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Plecoptera of Gunnison County, Colorado
Annotated Publication List

Updated 1 March 2024

You may search this page (or any other webpage) with your browser. Type Ctrl F and fill in your search term. Species names are links to pages on this website. Many older publications are available from the Biodiversity Library. The International Journal of Stonefly Research Illiesia, has recently stopped publishing new papers.

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Alexander,KD; Stewart,KW 1996a Description and theoretical considerations of mate finding and other adult behaviors in a Colorado population of Claassenia sabulosa (Plecoptera: Perlidae). Annals of the Entomological Society of America 89 2, 290-296.
They studied Claassenia in the Gunnison river near Almont.

Alexander,KD; Stewart,KW 1996b The mate searching behavior of Perlinella drymo (Newman) (Plecoptera: Perlidae) in relation to drumming on a branched system. Mitteilungen Der Schweizerischen Entomologischen Gesellschaft Bulletin de la Societe Entomologique Suisse 69, 121-126.

Alexander,KD; Stewart,KW 1997a Furthur considerations of mate searching behavior and communication in adult stoneflies (Plecoptera); first report of tremulation in Suwallia (Chloroperlidae). In: Ephemeroptera and Plecoptera: Biology-Ecology-Systematics. Eds: Landolt,P; Sartori,M MTL, Fribourg, 107-112.

Alexander,KD; Stewart,KW 1997b The importance of aedeagal characters in species delineation and revision of the stonefly tribe Suwalliini Surdick (Chloroperlidae). In: Ephemeroptera and Plecoptera: Biology-Ecology-Systematics. Eds: Landolt,P; Sartori,M MTL, Fribourg, 484-488.

Alexander,KD; Stewart,KW 1999 Revision of genus Suwallia Ricker (Plecoptera: Chloroperlidae). Transactions of American Entomological Society 125(3)185-250. Used for this website to develop Suwallia female key Suwallia male key

Allan, J.D. 1975a. The distributional ecology and diversity of benthic insects in Cement Creek, Colorado. Ecology 56(5) 1040-1053. Abstract

Allan,JD 1975b Faunal replacement: altitudinal zonation in an alpine stream. Verhandlungen der Internationale Vereinigung für Theoretische und Angewandte Limnologie 19:1015-1022.
A longitudinal survey of Cement Creek in the 1970s.

Allan,JD 1978 Diet of brook trout (Salvelinus fontinalis Mitchell) and Brown Trout (Salmo trutta L.) in an alpine stream. Internationale Vereinigung für Theoretische und Angewandte Limnologie Verhandlungen 20, 2045-2050.
    Allan studied drift and trout diets in August of 1975 in Cement Creek. Regarding Plecoptera, he found Alloperla spp (3.8% of total drift), Zapada haysi (2.1%), Kogotus modestus (.1%) and Pteronarcella badia (.1%) in the drift and in Brook and Brown Trout stomachs. Trout diets correlated with species available in the drift.

Allan,JD 1982 Feeding habits and prey consumption of three setipalpian stoneflies (Plecoptera) in a mountain stream. Ecology 63 1, 26-34. Abstract

Allan,JD 1982 The effects of reduction in trout density on the invertebrate community of a mountain stream. Ecology 63(5) 1444-1455. Abstract

Allan, J.D. 1983. Food consumption by trout and stoneflies in a Rocky Mountain stream, with comparison to prey standing crop. Pages 371-390 in Dynamics of Lotic Systems, eds. T.D. Fontaine and S.M. Bartell. Ann Arbor: Ann Arbor Publishers.

Allan,JD; Flecker,AS; McClintock,NL 1987a Prey preference of stoneflies: sedentary vs mobile prey. Oikos 49 3, 323-331.

Allan,JD; Flecker,AS; McClintock,NL 1987b Prey size selection by carnivorous stoneflies. Limnology and Oceanography 32 4, 864-872.

Allan, JD 1995 Stream Ecology: Structure and function of running waters. Chapman and Hall, London England. 400pages.

Armold,MT; Blomquist,GJ and Jackson,LL 1969 Cuticular lipids of insects—III. The surface lipids of the aquatic and terrestrial life forms of the big stonefly, Pteronarcys californica newport. Comparative Biochemistry and Physiology 31(5)685-692.


Banks, N. 1895 New Neuropterid Insects. Transactions of the American Entomological Society 22: 313-316.

Banks,N 1897 New North american neuropteroid insects. Transactions of the American Entomological Society 24:21-31.

Banks,N 1900 New genera and species of Nearctic Neuropteroid Insects. Transactions of the American Entomological Society 26:239-259.

Banks,N 1902 Notes and descriptions of Perlidae. Canadian Entomologist 34:123-125. PDF

Banks,N 1904 Neuropteroid insects from New Mexico. Transactions of the American Entomological Society 30:97-110.

Banks,N 1908 Neuropteroid insects - notes and descriptions. Transactions of the American Entomological Society 34:255-267.

Banks,N 1911 Descriptions of new species of North American Neuropterid Insects. Transactions of American Entomological Society 37, 335-360.

Banks,N 1920 New Neuropteroid insects. Bulletin of the Museum of Comparative Zoology. 64: 299-362.

Banks,N 1938 New native neuropteroid insects. Psyche (45) 72-79.

Barton, DR 1980 Benthic macroinvertebrate communities of the Athabasca river near Ft. Mackay, Alberta. Hydrobiologia 74(2) 151-160. Abstract

Baumann,RW 1973 Studies on Utah stoneflies (Plecoptera) Western North American Naturalist 33(2) 91-108. PDF

Baumann,RW 1975 Revision of the stonefly family Nemouridae (Plecoptera): A study of the world fauna at the generic level. Smithsonian Contributions Zoology 211, 1-74. PDF

Baumann,RW 1979 Nearctic stonefly genera as indicators of ecological parameters (Plecoptera: Insecta). The Great Basin Naturalist, 39(3) 241-244. PDF

Baumann,RW, Gaufin,AR and Surdick,RF 1977 The stoneflies (Plecoptera) of the Rocky Mountains. Memoirs of the American Entomological Society 31, 1-208. PDF
     They provide male and female keys, sometimes nymph keys as well as illustrations, geographic range, distribution and discussions. Gunnison County is in the Southern Rocky Mountains in this book. The main stonefly publication for the Rocky Mountains, you'll use it to key adults and sometimes larvae to species. A few of the species names have changed as the years go by and revisions of various genera have been published. This website (www.gunnisoninsects.org) quotes paragraphs about Gunnison County stoneflies in the references of many of our webpages.

Baumann,RW and Kondratieff, BC 2008a The Alloperla severa complex (Plecoptera: Chloroperlidae) of western North America. Illiesia 2008 4(6):66-75. Abstract PDF

Baumann,RW and Kondratieff,BC 2008b A review of the western North American genus Triznaka (Plecoptera: Chloroperlidae) with a new species from the Great Basin, USA. Proceedings of the Entomological Society of Washington 110(2)345-362

Baumann,RW and Kondratieff,BC 2009 Studies on the Holarctic subfamily Brachypterainae (Plecoptera: Taeniopterygidae) using the scanning electron microscope to study male terminalia. Aquatic Insects 31(S1)219-230.
     Abstract: "The epiproct complex and other details of the male terminalia of the Holarctic stonefly subfamily Brachypterainae are illustrated by scanning electron microscope micrographs or images. Selected species of eight of the 12 genera, Bolotoperla, Doddsia, Oemopteryx, Strophopteryx, Taenionema, Mesyatsia, Brachyptera, and Rhabdiopteryx are examined. Three images are provided for each species, which usually include a dorsal view of the male terminalia and close up views of the epiproct complex or other genital structures. These genera are organised in a zoogeographical format beginning with the Nearctic fauna and subsequently moving in a westerly direction around the globe, ending with the west Palearctic fauna. Locality data are given for all specimens illustrated. "

Bergey,EA and Ward,JV 1989 Upstream-downstream movements of aquatic invertebrates in a Rocky Mountain stream, Hydrobiologia, Volume 185( 1) 71-82. Abstract

Béthoux,O. 2005 Wing venation pattern of Plecoptera (Insecta: Neoptera). Illliesia 1: 52-81. PDF
     Abstract: " Some consider the order Plecoptera as the sister-group of all neopterous insects. Hence the interpretation of Plecoptera wing venation has critical implications for character polarization among basal neopterous taxa, i.e. polyneopterous insects, and especially for fossil taxa, mainly known after isolated wings. However, no consensus ever emerged from the previous interpretations, partly contradicting. This study provides a detailed morphological comparative study of the wing venation of the order Plecoptera, based on modern taxa. It reveals that 1) the arculus is not a posterior branch of the media but a secondarily strengthened crossvein, always present in hind wings and very generally in forewings; 2) the media is primitively two-branched in both wing pairs; 3) in hind wings the stems of the radius and the media are basally distinct, but a fusion of the posterior radius (RP) with the media (M) occurs distal to the wing base, both branches diverging further; and 4) the vannus is composed of branches belonging to the anterior analis sector (AA) only (i.e. the analis posterior and jugal areas are lacking). A new nomenclature is proposed for describing the branches of AA2. Character states presence of an arculus in both fore- and hind wings, media two-branched, and in hind wings, presence of fusion of RP with M are diagnostic of the order, based on outgroup comparison with other polyneopterous insects. Similarities are noticed between most basal Archaeorthoptera (i.e. insects close related to Orthoptera) and Plecoptera concerning the organization of the anal area, although an AP area is retained in the former and absent in the latter. Additionally, wing characters susceptible of being informative for the resolution of the inner phylogeny of the Plecoptera are proposed throughout the paper."

Borror,DJ; De Long,DM; Triplehorn,CA An Introduction to the Study of Insects. Saunders College Publishing, Philadelphia, PA.
     A classic, useful for all insect studies. The 7th edition has new authors Johnson and Triplehorn. This amazing book was affectionately referred to as "Boring and Too Long" by entomology wiseasses everywhere. It's long tenure as the general go-to bug reference means I had to leave the old title in this list of publications.

Bottorff,RL and Bottorff,LD 2007 Phenology and diversity of adult stoneflies (Plecoptera) of a small coastal stream, California. Illiesia 3(1):1-9 pdf 292 Kb

Bottorff,RL; Knight,AW 1987 Ectosymbiosis between Nanocladius downesi (Diptera: Chironomidae) and Acroneuria abnormis (Plecoptera: Perlidae) in a Michigan stream, USA) Entomol. Gener. 12: 97-113.

