IV-1996, 10: Spearfish Creek, near jet Redwater River, 24-111­ Iron Creek, 7-VI-1995, 1~, 23-1V-1996, 20 1~; 1982, 1 ~. Little Spearfish Creek, 23-1V-1996, 10; Spearfish DISCUSSION.-The genus lsocapnia is interest­ Creek, 25-IlI-1968, 40 (INHS), 7-VI-1995, 4~, 23­ ing because of its rarity. The specimen collected in IV-I996, 350 26~; Whitewood Creek, 20-V-1981, the Black Hills is of special interest because of the 2~. Pennington Co.: Deer Creek, 21-1V-1996, 16 hrachyptery of the female, a condition unreported 1~; Rapid Creek, 6-VI-1995, 10 2~, 21-IV-1996, for the genus. Males are required for specific deter­ 560 18~; South Fork Castle Creek, 21-IV-1996, mination. 40; creek in Sunday Gulch, 5-VI-1995, 3~, 19-1V­ 1996, 50 6~. Genus Paracapnia DISCUSSION.-This species is widespread and Paracapnia angulata Hanson common throughout the "Vest, occurring from Alaska south to New Mexico and east to Manitoba (Nelson SOUTH DAKOTA DISTRlBUTION.-Lawrence Co.: and Baumann 1989). Bear Butte Creek, 22-IV-1996, 10 6~; Boxelder 12 GREAT BASIN NATURALIST [Volume 59

Creek, 24-III-1968, 2<:1 (INHS), 22-IV-I996, 1 ~ .. Sunday Gulch, 5-VI-1995, Bel I?, 9-VII-1997, 10 Hay Creek, 22-IV-1996, 1 ~ .. Iron Creek, 7-Il-1995, 2? .. Willow Creek, 5-VI-1995, 240 9~, lO-VII­ 1<:1* 6~* .. Jim Creek, 6-Il-1995, 3N.. Little Elk 1997, 70 19~. Creek, 22-IV-1996, 1<:1 4~. Meade Co.: Elk Creek, DISCUSSION.-This species is the only leuctrid 22-IV-1996, 10<:1 18 ~ . Pennington Co.: Battle Creek, found in the Black Hills. Paraleuctra vershina is 8-Il-1995, 1 ~ '; Boxelder Creek, 22-IV-1996, 14<:1 found throughout the West, including the Coast, 23 ~ .. Burnt Fork, 21-IV-1996, 90 3 ~ ; Grizzly Bear Cascade, Rocky, and Sierra Nevada Mountains. The Creek, 20-IV-1996, 1<:1, Horse Creek, 18-III-1988, Black Hills population represents the easternmost 11<:1 13? .. Middle Fork Boxelder Creek, 22-IV-1996, range limit for this species. 1~ .. Palmer Creek, 5-VI-1995, 1~, 20-IV-1996, 20 1 ~ .. Rapid Creek, 21-IV-1996, 2<:1 I?; Spring Creek, Group Systellognatha 19-III-1988, 8~; creek in Sunday Gulch, 5-VI­ Family Perlodidae 1995, 4 ~ , 19-IV-1996, 6cl 5? . WYOMING DISTRIBUTlON.-Crook Co.: Beaver Genus lsogenoides Creek, 24-IV-1996, 27 <:1 44? . Isogenoides elongatus (Hagen) DrSGUSSION.-This eastern North American species has a remarkable geographic distribution. It SOUTH DAKOTA DISTRIBUTION.-Lawrence Co ..< occurs primarily from Quebec south to Tennessee; Spearfish Creek, Hwy 85, above Cheyenne Crossing, however, there are isolated populations reported in 7-II-1995, 3 ~ *. Pennington Co.: Rapid Creek, 6- II­ the Ozarks of Oklahoma, Rockies of Colorado and 1995,1<:1' ION, 6-VI-1995, 90 3?, 21-IV-1996, 9N. Wyoming, boreal streams ofManitoba and Saskatch­ DISCUSSION.-This species is typically found in ewan, and now in the Black Hills. larger creeks and rivers in the West. In the Black Hills, L elongatus was collected in only 2 of the Genus Utacapnia larger streams. Utacapnia lemoniana (Nebeker and Gaufin) Genus Isoperla Isoperla longiseta Banks SOUTH DAKOTA DISTRIBUTION,-Cusfer Co,: Wil­ low Creek, Hwy 89, N of Custer, 4-VI-1995, 1 ~, SOUTH DAKOTA DISTRIBUTION.-Butte Co.: Belle 19-IV-I996, 3<:1 7~. Lawrence Co.: tributary of Fourche River, Belle Fourche, 7-VI-1995, 2?; Newell Bear Butte Creek, Strawberry Picnic Area, 22-IV­ [probably Belle Fourche River], 19-VII-1923, 1<3 1996,10. Pennington Co.: Burnt Fork, rd 389, NW (INHS). Fall River Co.: Oral [probably Cheyenne of Hill City, 6-VI-1995, 1 ~, 21-IV-1996, 3<:1 5 ~ .. River], 4-VII-1953, 1<:1 (INHS). North Fork Castle Creek, rd 306, Mercedes Gulch, WYOMING DISTRIBUTION.-Crook Co.: Belle 21-IV-1996, 10 1 ~. Fourche River, Hwy 24, Hulett, 15-VII-1997, I? DISCUSSION.-This species occurs in the central DISCUSSION.-This species is one of the few Rocky Mountains. The Black Hills populations rep­ species common in the prairies (Ricker 1946, 1964). resent the easternmost range limit for this species. It is found throughout the West and Midwest in larger rivers. In the Black Hills, L longiseta is found Family Leuctridae in the 2 large rivers that drain the area.

