Stoneflies (Plecoptera) of Southern Utah with an Updated Checklist of Utah Species

Monographs of the Western North American Naturalist

Volume 1 Article 2

11-25-2002

Stoneflies (Plecoptera) of southern Utah with an updated checklist of Utah species

Ronald G. Call

Richard W. Baumann Brigham Young University, [email protected]

Follow this and additional works at: https://scholarsarchive.byu.edu/mwnan

Recommended Citation Call, Ronald G. and Baumann, Richard W. (2002) "Stoneflies (Plecoptera) of southern Utah with an updated checklist of Utah species," Monographs of the Western North American Naturalist: Vol. 1 , Article 2. Available at: https://scholarsarchive.byu.edu/mwnan/vol1/iss1/2

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STONEFLIES (PLECOPTERA) OF SOUTHERN UTAH WITH AN UPDATED CHECKLIST OF UTAH SPECIES

Ronald G. Call1,4 and Richard W. Baumann2,3

ABSTRACT.—Southern Utah comprises 4 major physiographic divisions: the Great Basin, Colorado Plateau, Central Rocky Mountains, and Southern Rocky Mountains, which have been partitioned into faunal regions. We discuss the uniqueness of southern Utah and the amount of land set aside for recreation and preservation, and we encourage the study and preservation of its water resources. The stonefly (Plecoptera) fauna of 13 counties in southern Utah was sur- veyed. We found 58 species representing 34 genera and 8 families. Three new state records for Utah and many new county records for southern Utah are presented, along with a discussion of distribution patterns and faunal affinities of each species. Distribution patterns indicate a historical connection between the Sevier River and the Colorado drainage and a separation of faunal areas within the Central Rocky Mountains at boundaries between the Wasatch Mountains, Wasatch Plateau, and the Southern High Plateau. The uniqueness of the isolated Abajo and LaSal Mountains, the Virgin River valley, and other faunal areas in southern Utah is presented.

Key words: Plecoptera, stoneflies, aquatic insects, Utah, species inventory.

Southern Utah is famous for its spectacular population growth. Washington County, in the outdoor scenery. The area has long been southwestern corner of the state, experienced appreciated and grows in popularity each year. the largest increase in population. St. George, Tourism in southern Utah is increasing, as are the largest city of Washington County, had the conservation efforts in the region. The designa- 2nd largest percentage population increase of tion of the 1.7-million-acre Grand Staircase– any city in the state for the past 6 years, over Escalante National Monument in southern 36% (Governor’s Office of Planning and Bud- Utah in 1997 (Maurer and Beath 1997) is per- get 1998). haps the greatest evidence of growing interest Utah is a treasure trove of natural beauty. in the preservation of the area. Greater under- Southern Utah contains all 5 of the state’s na- standing of southern Utah’s living resources is tional parks, 5 of its 7 national monuments, 4 of essential to wise management decisions in the the state’s 8 national forests, 1 national recre- face of increasing tourism and population ation area, 7 designated wilderness areas, and growth. 20 state parks (Utah Travel Council 1999). With Utah has the 2nd highest birth rate and the approximately 69% of its land federally owned 2nd lowest death rate of any state in the nation, (U.S. Department of the Interior, Bureau of which together contribute significantly to Land Management, Utah State Office 1997), Utah’s high population growth. With the addi- Utah ranks 2nd among states in amount of tion of a high rate of in-migration, Utah has federally owned land, next to Nevada, which one of the fastest growing populations of any lists 77% of its land as federally owned (U.S. state. Although currently about 77% of Utah’s Department of the Interior, Bureau of Land population lives along the Wasatch Mountain Management 1999). However, nearly 4.5% of Range in northern Utah, this condition is chang- Nevada’s land is federally managed under the ing. From 1980 to 1996, northern Utah, which Department of Defense (Nevada Division of includes the urban centers of the Wasatch State Lands 1995), while in Utah a higher per- Mountain Range, experienced an average annual centage of federal land is set aside as national population growth of 1.67%, while southern parks, national monuments, national recreation Utah experienced a 1.74% average annual areas, and national forests. Utah ranks highest

1Richfield High School, Richfield, UT 84701. 2Department of Integrative Biology, and Monte L. Bean Life Science Museum, Brigham Young University, Provo, UT 84602. 3Author to whom reprint requests should be sent. 4Present address: Madison High School, Rexburg, ID 83460.

65 66 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1 among the continental states in the proportion such as stoneflies. Stoneflies typically inhabit of its land designated for public recreation and cool streams and rivers. They are excellent in- preservation. While it is true that more feder- dicators of changing water conditions and qual- ally owned land, over 50 million acres, is spec- ity and can aid in understanding the diversity ified for preservation and recreation in Alaska and uniqueness of areas. Information relative than in Utah, which has approximately 11.7 to stonefly species inhabiting the area may be million acres, nearly 9 million visits are made useful in helping determine which areas are to public recreation sites in Utah annually com- most critical to preserve and in making man- pared to approximately 2 million annual visits agement decisions involving the use of water in Alaska. Utah ranks 8th among the states for resources. number of annual visits to recreational sites The species composition and distribution of managed by the National Park Service (U.S. stoneflies in Utah is perhaps the best known Department of the Interior, National Park Ser- of any state in the western United States (Tan- vice 1998). Southern Utah, in particular, is ner 1940, Gaufin et al. 1966, Baumann 1975, unique. Nearly 7.7 million of approximately 30 Baumann et al. 1977). Previous knowledge of million acres of land in southern Utah is pub- Plecoptera in Utah is primarily the result of licly owned as national parks, national monu- the work of the late Arden R. Gaufin and his ments, national recreation areas, national forests, students at the University of Utah. In 1966, and state parks. That is over 25% of the area of Gaufin listed 71 species, 18 genera, and 5 fam- southern Utah (U.S. Department of the Inte- ilies of Plecoptera as occurring in Utah. Since rior, Bureau of Land Management, Utah State that time many subgenera have been raised to Office 1997; U.S. Department of the Interior, generic status, and several subfamilies have National Park Service 1998; Utah Travel Coun- been elevated to family status (Illies 1966). cil 1999). No other area in the country has The most recent publication providing a state such a large proportion of land allocated for list of species occurring in Utah listed 75 species preservation and recreation by both state and in 34 genera and 8 families (Baumann 1973). federal agencies. These facts are presented to New records were added in 1977, document- emphasize that Utah has both a large amount ing the occurrence in Utah of 77 stonefly spe- of land designated for recreation and preser- cies in 35 genera and 8 families (Baumann et vation and an increasing number of people al. 1977). Stanger and Baumann (1993) de- visiting these unique sites. Despite these facts, scribed a new species from the state, Surdick little research has been done regarding the (1995) included the description of a near chlor- biological components of the varied ecosys- operlid from southeastern Utah, and House- tems of southern Utah. man and Baumann (1997) recorded 5 species Arid conditions throughout southern Utah previously unrecorded from the state. place a premium on freshwater. Virtually all Most knowledge of stoneflies in Utah comes municipal waterworks systems in Utah tap from sites in the Central Rocky Mountain, groundwater resources through springs and Uintah Basin, Columbia Plateau, and Great wells. Expanding human population and agri- Basin areas in the northern part of the state culture will place even greater demands on (Gaufin et al. 1966, Baumann and Gaufin 1969, southern Utah’s freshwater supplies in the Baumann et al. 1977, Houseman and Baumann future, groundwater as well as streams and 1997). The southern Utah stonefly fauna is rivers. The city of St. George, southern Utah’s poorly known compared to that of northern largest city, already pays the highest price for Utah. This has resulted from the major univer- water in the state (Governor’s Office of Plan- sities in Utah all being located in the northern ning and Budget 1998). A large proportion of part of the state: University of Utah, Salt Lake precipitation in southern Utah is received in City; Brigham Young University, Provo; and the form of winter snows, which form the largest Utah State University, Logan. These 3 institu- freshwater reserve in the area (Harper et al. tions have all been instrumental in providing 1994). Because of the importance and relative researchers and facilities to house insect col- rarity of freshwater in this area, it is important lections. Consequently, due to both time and to gain a greater understanding of all aspects budget constraints, much more research has of the area’s aquatic resources. One method of been conducted in northern Utah than in south- gaining this insight is by studying aquatic insects ern Utah. Distance between suitable stonefly 2002] STONEFLIES OF SOUTHERN UTAH 67 habitats, the remoteness of some areas, and the orado River and its tributaries, including the rugged terrain of some areas of southern Utah Green River, San Juan River, and Virgin River, have also been obstacles to stonefly research to the southwest (Fig. 1). in this part of the state. A large amount of information regarding stoneflies and other aquatic insects of Hunt- STUDY AREA ington Creek, a tributary of the San Rafael River, itself a tributary of the Green River, was Some authors have divided Utah into 3 compiled by R.N. Winget, E.C. Devenport, major physiographic provinces or physical and M.K. Reichert during a study from 1970 divisions (Harper et al. 1994). The western to 1977. The principal report (Winget 1972), portion of the state is occupied by the Great detailing the site and explaining the scope of Basin, a part of the Basin and Range Province. the study, was published early in the study, The northeastern portion of the state consists but data were recorded for the entire 7-year of a portion of the Central Rocky Mountains. study period. Huntington Creek has been The southeastern portion of the state is part of more extensively studied, relative to aquatic the Colorado Plateau. Other authors have con- insects, than any other drainage in southern cluded that some isolated mountain regions of Utah (Reichert 1975). The only other stream southern Utah are more correctly considered in southern Utah with aquatic invertebrates part of the Southern Rocky Mountain Prov- that have been studied to any degree is Pleas- ince (Durrant 1952, MacMahon and Wieboldt ant Creek, which flows through Capitol Reef 1978). These isolated mountains, such as the National Park. We are indebted to Colin Bram- LaSal and Abajo Mountains, more closely re- mer and Dr. John MacDonald for their study semble, faunally, the Southern Rocky Moun- of this creek (Brammer 1997). tains of Colorado than the surrounding plateaus. Durrant (1952) partitioned Utah into faunal A large portion of southern Utah is considered areas based on distributions of mammals. These part of the Colorado Plateau Physiographic divisions correlated fairly closely with the Province. This area consists of high-elevation physiographic provinces but were subdivided plateaus incised by canyons. Another major into smaller regions. The Middle or Central portion of southern Utah is the Great Basin, a Rocky Mountains he subdivided into the very arid region in the southwestern portion Wasatch Mountains, the Uinta Mountains, and with few high-elevation mountains. Water in the Northern and Southern High Plateau pro- the area is primarily ephemeral streams from vinces. The Southern Rocky Mountains were mountain runoff or desert rainstorms. divided into the Abajo subcenter and the As sensitive aquatic insects, stoneflies are LaSal subcenter. He divided the Colorado Pla- limited to freshwater lotic systems; therefore, teau into the Virgin River Valley Province and distributions of stoneflies reflect the presence the Canyon Lands Province, each of which of streams and rivers. Northern Utah contains was further subdivided. The Great Basin is the the Wasatch and Uinta Mountains, which only province Durrant did not divide into include many streams and rivers that provide smaller regions or zones, as the mammal fauna suitable habitat for stoneflies. Northern Utah of that region was uniform. contains 3 principal drainage areas. The Raft Our study area includes the following Utah River Mountains in the extreme northwestern counties (Fig. 2): Beaver, Emery, Garfield, corner of the state drain northward into the Grand, Iron, Kane, Millard, Piute, San Juan, Snake River of Idaho. The Wasatch Mountains Sanpete, Sevier, Washington, and Wayne and northwestern portion of the Uinta Moun- (approximately 37°–39°30′N latitude, 109°05′– tains drain into Utah Lake and the Great Salt 114°W longitude). The area covers approxi- Lake. The southeastern portion of the Uinta mately 30 million acres (Bureau of Land Man- Mountains drains into the Green River, which agement 2001) or 124,600 km2 (48,100 square empties into the Colorado River. Southern miles; DeLorme Mapping 1993). Utah contains 2 principal drainage basins. The Sevier River and Beaver River flow to the METHODS north and eventually dissipate in the desert near Sevier Dry Lake in Millard County. The Collections were made at over 100 sites other major drainage is the flow of the Col- within the study area. We chose collection sites 68 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