Bottorff,RL; Szczytko,SW; Knight,AW; Dimick,JJ 1990 Drumming behavior of four western nearctic Isoperla species (Plecoptera:Perlodidae). Annals of the Entomological Society of America 83 5, 991-997.

Branham,JM and Hathaway,RR 1975 Sexual differences in the growth of Pteronarcys californica Newport and Pteronarcella badia (Hagen) (Plecoptera). Canadian Journal of Zoology, 1975, 53:(5) 501-506.
     Abstract: "Two species of Plecoptera, Pteronarcys californica Newport and Pteronarcella badia (Hagen), were collected seasonally from a single site in the Provo River, Utah. Body weights of the living animals were used to distinguish four size classes in each sex of Pteronarcys californica, and one size class for each sex of Pteronarcella badia. Ovaries, fat bodies, and guts were dissected from Pteronarcys californica. Growth of whole animals and of the organs are discussed with regard to life cycle, sexual development, and age. The importance of weight measurements on insects to be used for physiological studies is discussed."

Buchwalter,DB; Cain,DJ; Martin,CA; Xie,L; Luoma,SN; Garland,JT 2008 Aquatic insect ecophysiological traits reveal phylogenetically based differences in dissolved cadmium susceptibility. Proceedings of the National Academy of Sciences 105 24, 8321-8326.


Canton,SP and Chadwick,JW 1983 Seasonal and longitudinal changes in invertebrate functional groups in the Dolores River, Colorado. Freshwater Invertebrate Biology, 41-47. PDF

Canton,SP and Ward,JV 1981 The aquatic insects, with emphasis on Trichoptera, of a Colorado stream affected by coal strip-mine drainage. Southwestern Naturalist 25 4, 453-460.
     They studied Trout Creek where it runs through the Edna Coal Mine in northwestern Colorado. The mine spoils were 30 meters from the edge of the creek (approximately a 100 foot buffer zone). They found the aquatic insect density (numbers per square meter) and biomass (weight in grams per square meter) did not change above and below the mine. The Shannon-Weaver Diversity index also showed no difference between sites. However the community structure (which species were present and proportions) did change. Since there were irrigation water and cattle influences at their downstream site, their results may reflect these additional water uses. They note the biggest visible change in riparian vegetation at this mine is the loss of willow and alder trees downstream of the mine. The caddisfly population changed the most between sites, shifting from a mix of families above the mine to dominance by Hydropsychidae and Glossosomatidae below the mine.
Prostoia besametsa was most common at the clean, forested, shaded, upstream site. It was probably more common here partly because it is a shredder and needs leaves to eat.

Cao,J; Guo,X; Guo,C; Wang,X; Wang,Y and Yan,F 2022 Complete mitochondrial genome of Malenka flexura (Plecoptera: Nemouridae) and phylogenetic analysis. Genes, 13(5), p.911. PDF

Carlisle,DM; Clements,WH 2003 Growth and secondary production of aquatic insects along a gradient of Zn contamination in Rocky Mountain streams. Journal North American Benthological Society 22(4), 582-597. Abstract and entire paper

Carlisle,DM; Clements,WH 2005 Leaf litter breakdown, microbial respiration and shredder production in metal-polluted streams. Freshwater Biology 50, 380-390.
     Discusses the effects of Zinc on several streams. Taenionema pallidum populations were reduced by metal pollution and this had a larger effect on the stream communities because no other insect filled Taenionemas shredder role in polluted streams.

Caruso,C and Wichard,W 2010 Overview and descriptions of fossil stoneflies (Plecoptera) in Baltic Amber. Entomologie Heute, (22) 85-97. PDF Cather,MR; Gaufin,AR 1975 Life history and ecology of Megarcys signata (Plecoptera:Perlodidae), Mill Creek, Wasatch Mountains, Utah. Great Basin Naturalist 35, 1.

Cather,MR and Arden R. Gaufin,AR 1976 Comparative ecology of three Zapada species of Mill Creek, Wasatch Mountains, Utah (Plecoptera: Nemouridae). American Midland Naturalist 95(2) 464-471. Abstract

Chen,ZT 2018 Females of the genus Podmosta (Plecoptera: Nemouridae): comparison of terminalia and a new female record in Baltic Amber. Zootaxa, 4407(2)293-297. PDF

Claassen,PW 1924 New species of North American Capniidae (Plecoptera). Canadian Entomologist 56, 43-48, 54-57.

Claassen,PW 1931 Plecoptera nymphs of America north of Mexico. Entomological Society of America 3, 1-199.

Claassen,PW 1936 New names for stoneflies (Plecoptera). Annals of the Entomological Society of America 29, 622-623.

Claassen,PW 1937 New species of stoneflies (Plecoptera). Canadian Entomologist 69, 79-82.

Clements,WH 1999 Metal tolerance and predator-prey interactions in benthic macroinvertebrate stream communities. Ecological Applications 9, 1073-1084.

Clements,WH; Carlisle,DN; Lazorchak,JM; Johnson,PC 2000 Heavy metals structure benthic communities in Colorado mountain streams. Ecological Applications 10(2)626-638. Abstract
     The authors discuss the EPA's Regional Environmental Monitoring and Assessment Program (REMAP) data on aquatic insects among a number of mine-polluted and clean streams and rivers in Colorado.

Clubb,RW; Gaufin,AR and Lords,JL 1974 Acute cadmium toxicity studies upon nine species of aquatic insects. Environmental Research 9(3) 332-341.
     Abstract: Continuous-flow bioassays were employed to determine 96-hour median tolerance limits (TLm), for the stonefly, Pteronarcella badia (Hagen) (TLm was 18.0 mg Cd/l) and the mayfly, Ephemerella grandis grandis Eaton (TLm was 28.0 mg Cd/l). Ninety-six hours TLm values for other species of aquatic insects tested were not determined, since these species were relatively insensitive to cadmium.
Insects exposed for four days in cadmium-containing water, then placed in tap water, show a linear rate of cadmium loss. This loss may lower or prevent mortality under ideal conditions.

Colburn,T 1982 Measurement of low levels of molybdenum in the environment by using aquatic insects. 29, 422-428.

Colburn,T 1986 The use of the stonefly Pteronarcys californica Newport as a measure of biologically available cadmium in a high altitude river system Colorado, USA. Water Quality Bulletin 11, 141-147.

Corkum LD and Clifford HF 1980 The importance of species associations and substrate types to behavioural drift. Pages 331-341 in Flannigan JF; Marshall KE. Advances in Ephemeroptera Biology. Plenum Press, New York. PDF

Courtney-Mustaphi,CJ; Steiner,E; von Fumetti,S and Heiri,O 2024 Aquatic invertebrate mandibles and sclerotized remains in Quaternary lake sediments. Journal of Paleolimnology, 71(1), pp.45-83. PDF
     Abstract: "Subfossil remains of aquatic invertebrates found in lacustrine sediments are useful paleoenvironmental indicators. Strongly scleroticized chitinous body parts from the exoskeleton or exuviae from invertebrates are often the most resistant to degradation during syn- and post-depositional processes. Invertebrate mandibles and body parts that superficially resemble mandibles, such as claw-like appendages and pygopodia, are frequently found in sieved Quaternary lacustrine, palustrine, and deltaic sediments. Guides, catalogs and atlases have been published that are well suited for the identification of subfossil remains for several invertebrate groups, such as chironomids, cladocerans, and ostracods, among others. However, aquatic invertebrate remains of several ecologically important invertebrate groups continue to be underused in paleoenvironmental studies, in part, because there are few visual keys or other documentation sources (e.g. descriptions, catalogs or atlases) that increase awareness and facilitate identification. Here we present sets of digital photomicrographs of pre-identified aquatic invertebrate specimens collected from streams, lakes and ponds that have been chemically cleared to preserve structures that are observed in subfossil remains in sieved sediment samples, commonly the > 100 μm size fractions. In addition, we present examples of these structures from Quaternary lake-sediment samples and cite the dispersed literature that demonstrate that these remains are preserved and remain identifiable in the fossil record. We document mandibles from several taxonomic groups that include Crustacea: Amphipoda, Isopoda, Ostracoda, and Notostraca; and Insecta orders: Coleoptera, Diptera, Ephemeroptera, Hemiptera, Odonata, Lepidoptera, Megaloptera, Plecoptera, and Trichoptera. The compilation of microphotographs also includes pygopodia and claw appendages of Plecoptera and Trichoptera, with additional images of other common invertebrate mouthpart and head remains. We describe several types of fossilizing structures that are, to our knowledge, not previously described in the paleoecological literature (e.g. mandibles of amphipods or plecopterans) but also show that some structures are considerably more variable than expected based on available descriptions, such as the mandibles of Ephemeroptera or Trichoptera, and that these can potentially be separated into different morphotypes useful for identification of subfossil material. We also discuss the potential of analyzing and interpreting the additional remains together with the remains of more commonly analyzed invertebrate groups (e.g. Chironomidae) to contribute to paleoenvironmental interpretations, which will allow assessments of functional groups (e.g. predators, shredders, grazers) or habitat types (e.g. littoral, profundal or lotic environments) that aquatic invertebrate remains originate from."

Crespo,JG 2011 A review of chemosensation and related behavior in aquatic insects. Journal of Insect Science, 11. PDF

Cui, Y; Béthoux, O; Kondratieff,B; Shih,C and Ren,D 2016 The first fossil salmonfly (Insecta: Plecoptera: Pteronarcyidae), back to the Middle Jurassic. BMC evolutionary biology, 16(1) 217 HTML

Cummins,KW; Wilzbach,MA; Gates,DM; Perry,JB; Taliaferro,WB 1989a Leaf litter that falls into streams influences communities of stream invertebrates. BioScience 39 1, 24-30.

Cummins,KW; Wilzbach,MA; Gates,DM; Perry,JB and Taliaferro,WB 1989 Shredders and riparian vegetation. BioScience, 39(1), pp.24-30. PDF


Delk,JK; Kilgore,MJ; Stark,BP 1998 Comparison of the epiproct structure of two closely related species, Sweltsa fidelis (Banks) and S. revelstoka (Jewett) (Plecoptera: Chloroperlidae). Great Basin Naturalist 58, 3.
     They decided you can't tell the epiprocts of S. fidelis and S. revelstoka apart. Luckily for us S. revelstoka doesn't live here. Has scanning electron photomicrographs of epiprocts. Female subgenital plates are very different so they don't propose synonymy.