Genus Paralew:;tra Isoperla plwlerota (Smith) Paraleuctra vershina Gaufin and Ricker SOUTH DAKOTA DlSTRIBUTlON.-Butte Co.: Red~ water River, 7-II-1995, IN. Ouster Co.: Beaver Creek, SOUTH DAKOTA DISTRIBUTION.-Custer Co.: U-VI-1975, 1cl2~ (PURC), 9-VII-1997, 1~ .. Grace Gr""e Coolidge Creek, 5-VI-1995, 140 12? .. Iron Coolidge Creek, 10-VIl-1997, 1 ~; Iron Creek, 6­ Creek, 5-VI-1995, 240 12?, 10-VII-1997, 7cl 6? VII-1968, 1<:1 1~, 5-VI-1995, 1<:1, 20-IV-1996, 3N, Meade Co.: Sturgis [probably Bear Butte Creek], 22-VIII-1996, 3N, 1O-VII-1997, 4~ .. Little Squaw 26-V-1951, 1 ~ (SDSU). Lawrence Co.: East Spear­ Creek, 15-IX-1970, 1<:1*, 1O-VII-1997, 2~. Lawrence fish Creek, 17-VI-1936, 1 ~ (SDSU) .. Iron Creek, 7­ Co.: Bear Butte Creek, 22-IV-1996, 3N.. Black Hills VI-1995, 2<:1 6?; Lead Creek [probably Whitewood Natl Forest near Savoy, [probably Spearfish Creek], Creek], 9-VI-1973, 5<:1 4? .. Spearfish Creek, 20-V­ 6-VII-1968, 1 ~ (INHS); Boxelder Creek, 6-1I-1995, 1981, 10, 7-VI-1995, 13<:1 18?, 13-VII-I997, I? .. 3N, 25-VIII-1996, IN; Cold Springs Creek, 20-V­ Whitetail Creek, 20-V-1981, 1 <:1 .. Whitewood Creek, 1981, 3N; East Spearfish Creek, 7-VI-1995, 1 <:1, 13­ 25-III-1982, IN. Pennington Co.: Elkhorn Spring, VII-1997, 1 ~ .. Elk Creek, 22-IV-1996, 7N; Engle­ 5-VI-1995, 65<:1 31~ .. Grizzly Bear Creek, 10-VII­ wood [probably Whitewood Creek], 18-VI-1925, 1997, 2 ~ .. Horse Creek, 6-VI-1995, 1 ~, Palmer 10 (INHS) .. Little Spearfish Creek, 17-III-1981, Creek, 5-VI-1995, 1cl, 10-VII-1997, 2?; Pine Creek, UN, 23-IV-1996, 3N, 13-VII-1997, 1<:1 1~ .. South 5-VI-1995, 450 17~, 10-VII-1997, 1 ~ .. creek in Fork Boxelder Creek, 22-IV-1996, 5N, 12-VIl-1997, 1999J STONEFLIES OF THE BLACK HILLS 13

19; South Fork Rapid Creek, 12-V11-1997, 10 41'; 1961, 19 (NDSU), 9-VI-1971, 30 (SDSU), 21-V­ Spearfish Creek, 18-VI-J966, 10 (SDSU), 9-VI­ 1981, 830 279, 16-VIJ-198I, 10, 24-111-1982, 1971,31' (SDSU), 17-II1-198I, 5N, 2O-V-198I, 10 30N, 7-11-1995, ION, 7-VI-1995, IN, 13-VII-1997, 39 5N, 6-X-198I, 7N, 9-XIl-198I, 17N, 25-1II­ 160 141'; Strawberry Creek [probably West Straw­ 1982, 19N, 7-[[-1995, 4N, 7-VI-I995, 19, 13·V[[­ berry Creek], 23-111-1982, 3N; tributalY of Bear 1997, 30 21'; Whitewood Creek, 19-V-198I, 10 Ilutte Creek, 12-VlI-1997, 19; Whitewood Creek, IN, 6-X-1981, 5N, 9-X11-198I, 5N, 25-II1-1982, 6N, 19-V-1981, IN, 23-111-1982, 3N, 14-VI-1982, 5N. Meade Co., Elk Creek, 22-IV-I996, 7N; Meadow Meade Co., Elk Creek, 22-IV-1996, 13N; Meadow Creek, 22-IV-1996, SN. Pennington Co., Battle Creek, Creek, 22-IV-1996, IN. PennulgWn Co., Battle 8-[[-1995,40' 21' '; Beaver Creek, 16-VlI-I997, Creek, 8-11-1995, 10; Beaver Creek, 16-V11-1997, 3 9 ; Boxelder Creek, 22-IV-I996, 3 ; Castle Creek, 20; Boxelder Creek, 22-lV-1996, 3 ; Deer Creek, 9-[[-1995, 2N; Deer Creek, 21-IV-I996, 29' 4N, 21-lV-I996, IN, ll-V11-1997, 460 499; Crizzly Bear 24-V1II-1996, IN; Grizzly Bear Creek, 20-IV-1996, Creek, IO-VII-1997, 20 19; Newtun Fork, U-VII­ IN; Middle Fork Boxelder Creek, 6-VI-1995, 20' 1997,260109; North Fork Castle Creek, IS-VIII­ 49', 22-IV-I996, 2N; Newton Fork, 23-VIII-1996, 1954, 19 (INHS); Palmer Creek, 1O-Vl1-1997, 19; 3N, ll-VII-I997, 39, North Fork Castle Creek, 24­ Rapid Creek, 6-11-1995, IN, 6-VI-1995, 20 29' VIII-1996, IN; Palmer Creek, 8-11-1995, 2N, 20-IV­ 9N, 21-IV-1996, 4N, 24-VllI-1996, II' 18N, ll-Vll­ 1996,2N, 1O-VII-1997, II'; Rapid Creek, 6-11-1995, 1997, 150 249; Spring Creek, 19-III-1988, IN, 19­ 6N, 6-VI-1995, 2N, 21-1V-1996, 4N, ll-VIl-1997, IV-I996, 14N, 21-VIII-1996, 40 II', ll-Vll-1997, 2E; Spring Creek, 19-1II-1988, 8N, 6-V1-1995, 50 310 191'; creek in Sunday Culeh, 5-VI-1995, IN; 59, 19-IV-1996, 3N, 21-VIII-1996, 4N; creek in stream in jenny Culch, II-Vll-1997, 10 19. Sunday Culch, 5-VI-I995, 10' 11", 19-IV-1996, WYOMINC mSTJUDUTION.