Fig. 1. Major drainages of Utah. 2002] STONEFLIES OF SOUTHERN UTAH 69

Fig. 2. Counties in southern Utah study area.

to achieve as much coverage of the area as specimens, we examined preserved specimens possible, attempting to obtain samples from from the following institutions: Monte L. Bean every major drainage and mountain range dur- Life Science Museum, Brigham Young Univer- ing each season of the year. Collections of sity, Provo, Utah (BYUC); Canadian National fresh specimens were made from 1997 to Collection, Ottawa, Ontario, Canada (CNC); 2000. We collected nymphs by disturbing the C.P. Gillette Museum of Arthropod Diversity, substratum of streams and catching specimens Colorado State University, Fort Collins, Colo- in kick nets held downstream. Adults were rado (CSUC); Dixie College, St. George, Utah collected principally by beating riparian vege- (DCC); Illinois Natural History Survey, Cham- tation with sticks while we held beating sheets paign, Illinois (INHS); Purdue Entomological onto which specimens would fall from the Research Collection, West Lafayette, Indiana vegetation. All specimens were preserved in (PERC); Utah State University, Logan, Utah 75% ethyl alcohol. In addition to field-collected (USUC). Specimens collected during the study 70 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1 are deposited in the Monte L. Bean Life Sci- corner of Utah are faunally interesting and dif- ence Museum. ferent from the rest of southern Utah. We feel Each specimen was identified to the lowest this area needs much more study. possible taxonomic level, usually to species. In The stonefly species Oemopteryx fosketti the case of some nymphs, identification was (Ricker) and Isoperla longiseta Banks are lim- possible only to genus. Locations of specimen ited in Utah to the large rivers of the Colorado collection sites were then plotted on maps of drainage, indicating a unique aquatic faunal the study area to reveal the distribution of region in southern Utah. Additionally, Acro- each species within the study area. We exam- neuria abnormis (Newman) and Isogenoides ined each species distribution and noted its colubrinus (Hagen) occur in Utah only in the pattern relative to physiographic provinces or Colorado drainage, with the exception of the faunal areas. Sevier River. We believe this indicates that the Sevier River was historically part of the Col- RESULTS orado drainage and that the large rivers of the Colorado drainage represent a unique faunal Fifty-eight species in 34 genera and 8 fami- region. lies of stoneflies were recorded from the south- Stonefly distributions indicate that the Wa- ern Utah study area. Three species, Isoperla satch Plateau, Southern High Plateau, Virgin phalerata (Needham), Malenka flexura (Claas- River valley and Zion National Park area of sen), and Sweltsa cristata Surdick, are new the Virgin River, Abajo and LaSal Mountains, state records. Many species listings are new and rivers of the Colorado drainage are dis- county records, which help to establish more tinct faunal regions (Fig. 3). Stonefly distribu- complete distributions for some species. tions appear to confirm, for the most part, the One of our most interesting findings was Utah faunal divisions of Durrant, which were documenting 2 different species, Acroneuria based on mammal distributions. Stonefly dis- abnormis (Newman) and Isogenoides colubri- tribution patterns confirm Durrant’s separa- nus (Hagen), in the Sevier River, which flows tion of the Central Rocky Mountains into the north into the Great Basin. This was particu- Wasatch Mountains, the Northern High Plateau larly noteworthy because these 2 species were (herein referred to as the Wasatch Plateau), and previously believed to occur in Utah only in Southern High Plateau, based on the separation the large rivers of the Colorado drainage. of ranges of Amphinemura banksi Baumann Also of much interest are isolated occur- and Gaufin and Amphinemura mogollonica rences of 3 species. Single specimens of both Baumann and Gaufin at the border between Isoperla phalerata (Needham) and Isoperla the Wasatch Mountains and the Wasatch Pla- pinta Frison are documented from high-eleva- teau. The distributions of Paraleuctra vershina tion, isolated areas in southern Utah. Although (Gaufin and Ricker) and Malenka californica the type locality of Utaperla sopladora Ricker (Claassen) in northern Utah, south to the bor- is in southern Utah, only a single specimen of der between the Wasatch Mountains and the this species in addition to the type specimen Wasatch Plateau, also support the concept that has ever been collected from southern Utah. these are separate faunal regions. Obviously, this species is isolated and rare in The separation between the Wasatch Plateau this area. and the Southern High Plateau, although less During the course of our study, it became definite, is indicated by the distribution of Ma- apparent how different the isolated Abajo and lenka coloradensis (Banks), which extends across LaSal Mountains in the southeast corner of Utah the Southern High Plateau to the edge of the are. Several stonefly species in these moun- Wasatch Plateau. tains occur nowhere else in southern Utah. The species Amphinemura banksi Baumann Zapada frigida (Claassen), Sweltsa cristata Sur- and Gaufin, Malenka flexura (Claassen), Zapada dick, and Malenka flexura (Claassen) occur frigida (Claassen), and Sweltsa cristata Surdick, nowhere else in the state. Additionally, Amphin- which are found at high elevations in the Abajo emura banksi (Baumann and Gaufin), Triznaka Mountains and LaSal Mountains, confirm Dur- pintada (Ricker), and Kogotus modestus (Banks) rant’s placement of these mountains as part of are found nowhere else in southern Utah. the Southern Rocky Mountains, similar to those Clearly, these mountains in the southeastern of Colorado (Durrant 1952). 2002] STONEFLIES OF SOUTHERN UTAH 71

Fig. 3. Southern Utah faunal areas indicated by stonefly distributions.