DelVecchia,AG; Stanford,JA and Xu,X 2016 Ancient and methane-derived carbon subsidizes contemporary food webs. Nature communications, 7 PDF

DeWalt,RE; Maehr,MD; Neu-Becker,U and Stueber,G 2017 Plecoptera Species File Online. Version 5.0/5.0. [5Aug2017]. http://Plecoptera.SpeciesFile.org.

DeWalt,RE and Ower,GD 2019 Ecosystem services, global diversity, and rate of stonefly species descriptions (Insecta: Plecoptera). Insects, 10(4), p.99. PDF

     Abstract: "Thirty-one stonefly species representing eight families were collected during the March 1987 to May 1990 study period. Genera represented by more than one species included Capnia, Utacapnia, Taenionema, Suwallia, Triznaka, Isogenoides, and Isoperla. Peak species richness was recorded on or near the summer solstice in 1988 and 1989. Climatic differences between years were reflected in nymphal development and emergence phenology of most species. New or important corroborative life history data are presented for 11 stonefly species of this assemblage. The hyporheic nymphal development of most chloroperlid species limited the number of early instars sampled and our capacity to interpret voltinism. Limited nymphal data suggested a univoltine-slow cycle for Plumiperla diversa (Frison). Adults of Suwallia pallidula (Banks) and S. wardi (Banks) were present for an extended summer period, but the bulk of their respective emergence times was temporally separated. Isogenoides zionensis Hanson, Pteronarcella badia (Hagen), and Pteronarcys californica Newport were all shown for the first time to have a 9–10-mo egg diapause, and all three species have a semivoltine life cycle. Skwala americana (Klapálek) and Isoperla fulva Claassen were further confirmed to have univoltine-slow cycles. Univoltine-fast and univoltine-slow life cycles are reported for the first time in I. phalerata and I. quinquepunctata, respectively. Regression analysis revealed that six of the eight abundant species had extended emergence patterns (slopes of <5%/d), while only two had synchronous patterns. Warmer spring and summer temperatures in 1989 increased the slopes for five of the eight species studied, but did not change their synchrony designation. Nine of 11 abundant species advanced their median emergence date in 1989 over 1988. This and the higher slope values are consistent with a hurried nymphal development and narrower emergence period due to the warmer thermal regime of 1989. "

DeWalt,RE and Stewart,KW 1995 Life histories of stoneflies (Plecoptera) in the Rio Conejos of southern Colorado. Great Basin Naturalist 55, 1-18. PDF DeWalt,RE; Stewart,KW; Moulton,SR; Kennedy,JH 1994 Summer emergence of mayflies, stoneflies, and caddisflies from a Colorado mountain stream. Southwestern Naturalist 39 (3) 249-256. PDF
     Abstract: The summer emergence patterns of mayfly, stonefly, and caddisfly species are described for a second order, southern Rocky Mountain Colorado stream. Frequent, standardized, sweepnetting and emergence trap samples provided 1,779 adults of 45 taxa consisting of 11 mayfly, 15 stonefly, and 19 caddisfly species. The five most dominant species were, in order of importance, Suwallia nr. lineosa, Sweltsa coloradensis, Oligophlebodes minutus, Paraleuctra vershina, and Baetis bicaudatus. Stoneflies contributed 70% of all adults collected. Peak species richness of all orders occurred near the summer solstice and maximum water temperatures. The range of slope values generated by simple linear regression of cumulative percentage catch revealed that emergence was extended (slopes < 4%/day) for five of the seven most abundant species. Two caddisflies, O. minutus and Rhyacophila pellisa, displayed a synchronous emergence pattern. This study adds 35 new records for the Rio Conejos drainage and Conejos County and provides a baseline of comparison against future changes in species richness among these orders.

Ding,S; Li,W; Wang,Y; Cameron,SL; Murányi,D and Yang,D 2019 The phylogeny and evolutionary timescale of stoneflies (Insecta: Plecoptera) inferred from mitochondrial genomes. Molecular Phylogenetics and Evolution, (135) 123-135.
     Abstract: "Phylogenetic analysis based on mitochondrial genomic data from 25 stonefly species recovered a well-supported tree resolving higher-level relationships within Plecoptera (stoneflies). The monophyly of both currently recognized suborders was strongly supported, concordant with previous molecular analyses of Plecoptera. The southern hemisphere suborder Antarctoperlaria formed two clades: Eustheniidae + Diamphipnoidae and Austroperlidae + Gripopterygidae; consistent with relationships proposed based on morphology. The largely northern hemisphere suborder Arctoperlaria also divided into two groups, Euholognatha and Systellognatha, each composed of the five families traditionally assigned to each infraorder (the placement Scopuridae by mt genome data remains untested at this time). Within Euholognatha, strong support for the clade Nemouridae + Notonemouridae confirmed the northern origin of the currently southern hemisphere restricted Notonemouridae. Other family level relationships within the Arctoperlaria differ from those recovered by previous morphology and molecular based analyses. A fossil-calibrated divergence estimation suggests the formation of two suborders dates back to the Jurassic (181 Ma), with subsequent diversification of most stonefly families during the Cretaceous. This result confirms the hypothesis that initial divergence between the suborders was driven by the breakup of the supercontinent Pangaea into Laurasia and Gondwanaland (commencing 200 Ma and complete by 150 Ma)."

Dodds,GS; Hisaw,FL 1925 Ecological studies on aquatic insects. IV. Altitudinal range and zonation of mayflies, stoneflies and caddisflies in the Colorado Rockies. Ecology 6(4), 380-390. Abstract PDF
     Widely cited early work on aquatic insects in Colorado. Taxonomy has changed a lot since this paper was written so here's a table with the name changes.

Figure 1 Taxonomy Translation (page 382)
Dodds and Hisaw Name 2004 Name
Alloperla pilosa same
Nemura sp. unknown
Perlodes signata Megarcys signata
Perla modesta Kogotus modestus
Alloperla lamba Sweltsa lamba
Nemura delicatula Podmosta delicatula
Leuctra occidentalis Paraleuctra occidentalis
Taenionema analis Taenionema pacificum
Paraperla frontalis same
Perla expansa Pictetiella expansa
Nemura venusta Amphinemura banksi
Arsapnia decepta Capnia decepta
Alloperla sp. unknown
Capnia brevicauda Eucapnopsis brevicauda
Taeniopteryx occidentalis Doddsia occidentalis
Perla sp. unknown
Perla modesta Kogotus modestus
Acroneuria pacifica Hesperoperla pacifica
Perla ebria Isoperla sobria
Alloperla borealis Sweltsa borealis
Isogenus elongatus Isogenoides elongatus
Alloperla pallidula Suwallia pallidula
Pteronarcella badia same
Isoperla punctata same
Alloperla signata Triznaka signata
Isoperla sp. unknown

Dosdall, LM 1991 Survival of selected aquatic insects exposed to methoxychlor treatment of the Saskatchewan River system. Water Quality Research Journal of Canada. 26(1) 27-40.

Dosdall,LM and Giberson,DJ 2014 Stoneflies (Plecoptera) of the Canadian Prairie Provinces. Arthropods of Canadian Grasslands, 3: 201–229. PDF
     Abstract: "Most Plecoptera (stonefly) species require cool, well-oxygenated water and are therefore not as well represented in prairie grassland habitats as they are in steep mountain streams or forested habitats. One hundred thirty-one species of Plecoptera have been recorded from the Prairie Provinces of Alberta, Saskatchewan, and Manitoba, including 61 that occur in or border aquatic habitats in the Prairies Ecozone. Alberta has the highest stonefly diversity because of its proximity to the mountains (108 species, 104 of which occur in the mountains), and the diversity in Saskatchewan is similar to that of Manitoba at 44 and 46 species, respectively. Only 2 of the 61 Prairies Ecozone spe.cies (Perlesta dakota and Pteronarcys pictetii) are restricted to this ecozone in Canada, and most of the species that occur in the Prairies Ecozone also occur in the Boreal Shield and Boreal Plains ecozones (40 species) and/or in the Montane Cordillera Ecozone (42 species). A list of species collected for each province and ecozone is provided, with references and probable refugial origins. Information on stonefly ecology, taxonomic works, zoogeography, and importance as monitoring tools pertinent to the region is also presented."

Dosdall, LM; Goodwin, LR; Casey, RJ; Noton, L 1997 The effect of ambient concentrations of chlorate on survival of freshwater aquatic invertebrates. Water Quality Research Journal of Canada. 32(4) 839-854.
      This paper is mentioned in some of the PAN pesticides database http://www.pesticideinfo.org/Index.html links scattered throughout this website and on the Ambient Water Quality Guidelines for Chlorate website from Government of British Columbia, Ministry of the Environment.

Dosdall,LM and Lehmkuhl,DM 1979 Stoneflies (Plecoptera) of Saskatchewan. Quaestiones Entomologicae.

Donald,DB 1980 Deformities in Capniidae (Plecoptera) from the Bow River, Alberta. Canadian Journal of Zoology 58, 682-686.

Downes,JA 1964 Arctic insects and their environment. Canadian Entomologist 96, 279-307.

Duffield,RM and Nelson,CH 1998 Stoneflies (Plecoptera) in the diet of brook trout (Salvelinus fontinalis Mitchell) in Libby Creek, Wyoming, USA. Hydrobiologia 380, 59-65.
     Abstract: "The stonefly component of a naturally reproducing population of brook trout (Salvelinus fontinalis) was studied by analyzing 216 stomach pump samples collected between May 15 and August 10, 1991 and 1992, from Libby Creek, an alpine stream in the Medicine Bow National Forest in Wyoming. Stoneflies constituted 10.3% of the total items recovered and were the second most abundant order in the samples. Nineteen species of Plecoptera representing five families were identified. Approximately 43% of all stoneflies were Sweltsa lamba; the next most prevalent species was Zapada haysi (12%). The other prevalent insect orders in the samples were Diptera (57.2%), Ephemeroptera (8.4%) and Trichoptera (6.1%). Terrestrial Coleoptera, Hymenoptera, and Hemiptera were most common in July and August."


Elmork,K; Saether,OR (1970) Distribution of invertebrates in a high mountain brook in the Colorado Rocky Mountains. University of Colorado Studies Series in Biology No 31.