--Crook Co.: Beaver IN; sb'eam in jenny Culch, ll-VIl-I997, II'. Creek, 24-lV-I996, 7N, 16-VII-I997, 50 II'; Beaver WYOMING DISTRI.BUTION.-Crook Co.: Beaver Creek, S-VI-I995, 70 79; Cold SPI;Dgs Creek, 16­ Creek, 24-JV-I996, 6N; Cold Springs Creek, 28­ Vll-1997,30 11'; Lame jones Creek, 16-VII-I997, VIII-1996, ION; Redwater Creek, 19-1Il-1981, 140 211'; Nurth Redwater Creek, 24-IV-I996, 6N; 2ON; Sand Creek, 19-III-198I, 6ON, 21-V-198I, 30 Redwater Creek, 19-I11-198I, 30N; Sand Creek, 59 15N, 7-X-1981, 12N, 8-Xll-1981, 40N, 24-IlI­ 19-IIl-198I, 50N, 21-V-1981, 350 141' 3N, 16-VII­ 1982, 12N, 24-IV-1996, 70 29. Weston Co.: Cold 1981, 10 8N, 8-Xll-198I, 20N, 24-Ill-1982, 25N, Creek, 23-IV-I996, 4N, 16-Vll-1997, 20 41'. 24-IV-1996, 15N, 28-VII1-1996, 70 149; Whitelaw DIscussrON.-The distribution of this species is Creek, 16-VII-1997, 170 249. Weston Co.: Cold interesting in that it is considered uncommon Creek, 16-VII-I997, 3029. throughout the West (SzczyLko and Stewart 1979) DISCUSSION.-This widespread western North but is common in the Black Hills. This may be due American species is common throughout the Black to a lack of competition from other medium-sized Hills. Szezytko and Stewart (1978) synonymized 1. predatory stoneflies. patricia Frison. originally described from Spearfish, South Dakota, with I. quinquep1mctata. lsoperla quinquepunctata (Banks) lsoperla tmHsmarina (Newman)

SOUTH DAKOTA DISTRIBUTION.-Butte Co.: Belle SOUTH DAKOTA DISTRIBUTION.-Pennington Co.: Fourche Rivet; 7-VI-199S, 1 <:?; Redwater River, 7­ Rapid Creek, Pactola Inlet, Silver City, 6-11-1995, [[-1995, 13N. Custer Co.: Battle Creek, 5-11-1995, lo,6-VI-1995,30 II'. 8N; Beaver Creek, 9-VIl-1997, 140 109; Flynn WYOMING DISTRIB1JTJON.-Crook Co.: Beaver Creek, 9-VII-1997, 20; French Creek, 9-VII-I997, Creek, ahove Cook Lake, Cook Lake Campgmund, 49; Crace Coolidge Creek, 5-II-1995, 3N, IO-VlI­ 16-VII-1997, 19. 1997,80 129; Little Squaw Creek, IO-VII-1997, DISCUSSION.-Thi.. spt.--cies is found in the north­ 8039; Iron Creek, 5-VI-1995, I ,IO-VII-I997, eastern United State.c; and across Canada (Stark et 110 181'; Spokane Creek, IO-VIJ-I997, 40 139; aI. 1986). The Black Hills population represents a tributary or Beaver Creek, 9-VI1-1997, 10 31'. southwest range extension for tbis species. Lawrence C()..· Boxelder Creek, 13-VI-1975, IN, 6­ 11-1995, 16N, 22-IV-I996, 2N, 12-Vll-1997, 310 Cenus SkWiM 269, Hay Creek, 22-lV-1996, IN, jim Creek, 11­ Skw,w, americana (Klapalek) Vll-1997, 140 189, Little Elk Creek, 22-IV-1996, 12N, 25-VIIl-1996, 10 39, 12-Vll-1997, 70 59; SOUTH DAKOTA DISTRIBUTfON.