Other subcenters designated by Durrant and Sweltsa lamba (Needham and Claassen), were not indicated by stonefly distributions. which are not found in the Zion National Park Durrant divided the Canyon Lands Province area of the Virgin River. One species, Meso- into the Kaiparowits, San Rafael, Grand Valley, capnia frisoni (Baumann and Gaufin), occurs San Juan, and Painted Desert subcenters. How- in Utah exclusively within the Pine Valley ever, stonefly species occurring within this Mountain area and the Virgin River Valley province appear to be widespread within it, in- Province, indicating a unique faunal region. dicating no divisions. This entire area is a desert Certainly, the Virgin River valley is a biologi- plateau through which flow rivers that appear cally unique area of southern Utah. to have been connected with each other in the Of southern Utah’s 4 physiographic prov- past as part of the Colorado drainage. inces, the Great Basin is the poorest in terms Durrant divided the Virgin River Valley of numbers of stonefly species. Every species Province into the St. George subcenter and present in the Great Basin occurs also in the the Beaver Dam Wash subcenter. This divi- bordering Central Rocky Mountains or is wide- sion, based on mammal distribution, is con- spread throughout southern Utah. Isoperla firmed by plant distributions (Durrant 1952). mormona Banks is the only species that is Our data do not indicate a division within the known in southern Utah to extend far into the Virgin River Valley Province at the boundary Great Basin. Indeed, few stoneflies are pres- of these 2 areas, but they may indicate a bound- ent deep in the Great Basin, reflecting its gen- ary between the Virgin River in the Zion eral unsuitability as stonefly habitat (Nelson National Park area and the remainder of the 1994). In fact, the delineation of the Great drainage. Amphinemura mogollonica (Baumann Basin as a faunal region is revealed by the lack and Gaufin) and Isogenoides zionensis Hansen of stoneflies in the region. occur in the Zion National Park area but are Boundaries between physiographic provinces not recorded from the Virgin River west of the in southern Utah appear to be barriers in the park. The Virgin River west of Zion National dispersal of some stonefly species. This study, Park contains Mesocapnia frisoni (Baumann along with previous records, indicates that the and Gaufin), Hesperoperla pacifica (Banks), edge of the Colorado Plateau in southern Utah 72 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1 is the eastern limit of Capnia utahensis Gaufin vicariance of ancestral populations. Given the and Jewett and that its range extends into the widely disjunct distributions of some species in Wasatch Mountains of northern Utah. Southern southern Utah such as Isoperla phalerata Utah appears to be the southern range limit of (Needham), Isoperla pinta Frison, and Uta- Utacapnia lemoniana (Nebeker and Gaufin) and perla sopladora Ricker, and the generally poor Utacapnia logana (Nebeker and Gaufin). dispersal ability of stoneflies, vicariance of an- Another result of our study is that several cestral populations seems the more likely ex- species previously unrecorded or recorded from planation. only a few localities in southern Utah have Fifty-eight species of stoneflies from 34 now been documented from several localities. genera and 8 families are reported from south- Alloperla severa (Hagen), previously unrecorded ern Utah. Three species represent new state from southern Utah, was recorded from sev- records, and many others represent new county eral localities. Previously listed from only 2 records. This publication enhances knowledge counties in southern Utah, Megarcys signata of stoneflies occurring in Utah by updating the (Hagen) is now documented by this study as state list of species. Also provided are distribu- being widespread in the area. The validity of tional data for each species occurring in south- the recently designated species Sweltsa cristata ern Utah. This study provides a higher degree Surdick from the isolated LaSal and Abajo of resolution of stonefly distributions in this Mountains was confirmed by examination of area. Distribution patterns indicate that some several excellent specimens. species are widespread throughout the region. Many species occur in the Central Rocky CONCLUSIONS Mountain region, while fewer species occur in the Southern Rocky Mountain and Colorado The number of species recorded from Plateau regions. southern Utah is less than that recorded from Distributions of stonefly species indicate northern Utah (Call 1999). The Wasatch and the division of the Central Rocky Mountains Uintah Mountains in northern Utah form a region into the Wasatch Mountains, Wasatch fairly contiguous band of high-elevation moun- Plateau, Southern High Plateau, and Virgin tains with suitable stonefly habitat. Only the River valley as one moves from north to south Raft River Mountains in the northwestern cor- within the Central Rocky Mountains of Utah. ner of the state are an isolated range in the Distribution patterns also reveal the Abajo and north. In contrast, southern Utah has more LaSal Mountains are unique faunal regions isolated, high-elevation mountains, such as the with close affinities to the nearby Southern LaSal, Abajo, and Henry Mountains. Rocky Mountains of Colorado. Although fewer species occur in southern The presence of stonefly species in the Utah than in northern Utah, 58 species, which Sevier River, otherwise known only from the is over 70% of the total number of species in Colorado drainage, reveals that the Sevier the state, occur in southern Utah. Possible River was connected to the Colorado drainage explanations of species diversity in the area in the geologic past and flowed south rather include speciation occurring in isolated areas, than north as it does today. Additional study of migration into the area from outside, and vicar- the Sevier River is needed desperately. This iance of ancestral populations. Only one species, river is a major source of freshwater for agri- Sweltsa cristata Surdick, is endemic to south- culture in southern Utah. With continued ern Utah. Lack of endemic species, especially population growth throughout southern Utah, given the isolated mountain ranges, argues this river will be depleted even further. Little against speciation having occurred recently or is known about the stoneflies and other aquatic frequently in the area. Virtually all species that macroinvertebrates of this river, but from pre- occur in southern Utah also occur outside vious collecting efforts this river appears strange southern Utah. Even Sweltsa cristata Surdick, indeed because it harbors so few stoneflies the only species currently endemic to south- compared with other rivers of similar size, ern Utah, may in the future be found to occur depth, and flow rate. Certainly, to preserve this in nearby mountains of Colorado. The distri- unique river, we need to understand it better. bution pattern for southern Utah is the result Our data also reveal that the Virgin River of colonization of the area from the outside or valley and the Zion National Park region of 2002] STONEFLIES OF SOUTHERN UTAH 73 the Virgin River are unique faunal regions. * C. vernalis Newport This river and its tributaries face increased ¦ Capnura intermontana Nelson and pressure as the human population of southern Baumann Utah continues to expand. Efforts to preserve o C. wanica (Frison) and study this river should be increased. * Eucapnopsis brevicauda (Claassen) * Isocapnia crinita (Needham and The occurrence of the isolated species Iso- Claassen) perla phalerata (Needham) and Isoperla pinta ¦ I. grandis (Banks) Frison from Panguitch Creek and Podunk ¦ I. hyalita Ricker Creek, respectively, illustrate how little we ¦ I. missourii Ricker know about which species occur in southern * I. vedderensis (Ricker) Utah. Undoubtedly, there are more species o Mesocapnia frisoni (Baumann and that occur in isolated areas that we do not yet Gaufin) know exist. These species also demonstrate * Utacapnia lemoniana (Nebeker and the fragility of freshwater ecosystems. Podunk Gaufin) Creek is a perfect example. An extremely *U. logana(Nebeker and Gaufin) ¦ U. poda (Nebeker and Gaufin) shallow, narrow creek meandering through a Family Leuctridae meadow used for cattle pasture, it already has ¦ Paraleuctra jewetti Nebeker and steeply cut banks typical of erosion due to Gaufin grazing. How typical this situation is in south- * P. occidentalis (Banks) ern Utah where many small creeks flow through * P. rickeri Nebeker and Gaufin land used by grazing cattle. It certainly did not * P. vershina Gaufin and Ricker look unique at first, and we easily could have * Perlomyia utahensis Needham and missed the single specimen of Isoperla pinta Claassen Frison had we come at a different time of year. Family Nemouridae How many more seemingly uninteresting creeks * Amphinemura banksi Baumann and Gaufin in southern Utah harbor hidden biological o A. mogollonica Baumann and Gaufin treasures? We fear many undiscovered organ- ¦ Malenka californica (Claassen) isms will be lost by increased demands for * M. coloradensis (Banks) water. Surely, more research needs to be done o M. flexura (Claassen) documenting and understanding species of ¦ M. tina (Ricker) stoneflies and other organisms inhabiting fresh- ¦ Podmosta decepta (Frison) water sources if we are to preserve our re- * P. delicatula (Claassen) sources in the face of increased tourism and * Prostoia besametsa (Ricker) * Zapada cinctipes (Banks) population growth in southern Utah. It is our ¦ hope that this study will enhance understand- Z. columbiana (Claassen) o Z. frigida (Claassen) ing of the biological properties of this unique *Z. haysi(Ricker) area of the world and enable wiser decisions Family Taeniopterygidae regarding conservation and management. * Doddsia occidentalis (Banks) * Oemopteryx fosketti (Ricker) LIST OF STONEFLIES OF UTAH (1999) * Taenionema pacificum (Banks) * T. pallidum (Banks) o occurs only in southern Utah ¦ T. uinta Stanger and Baumann ¦ occurs only in northern Utah ¦ Taeniopteryx nivalis (Fitch) * occurs in both northern and southern Utah Group Systellognatha Family Chloroperlidae Order Plecoptera * Alloperla severa (Hagen) Suborder Arctoperlaria ¦ Paraperla frontalis Banks Group Euholognatha * Plumiperla diversa (Frison) Family Capniidae ¦ Suwallia lineosa (Banks) ¦ Capnia coloradensis Claassen * S. pallidula (Banks) * C. confusa Claassen * Sweltsa borealis (Banks) * C. gracilaria Claassen * S. coloradensis (Banks) ¦ C. nana Claasen o S. cristata Surdick ¦ C. petila Jewett ¦ S. gaufini Baumann ¦ C. uintahi Gaufin *S. lamba(Needham and Claassen) * C. utahensis Gaufin and Jewett o Triznaka pintada (Ricker) 74 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

* T. signata (Banks) Raft River Mountains of northwestern Utah but * Utaperla sopladora Ricker was unreported by Houseman and Baumann Family Perlidae (1997). Triznaka diversa (Frison) was reassigned * Acroneuria abnormis (Newman) to the genus Plumiperla (Surdick 1985). Isoperla * Claassenia sabulosa (Banks) ebria has been synonymized under Isoperla ¦ Doroneuria sp. A * Hesperoperla pacifica (Banks) sobria (Hagen) (Szczytko and Stewart 1979), Family Perlodidae removing the former and adding the latter to * Cultus aestivalis (Needham and the state list. Skwala parallela has been syn- Claassen) onymized under the older name Skwala amer- * Diura knowltoni (Frison) icana (Needham and Claassen) (Zwick 1989). * Isogenoides colubrinus (Hagen) The following species were recorded from ¦ I. elongatus (Hagen) Utah in previous publications and are listed * I. zionensis Hanson here in this updated state list: Capnia petila * Isoperla fulva Claassen Jewett, Capnura intermontana Nelson and Bau- * I. longiseta Banks * I. mormona Banks mann, Doroneuria sp., Malenka tina (Ricker), * I. petersoni Needham and Claassen Taeniopteryx nivalis (Fitch) (Houseman and o I. phalerata (Needham) Baumann 1997), Suwallia lineosa (Banks), Acro- * I. pinta Frison neuria abnormis (Newman), and Perlesta decip- * I. quinquepunctata (Banks) iens (Hagen) (Baumann et al. 1977). The iden- * I. sobria (Hagen) tity of the Doroneuria species is unknown * Kogotus modestus (Banks) because to date only nymphs have been col- * Megarcys signata (Hagen) lected, and specific identification is possible ¦ Pictetiella expansa (Banks) only with adult males. This record is D. bau- * Skwala americana (Needham and Claassen) manni, D. theodora, or an undescribed species. Family Pteronarcyidae Capnia coloradensis Claassen occurs in the * Pteronarcella badia (Hagen) Raft River Mountains of northwestern Utah * Pteronarcys californica Newport but was omitted from the previous publication ¦ P. princeps Banks (Houseman and Baumann 1997). Baumann et TOTAL NUMBER OF SPECIES 81 al. (1977) record Perlesta placida as occurring in Uintah County. This species was synonym- EXPLANATION OF UPDATED RECORDS.—The ized under Perlesta decipiens (Hagen) (Stark following species have been synonymized 1989). We examined the Perlesta nymph from or reassigned in classification: Capnia cygna Sweetwater County, Wyoming, and are of the (Jewett), listed as occurring in the state (Bau- opinion that it is probably Perlesta decipiens mann 1973), does not occur in Utah and is (Hagen). This species is recorded from the removed from the state list. Capnia wanica Green River, Sweetwater County, Wyoming, (Frison) was reassigned to the genus Capnura and was thought to occur in Uintah County, (Nelson and Baumann 1987). Records previ- where the Green River flows from Wyoming ously listed as Paraleuctra sara are now listed into Utah. Baumann et al. (1977) record this as Paraleuctra vershina (Gaufin and Ricker) species as occurring in Uintah County; how- (Gaufin and Ricker 1974). The former species ever, there is no confirmed record of Perlesta has an eastern North American distribution from Uintah County. Dr. Boris Kondratieff, while the latter has a western North American who has collected extensively from the Green distribution. Taenionema nigripennis (Banks) River just over the Utah border in Colorado, has been synonymized under Taenionema pal- told us that he has not collected Perlesta from lidum (Banks) (Stanger and Baumann 1993), there. Consequently, we remove this species removing the former and adding the latter to from the Utah state list of stoneflies and explain the state list. Taenionema pacifica and Tae- the previously published record as an error. nionema pallida have been corrected to T. New state records added or confirmed from pacificum (Banks) and T. pallidum (Banks), this study are Isoperla phalerata (Needham), respectively (Steyskal 1976). Stanger and Bau- Malenka flexura (Claassen), and Sweltsa cristata mann (1993) list Taenionema uinta Stanger and Surdick (Surdick 1995). A single female of Baumann as occurring in several counties in Isoperla phalerata (Needham) collected by northern Utah. This species also occurs in the George Knowlton from Panguitch Creek is the 2002] STONEFLIES OF SOUTHERN UTAH 75 only specimen of this species in Utah. After f Pine Creek examining this specimen, we concur that it is g Pine Lake Campground seep I. phalerata (Needham). The distribution of h Poison Creek this species in other localities in the Central 2 Awapa Plateau (Wayne Co.) and Southern Rocky Mountains, however, does a Fremont River, Bicknell 3 Beaver Valley (Beaver Co.) make its presence in Utah reasonable. a Beaver Creek, Beaver Malenka flexura (Claassen) was recorded by b Birch Creek, SE Beaver Gaufin et al. (1966) as occurring in Utah. This c South Creek, S Beaver record was based on a single female collected 4 Boulder Mountain (Garfield Co.) at Huntsville, Weber County. After examining a Boulder Creek the specimen, Baumann (1973) found it to be- b Calf Creek long to M. californica (Claassen) and thus re- c Carcass Creek moved this species from the state list. We report d Durfey Creek 1 male specimen and 1 female specimen from e Pleasant Creek f Singletree Creek Lake Oowah, San Juan County, reinstating M. g Spruce Spring flexura (Claassen) to the state list of stonefly 5 Bull Valley Mountains (Washington Co.) species. a Magotsu Creek, N Veyo Sweltsa cristata Surdick, recently described b Pine Park Creek, Pine Park (Surdick 1995), is known only from the Abajo Campground and LaSal Mountains of southern Utah. Cur- c Slaughter Creek rently, this is the only species endemic to 6 Harmony Mountains (Iron Co.) southern Utah. Sweltsa cristata Surdick speci- a Iron Creek, W Cedar City mens from this area of southern Utah were 7 Hurricane Cliffs (Iron Co., Washington Co.) listed under S. lamba (Needham and Claassen) a Ash Creek in the past (Baumann et al. 1977). b Coal Creek, E Cedar City c LaVerkin Creek ANNOTATED LIST OF STONEFLY SPECIES 8 Markagunt Plateau (Iron Co.) OF SOUTHERN UTAH a Asay Creek, S Hatch b Bowery Creek, Parowan Canyon Distributional data are presented below in c Cedar Canyon Campground, near the following format: major physiographic Cedar Breaks National Monument province, physiographic region, geographic d Center Creek area, locality. Code numbers are assigned to e Clear Creek, above Panguitch Lake f Duck Creek, near Aspen Mirror Lake geographic areas, code letters to localities with- g Lowder Creek, E Cedar Breaks in the areas. Each species is then presented National Monument with a summary of distribution based on these h Mammoth Creek codes. Counties in which the geographic areas i Panguitch Creek, below Panguitch occur are presented in parentheses alongside Lake geographic areas to facilitate finding the local- j Parowan Creek, Parowan Canyon ities on maps of the areas (Fig. 4). Presentation k Red Creek, above Red Creek of the data is given in this format to save space. Reservoir, E Paragonah Anyone interested in the full data, available in l Yankee Lake Stream, above Parowan 9 Pahvant Range (Millard Co., Sevier Co.) an unpublished thesis (Call 1999), can contact a Chalk Creek, above Fillmore the authors, who will gladly provide the infor- b Clear Creek, Clear Creek Canyon mation. c Corn Creek, above Kanosh d Ivie Creek, above Scipio Lake Central Rocky Mountains e Oak Creek, above Oak City SOUTHERN HIGH PLATEAUS f Pole Creek, Clear Creek Canyon 1 Aquarius Plateau (Garfield Co.) g Willow Creek, W Salina a Antimony Creek 10 Pansaugaunt Plateau (Garfield Co., Kane b Birch Creek, Main Canyon, NW Co.) Escalante a Badger Creek c Center Creek, Osiris b Blubber Creek d Clay Creek, above Pine Lake c East Fork Creek e North Creek, NW Escalante d East Fork Sevier River 76 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