Finn,DS and Poff,NL 2008 Emergence and flight activity of alpine stream insects in two years with contrasting winter snowpack. Artic, Antarctic, and Alpine Research 40(4)638-646. PDF

Fochetti,R and De Figueroa,JMT 2007 Global diversity of stoneflies (Plecoptera; Insecta) in freshwater. In Freshwater Animal Diversity Assessment (pp. 365-377). Springer, Dordrecht. PDF

     Abstract: Plecoptera, or stoneflies, is a small order of hemimetabolous insects: according to our data, more than 3,497 species have been described so far in the world. The total number of species has enormously increased in the last 30 years (2,000 species estimated in 1976) and, if the trend continues, then it will nearly double in the near future. The order is divided into the suborders Arctoperlaria and Antarctoperlaria, and includes 16 families: 12 arctoperlarian and 4 antarctoperlarian. The Arctoperlaria account for a total number of 3,179 species, and Antarctoperlaria, only 318 species. The total number of genera is 286. We give in this article the estimated number of species for each family. The fauna and diversity of stonefly in North America (650 species reported) and Europe (426 species) are best known. Nevertheless, in the last 25 years, a mean of 2.6 Plecoptera species per year were described in Europe. Stonefly-faunas of Australia (191 species, Tasmania included) and New Zealand (104 species) are relatively well-known, while our knowledge of the Plecoptera of Central and South America (95 and 378 species respectively) is poor and still not representative of the real diversity. Africa has a reduced stonefly fauna (126 species). Asian stonefly diversity (approximately 1,527 species) is much greater than that of Europe or North America despite the fact that, except for Japan and Asiatic Russia that have been well studied, our knowledge of the remaining Asiatic areas is extremely poor. Even though our data indicate the Holarctic Region as the diversity hot-spot for the order, the analysis of the specific diversity divided by family suggests also an important role of tropical stoneflies.

Freilich,JE 1991 Movement patterns and ecology of Pteronarcys nymphs (Plecoptera): observations of marked individuals in a Rocky Mountain stream. Freshwater Biology 25, 379-394.

Frison,TH 1935 New North American species of the genus Alloperla (Plecoptera: Chloroperlidae). Transactions of the American Entomological Society, 61:331-344.

Frison,TH 1937 II. Descriptions of Plecoptera with special reference to the Illinois species. Illinois Natural History Survey Bulletin 21 (3) 78-99. Click here, then scroll down, click on this publication for instructions on how to order it.

Frison,TH 1942a Descriptions, records and systematic notes concerning western North American stoneflies (Plecoptera). Pan-Pacific Entomologist, 18(1) 9-16.

Frison,TH 1942b Studies of North American Plecoptera, with special reference to the fauna of Illinois. Bulletin of the Illinois Natural History Survey 22: 235-355. PDF

Fuller,RL; Stewart,K,W 1977 The food habits of stoneflies (Plecoptera) in the Upper Gunnison River, Colorado. Environmental Entomology 6, 293-302.
     They studied the Gunnison River Plecoptera fauna at Lost Canyon Resort for 3 years in the early 1970's. Article from the Troutfitter website about this paper and Dr. Stewart: http://www.troutfitter.com/index.asp?PageAction=Custom&ID=4.

Fuller,RL and Stewart,KW 1979 Stonefly (Plecoptera) Food habits and prey preference in the Dolores River, Colorado. American Midland Naturalist, 101(1) 170-181. First page
     Abstract: Gut contents of 1013 stonefly nymphs, comprising nine species, from the Dolores River, Colorado, were analyzed from December 1974 - October 1975 and compared with food availability. Pteryonarcyids ingested large quantities of detritus and some moss, moss being a substantial food item in later instar Pternarcella badia nymphs. Perlodids fed primarily on chironomids and simuliids, although Isoperla fulva also ingested mayflies in June. Claassenia sabulosa remained carnivorous throughout development: young nymphs ingested chironomids and small mayflies and larger nymphs ingested caddisflies and mayflies. Horn's Coefficient of Dietary Overlap indicated significant overlap between all perlodids and chloroperlids. It also showed significant overlap between small and large C. sabulosa nymphs, yet selection of different prey sizes indicated resource partitioning. A comparison of food habits with the Gunnison River stoneflies indicated differences between the diets of large and small Claassenia sabulosa, with chironomids comprising large percentages of the diet for both size classes in the Gunnison River and smaller nymphs in the Dolores. Mayflies were important prey for larger individuals in the Dolores River. These differences could be attributed to different prey populations in each river and/or to availability of prey in the particular size that each predator preferred. In both rivers, Chironomidae and Simuliidae larvae were the major prey in the guts of Cultus aestivalis and Isoperla fulva. This prey specificity may have been due to decreased availability of smaller individuals in the other major prey groups or a difficulty in capture of larger prey organisms.

Fullington,KE and Stewart,KW 1980 Nymphs of the stonefly genus Taeniopteryx (Plecoptera: Taeniopterygidae) of North America. Journal of the Kansas Entomological Society, pp.237-259.
     Abstract: "Nymphs of the nine Nearctic Taeniopteryx species were reared and studied during 1976-1978. Nymphal associations of the 839 specimens examined corresponded with the two established adult groupings, Taeniopterys burksi-maura, and T. lita-lonicera-starki complexes. A key separating seven of the nine species, based primarily upon pigment patterns and abdominal setal arrangements was constructed. Taeniopteryx lita and T. starki were indistinguishable; T. burksi can be separated from T. maura when no developing femoral spur is present. Mouthparts were not species-diagnostic. Detailed habitus illustrations were made for six species. SEM study revealed that eggs of three species were 1.2-1.4 mm in diameter, with a highly sculptured chorion, generally resembling a Maclura fruit; micropyles were scattered. Taeniopteryx lita, lonicera, starki and ugola nymphs were reared and described for the first time. Previously unreported nymphal and adult distribution records are included."
So far Taeniopteryx species have only been reported from the front range and plains of Colorado, but I included this reference to be complete.


Gaufin,AR 1964 Systematic list of Plecoptera of intermountain region. Proceedings of the Utah Academy of Science, Arts and Letters 41, 221-227.
      When published this was a useful Western United States list, now you may want to refer to the 2009 Valid Stonefly Names for North America PDF More details about individual species may be found at Plecoptera species files

Gaufin,AR 1970 Type species designation for the subgenus Utacapnia (Plecoptera: Capniidae). Entomological News 81:197.
     Describes the genus Utacapnia.

Gaufin,AR 1973 Water quality requirements of aquatic insects (Vol. 4). USGPO. html

Gaufin,AR and Ricker,WE 1974 Additions and corrections to a list of Montana stoneflies. Entomological News 85: 285-288.

Gaufin,AR; Ricker,ER; Miner,M; Milam,P and Hays,RA 1972 The stoneflies (Plecoptera) of Montana. Transactions of the American Entomological Society 98(1):1-161.

Gehrken,U; Somme,L. 1987 Increased cold hardiness in eggs of Arcynopteryx compacta (Plecoptera) by dehydration. Journal of Insect Physiology 33(12) 987-991.

Gibert,J Danielopol,D and Stanford,JA 1994 Groundwater ecology. Academic Press 571 pages.

Gill,BA; Harrington,RA; Kondratieff,BC; Zamudio,KR; Poff,NL and Funk,WC 2014 Morphological taxonomy, DNA barcoding, and species diversity in southern Rocky Mountain headwater streams. Freshwater Science 33(1) 288-301

Gill,BA Kondratieff,BC and Sandberg,JB 2015 Evaluation of the morphological species concepts of 16 western Nearctic Isoperla species (Plecoptera: Perlodidae) and their respective species groups using DNA barcoding. Illiesia: International Journal of Stonefly Research, 11(11), 130-146. PDF

Gill,BA; Sandberg,JB and Kondratieff,BC 2015 Evaluation of the morphological species concepts of 16 western Nearctic Isoperla species (Plecoptera: Perlodidae) and their respective species groups using DNA barcoding. Illiesia, 11(11) 130-146. PDF

Goodyear,KL and McNeill,S 1999 Bioaccumulation of heavy metals by aquatic macro-invertebrates of different feeding guilds: a review. Science of the Total Environment, 229(1) 1-19. PDF

Gray,LJ and Ward,JV 1979 Food habits of stream benthos at sites of differing food availability. American Midland Naturalist 102 1, 157-167.

Gregory,J.S., Beesley,S.S. and Van Kirk,R.W. 2000 Effect of springtime water temperature on the time of emergence and size of Pteronarcys californica in the Henry's Fork catchment, Idaho, U.S.A. Freshwater Biology 45(1) 75 Abstract

Grubbs,SA; Baumann,RW; DeWalt,RE and Tweddale,T 2014. A review of the Nearctic genus Prostoia (Ricker)(Plecoptera, Nemouridae), with the description of a new species and a surprising range extension for P. hallasi Kondratieff and Kirchner. ZooKeys, 401: 11-30. html


Hagen,HA, 1874 Report on the Pseudo-neuroptera and Neuroptera collected by Lieut. W.L. Carpenter in 1873 in Colorado. Annual Report of the U.S. Geological and Geographical Survey of the Territories, embracing Colorado, 7: 571-577. Google Books

Hanson,JF 1946 Comparative morphology and taxonomy of the Capniidae (Plecoptera) American Midland Naturalist 35(1) 193-249 first page

Harper,PP; Lauzon,M; Harper,F 1991 Life cycles of 12 species of winter stoneflies from Quebec (Plecoptera, Capniidae and Taeniopterygidae). Canadian Journal of Zoology 69 3, 787-796.

Harper,PP; Ricker,WE 1994 Distribution of Ontario Stoneflies (Plecoptera). Proceedings of the Entomological Society of Ontario 125, 43-66.