-Custer Co.: Iron Little Spearfish Creek, 17-111-1981, 2N, 7-VI-I995, Creek, 20-lV-1996, 30 12N, 22-VllI-I996, llN. IN, 23-IV-1996, 2N, 13-VII-I997, 80 29; Redwa­ Lawrence Co., Boxeldcr Creek, 6-11-1995, ION, 22­ ter River, 13-Vll-1997, 660 201'; South Fork Box­ lV-I996, IN, 25-VIII-I996, ION; Elk Creek, 22-lV­ elder Creek, 12-Vll-1997, 280 199; Spearfish 1996, II' 6N; Iron Creek, 23-lV-I996, 11'; North Creek, 27-VII-I940, 270 139 19 (I HS),I-VI- Boxelder Creek, 22-lV-I996, 2 ; South Fork Box- 14 GREAT BASIN NATURALIST [Volume 59 elder Creek, 25-VIII-1996, IN; Spearfish Creek, has a disjunct population in Quebec (Stark et al. 27-Vll-1940, 3E (lNHS), 17-111·1981, 2N, 21·Y·I981, 1986). Its distribution in the Black Hills is limited. IE, 16-Y11-1981, IN, 23-IX-1995, 20N (SDSU), 23­ IY-1996, 90 11 ~, 26-YIII-1996, 19N; Strawberry Genus Hesperoperla Creek [probably West Strawberry Creek], lO·XII­ Hesperope,.za pacifwa (Banks) 1981, 6N; tributary of Elk Creek, 24-1II-1968, IN (INHS); Whitetail Creek, 9-XlI-1981, IN; White­ SOUTH DAKOTA DISnUBlJflON.-Custer Co.: Iron wood Creek, 18-1II-19Bl, 5N, 8-X-1981, 2N, 9·XII­ Creek, 5-YI-1995, 6N, 2O-IV-1996, 9N, 22-YIlI­ 1981, 2N, 25·1II-1982, IN, 7·11-1995, 7N, Penning· 1996, 4N; Little Squaw Creek, 15-L,(-1970, IN, 10­ ton Co... Battle Creek, 8-11·1995, 50' 2N; Boxelder VlI-1997, 30 3~. Lawrence Co... Boxelder Creek, Creek, 22·IY-1996, IN; Bumt Furk, 21-IY-I9'J6, 19, 13-YI-197S, 10 3N (PURC), 6-I1-1995, 3N, 22-1Y­ 23·VlIl-l996, IN; Castle Creek, 9-lI-l995, 7N; Criz­ 1996, IN, 25-VlIl-1996, 3N; East Spearfish Creek, z1y Bear Creek, 2O-IY-1996, 90, 22-YIII-1996, 8N; 20-Y-I981, IN, 7-II-1995, 4N, 7-YI-I995, 3N, 25­ Horse Creek, 18-1II-198B, 4N; Nortll Fork Castle Vlll-I996, IN; Elk Creek, 25-YIIl-1996, 6N; Hay Creek, 24-YlII-1996, IN; Rapid Creek, 6-Il-1995, Creek, 22-IY-1996, 2N; Iron Creek, 7-I1-1995, 5N, 10'; Spring Creek, 9-1I-1995, 8N, 19-IY-1996, 10' 7-YI-1995, 30 29 5N, 23-IY-1996, 4N, 26-YIlI­ 5~* 10 17N, 21-YIII-1996, 4N; creek in Sunday 1996, 3N, 13-YII-1997, 20 2 ~; Jim Creek, 13-YI­ Gulch, 5-V1-1995, IN, 19-IY-1996, 140 3N. 1975, 7N (PURe), 6-I1-1995, IN; Little Elk Creek, WYOMING DISTRIBUTlON,-Crook Co.: Beaver 25-YIIl-1996, 3N; Little Spearfisb Creek, 17-IIl­ Creek, 27-YIII-I996, 2N; Whitelaw Creek, 27-VllI­ 1981, 3N, 21·Y-1981, IN, 15·X-1994. 9N (SDSU), 1996,3N. 23-IY-1996. 3N, 7-YI-I995. 3N, 26·VllI-1996, 4N, DrSCUSSION.-A widespread species ofthe Rock)', 13-YlI-1997, 3029; South Fork Boxelder Creek, Cascade, Coast, and Sierra Nevada Mountains (Bau­ 22-IY-1996, 2N, 12-Vll-1997, 10 29; South Fork mann et al. 1977), S. anu..7'icana is the eommon large Rapid Creek, 4-YlII-1967, 10 19 (SDSU); Spearfish predatory stonefly in the Black Hills. This species Creek, 27-YII-1940, 32N (INHS), 19-I!I-1981, 8N, was previously known as S. parallela (Frison), 2O·Y-1981, 7N, 16·YlI·1981, 7N, 6·X·1981, 5N, 9· XlI-1981, 8N, 25-IIl-1982, lSN, 7-11-1995, 12N, 7­ Family YI-1995, 2N, 23-lV-1996, 2N, 26-YllI-1996, 9N; Strawbeny Creek [probably West Strawbeny Creek], Genus Acroneuria IO-XlI-1981, 2N, 23-IlI-1982, 3N; tribulaty of Bear Acroneuria abnormis (Newman) Butte Creek, 22-1Y.1996, 6N, 24-YIIl-1996. 5N. 12­ YlI-1997, 10; tributary of Whitewood Creek, l2-Yll­ SOUTH DAKOTA DlSTRIBUTlON.-Butte Cu.: BeUe 1997,10; Whitetail Creek, 19-11I-1981, 4N, 16-VII­ Fourcbe River, Bene Fourche, 23-lV-1996, 4N. 1981, 2N, 8-X-1981, IN, 9-X11-1981, IN; White­ WYOMING DISTRIBtrrION.-Crook Co.: Belle wood Creek, 19-1lI-1981, 50N, 19-Y-1981, 10 2 ~ Fourche River, Hwy 24, Hulett, 7-1I-1995, IN. 24N, 15-YII-1981, 27N, 7-X-1981, 21N, 8-XlI-1981, DISCUSSION.