Fig. 4. Geographic areas of southern Utah collection sites. 2002] STONEFLIES OF SOUTHERN UTAH 77

e Kanab Creek, above Alton g Griffeth Creek, NE Beaver f Podunk Creek, S Tropic Reservoir h Lott Creek, near Fremont Indian State gSevier River Park 11 Pine Valley Mountains (Washington Co.) i Kents Lake a Ash Creek, near Sawyer Spring j Merchant Creek b Santa Clara River, Pine Valley k Mill Creek, SW of Castle Rock Mountains Campground c Leap Creek, near Pintura l Mud Lake, NE Beaver d Leeds Creek, Leeds Canyon m Pine Creek, Marysvale 12 Sanpitch Mountains (Sanpete Co.) n Pine Grove Creek, NE Beaver a Fountain Green Spring, Fountain o Poison Creek, S Mud Lake, NE Green Beaver b Maple Canyon, near Freedom p Puffer Lake Campground stream c Box Canyon, off Maple Canyon, near q Sevier River, Marysvale Canyon Freedom r Shelly Baldy Creek, Big John Flat, NE 13 Sevier Plateau (Garfield Co., Sevier Co.) Beaver a Abes Creek, above Gooseberry s Shingle Creek, Clear Creek Canyon Campground WASATCH PLATEAU b Bowery Creek, Fish Lake 15 Wasatch Plateau (Emery Co., Sanpete Co.) c Dairy Creek, Fish Lake a Becks Creek, above junction Seely d Fremont River, Mill Meadow Creek Reservoir b Bougler Creek e Fremont River, Zedds Meadow c Cedar Creek, NE Spring City f Gates Creek, below Gooseberry d Cottonwood Creek, above Fairview Campground e Ephraim Creek, Ephraim Canyon g Gooseberry Creek f Fairview Creek, Fairview Canyon h Ivie Creek g Ferron Creek i Little Lost Creek h Flat Canyon (Flat Canyon Spring, j Lost Creek O’dell Spring, Stanley Spring) k Mamoits Spring, near Fremont River i Gooseberry Creek, Flat Canyon l Monkey Fork, Sawmill Flat, SE j Huntington Creek, Huntington Monroe Canyon m Monroe Creek k Joes Valley Reservoir n Niotche Creek, above Gooseberry l Millstream, above Willow Lake Campground m Quitchupah Creek, Convulsion o Otter Creek, above Koosharem Canyon Reservoir n San Pitch River, Milburn p Peterson Creek, Kings Meadow o Sixmile Creek, E Sterling Canyon p Seely Creek, Olsen Canyon q Quitchupah Creek, near Emery q Thistle Creek, Coyote Springs near r Salina Creek Indianola s Sanford Creek, NE Panguitch Southern Rocky Mountains t Sawmill Creek, tributary of Sevenmile 16 Abajo Mountains (San Juan Co.) Creek, N Fish Lake a Dalton Springs u Sevenmile Creek, N Fish Lake b Johnson Creek, N Blanding v South Last Chance Creek c North Creek, above Dalton Springs w Twin Creeks, Fish Lake Campground x UM Creek, N Fish Lake 17 LaSal Mountains (Grand Co., San Juan y Water Creek, S Glenwood Fish Co.) Hatchery a Bear Creek, above Hidden Lake 14 Tushar Mountains (Beaver Co., Piute Co., b Castle Creek, above LaSal Mountain Sevier Co.) Loop Road a Beaver Creek c Fisher Creek b Bullion Canyon Creek, Marysvale d Horse Creek c City Creek, City Creek Campground e Mill Creek, below Oowah Lake d Cottonwood Creek f Oowah Lake, springs e Deer Creek, near Big Rock Candy g Pack Creek, Pack Creek Campground Mountain h Placer Creek f Fish Creek, Clear Creek Canyon i Warner Campground 78 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

Colorado Plateau 27 Vermillion Cliffs (Kane Co.) GRAND VALLEY SUBCENTER a Kanab Creek, Kanab 18 Grand Valley (Grand Co.) BEAVER DAM WASH SUBCENTER a Colorado River, near Moab 28 Beaver Dam Wash (Washington Co.) b Dolores River a Beaver Dam Wash, below Slaughter KAIPAROWITS PLATEAU Creek 19 Escalante River Drainage (Garfield Co.) b Beaver Dam Wash, East Fork a Calf Creek c Beaver Dam Wash, West Fork b Deer Creek Great Basin c Escalante River 29 Great Basin Desert (Iron Co.) d Main Creek a Beaver Creek, Milford e Steep Creek b Duncan Creek, E Newcastle f Birch Creek 20 Henry Mountains (Garfield Co.) Species are presented in alphabetical order a Crescent Creek by family, genus, and species. The classifica- b Dark Canyon tion system follows that used by Stewart et al. c Dugout Creek (1988) as outlined by Stark et al. (1986). d Slate Creek e South Copper Creek f Sweetwater Creek Order Plecoptera g Trachyte Creek Suborder Arctoperlaria h Willow Spring Group Euholognatha 21 Paria River Drainage (Kane Co.) Family Capniidae a Cottonwood Creek bParia River c Willis Creek Capnia confusa Claassen d Henrieville Creek GEOGRAPHIC RANGE.—Coast, Cascade, and SAN JUAN SUBCENTER Rocky Mountains. 22 Canyonlands (San Juan Co.) DISTRIBUTION IN SOUTHERN UTAH.—Aquar- a Big Springs, Canyonlands National ius Plateau: e, Beaver Valley: a, Boulder Moun- Park tain: ab, Harmony Mountains: a, Markagunt b Cave Springs, Canyonlands National Plateau: abh, Pahvant Range: be, Pine Valley Park c Indian Creek, Newspaper Rock State Mountains: b, Sevier Plateau: m, Tushar Moun- Park tains: aem, Wasatch Plateau: fj, Abajo Moun- d Squaw Flat, Canyonlands National tains: b, LaSal Mountains: egh, Capitol Reef: ce. Park DISCUSSION.—This species is the most wide- 23 Painted Desert (Kane Co., San Juan Co.) spread species of the genus in North America a Colorado River (Nelson and Baumann 1989). It occurs com- b Lime Creek, near Mexican Hat monly throughout Utah and has been found in c San Juan River all physiographic regions in southern Utah SAN RAFAEL SUBCENTER except the Great Basin. 24 Capitol Reef (Wayne Co.) a Carcass Creek, S Grover Capnia gracilaria Claassen b Deep Creek c Fish Creek GEOGRAPHIC RANGE.—Coast, Cascade, and d Fremont River, Capitol Reef National Rocky Mountains, and Northern Great Plains. Park DISTRIBUTION IN SOUTHERN UTAH.—Aquar- e Pleasant Creek ius Plateau: e, Beaver Valley: b, Boulder Moun- f Singletree Creek tain: cfg, Bull Valley Mountains: ab, Markagunt g Sulphur Creek Plateau: bej, Pahvant Range: a, Pine Valley 25 Green River (Emery Co. and Uintah Co.) Mountains: b, Sevier Plateau: gmsx, Tushar a Green River, near Green River town Mountains: bemq, Wasatch Plateau: dfjl, Abajo b Green River, Jensen Mountains: ab, LaSal Mountains: bg, Virgin ST. GEORGE SUBCENTER 26 Virgin River (Kane Co., Washington Co.) River: c. a Emerald Pools, Zion National Park DISCUSSION.—A common species through- b Virgin River, East Fork out western North America, it is widespread c Virgin River, North Fork, Zion throughout southern Utah except in the Great National Park Basin. 2002] STONEFLIES OF SOUTHERN UTAH 79