Hassage,RL 1989 Life histories, behavior and space partitioning in selected species of western North American Plecoptera. pHd Dissertation, University of North Texas. 105pgs. PDF
     Abstract: "Five species of stoneflies (Zapada haysi, Plumiperla diversa, Taenionema pacificum, Isoperla petersoni, Arcynopteryx compacta) from the North Slope and Interior of Alaska were examined for seasonal patterns of emergence of adults and growth of nymphs. Generally growth was retarded during the winter in this region, and all species except I. petersoni completed growth prior to January. The life cycles of six stonefly species (Prostoia besametsa, Triznaka signata, Sweltsa coloradensis, Isoperla fulva, Skwala parallela, Claassenia sabulosa) are described from northern New Mexico. In this region growth was generally less retarded during the winter than in Alaska; P. besametsa completed all nymphal growth during late fall and winter. Drumming behavior of a Colorado population of Pteronarcella badia was described using an evolutionary framework to explain the maintenance of signal variation in this species. Laboratory experiments were used to explore the effect of intraspecific and interspecific interactions on spatial partitioning in P. badia and Claassenia sabulosa. P. badia exhibited clumping and distributed itself as the surface area of substrate in low densities; however, in the presence of C. sabulosa its distribution was random and different from available surface area. A field study was used to examine spatial partitioning by three New Mexico stonefly species (I. fulva, P. besametsa, T. signata) and to ascertain patterns of microdistribution relating to several abiotic and biotic factors. Generally, there was an interaction of the measured abiotic parameters (current, water temperature, time) with nymphal size. Additionally, void space and sample volume were successfully used to compare biotic densities among leaf and mineral substrates, which were higher in leaf packs than in mineral substrates."

Hassage,RL and Stewart,KW 1990 Growth and voltinism of five stonefly species in a New Mexico mountain stream. The Southwestern Naturalist, 35 (2)130-134. Abstract and first page

Hawkins,CP 2009 Revised invertebrate RIVPACS model and O/E index for assessing the biological condition of Colorado streams. Prepared by Western Center for Monitoring and Assesment of Freshwater Ecosystems, Department of Watershed Sciences, Utah State University for Colorado Department of Public Health and Environment, Water Quality Control Division-Monitoring Unit. PDF

Heino,J 2011 A macroecological perspective of diversity patterns in the freshwater realm. Freshwater Biology, 56(9), pp.1703-1722.
     Abstract: "The present distributions of stonefly genera in North America, their occurrence as endemics, or as shared with the Far East, Europe and South America, are considered in conjunction with geological history. It is concluded that the Plecoptera of North America have four sources of origin.There was an ancient eastern fauna shared with Europe before the formation of the North Atlantic Ocean. A second group moved in from the west during the formation of the western mountains. After formation of the isthmus one genus moved northward from South America. After the Pleistocene period several species migrated from the Bering Strait region, possibly from an Alaskan refugium. Some of these have clearly moved eastward, but a few may have moved westward."

Heinold,B 2010 The mayflies (Ephemeroptera), stoneflies (Plecoptera), and caddisflies (Trichoptera) of the South Platte River Basin of Colorado, Nebraska, and Wyoming. M.S. Thesis, Colorado State University, Fort Collins, CO 375 pages. 148 distribution maps. PDF

Hering,D and Plachter,H 1997 Riparian ground beetles (Coeloptera, Carabidae) preying on aquatic invertebrates: a feeding strategy in alpine floodplains. Oecologia 111(2):261-270. Abstract
     Quote from the abstract: "Aquatic invertebrates composed 89% of the potential prey for carnivorous terrestrial insects along the Isar. Besides aquatic organisms washed ashore, stoneflies emerging on land are of considerable importance as potential prey for terrestrial predators."

Houston,DD; Satler,JD; Stack,TK; Carroll,HM; Bevan,AM; Moya,AL and Alexander,KD 2022 A phylogenomic perspective on the evolutionary history of the stonefly genus Suwallia (Plecoptera: Chloroperlidae) revealed by ultraconserved genomic elements. Molecular Phylogenetics and Evolution, 166, p.107320. PDF

Hynes,HBN 1976. The biology of Plecoptera. Annual Review of Entomology 21: 135-153.

Hynes,HBN 1988 Biogeography and origins of the North American stoneflies (Plecoptera). The Memoirs of the Entomological Society of Canada, 120(S144), pp.31-37.


Illies,J 1965 Phylogeny and zoogeography of the Plecoptera. Annual Review of Entomology (10) 117-140. PDF

Illies,J 1966 Katalog der rezenten Plecoptera. Das Tierreich - Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen. (Das Tierreich) 82:435


Jacobi,GZ; Cary,SJ 1996 Winter stoneflies (Plecoptera) in seasonal habitats in New Mexico, USA. Journal of the North American Benthological Society 15 4, 690-699.

Johnson, NF; Triplehorn, CA 2004 Borror and DeLong's Introduction to the Study of Insects 7th edition, Brooks Cole. 864 pages.


Kauwe,JSK and Shiozawa,DK, 2004 Phylogeographic and nested clade analysis of the stonefly Pteronarcys californica (Plecoptera: Pteronarcyidae) in the western USA. Journal of the North American Benthonlogical Society 23(4)824-838. PDF

Klapálek,F 1905 Conspectus Plecopterorum Bohemiae. Casopis Ceskoslovenské Spolecnosti Entomologické 2: 27-32.

Klapálek,F 1909 In Klapálek and Grunberg. Hft. 8. Ephemerida, Plecoptera, Lepidoptera. In Brauer, A. Die Süsswasserfauna Deutschlands. Eine Exkursionsfauna 39

Klapálek, Frantisek 1912 Plécoptères. I. Fam. Perlodidae; [monographische Revision. II. Fam. Perlidae; Subfam. Perlinae, Subfam. Neoperlinae; mongraphische Revision] Series Sélys-Longchamps, Edmond de, baron, 1813-1900. Collections zoologiques; catalogue systematique et descriptif, fasc. 4, pt. 1-2.

Kiffney,PM 1996 Main and interactive effects of invertebrate density, predation, and metals on a Rocky Mountain stream macroinvertebrate community. Can. J. Fish. Aquat. Sci. 53(7): 1595-1601 .

Kiffney,PM; Clements,WH 1993 Bioaccumulation of heavy metals by benthic invertebrates at the Arkansas River, Colorado. Environmental Toxicology and Chemistry 12, 1507-1517.

Kiffney,PM; Clements,WH 1994 Effects of heavy metals on a macroinvertebrate assemblage from a Rocky Mountain stream in experimental microcosms. Journal of the North American Benthological Society 13 4, 511-523.
     Quote from page 519-520: "Our results were similar to other experiments (Clements et al. 1988a, 1988b, Leland et al. 1989, Kiffney and Clements 1994) and field studies (Chadwick et al. 1986, Clements 1994; Clements, unpublished results) that have examined the effects of metals on stream macroinvertebrates. Specifically, mayflies and some stoneflies were sensitive, and caddisflies and chironomids were relatively tolerant to metal exposure. However, the sensitivity to metals differed within families, genera and across lifestages."

Knight,AW and Gaufin,AR 1966 Altitudinal distribution of stoneflies (Plecoptera) in a Rocky Mountain drainage system. Journal of the Kansas Entomological Society 39 4, 668-675. First page
      Abstract: "Stoneflies collected between the altitude of 6,760 and 10,770 feet in the Gunnison River drainage, Colorado, show definite altitudinal distribution. The effect of altitude on stonefly distribution is largely due to temperature and, in part, food supply. The greatest number of stonefly species was recorded between 7,000 and 9,000 feet. Carnivorous stonefly species were dominant at all elevations and reached high dominance values at the higher elevations."
      Stonefly names have changed quite a bit since this publication.
Figure 1. Synonyms (Taxonomy Translation) page 669
Knight and Gaufin Name 2023 Name (Names that have stayed the same are not listed)
Alloperla pintada Triznaka_pintada
Nemoura oregonensis Zapada oregonensis
Brachyptera pallidula Suwallia pallidula
Capnia poda Utacapnia poda
Capnia logana Utacapnia logana
Capnia limita Capnia vernalis
Nemoura besametsa Prostoia besametsa
Nemoura cinctipes Zapada cinctipes
Nemoura coloradensis Capnia coloradensis
Brachyptera pacifica Hesperoperla pacifica
Isogenus modesta Kogotus modestus
Alloperla borealis Sweltsa borealis
Leuctra sara Paraleuctra vershina
Isogenus aestivalis Cultus aestivalis
Arcynopteryx signata Megarcys signata
Arcynopteryx parallela Skwala americana
Alloperla signata Triznaka signata
Isogenus elongatus Isogenoides elongatus
Isoperla patricia Isoperla quinquepunctata
Alloperla coloradensis Sweltsa coloradensis
Alloperla pallidula Suwallia pallidula
Acroneuria pacifica Hesperoperla pacifica

Knight,AW and Gaufin,AR 1967 Stream type selection and associations of stoneflies (Plecoptera) in a Colorado River drainage system. Journal of the Kansas Entomological Society, 40(3) 347-352.
     Abstract: "A stream classification system for the Gunnison River drainage, Colorado is given. The stonefly fauna has been placed into appropriate stream types. In order to correlate the occurrence of a given species with any other species, an association diagram has been constructed. The stream type classification of stoneflies indicates the association of stoneflies according to stream types; the stonefly association diagram reveals the local association of stoneflies."
This and the previous paper by Knight and Gaufin are interesting partly because they were conducted before the Curecanti storage unit was built on the Gunnison river. The Curecanti storage unit (also called the Aspinall unit) consists of the Blue Mesa and Morrow Point dams.

Kondratieff,BC and Baumann,RW 2002 A review of the stoneflies of Colorado with description of a new species of Capnia (Plecoptera: Capniidae). Transactions of American Entomological Society 128 (3) 385-401.
     Abstract:"Eighty-six species of stoneflies are reported from Colorado, including a new species of Capnia. Three new state records are reported, Bolshecapnia milami (Nebeker and Gaufin), Capnura fibula (Claassen), and Isoperla marlynia (Needham and Claassen). Ninety percent of all Colorado stoneflies are typical western North American species, with seven species, Paracapnia angulata Hanson, Taeniopteryx burksi Ricker and Ross, T. parvula Banks, I. marlynia, Acroneuria abnormis (Newman), Perlesta decipiens (Walsh), and Isoperla bilineata (Say) having eastern affinities. Arcynopteryx compacta (McLachlan) is considered circumpolar species."
Overview of Plecoptera in Colorado. Briefly refers to some nomenclature changes, habitat, distribution, other comments and useful publications about Colorado Stoneflies. This website (www.gunnisoninsects.org) quotes short paragraphs about species found in Gunnison County in the reference section of each species webpage.


Larson,EI; Poff,NL; Atkinson,CL and Flecker,AS 2018 Extreme flooding decreases stream consumer autochthony by increasing detrital resource availability. Freshwater Biology, 63(12), pp.1483-1497. PDF

Leiser,E and Boyle,R 1990 Stoneflies for the Angler: How to Know Them, Tie Them, and Fish Them. Stackpole Books 174 pages.