-This is one of the most wide· 26N, 23-1lI-1982, 13N, 14-Vl-1982, 9N, 7-Yl-1995, spread species in all of North America (Stark et a1. 4N. Pennington Co... Battle Creek, 8-11-1995, IN; 1986). It is generally found in larger streams. It was Beaver Creek, 23-YIlI-1996, 2N, 16-YII-1997, 29; not found in the Black Hills proper but was found Castle Creek, 9-11-1995, 8N, 23-YIIl-1996, 3N; Ditch in the Belle Fourche River, which drains the north~ Creek, 23-YIlI-1996, 4N; Grizzly Bear Creek, 5-Yl­ ern section of the Black Hills and the Bear Lodge 1995, IN, 20-IY-1996, 8N, 22-YIIl-1996, 3N, 10­ Mountains. YII-1997, 1 ~; Newton Fork, 9-Il-1995, 8N, 6-YI­ 1995, 7N, 21-lV-1996, 4N, 11-Y11-1997, 29; Rapid Genus Claassenia Creek, 13-YI-1975, IN (PURe). 6-11-1995, 6N, 21­ Claassenia sabtJosa (Banks) lV-l996, 2N, 24-VlII-1996, 4N; Slate Creek, 21-lV­ 1996, 2N; Soutb Fork Castle Creek, 21-1Y-1996, SOUTH DAKOTA DISTRfBUTION.-Lawrence Co.: 3 ; creek in Sunday Gulch, 8-11-1995, 6N, 5-Yl­ Boxelder Creek, Boxelder Forks Campground, 13­ 1995, 2N, 19-IY-1996, 3N, 21-Ylll-1996, 3N, 9-YII­ Vl·1975, IN (PURe), 6-I1-1995, IN, 12-Yll-1997, 1997,10. 10; Spearfish, 25-YII-1924, 10 (lNIlS), 27-YII­ WYOMING DlSTRIBUTION.-Crook Co.: Beaver 1940, 3E (INHS); Spearfish Creek, Hwy 14A, Creek, 27-YllI-19'J6, 4N; Lytle Creek, 27-Ylll-1996, Botany Bay Picnic Area, 26-YlIl-1996, IN: White­ IN; Sund Creek, 19-IlI-1981, 19N, 21-Y-1981, 10, wood Creek, near confluence Whitetail Creek, 19 w 24-IY-1996, IN, 28-Ylll-1996, 2N; Whitelaw Creek, 1lI-1981, 4N, 7-X-1981, 2N, 25-1II-1982, IN; Rapid 27-YllI-1996, 2N. Weston Co ... Beaver Creek, 23­ Creek, Hwy 44, near Big Bend, 6-11-1995, 9N; IY-l996, 3N; Cold Springs Creek, 27-Y-1959, IN Rapid Creek, Pactola Inlet, Silver City, 6-Vl-l995. (NDSU).. 2N, 21·IV-1996, IN, 24-YIII,1996, 7N. DISCUSSION.-A typical western North Ameri~ DlscussroN.-This large, active perlid is <.'Ommon can species, it occurs from Alaska south to New throughout the West in larger streams, but it also Mexie<> (Stark et aI. 1986). It is also ooe of the most 1999] STO"lEFUES OF THE BL'CK HIllS 15 common species in the Black Hills, found in very 1995, 10 19; Spearfish Creek, Hw)' 14A, helow small to very large streams. The Black Hills repre­ Bridal Veil Falls, 7-VI-1995, 19; Spe.rfish Creek, sent the easternmost range limit for this species. Hwy 85, abuve Cheyenne Cl"Ossing, 7-VI-1995, 20 1 ~; Spearfish Creek, Hwy 14A, Botany Bay Picnic Genus Perlesta Area, 13-VII-1997, 1 ~; trihutary ofBear Butte Creek, Pedesta decipiem (Walsh) Strawherry Picnic Area, 12-VlI-1997, 20 2 ~. Pen­ nington Co.: Beaver Creek, rd 111, Beaver Creek SOUTH DAKOTA D1STRI6UTION.-Butte Co.: Belle Cmpgd, lo-VlI-I997, 19; Pine Creek, rd 244, above Fourche River, Belle Fourche, 13-VII-1997, 2<2. "au lIorsethief Lake, 5-VI-I995, l'i, 10-YII·I997, 29. River Co.: Cheyenne Hiver, Angostura Reservoir, DISCUSSION.-l1Iis species is found in the Cas­ Angostura State Park, Ii-VII-1997, 10 2 ~; Cheyenne cade, Coast, Rocky, and Sierra _ evada Mountains River, Oral, below Angostura Reservoir. 8·VIT-1997, (Baumann et al. 1977). Sweltsa borealis seems to 150 99. prefer small streams io the Black lUlls, The Black DISCUSSJON.