Capnia utahensis Gaufin and Jewett Isocapnia crinita (Needham and Claassen) TYPE LOCALITY.—Beaver Creek, Beaver Co., GEOGRAPHIC RANGE.—Rocky Mountains. Utah. DISTRIBUTION IN SOUTHERN UTAH.—Pahvant GEOGRAPHIC RANGE.—Basin and Range. Range: b, Wasatch Plateau: j. DISTRIBUTION IN SOUTHERN UTAH.—Beaver DISCUSSION.—This rare species is known Valley: abc, Markagunt Plateau: bjl, Pahvant from only 2 areas in southern Utah. Full data are Range: abef, Pine Valley Mountains: abcd, presented for this species. Sevier Plateau: agmpz, Tushar Mountains: Emery Co.: Huntington Creek, between befhmq, Virgin River: a, Beaver Dam Wash: a, North and South Hughes Canyons, 2-V-1978, Great Basin Desert: b. E.C. Devenport and R.N. Winget, 2 Ɋ; Hunt- DISCUSSION.—This species ranges from the ington Creek, Left Fork, 4-V-1971, E.C. southern Sierra Nevada across the Great Basin Devenport and R.N. Winget, 2 ɉ 1 Ɋ; Hunt- of Nevada and California to the edge of the ington Creek, Stuart Station, 14-IV-1977, R.W. Colorado Plateau in central Utah, which Baumann and R.N. Winget, 5 ɉ 1 Ɋ 1 nymph; appears to be the eastern distributional limit. 4-V-1978, R.W. Baumann and R.N. Winget, 6 ɉ 7 Ɋ. Sevier Co.: Clear Creek, Hwy 89, Capnia vernalis Newport Sevier, 29-III-1987, R.W. Baumann, 5-IV-1968, 2 ɉ 1 Ɋ; R.W. Baumann and P. Zwick, 1 ɉ 1 Ɋ. GEOGRAPHIC RANGE.—Northern North America south to New Mexico. Isocapnia vedderensis (Ricker) DISTRIBUTION IN SOUTHERN UTAH.—Pahvant Range: b, Sevier Plateau: v. GEOGRAPHIC RANGE.—Cascade and Rocky DISCUSSION.—This northern species extends Mountains. as far south as New Mexico. It occurs in DISTRIBUTION IN SOUTHERN UTAH.—Wasatch northern and southern Utah. Nelson and Bau- Plateau: j, Capitol Reef: deg. mann (1989) list this species as occurring in DISCUSSION.—This rare species is known Grand County in addition to the regions listed from only 2 areas in southern Utah. Wayne herein. County records have not previously been pub- lished and are given herein. Capitol Reef is Capnura wanica (Frison) the southern limit of its distribution. Wayne Co.: Fremont River, jct Sulphur GEOGRAPHIC RANGE.—Southern Rocky Creek, Capitol Reef National Park, 18-III-1989, Mountains, Colorado Plateau, and Great Basin. R.W. Baumann, 8 ɉ 6 Ɋ; Pleasant Creek, Lower DISTRIBUTION IN SOUTHERN UTAH.—Beaver Canyon, Capitol Reef National Park, 4-IV- Valley: abc, Boulder Mountain: ab, Bull Valley 1994, J.F. MacDonald and C. Brammer, 1 Ɋ Mountains: b, Hurricane Cliffs: c, Pahvant (PERC). Range: b, Pine Valley Mountains: bc, Escalante River Drainage: c, Henry Mountains: acfg, Mesocapnia frisoni (Baumann and Gaufin) Escalante River Drainage: ace, Kaiparowits TYPE LOCALITY.—Ash Creek, Anderson Junc- Plateau: a, Paria River Drainage: abc, Virgin tion, Washington County, Utah. River: b, Beaver Dam Wash: ab, Great Basin GEOGRAPHIC RANGE.—Southwestern United Desert: b. States. DISCUSSION.—Southern Utah is in the mid- DISTRIBUTION IN SOUTHERN UTAH.—Bull dle of the total range of this species. Valley Mountains: a, Hurricane Cliffs: c, Pine Valley Mountains: abd, Beaver Dam Wash: a. Eucapnopsis brevicauda (Claassen) DISCUSSION.—Southern Utah represents the GEOGRAPHIC RANGE.—Coast, Cascade, and northern limit of this species. It occurs along Rocky Mountains, and Sierra Nevada. the Colorado Plateau and Southern Rocky Moun- DISTRIBUTION IN SOUTHERN UTAH.—Beaver tain portions of Washington County, but not Valley: b, Markagunt Plateau: h, Pahvant Range: into the Great Basin. This is the only species a, Sevier Plateau: m, Tushar Mountains: a, Wa- in southern Utah that is unique to the Virgin satch Plateau: j, Abajo Mountains: ab. River Valley Province, revealing that this prov- DISCUSSION.—This species is widespread ince is a distinct faunal area. Other species but uncommon in southern Utah. occurring in the area are either widespread 80 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1 throughout southern Utah or show affinities DISTRIBUTION IN SOUTHERN UTAH.—Sevier with the Central Rockies. Plateau: m, Abajo Mountains: bc. DISCUSSION.—This species is widespread Utacapnia lemoniana (Nebeker and Gaufin) throughout the mountains of western North GEOGRAPHIC RANGE.—Central Rocky Moun- America. It occurs in the Central and South- tains. ern Rocky Mountain regions of southern Utah. DISTRIBUTION IN SOUTHERN UTAH.—Aquar- Paraleuctra females have been reported from ius Plateau: ef, Beaver Valley: b, Markagunt Pla- Pleasant Creek, Boulder Mountain (Brammer teau: e, Pahvant Range: b, Wasatch Plateau: jn. 1997); these are probably P. occidentalis. How- DISCUSSION.—Southern Utah is the south- ever, males are needed to confirm the species. ern limit of this species, which is common in Because of the few available records from the northern Utah, but does not extend into the area, full data are given below. Southern Rocky Mountains, Colorado Plateau, San Juan Co.: Johnson Creek, 14 mi NW of or Great Basin regions of southern Utah. Blanding, 21-IV-1986, R.W. Baumann and S.A. Wells, 4 ɉ 2 Ɋ; North Creek, above Dalton Utacapnia logana (Nebeker and Gaufin) Springs Campground, 7 mi W of Monticello, GEOGRAPHIC RANGE.—Central and Southern 11-V-1976, K.W. Stewart and S.W. Szczytko, 5 Rocky Mountains. ɉ 4 Ɋ; 5-II-1976, 1 N. Sevier Co.: Monroe DISTRIBUTION IN SOUTHERN UTAH.—Aquar- Creek, Monrovian Park, 29-IV-1998, R.G. Call, ius Plateau: ef, Boulder Mountain: ab, Harmony 1 ɉ. Mountains: a, Hurricane Cliffs: b, Pahvant Range: de, Sevier Plateau: grs, Wasatch Plateau: Paraleuctra rickeri Nebeker and Gaufin dfj, LaSal Mountains: eg, Escalante River Drainage: d, Virgin River: c. GEOGRAPHIC RANGE.—Rocky Mountains and DISCUSSION.—This species was previously Alaska. recorded from few locations in southern Utah, DISTRIBUTION IN SOUTHERN UTAH.—Capitol but it is now known to be widespread in this Reef: e. area. New county records with full data are Discussion.—This is the only known record given below. of this species in southern Utah. Full data are Garfield Co.: Main Creek, Main Canyon, given herein. 18-II-2000, R.W. Baumann, 1 Ɋ; North Creek, Garfield Co.: Pleasant Creek, 24-VI-1995, above North Creek Reservoir, 18-II-2000, J.F. MacDonald and C. Brammer, 2 ɉ 2 Ɋ R.W. Baumann, 1 Ɋ; Pine Creek, Death Box (PERC). Hollow Trailhead, 18-II-2000, R.W. Baumann, ɉ Ɋ 4 20 . Grand Co.: Mill Creek, Hwy 191, Paraleuctra vershina Gaufin and Ricker Moab, 4-II-1985, R.W. Baumann and C.R. Nelson, 1 ɉ 3 Ɋ. Kane Co.: Virgin River, North GEOGRAPHIC RANGE.—Coast, Cascade, and Fork, above Zion Narrows, 5-II-1977, R.W. Rocky Mountains, and Sierra Nevada. Baumann, 24 ɉ 11 Ɋ. San Juan Co.: Pack DISTRIBUTION IN SOUTHERN UTAH.—Wasatch Creek, Pack Creek Campground, LaSal Moun- Plateau: q. tains, 5-II-1985, R.W. Baumann and C.R. Nel- DISCUSSION.—Baumann et al. (1977) list son, 2 ɉ. Sanpete Co.: Cottonwood Creek, this species as occurring abundantly in creeks above Fairview, 19-III-1988, S.A. Wells, 21 ɉ and rivers; however, it is documented from 11 Ɋ; Fairview Creek, Fairview Canyon, 30- only a single locality in southern Utah. Its III-1978, R.W. Baumann, 7 ɉ 1 Ɋ. Washing- absence from southern Utah is perplexing in ton Co.: Virgin River, North Fork, below Zion light of its presence in the Central Rockies of Narrows, Zion National Park, 20-II-1972, R.W. northern Utah and the Southern Rockies of Baumann and R.N. Winget, 1 Ɋ. Colorado and New Mexico. Its distribution pattern reveals a border where the Wasatch Family Leuctridae Mountain Range and the Wasatch Plateau meet. Full data are presented. Paraleuctra occidentalis (Banks) Sanpete Co.: Thistle Creek, Coyote Springs GEOGRAPHIC RANGE.—Coast, Cascade, and stream, 10-VI-1986, S.M. Clark and B.S. Clark, Rocky Mountains, and Sierra Nevada. 5 ɉ 4 Ɋ. 2002] STONEFLIES OF SOUTHERN UTAH 81

Perlomyia utahensis Needham and Claassen DISCUSSION.—This species, although wide- GEOGRAPHIC RANGE.—Coast, Cascade, and spread in southern Utah, is not recorded from Rocky Mountains. northern Utah. It occurs in Southern Rocky DISTRIBUTION IN SOUTHERN UTAH.—Pahvant Mountain faunal regions as well as Colorado Range: c, Sevier Plateau: p. Plateau, Great Basin, and Central Rocky Moun- DISCUSSION.—This species, previously known tain faunal regions. It is unclear why it occurs only from Millard County in southern Utah, is in the Central Rockies of southern Utah but now known from Sevier County as well. Full does not extend into the Central Rockies of data for these 2 localities are given. northern Utah. The northern limit of its range Millard Co.: Kanosh Canyon, 27-V-1939, in Sanpete County coincides with the transi- G.F. Knowlton and F.C. Harmston, 1 ɉ 1 Ɋ. tion from the Wasatch Plateau to the Wasatch Sevier Co.: Monroe Creek, Monrovian Park, Mountain Range, providing further evidence 20-VI-1998, R.G. Call, 2 Ɋ. of a faunal division between these 2 areas.

Family Nemouridae Malenka coloradensis (Banks) GEOGRAPHIC RANGE.—Central and South- Amphinemura banksi Baumann and Gaufin ern Rocky Mountains. GEOGRAPHIC RANGE.—Rocky Mountains. DISTRIBUTION IN SOUTHERN UTAH.—Aquar- DISTRIBUTION IN SOUTHERN UTAH.—Abajo ius Plateau: cdg, Boulder Mountain: deg, Mark- Mountains: abd, LaSal Mountains: ae. agunt Plateau: cj. Pansaugaunt Plateau: ef, Paria DISCUSSION.—This species is limited to the River Drainage: d, Pine Valley Mountains: bd, Abajo and LaSal Mountains in southern Utah. Sevier Plateau: kw, Tushar Mountains: gi, Abajo This pattern agrees with the placement of the Mountains: ab, LaSal Mountains: gi, Henry Abajo and LaSal Mountains as a part of the Mountains: ae, Virgin River: b. Southern Rocky Mountain faunal region con- DISCUSSION.—This species is widespread nected to the Southern Rockies of Colorado. in southern Utah in the Colorado Plateau, Although both this species and its sister species Great Basin, Central Rocky Mountain, and A. mogollonica (Baumann and Gaufin) occur Southern Rocky Mountain faunal regions. It in southern Utah (Baumann and Gaufin 1972), does not occur in northern Utah (Baumann their ranges do not overlap. Amphinemura 1970). Gaufin et al. (1966) incorrectly reported (Baumann and Gaufin) occurs commonly banksi it as occurring in Weber County and Salt in Colorado and sporadically in northern Utah, Lake County as did Baumann et al. (1977) in but it is conspicuously absent from all of south- Duchesne County. Instead, M. californica ern Utah except the Abajo and LaSal Moun- (Claassen) occurs commonly throughout north- tains. The presence of A. banksi (Baumann ern Utah, while M. coloradensis (Banks) is lim- and Gaufin) in the Abajo and LaSal Mountains ited to the southern part of the Central Rock- of southern Utah and its absence from the ies. There is a division between the 2 species Central Rocky Mountains in the remainder of southern Utah indicate that this species origi- within the Central Rocky Mountains. This dis- nally occurred in the Abajo and LaSal Moun- tribution pattern indicates separate faunal tains or migrated there from the southern regions for the Northern High Plateau sub- Rocky Mountains of Colorado rather than from center and the Southern High Plateau subcen- northern Utah. ter within the Central Rockies. Amphinemura mogollonica Baumann and Malenka flexura (Claassen) Gaufin GEOGRAPHIC RANGE.—Coast, Cascade, and GEOGRAPHIC RANGE.—Southern Rocky Rocky Mountains. Mountains. DISTRIBUTION IN SOUTHERN UTAH.—LaSal DISTRIBUTION IN SOUTHERN UTAH.—Aquar- Mountains: f. ius Plateau: beg, Awapa Plateau: a, Boulder DISCUSSION.—This species is a new state Mountain: de, Markagunt Plateau: fhj, Pah- record for Utah. It is rare and has been col- vant Range: cg, Pansaugaunt Plateau: e, Paria lected only from the LaSal Mountains. Full River Drainage: d, Sevier Plateau: begimoq- data are presented below. Its distribution pat- tuwxy, Tushar Mountains: cks, Wasatch Plateau: tern agrees with the placement of the LaSal dfj, Virgin River: bc. Mountains as a part of the Southern Rocky 82 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