Lyon,ML; Stark,BP 1997 Alloperla (Plecoptera: Chloroperlidae) of western North America. Entomological News 108 5, 321-334
     The key for Alloperla, has scanning electron photomicrographs of eggs and pertinent taxonomic characters.


Maketon,M;Stewart,KW; Kondratieff,BC and Kirchner,RF 1988 New descriptions of drumming and evolution of the behavior in North American Perlodidae (Plecoptera). Journal of the Kansas Entomological Society. 61(2) 161-168 Abstract and first page

Malison,RL; DelVecchia,AG; Woods,HA; Hand,BK; Luikart,G and Stanford,JA 2020 Tolerance of aquifer stoneflies to repeated hypoxia exposure and oxygen dynamics in an alluvial aquifer. Journal of Experimental Biology, 223(16). PDF

Malison,RL; Ellis,BK; DelVecchia,AG; Jacobson,H; Hand,BK; Luikart,G; Woods,HA; Gamboa,M; Watanabe,K and Stanford,JA 2020 Remarkable anoxia tolerance by stoneflies from a floodplain aquifer. Ecology, 101(10), p.e03127. PDF
     Abstract: "Alluvial aquifers are key components of river floodplains and biodiversity worldwide, but they contain extreme environmental conditions and have limited sources of carbon for sustaining food webs. Despite this, they support abundant populations of aquifer stoneflies that have large proportions of their biomass carbon derived from methane. Methane is typically produced in freshwater ecosystems in anoxic conditions, while stoneflies (Order: Plecoptera) are thought to require highly oxygenated water. The potential importance of methane-derived food resources raises the possibility that stonefly consumers have evolved anoxia-resistant behaviors and physiologies. Here we tested the anoxic and hypoxic responses of 2,445 stonefly individuals in three aquifer species and nine benthic species. We conducted experimental trials in which we reduced oxygen levels, documented locomotor activity, and measured survival rates. Compared to surface-dwelling benthic relatives, stoneflies from the alluvial aquifer on the Flathead River (Montana) performed better in hypoxic and anoxic conditions. Aquifer species sustained the ability to walk after 4-76 h of anoxia vs. 1 h for benthic species and survived on average three times longer than their benthic counterparts. Aquifer stoneflies also sustained aerobic respiration down to much lower levels of ambient oxygen. We show that aquifer taxa have gene sequences for hemocyanin, an oxygen transport respiratory protein, representing a possible mechanism for surviving low oxygen. This remarkable ability to perform well in low-oxygen conditions is unique within the entire order of stoneflies (Plecoptera) and uncommon in other freshwater invertebrates. These results show that aquifer stoneflies can exploit rich carbon resources available in anoxic zones, which may explain their extraordinarily high abundance in gravel-bed floodplain aquifers. These stoneflies are part of a novel food web contributing biodiversity to river floodplains."

Mani,MS 1968 Ecology and biogeography of high altitude insects (Vol. 4). Springer-Verlag New York 541 pages.

Marden,JH and Kramer,MG 1994 Surface-skimming stoneflies: a possible intermediate stage in insect flight evolution. Science, New Series, 66(5184) 427-427. PDF
     Abstract: " lnsect wings appear to have evolved from gills used by aquatic forms for ventilation and swimming, yet the nature of intermediate stages remains a mystery. Here a form of nonflying aerodynamic locomotion used by aquatic insects is described, called surface skimming, in which thrust is provided by wing flapping while continuous contact with the water removes the need for total aerodynamic weight support. Stoneflies surface skim with wing areas and muscle power output severely reduced, which indicates that surface skimming could have been an effective form of locomotion for ancestral aquatic insects with small protowings and low muscle power output."

Marden,JH; O’Donnell,BC; Thomas,MA and Bye,JY 2000 Surface-skimming stoneflies and mayflies: The taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion. Physiological and Biochemical Zoology, 73(6), 751-764. PDF

McCafferty, WP 1983 Aquatic Entomology: The Fishermens Guide and Ecologists Illustrated Guide to Insects and Their Relatives. Jones and Bartlett Publishers, Inc. 480 pages.

Merritt,RW; Cummins,KW (Eds.) 1996 An Introduction to the Aquatic Insects of North America. 3rd ed. Kendall/Hunt Publishing Company, Dubuque, Iowa. 862 pages.
     The best all around aquatic insect key and general reference for North America until the 2008 edition came out.

Merritt,RW; Cummins,KW and Berg,MB (Eds.) 2008 An Introduction to the Aquatic Insects of North America. 4th ed. Kendall/Hunt Publishing Company, Dubuque, Iowa. 1158 pages.
     The latest edition of a classic aquatic entomology key. Required for all serious aquatic insect identification in America.

McCafferty, WP 1983 Aquatic Entomology: The Fishermens Guide and Ecologists Illustrated Guide to Insects and Their Relatives. Jones and Bartlett Publishers, Inc. 480 pages.

Milner,AM 1987 Colonization and ecological development of new streams in Glacier Bay National Park, Alaska. Freshwater Biology, 18(1), pp.53-70.

Misof,B; Liu,S; Meusemann,K; Peters,RS; Donath,A; Mayer,C; Frandsen,PB; Ware,J; Flouri,T; Beutel,RG; Niehuis,O; et al. 2014 Phylogenomics resolves the timing and pattern of insect evolution. Science, 346(6210), pp.763-767. PDF
     Abstract: "Insects are the most speciose group of animals, but the phylogenetic relationships of many major lineages remain unresolved. We inferred the phylogeny of insects from 1478 protein-coding genes. Phylogenomic analyses of nucleotide and amino acid sequences, with site-specific nucleotide or domain-specific amino acid substitution models, produced statistically robust and congruent results resolving previously controversial phylogenetic relationships. We dated the origin of insects to the Early Ordovician [~479 million years ago (Ma)], of insect flight to the Early Devonian (~406 Ma), of major extant lineages to the Mississippian (~345 Ma), and the major diversification of holometabolous insects to the Early Cretaceous. Our phylogenomic study provides a comprehensive reliable scaffold for future comparative analyses of evolutionary innovations among insects."

Molles,MC and Pietruszka,RD 1983 Mechanisms of prey selection by predaceous stoneflies: roles of prey morphology, behavior and predator hunger. Oecologia 57(1) 25-31. Abstract

Mogren,CL and Trumble,JT 2010 The impacts of metals and metalloids on insect behavior. Entomologia Experimentalis et Applicata, 135: 1-17. Full Text

Muchow,CL and Richardson,JS 1999 Unexplored diversity: macroinvertebrates in coastal British Colombia headwater streams. In Proceedings of a Conference on the Biology and Management of Species and Habitats at Risk, Kamloops, BC (2) 503-506. PDF
     Abstract: " At the very tips of most drainage networks are small stream channels, many of which have no flow at times during the summer. These small, zero-order streams contribute a significant portion of cumulative stream length in a watershed, but are largely unexplored and receive little protection under current legislation. We studied the aquatic macroinvertebrate assemblage of 7 small coastal streams with a range of permanence of flow. Three streams have no detectable surface flow for periods in summer (i.e., intermittent flow). We hypothesize that the invertebrate assemblage in these zero-order streams does not completely overlap that of larger streams. Stream sites were sampled using a combination of emergence traps emptied every 14 days throughout the year. In even the smallest streams (<0.5 m bankfull width) with intermittent flow, true aquatic insects with 1-year life cycles were found emerging, even in periods when no flow was perceptible. Species richness in intermittent and continuous streams was approximately equal, while intermittent streams appeared to produce as much as twice the number of adult stoneflies as continuous streams. This study shows that even intermittent streams harbour a true aquatic fauna and, potentially, species for which little is known, and indicates that these unique habitats make an important contribution to their ecosystems."

Mutch,RA and Pritchard,G 1986 Development rates of eggs of some Canadian stoneflies (Plecoptera) in relation to temperature. Journal of the North American Benthological Society 5(4)272-277. Abstract and first page


Neave,F 1934 Stoneflies from the Purcell Range, B.C. The Canadian Entomologist 66(1):1-6.

Nebeker,AV and Gaufin,AR 1965 The Capnia columbiana complex of North America (Capniidae: Plecoptera). Transactions of the American Entomological Society 91:467-487.

Nebeker,AV and Gaufin,AR 1966 New Paraleuctra from the Rocky Mountains (Plecoptera: Leuctridae). Entomological News 77:255-259.

Nebeker,AV and Gaufin,AR 1967 Geographic and seasonal distribution of the family Capniidae of western North America (Plecoptera). Journal of the Kansas Entomological Society 40(3)415-421 Abstract and first page

Nebeker,AV and Gaufin,AR 1967 Geographic and seasonal distribution of the family Capniidae of Western North America (Plecoptera). Journal of the Kansas Entomological Society 40(3) 415-421. Abstract and first page

Needham,JG and Claassen,PW 1925 A Monograph of the Plecoptera of North America. Entomological Society of America, Lafayette, Indiana. 397 pages.

Nelson,CH 2009 Surface ultrastructure and evolution of tarsal attachment structures in Plecoptera (Arthropoda: Hexapoda). Aquatic Insects, (31)523-545. PDF
     Abstract: "The plantar surfaces of the stonefly tarsomeres and pretarsus are examined chiefly using scanning electron microscopy (SEM) of 39 Plecoptera exemplar species representing each family within the two suborders Arctoperlaria and Antarctoperlaria. Features examined include the shape and size of the tarsomeres as well as external features of the plantar surfaces of the tarsomeres, unguitractor plate, claws, arolium and orbicula. Characteristics presumed to be part of the stonefly ground pattern were determined by mapping these features on a cladogram of stonefly phylogeny (Zwick 2000 Zwick, P. 2000. ‘Phylogenetic system and zoogeography of the Plecoptera’. Annual Review of Entomology, 45: 709–746. [Crossref], [PubMed], [Web of Science ®], , [Google Scholar], Annual Review of Entomology 45:709–746). This analysis indicates that the following characters are part of the stonefly ground pattern: (1) cylindrical shape of the tarsomeres, (2) an elongate tarsomere 1, (3) a short tarsomere 2, (4) presence of hairy euplantulae on the plantar surfaces of tarsomeres 1 and 2, (5) absence of specialised attachment surfaces on the plantar area of tarsomere 3, (6) absence of setae on the arolium plantar surface, and (7) presence of setae on the orbicula surface. Hairs covering the euplantulae of tarsomeres 1 and 2 and the median longitudinal unscelerotised region of tarsomeres 1–3 are non-setal cuticular projections and appear to be acanthae. Hairy euplantulae are unusual and are only present in one other insect order, the Mantophasmatodea. Specialised tenent setae were not found. The presence of hairy euplantulae as part of the stonefly ground pattern contradicts the recent view (Beutel and Gorb 2006 Beutel, R. G. and Gorb, S. N. 2006. ‘A revised interpretation of the evolution of attachment structures in Hexapoda with special emphasis on Mantophasmatodea’. Arthropod Systematics & Phylogeny, 64: 3–25. [Google Scholar], Arthropod Systematics & Phylogeny 64:3–25) that stoneflies lack pad-like euplantulae and that the ancestral condition for tarsomeres 1 and 2 may be a narrow median longitudinal unsclerotised band such as that found in representatives of the Austroperlidae. The hypothesis that Plecoptera are the sister clade to the remaining Neoptera suggests that euplantulae might be a synapomorphy of both clades. On the other hand, placement of Plecoptera within a monophyletic Polyneoptera clade within the Neoptera suggests that euplantulae could be an autapomorphy of this subordinate clade."