-Perlesta decipiens has a widespread Hills represent the eastern ran~c limit for this distribution, ranging from Virginia to Colorad() and species. from Wisconsin to Texas (Stark 1989). In the Black Hills, it was found in the 2 large rivers that drain Sweltsa coloradensis (Banks) the area but not in the Black Hills proper. SOUTH DAKOTA DI~IRIBVTION.--Custer Co.: Grace Family Chloroperlidne Coolidge Creek, 5-VI-I995, 50, 1O-VII-I997, 929; Iron Creek, 6-VII-1968, 29 (INlIS), 5-VI-I995, 250 Genus Pamperla 69, 1O-VII-I997, 130 759; Little Squaw Creek, Paraperk>!ro,rwlis (Banks) 10-VIl·1997, 159; Spokane Creek, IO-VII-1997, 3 ~ ; Willow Creek, 9-VII-I997, 8~. Lawrence Co.: Black SOUTH DAKOTA DlSfRIBUTION.---Custer Co.: Grace lIills National Forest near Savoy Iprohahly Spearfish Coolidge Creek, 5-VI-1995, 10; Iron Creek, 5-Vl­ Creek], 6-VlI-1968, 20 79 (INHS); Boxelder 1995,30 29. Lawrence Co" Black Hills National Creek, l2-VII-I997, 59; East Spearfish Creek, 20­ Forest neur Savoy [probably Spearfish Creek], 6­ V-1981, 1 ~, 7-VI-I995, 300 279, 13-YII-1997, 10 VIl-1968, 1 ~ (INHS); East Spearfish Creek, 7-VI­ 64 ~; Englewood [probahly Whitewood Creek], 18­ 1995, 20; Spearfish Creek, 21-V-1981, l'i, 7-Vl­ VI-1925, 20 (INHS); Iron Creek, 7-VI-1995, 30 1995, 39; Whitewood Creek, 18-IlI-1981, 6N, 19­ 6 'i, 13-VII-1997, 8 ~; Little Spearfish Creek, 21-Y­ Y-1981, 2N. Pennington Co.; creek in Sunday Gulch, 1981,20, 7-VI-1995, 60 l 2, 13-VII-1997, 79; South 5-Vl-I995,20 12; Grizzly Bear Creek, 5-VI-I995, Fork Boxelder Creek, 12-VlI-I997, 179; South Fork 10; Pine Creek, 5-VI-1995, 130 29, 1O-VII-1997, Rapid Creek, 12-VlI-1997, 3~; Spearfish Creek, 19; Rapid Creek, i>-Vl-I995, 20. 21-Y-1981, 3d, 5-Vl-1985, 50 3~ (I lIS),7-VI­ DISCUSSION.-Althougb this species is not com­ 1995, 150 21~, 13-VlI-l997, 10 22~; Whitetail mon., it occurs in the Coast, Cascade, Sierra Nevada, Creek, 20-V-I981, 220 4 ~; Whitewood Creek, 19-V­ and Rocky Mountains (Baumann et al. 1977). The 1981,30 19. Penningf-Vll-I968, 280519 (II liS), 16 GREAT BASIN NATURALIST [Volume 59

10-VII-1997, 193 71 ~; Little Squaw Creek, lS-IX­ State Park are much appreciated, Finally, spe­ 1970, 13* 2~*, 10-VII-1997, 33 18~; Spokane cial thanks are given to the following col­ Creek, 10-VlI-1997, 163 42 ~ ; Sylvan Lake, 20-VI­ leagues who helped introduce the second 1940,23 1 ~ (INHS); stream entering Sylvan Lake, author to the Black Hills during a study of 9-VlI-1997, 313 21 ~; tributary of Beaver Creek, 9­ Whitewood Creek in the early 1980" Robert VII-1997, 8 ~; Willow Creek, 9-VlI-1997, 83 28 ~, Lawrence Co,; Boxelder Creek, 12-VlI-1997, 40 Winget, John Unzicker, Joseph Furnish, and 9 ~; Englewood [probably Whitewood Creek], 18­ Edward Herricks, Vl-192S, 20 (lNHS, SDSU); Iron Creek, 13-VlI­ 1997, 30 10~; Jim Creek, ll-VlI-1997, 330 30~; LITERATURE CITED Little Elk Creek, 12-VII-1997, 10 3~; Little Spear­ fish Creek, 13-VlI-1997, 13 2~; South Fork Box­ BAUMANN, RW 1979. Nearctic stonefly genera as indica­ elder Creek, 12-VlI-1997, 23 8 ~ ; Spearfish Creek, tors of ecological parameters (Plecoptera: Insecta). 13-VII-1997, 1 ~; tributary of Bear Butte Creek, 12­ Great Basin Naturalist 39:241-244. VlI-1997, 63 3n; tributary of Whitewood Creek, BAUMANN, R.W., A.R GAUFIN, AND R.E SURDICK. 1977. ~, The stoneflies (Plecoptera) of the Rocky Mountains. 