Mountains faunal region connected to the the Southern Rocky Mountains of Colorado. Southern Rockies of Colorado. Complete data are given for the only known San Juan Co.: springs entering Lake specimens. We regret having neither the time Oowah, LaSal Mountains, 20-VI-1987, C.R. nor resources to collect from this area dur- Nelson, 1 ɉ 1 Ɋ. ing our study to verify that this species is still present. Given the size of our study area and Podmosta delicatula (Claassen) the fact that many stonefly species emerge GEOGRAPHIC RANGE.—Coast, Cascade, and as adults during a brief period in the spring, Rocky Mountains. we simply could not be everywhere we wanted DISTRIBUTION IN SOUTHERN UTAH.—Wasatch to be. Plateau: bj. San Juan Co.: Horse Creek, LaSal Moun- tains, 9-VI-1954, Ross and Ross, 1 ɉ (INHS); DISCUSSION.—This species’ distribution pat- Johnson Creek, 19 mi N of Blanding, Abajo tern in southern Utah indicates this area is ɉ part of the Central Rocky Mountain faunal Mountains, 18-VI-1946, S. and D. Muliak, 1 . region. Zapada haysi (Ricker) Prostoia besametsa (Ricker) GEOGRAPHIC RANGE.—Coast, Cascade, and Rocky Mountains, and Sierra Nevada. GEOGRAPHIC RANGE.—Coast, Cascade, and Rocky Mountains, and Sierra Nevada. DISTRIBUTION IN SOUTHERN UTAH.—Aquar- ius Plateau: d, Beaver Valley: a, Markagunt DISTRIBUTION IN SOUTHERN UTAH.—Beaver Valley: b, Boulder Mountain: e, Bull Valley Plateau: hj, Sevier Plateau: jlmx, Tushar Moun- tains: p, Wasatch Plateau: jq, Abajo Mountains: Mountains: b, Markagunt Plateau: ehj, Pahvant b, Capitol Reef: e. Range: ac, Sevier Plateau: gmv, Tushar Moun- DISCUSSION.—This species is widely dis- tains: aemp, Wasatch Plateau: bdej, Abajo tributed across southern Utah but is not com- Mountains: b, LaSal Mountains: g, Virgin mon. Brammer (1997) reports females from River: c. Pleasant Creek, Boulder Mountain. Without DISCUSSION.—This species is common associated males, the species identification is throughout its range including southern Utah. uncertain. Zapada cinctipes (Banks) GEOGRAPHIC RANGE.—Coast, Cascade, and Family Taeniopterygidae Rocky Mountains, and Sierra Nevada. Doddsia occidentalis (Banks) DISTRIBUTION IN SOUTHERN UTAH.—Aquar- GEOGRAPHIC RANGE.—Western North Amer- ius Plateau: cd, Beaver Valley: b, Boulder ica. Mountain: e, Markagunt Plateau: ce, Pahvant DISTRIBUTION IN SOUTHERN UTAH.—Har- Range: ade, Pine Valley Mountains: bd, San- mony Mountains: a, Sevier Plateau: m, Tushar pitch Mountains: ab, Sevier Plateau: acgmpxz, Mountains: aem. Tushar Mountains: hmnop, Wasatch Plateau: DISCUSSION.—Although recorded in north- abdhjlq, Abajo Mountains: abc, LaSal Moun- ern Utah, this species was previously tains: bg, Capitol Reef: e, Virgin River: bc. unknown from southern Utah. We document DISCUSSION.—This species is widespread it in southern Utah with the following new across southern Utah and can be very common. county records. The distribution pattern shows a Central Rocky Mountain faunal affin- Zapada frigida (Claassen) ity at the edge of the Great Basin. GEOGRAPHIC RANGE.—Coast, Cascade, and Iron Co.: Iron Creek, 10 mi above Cedar Rocky Mountains, and Sierra Nevada. City, 29-III-1987, R.W. Baumann and P. Zwick, DISTRIBUTION IN SOUTHERN UTAH.—Abajo 9 ɉ 4 Ɋ 1 exuvium. Piute Co.: Beaver Creek, Mountains: b, LaSal Mountains: d. 21-IV-1977, R.D. Williams, 2 nymphs; Deer DISCUSSION.—This uncommon species is Creek, Hwy 89, Big Rock Candy Mountain, 6- known in Utah only from the Abajo and LaSal IV-1968, R.W. Baumann, 1 Ɋ; Pine Creek, Mountains, but it occurs commonly in nearby Marysvale, 28-III-1987, R.W. Baumann and P. mountains of Colorado, showing that these Zwick, 1 ɉ. Sevier Co.: Monroe Creek, Mon- southern Utah mountains are faunally part of rovian Park Campground, 21-II-1986, R.W. 2002] STONEFLIES OF SOUTHERN UTAH 83

Baumann and C.R. Nelson, 14 nymphs; 28-III- L.L. Wu, 1 ɉ (USUC). Iron Co.: Parowan 1987, R.W. Baumann and P. Zwick, 9 ɉ 13 Ɋ. Creek, 2 mi N of Brian Head Ski Area, 2-VI- 1998, R.G. Call, 1 ɉ 1 Ɋ. Sanpete Co.: Cot- Oemopteryx fosketti (Ricker) tonwood Creek, 6 mi above Fairview, 29-V- ɉ Ɋ GEOGRAPHIC RANGE.—Western North Amer- 1981, S.M. Clark and J.A. Stanger, 10 9 ; ica. Thistle Creek, Coyote Springs, 25-V-1981, Ɋ DISTRIBUTION IN SOUTHERN UTAH.—Grand S.M. Clark, 1 . Valley: ab, Green River: a. DISCUSSION.—This species is limited in Group Systellognatha southern Utah to large rivers. Family Chloroperlidae Alloperla severa (Hagen) Taenionema pacificum (Banks) GEOGRAPHIC RANGE.—Coast, Cascade, and GEOGRAPHIC RANGE.—Western United States Rocky Mountains. and Canada. DISTRIBUTION IN SOUTHERN UTAH.—Mark- DISTRIBUTION IN SOUTHERN UTAH.—Boul- agunt Plateau: k, Pahvant Range: b, Pine Val- der Mountain: b, Pahvant Range: b, Pine Valley ley Mountains: d, Sanpitch Mountains: dku, Mountains: d, Tushar Mountains: q, Wasatch Wasatch Plateau: j. Plateau: j, Grand Valley: c, Escalante River DISCUSSION.—This species was previously Drainage: abce, Paria River Drainage: b, Capi- unrecorded from southern Utah. Previously tol Reef: dg, Virgin River: abc. unreported records and specimens we col- DISCUSSION.—Several new records show lected from several localities reveal its occur- this to be an abundant and widespread species rence in the Central Rocky Mountain region in rivers and larger creeks of southern Utah. of southern Utah. Its presence in the Pine Val- ley Mountains of Washington County in the Taenionema pallidum (Banks) southwestern corner of the state confirms that GEOGRAPHIC RANGE.—Western United States these mountains are a continuation of the Cen- and Canada. tral Rockies. Because these records are new DISTRIBUTION IN SOUTHERN UTAH.—Boulder county records, the complete data are presented. Mountain: e, Markagunt Plateau: hj, Tushar Emery Co.: Huntington Creek, Hunting- Mountains: p, Wasatch Plateau: djkq, LaSal ton Canyon, below Electric Lake, 12-VI-1972, Mountains: df. E.C. Devenport and R.N. Winget, 2 ɉ; 20-VI- DISCUSSION.—This species was previously 1972, 1 ɉ 1 Ɋ; Huntington Creek, upper, 20- recorded in southern Utah only from San Juan VI-1972, E.C. Devenport and R.N. Winget, 3 County. We collected this species from several ɉ 1 Ɋ. Iron Co.: Red Creek, above Red Creek new localities. Using these new records plus Reservoir, 15-VII-1998, B.O. Huntsman and those of previously unreported specimens K.L. Huntsman, 1 Ɋ. Sevier Co.: Clear Creek, which we verified, we now report this species Hwy 4 at Hwy I-70, 27-VI-1998, R.G. Call, 2 as occurring in several other southern Utah ɉ 2 Ɋ; Fremont River, above Mill Meadow counties. New county records are included Reservoir, 29-VI-1989, R.W. Baumann, 4 ɉ 10 below with full data. Ɋ; Mamoits Spring, jct Fremont River, 24- Beaver Co.: Puffers Lake, Beaver Canyon, VIII-1984, R.W. Baumann, 1 Ɋ; Sevenmile 11-VI-1943, G.F. Knowlton, 1 ɉ 3 Ɋ (INHS); Creek, Hwy 36, above Johnson Valley Reser- 1 ɉ 2 Ɋ (INHS). Emery Co.: Huntington voir, 29-VI-1989, R.W. Baumann, 4 ɉ 3 Ɋ. Creek, Coal Canyon, 24-VI-1975, M.K. Reichert Washington Co.: Leeds Canyon, 16-VI-1978, and R.N. Winget, 4 ɉ 1 Ɋ; Olsen Canyon, W.J. Hanson and G.F. Knowlton, 1 ɉ 5 Ɋ above Joes Valley Reservoir, 30-VI-1992, R.W. (USUC); Leeds Creek, jct Twin Pine Creek, Baumann, 1 Ɋ. Garfield Co.: Mammoth Creek, 31-V-1985, C.R. Nelson, 2 Ɋ. Mammoth Springs Campground, 2-VI-1998, R.G. Call, 7 ɉ 3 Ɋ; Pleasant Creek, 7 km W of Plumiperla diversa (Frison) Pleasant Creek Campground, Hwy 12, Boul- GEOGRAPHIC RANGE.—Coast, Cascade, and der Mountains, 17-VI-1994, J.F. MacDonald, 6 Rocky Mountains, and Sierra Nevada. Ɋ (PERC); 24-VI-1995, J.F. MacDonald and DISTRIBUTION IN SOUTHERN UTAH.—Boulder C.A. Brammer, 6 ɉ 26 Ɋ (PERC). Grand Co.: Mountain: e, Sevier Plateau: x, Abajo Moun- Lake Oowah, LaSal Mountains, 10-VI-1963, tains: b, LaSal Mountains, ef. 84 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