Nelson,RC; Baumann,RW 1989 Systematics and distribution of the winter stonefly genus Capnia (Plecoptera: Capniidae) in North America. Great Basin Naturalist (49) 289-363. PDF

Nelson,CH and Hanson,JF 1971 Contribution to the anatomy and phylogeny of the family Pteronarcidae (Plecoptera). Transactions of the American Entomological Society (1890-), 97(1), pp.123-200.

Nelson,CH and Hanson,JF 1973 The genus Perlomyia (Plecoptera: Leuctridae). Journal of the Kansas Entomological Society 46:187-199.

Nelson,CH and Hanson,JF 1968 Two New Species of Alloperla (Plecoptera: Chloroperlidae) from China. Journal of the Kansas Entomological Society (41)4 425-428. Abstract and First Page

Nelson,RC and Kondratieff,BC 1988 A new species of Capnia (Plecoptera: Capniidae) from the Rocky Mountains of Colorado. Entomological News 99 2, 77-80.

Nelson,SM and Roline,RA 1999 Relationships between metals and hyporheic invertebrate community structure in a river recovering from metals contamination. Hydrobiologia 397, 211-226. Abstract

Nehring,RB 1976 Aquatic insects as biological monitors of heavy metal pollution. Bulletin of Environmental Contamination and Toxicology 15 2, 147-154.

Newport,G 1848 On the anatomy and affinities of Pteronarcys regalis Newman; with a postscript containing descriptions of some American Perlidae, together with notes on their habits. Transactions Linnean Society of London 20: 425-452.



Peckarsky,BL 1980 Predator-prey interactions between stoneflies and mayflies: behavioral observations. Ecology 61(4) 932-943. Abstract

Peckarsky,BL 1986 Colonization of natural substrates by stream benthos. Canadian Journal of Fisheries and Aquatic Sciences 43, 700-709.

Peckarsky,BL 1987b Mayfly cerci as defense against stonefly predation: deflection and detection. Oikos 48 2, 161-170.

Peckarsky,BL 1990 Habitat selection by stream-dwelling predatory stoneflies. Canadian Journal of Fisheries and Aquatic Sciences 48, 1069-1076.

Peckarsky,BL 1991 A field test of resource depression by predatory stonefly larvae. Oikos 61 1, 3-10.

Peckarsky,BL 1991 Is there a coevolutionary arms race between predators and prey? A case study with stoneflies and mayflies. Advances in Ecology 1, 167-180.

Peckarsky,BL 1991 Mechanisms of intra- and interspecific interference between larval stoneflies. Oecologia 85(4) 521-529. Abstract

Peckarsky,BL 1996 Alternative predator avoidance syndromes of stream-dwelling mayfly larvae. Ecology 77(6), 1888-1905. Abstract

Peckarsky,BL and Cowan,CA 1991 Consequences of larval intraspecific competition to stonefly growth and fecundity. Oecologia 88, 277-288.

Peckarsky,BL and Cowan,CA 1995 Microhabitat and activity periodicity of predatory stoneflies and their mayfly prey in a western Colorado stream. Oikos, 513-521. PDF

Peckarsky,BL; Cowan,CA and Anderson,CR. 1994 Consequences and plasticity of the specialized predatory behavior of stream-dwelling stonefly larvae. Ecology 75(1) 166-181. Abstract

Peckarsky,BL; Cowan,CA; Penton,MA and Anderson,CR 1993 Sublethal consequences of stream-dwelling predatory stoneflies on mayfly growth and fecundity. Ecology 74(6):1836-1846. Abstract

Peckarsky,BL and Dodson,SI 1980. Do stonefly predators influence benthic distributions in streams? Ecology 61(6) 1275-1282. Abstract

Peckarsky,BL and Dodson,SI 1980. An experimental analysis of biological factors contributing to stream community structure. Ecology 61:1283-1290. Abstract

Peckarsky,BL, Kerans,B; Taylor,BW and McIntosh,AR 2008 Predator effects on prey population dynamics in open systems. Oecologia 156(2) 431-440. PDF

Peckarsky,BL and McIntosh,AR 1998 Fitness and community consequences of avoiding multiple predators. Oecologia 113, 565-576. Abstract

Peckarsky,BL and Penton,MA 1985 Is predaceous stonefly behavior affected by competition? Ecology 66(6) 1718-1728. Abstract

Peckarsky,BL and Penton,MA 1988 Why do Ephemerella nymphs scorpion posture: a "ghost of predation past"? Oikos 53:185-193.

Peckarsky,BL and Penton,MA 1989 Early warning lowers risk of stonefly predation for a vulnerable mayfly. Oikos 54:301-309.

Peckarsky,BL and Penton,MA 1989 Mechanisms of prey selection by stream-dwelling stonefly nymphs. Ecology 70(5) 1203-1218. Abstract

Peckarsky,BL and Penton,MA 1990 Effects of enclosures on stream microhabitat and invertebrate community structure. Journal of the North American Benthological Society 9:249-261.

Peckarsky,BL and Wilcox,RS 1989 Stonefly nymphs hydrodynamic cues to discriminate between prey. Oecologia 79:265-270.

Peckarsky,BL; Cowan,CA; Penton,MA and Anderson,C 1993 Sublethal consequences of stream-dwelling predatory stoneflies on mayfly growth and fecundity. Ecology 74(6):1836-1846. Abstract

Peckarsky,BL; Cowan,CA and Anderson,CR 1994 Consequences and plasticity of the specialized predatory behavior of stream-dwelling stonefly larvae. Ecology 75(1):166-181. Abstract

Peckarsky,BL; Dodson,SI and Conklin,DJ 1985 A key to the aquatic insects of streams in the vicinity of the Rocky Mountain Biological Lab, including chironomid larvae from streams and ponds. Colorado Division of Wildlife, Denver CO. 47 pages.
     This was one of the publications that inspired me to start developing www.gunnisoninsects.org :-) Do not use the Chloroperlidae section of this key.

Peckarsky,BL; Fraissinet,PR; Penton,MA and Conklin Jr.,DJ 1990 Freshwater Macroinvertebrates of Northeastern North America. Cornell University, Ithaca, NY. 442 pages.
     While not directly aimed at our area, it works pretty well to the genus level, useful as another tool on the shelf when identifying nymphs.

Peckarsky,BL; Horn,SC and Statzner,B 1990. Stonefly predation along a hydraulic gradient: a test of the harsh-benign hypothesis. Freshwater Biology 24:1503-1514.

Peckarsky,BL; Taylor,B and Caudill,CC 2000 Hydrologic and behavioral constraints on oviposition of stream insects: implications for adult dispersal. Oecologia 125:186-200. Abstract

Pennack,RW 1978 Fresh-water Invertebrates of the United States. 2nd ed. Wiley-Interscience, New York, NY. 803 pages.
     Now mostly outdated, see Merritt and Cummins 1996. Has wonderful illustrations and discussions. The current edition no longer has keys to aquatic insects.

Pennack,RW and Ward,JV 1986 Interstital faunal communities of the hyporheic and adjacent groundwater biotopes of a Colorado mountain stream. Archiv für Hydrobiologie Suppl. 74 3, 356-396.

Perry,JA and Schaeffer,DJ 1987 The longitudinal distribution of riverine benthos: A river dis-continuum?. Hydrobiologia, 148(3) 257-268.
     They studied Tomichi Creek in Gunnison County.

Petrin,Z 2011 Species traits predict assembly of mayfly and stonefly communities along pH gradients. Oecologia, 167(2), 513-524. Abstract
     Abstract: " Much recent ecological research has centred on the interrelations between species diversity and ecological processes. In the present study, I show how species traits may aid in comprehending ecology by studying the link between an environmental variable and functional traits. I examined the composition of species traits with a theoretically underpinned relationship to ecological processes along a pH gradient. I focused on body size, reproductive output, life cycle length and feeding habit of mayflies and stoneflies. In mayfly assemblages, I found smaller body size, greater reproductive output, faster life cycles and a larger proportion of gathering collectors and scrapers with increasing pH. In stonefly assemblages, I found smaller body size, greater reproductive output and faster life cycles at sites with a history of long-term natural acidification, but no clear trends in feeding habits and in most traits where acidification is anthropogenic. The results suggest that mayflies and stoneflies exhibit different ecological functions following different ecological strategies. Mayflies follow an opportunistic strategy relative to stoneflies, likely facilitating high rates of ecological processes with respect to the autotrophic resource base at neutral sites. Relative to mayflies, stoneflies follow an equilibrium strategy contributing to ecological functioning in heterotrophic ecosystems and likely maintaining heterotrophic processes despite the erosion of species diversity in response to acidification. The rules governing an ecological community may be more readily revealed by studying the distribution of species traits instead of species diversity; by studying traits, we are likely to improve our understanding of the workings of ecological communities. "



Rader,RB and Belish,TA 1999 Influence of mild to severe flow alterations on invertebrates in three mountain streams. Regulated Rivers: Research & Management. 15(4)353 - 363.
     Quote: "Water abstraction (extent and timing of diversion) could be managed to minimize risks to downstream ecological resources."