12-VlI-1997, 1 Pennington Co,; Burnt Fork, 11­ Memoirs of the American Entomological Society VlI-1997, 133 26~; Grizzly Bear Creek, 10-VII­ 3101-207, 1997, 12~; Middle Fork Boxelder Creek, 6-VI-199S, CROSS, RB., RL. MAYDEN, AND ].D. STEWART. 1986. Fishes 1 ~; Newton Fork, ll-VII-1997, 213 4S~; Palmer in the western Mississippi Basin (Missouri, Arkansas, Creek, 10-VlI-1997, 333 31 ~; Pine Creek, 10-VII­ and Red Rivers). Pages 363-412 in C.H. Hocutt and 1997, 10 1 ~ ; Spring Creek, ll-VII-1997, 2 ~; creek E.O. Wiley, editors, The zoogeography of North in Sunday Gulch, 9-\>'11-1997, ISo 17~; stream in American freshwater fishes. John Wiley & Sons, Jeuny Gulch, ll-VII-1997, 13 1~; Willow Creek, New York. 1O-VII-1997, 103 8 ~, DOSDALL, L., AND D.M. LEMKUHL. 1979. Stoneflies (Ple­ coptera) of Saskatchewan. Quaestiones Entomologi­ WYOMING DISTRIBUTION.-Crook Co.: Blacktail cae 15:3-116. Creek, 16-VII-1997, 13 1O~; T52N R63W [probably DRAKE, Y.A., AND RA. FARROW. 1988. The influence of ~; Houston Creek], 22-VlI-1982, 13 3 Whitelaw atmospheric structure and motions on insect migra­ Creek, 16-VlI-1997, 183 43 ~, Weston Co,; Cold tion. Annual Review of Entomology 33;183-210. Creek, 15-VII-1997, 53 7~, FLANNAGAN, P.M., AND J.E FLANNAGAN. 1982. Present dis­ DISCUSSION.-This species, often confused with tribution and the post-glacial origin ofthe Ephemer­ T signata, occurs in the Coast, Cascade, Sierra optera, Plecoptera, and Trichoptera of Manitoba. Nevada, and Rocky Mountains, but it has not been Manitoba Department of Natural Resources and reported from Canada (Baumann et al. 1977, Stark Fisheries Technical Report 82:1-79. et al. 1986). Triznaka pintada is very common in the FRISON, T.H. 1929. Fall and winter stoneflies, or Plecoptera, Black Hills, with this area representing the eastern­ of Illinois. Illinois Department of Registration and Education Bulletin 18:345--409. most range limit for this species. GRIFFITH, M.B., E.M. BARROWS, AND S.A. PERRY. 1998. Lateral dispersal ofadult aquatic insects (Plecoptera, ACKNOWLEDGMENTS Trichoptera) following emergence from headwater streams in forested Appalachian catchments. Annals We thank Oliver Flint, Jr, (United States ofthe Entomological Society ofAmerica 91:195-201. National Museum, Smithsonian Instituition), HARDEN, P.H., AND c.E. MICKEL. 1952. The stoneflies of Minnesota (Plecoptera). University of Minnesota Paul J. Johnson (South Dakota State University), Technical Bulletin 20Ll-84. Arwin V Provonsha (Purdue University), Brett Hayward, H.E. 1928. Studies of plants in the Black Hills C. Ratcliffe (University of Nebraska), David A, of South Dakota. Botanical Gazette 85:353-412. Rider (North Dakota State University), and HOFFMAN, RS., AND J.K. JONES, JE. 1970. Influence of Donald W Webb (Illinois Natural History Late-glacial and Post-glacial events on the distribu­ tion of Recent mammals on the Northem Great Plains. Survey) for the loan of specimens from South Pages 355-394 in W Dart, Jr., and J.K. Jones, Jr., Dakota, We appreciate Stanley W Szczytko, editors, Pleistocene and Recent environments of the University of Wisconsin, Stevens Point (Isop­ Central Great Plains. Kansas University Press, erla), and Kevin D Alexander, Wayne State Col­ Lawrence. lege (SuwaUia), for verifying some determina­ HOUSEMAN, R.M., AND RW. BAUMANN. 