DISCUSSION.—Garfield, Grand, and San Juan States, including the mountains of southern are new county records with the full data pre- Utah. sented. Together with previous records from Sevier County, the data reveal this species is Sweltsa coloradensis (Banks) widely distributed throughout southern Utah. GEOGRAPHIC RANGE.—Coast, Cascade, and It is found in the Central Rockies, Southern Rocky Mountains, and Sierra Nevada. Rockies, and the Colorado Plateau regions. DISTRIBUTION IN SOUTHERN UTAH.—Boulder Garfield Co.: Panguitch Creek, below Pan- Mountain: e, Pahvant Range: be, Sanpitch guitch Lake, 16-VII-1959, A.R. Gaufin, 1 ɉ 4 Ɋ; Mountains: ab, Sevier Plateau: jmuy, Tushar Pleasant Creek, Pleasant Creek Campground, Mountains: kp, Wasatch Plateau: jq, LaSal Hwy 12, Boulder Mountains, 20-VI-1986, R.W. Mountains: acd. Baumann, 3 ɉ 12 Ɋ; 24-VI-1995, J.F. Mac- DISCUSSION.—Widespread throughout the Donald and C.A. Brammer, 3 ɉ (PERC). Grand mountains of the western United States, this Co.: Oowah Campground, LaSal Mountains, species is the most common and widespread 11-VI-1963, W.J. Hanson, 1 ɉ (USUC). San chloroperlid species in southern Utah. Juan Co.: Mill Creek, LaSal Mountains, 9-VI- 1954, 2 ɉ (INHS); LaSal Mountains (probably Sweltsa cristata Surdick Mill Creek), 9-VI-1954, 1 ɉ (INHS); 20 miles TYPE LOCALITY.—Johnson Creek, 19 miles SW of Monticello, 9-VII-1970, J.L. Petty, 2 Ɋ north of Blanding, San Juan County, Utah. (USUC). GEOGRAPHIC RANGE.—Abajo and LaSal Mountains, Utah. Suwallia pallidula (Banks) DISTRIBUTION IN SOUTHERN UTAH.—Abajo Mountains: b, LaSal Mountains: f. GEOGRAPHIC RANGE.—Coast, Cascade, and Rocky Mountains, and Sierra Nevada. DISCUSSION.—This species described by Surdick (1995) is extremely limited in distri- DISTRIBUTION IN SOUTHERN UTAH.—Beaver Valley: ab, Boulder Mountain: e, Markagunt bution. The only known specimens have been Plateau: j, Sevier Plateau: lmuxy, Tushar Moun- collected from the Abajo and LaSal Mountains tains: jloprs, Wasatch Plateau: deijkp. of southern Utah. A series of 40 male and 20 female specimens collected by C.R. Nelson DISCUSSION.—This is a very widespread near Lake Oowah were in excellent condition, species in the western United States, occur- allowing us to examine the range of morphol- ring in the Central Rocky Mountain and Col- ogy exhibited in the species, particularly the orado Plateau regions of southern Utah. In male epiproct and lamella on the aedeagus. addition to our findings, Baumann et al. (1977) We confirm S. cristata as a valid species. Fur- list this species as occurring in San Juan ther collecting in nearby mountains of Col- County; however, we could not find specimens orado may reveal its presence there, which to confirm this record and did not collect any would be strong evidence of previous water new specimens from this area during our connections between these isolated mountains study. Bret Huntsman assisted us with collect- of southern Utah and nearby mountains of ing and obtained a single adult male from Iron Colorado. Sweltsa cristata specimens from this County, resulting in a new county record for area of southern Utah were listed under S. which full data are presented. lamba in the past (Baumann et al. 1977). Iron Co.: unnamed spring entering Parowan Creek, Hwy 143 jct Hwy 49, 14-VII-1998, Sweltsa lamba (Needham and Claassen) B.O. Huntsman, 1 ɉ. GEOGRAPHIC RANGE.—Coast, Cascade, and Rocky Mountains. Sweltsa borealis (Banks) DISTRIBUTION IN SOUTHERN UTAH.—Beaver GEOGRAPHIC RANGE.—Coast, Cascade, and Valley: b, Pine Valley Mountains: bd, Henry Rocky Mountains, and Sierra Nevada. Mountains: bd, Virgin River: b. DISTRIBUTION IN SOUTHERN UTAH.—Aquar- DISCUSSION.—This species is common in ius Plateau: d, Pine Valley Mountains: bd, San- the Central Rocky Mountain region of north- pitch Mountains: b, Sevier Plateau: eklx, Tushar ern Utah. It also occurs in the Beaver Moun- Mountains: go, LaSal Mountains: ab. tains, Pine Valley Mountains, Virgin River, and DISCUSSION.—This species is widespread Henry Mountains of southern Utah. This dis- throughout the mountains of the western United tribution reveals the continuity of the Central 2002] STONEFLIES OF SOUTHERN UTAH 85

Rocky Mountains in Utah. It also shows that Beaver Co.: Puffers Lake, 11-VI-1943, G.F. the Henry Mountains are faunally more simi- Knowlton and P.E. Telford, 1 ɉ (holotype). lar to the rest of southern Utah than to the Sanpete Co.: Thistle Creek, Coyote Springs, Abajo and LaSal Mountains. Baumann et al. stream, E of Indianola, 10-VI-1986, S.M. (1977) list the occurrence of this species in Clark and B.S. Clark, 1 Ɋ. San Juan County; however, this listing was from specimens that are now included in Family Perlidae Sweltsa cristata (Surdick 1995). Acroneuria abnormis (Newman) Triznaka pintada (Ricker) GEOGRAPHIC RANGE.—United States and Canada. GEOGRAPHIC RANGE.—Coast, Cascade, and DISTRIBUTION IN SOUTHERN UTAH.—Sevier Rocky Mountains, and Sierra Nevada. Plateau: v, Grand Valley: a, Green River: a. DISTRIBUTION IN SOUTHERN UTAH.—Abajo DISCUSSION.—The distribution of this Mountains: b, LaSal Mountains: ac. species in southern Utah presents one of the DISCUSSION.—This species occurs in the most interesting findings of our study. It is Southern Rocky Mountains of Colorado, but uncommon in southern Utah, having been in Utah it is limited to the Abajo and LaSal recorded previously only from the large rivers Mountains, providing another example of the of the Colorado drainage, the Colorado, Green, faunal similarities between these isolated and White rivers, which all flow to the south- mountains of southern Utah and the Southern west. We record it from the Sevier River, a Rocky Mountains of Colorado. much smaller river that flows north. The pres- ence of A. abnormis in the Sevier River indi- Triznaka signata (Banks) cates that it was previously connected to the GEOGRAPHIC RANGE.—Coast, Cascade, and Colorado drainage. Full data for the Sevier Rocky Mountains. River specimens are given. DISTRIBUTION IN SOUTHERN UTAH.—Beaver Sevier Co.: Sevier River, Hwy 89, Big Rock Valley: ab, Boulder Mountain: a, Markagunt Candy Mountain, 7-VII-1999, R. Larsen, 1 Ɋ; Plateau: hi, Pahvant Range: be, Pansaugaunt Sevier River jct Elsinore canal head, 17-IV- Plateau: g, Pine Valley Mountains: bd, Sevier 1999, J. Paul, 1 nymph. Plateau: dej, Tushar Mountains: f, Wasatch Pla- (Banks) teau: j, LaSal Mountains: g, Canyonlands: c. Claassenia sabulosa DISCUSSION.—This species is widespread GEOGRAPHIC RANGE.—Western North Amer- in southern Utah. ica and eastern Canada. DISTRIBUTION IN SOUTHERN UTAH.—Pahvant Utaperla sopladora Ricker Range: b, Sevier Plateau: dj, Tushar Moun- tains: f, Grand Valley: a. TYPE LOCALITY.—Puffers Lake, Beaver Co., DISCUSSION.—Nymphs of this species can Utah. be abundant in larger creeks of southern Utah. GEOGRAPHIC RANGE.—Coast, Cascade, and The species occurs sporadically in both the Rocky Mountains. Central and Southern Rocky Mountains. DISTRIBUTION IN SOUTHERN UTAH.—Tushar Mountains: p, Wasatch Plateau: q. Hesperoperla pacifica (Banks) DISCUSSION.—This species is uncommon in GEOGRAPHIC RANGE.—Western North Amer- Utah. The complete data are given for the only ica. 2 specimens recorded from southern Utah. DISTRIBUTION IN SOUTHERN UTAH.—Aquar- Other than the type specimen, the Sanpete ius Plateau: c, Beaver Valley: b, Boulder Moun- County record is the first time this species has tain: e Pahvant Range: abe, Pine Valley Moun- been collected in southern Utah. It was col- tains: bd, Sanpitch Mountains: a, Sevier Plateau: lected in the Raft River Mountains (House- ackmosuxz, Tushar Mountains: aehks, Wasatch man and Baumann 1997), and it occurs more Plateau: gj, Abajo Mountains: a, LaSal Moun- commonly in northern Utah, particularly the tains: bdeg, Capitol Reef: e. Uinta Mountains, showing Central Rocky DISCUSSION.—This species is the most wide- Mountain affinities. Southern Utah is the south- ly distributed stonefly species in the Rocky ern limit of its range. Mountains (Baumann et al. 1977). 86 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