1997. Zoogeo­ graphic affinities of the stoneflies (Plecoptera) of the tions, The authors thank Mark C. Belk for his Raft River Mountains, Utah. Great Basin Naturalist assistance in the statistical analysis of dispersal 57,209-219, ability, Randy Lawson at Chadron State Col­ HYNES, H.B.N. 1988. Biogeography and origins of the lege, Chadron, Nebraska, provided help and North American stoneflies (Plecoptera). Memoirs of encouragement The help of Daniel Uresk and the Entomological Society ofCanada 144:31-37. ILLIEs, J. 1966. Katalog del' rezenten Plecoptera. Das personnel of the USFS, Black Hills National Tierreich. Lieferung 82. Walter de Gruyter & Com­ Forest, and Ronald Walker and staff at Custer pany, Berlin. 1999J STO EFLIES OF THE BLACK HILLS 17

JOHNSON, e.C. 1969. Migration and dispersa.l of instlCts by . t998. Stonelly li~t as of 5 January 1998. World Right. Methuen & Co., London. ----;''''V;ide Web URL http://www.mc.cdu/-slllrk!slonefly. JOSEI'HSON, R,K. 1981. lempemture and the mechanical hIm!. perlonnance ofinscct muscle. Pages 19-44 in B. Hein­ STARK, B.P., AKD A.R Gi\UFIN. 1976. The Nearetic species rich, editor, Insect thermoregulation. John Wiley & or Acnmeuria (Plecoptera: Perlidae). Journal of the Sons, New York. Kamas Entomological Society 49:221-253. Kos:crARAB, M., AND C.W SCHAEFER. 1990. SyHematics of STARK, D.P., B.R. OBLALJ, AND A.H.. CAunN. 1973. An North American insects and aruchnick status and annohlted list of the stonellies (Pleeoptenl) of Col­ ne~ds. Virginia Agricultural Experimental Station orado. Entomological News 84:269-277. 30.1-305. Information Series 90-1. STARK, B.P.. S.W SZCZYTKO, AND RW Di\lJMhNN. 1986. M...... ROF.N, J.H., ANO M.G. KRAMEIl. 1994. Surfucc..skimming North American stoneflies {l'lccoptera): systematics, stoneflies: a possible intennroiate stage in insect flight distribution, and taxonomk references. Great Basin evolution. Sciefll.'e 266:427-430. Naturalist 46:383-397. MCCAFFEIITY, WP. 1990. Biogeogr-J.phic affinities of the STEWAHT, K.W, AND B.P. STARK. 1988. Nymphs of North Ephcmeropter-a of the Black HiJls. South Dakota. American stonelly generd (Plecoptel'a). Entomologi­ Entomological News 101 :193-199. cal Soc'iely of America, Thomas Say Foundation McINTOSH, A.J. 1931. A butanie A.R. C"UI~IN. 1978. Environmental Hills: the status of our knowledge. United States I'~quirement" and pollution tolemnce of PIe<.."Optera. Department ofAgriculture, 'Forest SeIVice Hcscarcb U.S. Environmental Protection Agency Report EPA­ Paper l:\M-141. 60014-78-062. POULTON, B.B., AND K.W STEWART. 1991. The stoncflies of SZCZ\'TKO, S.W, A,,"O K.W STEWART. 1979. The genus the Ozark :md Ouachita Mountains (Plecoptera). Isoperla (Plecoptera) ofwestern North America: halo­ Memoirs or the American Entomological Soci.ety lnorphology and systematics, and a new stonelly 38,1-1l6. genus Casau1o'Jerfa. Memoirs or the American ETlto~ RIcKEn, WE. ]946. Some prairie stonefiies (Plecoptera). mological Society 32:1-120. Transacl'ions ofthe Royal C of uralist 36:323-331. the Central Grear Plains. Kansas University Press, SAS. 1990. SAS user's guide: statistic,;. Version 6 edition. Lawrence. SAS Institute Inc., Cary, !'JC. STARK, B.p. 1989. Perlesta placida (Hagen), an eastern Received 24 August 1998 Nearctic species compl~.'( (Plecoptera: Perlidae). Accepted 3 November 1998 ElltomologicD Scandinavka 20:263-286.