Family Perlodidae DISTRIBUTION IN SOUTHERN UTAH.—Hurri- Cultus aestivalis (Needham and Claassen) cane Cliffs: b, Markagunt Plateau: h, Sanpitch Mountains: a, Sevier Plateau: fgjopqrsuv, Wa- GEOGRAPHIC RANGE.—Coast, Cascade, and satch Plateau: gjm, Escalante River Drainage: Rocky Mountains. bc, Canyonlands: cd, Capitol Reef: de, Virgin DISTRIBUTION IN SOUTHERN UTAH.—Pan- River: bc. saugaunt Plateau: dg, Sevier Plateau: dk. DISCUSSION.—This species is common and DISCUSSION.—This species is uncommon in widespread throughout southern Utah. It is southern Utah. Several nymphs, which we have interesting to note its abundance and wide- been unable to identify to species, were col- spread occurrence throughout the Virgin River lected from Clear Creek, Monroe Creek, and drainage but its absence from the Pine Valley Niotche Creek. These nymphs are from Sevier Mountains and Santa Clara River drainage. County, where adults are available, indicating There is a definite faunal division between that these nymphs represent C. aestivalis. The these 2 regions. Common in the Virgin River species ranges in the Central Rocky Moun- drainage to the south, it ranges north to the tains. Sevier and Wasatch plateaus and even as far north as the Spanish Fork River in northern Diura knowltoni (Frison) Utah. Note its conspicuous absence from the GEOGRAPHIC RANGE.—Coast, Cascade, and Sevier River and Fremont River of Capitol Rocky Mountains, and Sierra Nevada. Reef National Park. The presence of I. zionen- DISTRIBUTION IN SOUTHERN UTAH.—Beaver sis rather than I. colubrinus in the Virgin River Valley: b, Markagunt Plateau: h, Sevier Plateau: indicates that the Virgin River was not always m, Tushar Mountains: a, Wasatch Plateau: jk, part of the Colorado drainage. Abajo Mountains: b, LaSal Mountains: g. DISCUSSION.—This species is widespread Isoperla fulva Claassen but uncommon throughout the high moun- tains of southern Utah. GEOGRAPHIC RANGE.—Coast, Cascade, and Rocky Mountains, and Sierra Nevada. Isogenoides colubrinus (Hagen) DISTRIBUTION IN SOUTHERN UTAH.—Aquar- ius Plateau: c, Beaver Valley: a, Boulder Moun- GEOGRAPHIC RANGE.—Coast, Cascade, and tain: c, Markagunt Plateau: af, Pahvant Range: Rocky Mountains. bc, Pansaugaunt Plateau: g, Pine Valley Moun- DISTRIBUTION IN SOUTHERN UTAH.—Pan- tains: b, Sevier Plateau: aegmnpuy, Tushar saugaunt Plateau: dg, Grand Valley: ab. Mountains: fs, Wasatch Plateau: ijq, Abajo DISCUSSION.—This species occurs in the Mountains: b, LaSal Mountains: bcd, Escalante large rivers of the Colorado drainage, but it is also found in the Sevier River, indicating that River Drainage: c, Capitol Reef: d, Virgin the Sevier River was connected in the past to River: c. the Colorado drainage. Baumann et al. (1977) DISCUSSION.—This species and I. quinque- list this species from San Juan County as well. punctata are the most commonly collected We have confirmed the specimens from Grand species of the genus in southern Utah. County and Garfield County but were unable to find specimens from San Juan County. Isoperla longiseta Banks However, given its presence in the other large GEOGRAPHIC RANGE.—Great Plains to Rocky rivers of the Colorado drainage, we are certain Mountains. it exists in San Juan County. The Sevier River DISTRIBUTION IN SOUTHERN UTAH.—Grand record is given below. Valley: ab, Canyonlands: abd, Painted Desert: Garfield Co.: Sevier River, Hwy 89 and abc, Green River: ab. Bryce Canyon Road, 12-VI-1950, W.E. Ricker, DISCUSSION.—In southern Utah this species 3 ɉ 2 Ɋ (CNC). occurs only in the large rivers of the Colorado drainage. Isogenoides zionensis Hanson Type locality.—Zion National Park, Utah. Isoperla mormona Banks GEOGRAPHIC RANGE.—Rocky Mountains and GEOGRAPHIC RANGE.—Coast, Cascade, and Alaska. Rocky Mountains, and Sierra Nevada. 2002] STONEFLIES OF SOUTHERN UTAH 87

DISTRIBUTION IN SOUTHERN UTAH.—Pah- DISTRIBUTION IN SOUTHERN UTAH.—Pan- vant Range: b, Sevier Plateau: dkv, Tushar saugaunt Plateau: f. Mountains: mq, LaSal Mountains: g, Capitol DISCUSSION.—We collected only a single Reef: dg, Virgin River: b, Great Basin Desert: specimen of this species from a remote site a, Escalante River Drainage: b. during our study, the full data for which are DISCUSSION.—This species is widespread given below. It is also reported from Garfield and can be common in southern Utah. It is County (Baumann et al. 1977), but we were un- one of the few species commonly found in the able to obtain and examine these specimens. Great Basin, being more tolerant of warmer Kane Co.: Podunk Creek, Hwy 99, S of and siltier lotic systems. Tropic Reservoir, R.W. Baumann and R.G. Call, 1 Ɋ. Isoperla petersoni Heedham and Christenson GEOGRAPHIC RANGE.—Alaska, Coast, Cas- Isoperla quinquepunctata (Banks) cade, and Rocky Mountains. GEOGRAPHIC RANGE.—Coast and Rocky DISTRIBUTION IN SOUTHERN UTAH.—Wasatch Mountains, and Sierra Nevada. Plateau: hi. DISTRIBUTION IN SOUTHERN UTAH.—Aquar- DISCUSSION.—The complete data for this ius Plateau: a, Boulder Mountain: e, Bull Valley species in southern Utah are given. Mountains: a, Markagunt Plateau: bij, Pahvant Sanpete Co.: Flat Canyon Spring, 18-VII- Range: acde, Pine Valley Mountains: bd, San- 1999, R. Hansen, 44 nymphs; O’dell Spring, pitch Mountains: a, Sevier Plateau: mpqry, Flat Canyon, 17-VII-1999, R. Hansen, 4 Tushar Mountains: en, Wasatch Plateau: ijq, nymphs; Stanley Spring, Flat Canyon, 19-VII- LaSal Mountains: g, Grand Valley: a, Henry 1999, R. Hansen, 12 nymphs; Gooseberry Mountains: ach, Capitol Reef: f, Virgin River: c. Creek, Flat Canyon, 19-VII-1999, R. Hansen DISCUSSION.—This species is widespread and S. Hansen, 2 nymphs. throughout southern Utah.

Isoperla phalerata (Needham) Isoperla sobria (Hagen) GEOGRAPHIC RANGE.—Coast, Cascade, and GEOGRAPHIC RANGE.—Coast, Cascade, and Rocky Mountains. Rocky Mountains, and Sierra Nevada. DISTRIBUTION IN SOUTHERN UTAH.—Mark- DISTRIBUTION IN SOUTHERN UTAH.—Boulder agunt Plateau: i. Mountain: e, Wasatch Plateau: j, Abajo Moun- DISCUSSION.—Complete data for this record tains: c, Grand Valley: fj, Capitol Reef: e, Mark- are included. This species is unrecorded else- agunt Plateau: f. where in Utah. Although we attempted to ver- DISCUSSION.—This species is widely dis- ify its presence in Panguitch Creek, we did tributed, but uncommon in southern Utah. not succeed in collecting any specimens. The distribution of this species in other localities Kogotus modestus (Banks) in the Central and Southern Rocky Moun- GEOGRAPHIC RANGE.—Coast, Cascade, and tains, however, does make its presence in Utah Rocky Mountains. plausible. We examined the specimen reported DISTRIBUTION IN SOUTHERN UTAH.—LaSal and concur that it is I. phalerata. We therefore Mountains: df. believe it to be present in this area and DISCUSSION.—This species is restricted in include it in the list of stoneflies occurring in southern Utah to the LaSal Mountains. It also Utah. Because it is the only record for the state, occurs in the Central Rocky Mountain region full data are given. of northern Utah and the Southern Rocky Garfield Co.: Panguitch Creek, between Mountains of Colorado. The distribution of Panguitch and Panguitch Lake, 27-VI-1967, this species is another example of the unique- Ɋ G.F. Knowlton, 1 (INHS). ness of this area compared with the rest of southern Utah. In the past these mountains Isoperla pinta Frison must have been connected to either the moun- GEOGRAPHIC RANGE.—Coast, Cascade, and tains of Colorado or the mountains of northern Rocky Mountains. Utah. 88 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1

Megarcys signata (Hagen) b, Pine Valley Mountains: b, Sevier Plateau: j, GEOGRAPHIC RANGE.—Coast, Cascade, and Tushar Mountains: f, Wasatch Plateau: j, Capi- Rocky Mountains. tol Reef: ade. DISTRIBUTION IN SOUTHERN UTAH.—Boul- DISCUSSION.—This species is sporadic and der Mountain: e, Hurricane Cliffs: b, Marka- disjunct in its distribution throughout south- gunt Plateau: hj, Sevier Plateau: gjmnotu, ern Utah. Tushar Mountains: gkop, Wasatch Plateau: ehjp, Abajo Mountains: b. ACKNOWLEDGMENTS DISCUSSION.—Prior to our study this species was known from only 3 counties in southern We thank Riley Nelson, Duane Jeffrey, and Utah. Several new records from our study Bruce Schaalje for their assistance. Thanks to show that this species is widespread in south- the Department of Zoology for the awards that ern Utah. We document it from 7 counties. have made this undertaking financially possi- ble. Special thanks to D.E. Bright and E.C. Skwala americana (Needham and Claassen) Becker (Canadian National Collection), Boris GEOGRAPHIC RANGE.—Western North Amer- C. Kondratieff (Colorado State University), ica. Andrew H. Barnum (Dixie College), Ed Dewalt DISTRIBUTION IN SOUTHERN UTAH.—Wasatch and Donald W. Webb (Illinois Natural History Plateau: j. Survey), John A. MacDonald and Colin A. DISCUSSION.—This rare species is known Brammer (Purdue University), Robert Hansen from only a single locality in southern Utah, as (Austin, Texas), Wilford J. Hanson (Utah State we define it. This record exemplifies the pit- University), Nancy Adams and Oliver S. Flint, falls of using political boundaries in a geo- Jr. (United States National Museum), S.M. graphical study. It actually occurs at the south- Clark (Charleston, West Virginia), Robert N. ern end of the Wasatch Mountain range or Winget (Brigham Young University–Hawaii), northern edge of the Wasatch Plateau, showing Bret O. Huntsman, Karen Rohner (Mammoth a faunal division at this boundary. This bound- Creek Fish Hatchery, Utah Department of ary is likely the southern limit of its range in Fish and Game), and Brent Farnsworth (Utah Utah. Division of Wildlife Resources) for providing Emery Co.: Huntington Creek, 15-VI-1948, records or specimens used in this study or collector ?, 1 ɉ 4 Ɋ. otherwise assisting in the project. Special thanks go to Harry Barber of the Grand Stair- Family Pteronarcyidae case–Escalante National Monument for help- ing us find isolated localities in the vast monu- Pteronarcella badia (Hagen) ment. We especially thank our families for their GEOGRAPHIC RANGE.—Rocky Mountains and support throughout this project. Alaska. DISTRIBUTION IN SOUTHERN UTAH.—Aquar- LITERATURE CITED ius Plateau: a, Beaver Valley: ab, Boulder Mountain: e, Escalante River Drainage: f, Mark- BAUMANN, R.W. 1970. The genus Nemoura (Plecoptera) of agunt Plateau: fhj, Pansaugaunt Plateau: dg, the Rocky Mountains. Unpublished doctoral disser- Pahvant Range: bce, Sevier Plateau: afgjmor, tation, University of Utah, Salt Lake City. 172 pp. ______. 1973. Studies on Utah stoneflies (Plecoptera). Tushar Mountains: aefhkms, Wasatch Plateau: Great Basin Naturalist 33:91–108. gjopq, LaSal Mountains: bg, Capitol Reef: ade, ______. 1975. Revision of the stonefly family Nemouridae Virgin River: c. (Plecoptera): a study of the world fauna at the DISCUSSION.—This species is widespread generic level. Smithsonian Contributions to Zoology and common throughout southern Utah in all 211:1–74. BAUMANN, R.W., AND A.R. GAUFIN. 1969. The stoneflies geographic provinces. (Plecoptera) of the Wasatch Mountains, Utah. Utah Academy Proceedings 46:106–113. Pteronarcys californica Newport ______. 1972. The Amphinemura venusta complex of GEOGRAPHIC RANGE.—Coast, Cascade, and western North America (Plecoptera: Nemouridae). Los Angeles County Museum Contributions in Sci- Rocky Mountains, and Sierra Nevada. ence 226:1–16. DISTRIBUTION IN SOUTHERN UTAH.—Beaver BAUMANN, R.W., A.R. GAUFIN, AND R.F. SURDICK. 1977. The Valley: a, Markagunt Plateau: h, Pahvant Range: stoneflies (Plecoptera) of the Rocky Mountains. 2002] STONEFLIES OF SOUTHERN UTAH 89

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