Rader,RB and Ward,JV 1988 Influence of regulation on environmental conditions and the macroinvertebrate community in the upper Colorado River. Regulated Rivers: Research and Management 2:597-618. PDF

Radford,DS and Hartland-Rowe,R 1971 Emergence patterns of some Plecoptera in two mountain streams in Alberta. Canadian Journal of Zoology, 49(5), 657-662.

Richardson,JW and Gaufin,AR 1971 Food habits of some western stonefly nymphs. Transactions of American Entomological Society 97, 91-121.
     They studied a number of rivers and streams in the Gunnison basin and Utah, providing detailed dietary information for the most common species.

Ricker,WE 1943. Stoneflies of Southwestern British Columbia. Indiana University Publications, Science Series 12 145 pages, Bloomington, Indiana.

Ricker,WE 1952. Systematic studies in Plecoptera. Indiana University Publications, Science Series 18, 200 pages, Bloomington, Indiana. PDF

Ricker,WE 1959 The species of Isocapnia Banks (Insecta, Plecoptera, Nemouridae). Canadian Journal of Zoology, 37(5), 639-653.

Ricker,WE 1965 New records and descriptions of Plecoptera (Class Insecta). Journal of the Fisheries Board of Canada, 22(2) 475-501.

Ricker,WE 1992 Origin of stonefly names proposed by Ricker and collaborators. Perla, 18(1) 12 pages. PDF

Robinson,CT and Minshall,GW 1986 Effects of disturbance frequency on stream benthic community structure in relation to canopy cover and season. Journal of the North American Benthological Society, 237-248. PDF

Roline,R 1988 The effects of heavy metals pollution of the upper Arkansas River on the distribution of aquatic macroinvertebrates. Hydrobiologia 160: 3-8.
     They sampled the Arkansas River upstream and downstream of mine drainage and clean water inputs in 1979 and 1980. After compositing 3 surber samplers in the field, they took the samples back to the lab and identified the macroinvertebrates to genus level. The author used a diversity index to evaluate the health of the macroinvertebrate community. Higher diversity is better. Diversity decreased downstream of heavy metal pollution from the Leadville Drain and California Gulch and increased downstream of clean water inputs.

del Rosario,RB; Betts,EA and Resh,VH. 2002 Cow manure in headwater streams: tracing aquatic insect responses to organic enrichment. Journal of the North American Benthological Society 21: 278-289.

Ruse,LP and Herrmann,SJ 2000 Plecoptera and Trichoptera species distribution related to environmental characteristics of the metal-polluted Arkansas River, Colorado. Western North American Naturalist 60 (1) 57-65. PDF


Sánchez-Bayo,F and Wyckhuys,KA 2019 Worldwide decline of the entomofauna: A review of its drivers. Biological conservation, 232, pp.8-27. PDF
     Abstract: "Biodiversity of insects is threatened worldwide. Here, we present a comprehensive review of 73 historical reports of insect declines from across the globe, and systematically assess the underlying drivers. Our work reveals dramatic rates of decline that may lead to the extinction of 40% of the world's insect species over the next few decades. In terrestrial ecosystems, Lepidoptera, Hymenoptera and dung beetles (Coleoptera) appear to be the taxa most affected, whereas four major aquatic taxa (Odonata, Plecoptera, Trichoptera and Ephemeroptera) have already lost a considerable proportion of species. Affected insect groups not only include specialists that occupy particular ecological niches, but also many common and generalist species. Concurrently, the abundance of a small number of species is increasing; these are all adaptable, generalist species that are occupying the vacant niches left by the ones declining. Among aquatic insects, habitat and dietary generalists, and pollutant-tolerant species are replacing the large biodiversity losses experienced in waters within agricultural and urban settings. The main drivers of species declines appear to be in order of importance: i) habitat loss and conversion to intensive agriculture and urbanisation; ii) pollution, mainly that by synthetic pesticides and fertilisers; iii) biological factors, including pathogens and introduced species; and iv) climate change. The latter factor is particularly important in tropical regions, but only affects a minority of species in colder climes and mountain settings of temperate zones. A rethinking of current agricultural practices, in particular a serious reduction in pesticide usage and its substitution with more sustainable, ecologically-based practices, is urgently needed to slow or reverse current trends, allow the recovery of declining insect populations and safeguard the vital ecosystem services they provide. In addition, effective remediation technologies should be applied to clean polluted waters in both agricultural and urban environments."

Sandberg,JB 2009 Vibrational communication (drumming) of the western nearctic stonefly genus Hesperoperla (Plecoptera: Perlidae). Illiesia 2009 5(13):146-155. PDF

Sandberg,JB 2011 Vibrational communication of Isoperla Banks from California and Oregon (Plecoptera: Perlodidae). Illiesia 2011 7(1):1-23. PDF

Sandberg,JB and Kondratieff,BC 2013 The Isoperla of California (Plecoptera: Perlodidae); Updated male descriptions and adult keys for 18 western Nearctic species. Illiesia (9)34-64. PDF

Sandberg,JB and Stewart,KW 2001 Drumming behavior and life history notes of a high-altitude Colorado population of the stonefly Isoperla petersoni Needham & Christenson (Plecoptera:Perlodidae). Western North American Naturalist 61 4, 445-451.

Sandberg,JB; Stewart,KW 2003 Continued studies of drumming in North American Plecoptera; Evolutionary implications. In: Research Update on Ephemeroptera and Plecoptera. Ed: Gaino,E University of Perugia, Perugia, Italy, 73-81.
Discusses stonefly communication via drumming for a number of species including Isoperla fulva , Isoperla mormona, Isoperla sobria and Paraperla frontalis. Also has sonograms of male and female stonefly courting duets. Discusses the evolutionary implications of recent studies of drumming behavior.

Sandberg, JB; and Stewart, KW 2005 Life history of the stonefly Isogenoides zionensis (Plecoptera:Perlodidae) from the San Miguel River, Colorado. Illiesia 1(4)21-32.     http://www2.pms-lj.si/illiesia/Illiesia01-04.pdf

Sandberg,JB and Stewart,KW 2005a Vibrational communication (Drumming) of the nearctic stonefly genus Isogenoides (Plecoptera:Perlodidae). Transactions of American Entomological Society 131 1+2, 111-130. PDF

Sandberg,JB; Stewart,KW 2005b Holomorphology and systematics of the stonefly genus Isogenoides (Plecoptera:Perlodidae). Transactions of American Entomological Society 131 3+4, 269-345.

Sandberg, JB; Stewart, KW 2006 Continued Studies of Vibrational Communication (Drumming) Of North American Plecoptera. Illiesia 2006 2(1):1-14. http://illiesia.speciesfile.orgpapers/Illiesia02-01.pdf (227 Kb)
     Has sonograms of a number of species including the local I. quinquepunctata.

Sanders,HO and Cope,OB 1968 The relative toxicities of several pesticides to naiads of three species of stoneflies. Limnology and Oceanography 13(1) 112-117. First page
     Abstract: Static bioassays were conducted to determine the relative acute toxicities of some insecticides, herbicides, fungicides, a defoliant, and a molluscicide to the naiads of three species of stonefly, Pteronarcys californica, Pteronarcella badia, and Claassenia sabulosa. Toxic effects were measured by determination of median lethal concetration (LC50) for 24-, 48-, and 96-hr exposures, at 15.5C. Endrin and dieldrin were the most and DDT the least toxic of the chlorinated hydrocarbon insecticides tested. Parathion was the most toxic organophosphate insecticide to P. californica naiads, but Dursban was the most toxic to P. badia and C. sabulosa naiads. Trichlorofon (Dipterex) was the least toxic to all three species. P. badia, the species of smallest size, was the species most susceptible to most pesticides, followed in descending order of sensitivity by C. sabulosa and P. californica. Smaller specimens of P. californica naiads were consistently more susceptible to some insecticides than larger specimens of the same species.

Scudder, GE 2006 Bill Ricker's Entomological Contributions. Environmental Biology of Fishes 75(1) 111-117. Abstract

Shapas,TJ; Hilsenhoff,WL 1976 Feeding habits of Wisconsin's predominant lotic Plecoptera, Ephemeroptera and Trichoptera. Great Lakes Entomologist 9, 175-188.

Sheldon,AL 1969 Comparative ecology of Arcynopteryx and Diura (Plecoptera) in a California stream. Archiv fur Hydrobiologie 69, 521-546.

Sheldon,AL 1999 Emergence patterns of large stoneflies (Plecoptera: Pteronarcys, Calineuria, Hesperoperla) in a Montana river. Great Basin Naturalist 59: 169-174. PDF

Shepard,WD and Stewart,KW 1983 Comparative Study of Nymphal Gills in North American Stonefly Genera and a New, Proposed Paradigm of Plecoptera Gill Evolution. Miscellaneous Publications of the Entomological Society of America 13:1-57
     Reviews history of describing and naming gills on the stoneflies of North America. Standarizes the terminology of gills. Suggests since function of the things we call gills appears to really be for osmoregulation that we call most of the gills on stonefly nymphs osmobranchiae. The only truly respiratory gills on stoneflies are the gills variously referred to as anal, caudal (historic terms) or subanal lobe (SL) gills found on Perlidae and Pteronarcyidae nymphs. Perlid and Pteronarcid nymphs grow more gills and more branches on their gills with sucessive instars. Many other gilled stoneflies gain gills as they grow making identifications based on gills problematic for immature specimens.

Short,RA and Ward,JV 1980 Life cycle and production of Skwala parallela (Frison) (Plecoptera:Perlodidae) in a Colorado montane stream. Hydrobiologia 69 3, 273-275.
     Now referred to as Skwala americana, Short and Ward provide life history and production data from Trout Creek in the South Platte River drainage of Colorado's Front Range.

Short,RA and Ward,JV 1980 Macroinvertebrates of a Colorado high mountain stream. The Southwestern Naturalist, 23-32. PDF

Short,RA and Ward,JV 1981 Trophic ecology of three winter stoneflies (Plecoptera). American Midland Naturalist 105, 341-347.
They studied Zapada oregonensis, Z. cinctipes and Capnia confusa in Little Beaver Creek in North Central Colorado. Diet analysis of field caught nymphs showed that they mostly were shredders who chewed up leaves into smaller pieces.

Smith,LW 1917 Studies of North American Plecoptera (Pteronarcinae and Perlodini) Transactions of the American Entomological Society 43(4):433-489. PDF

South,EJ; Skinner,RK; DeWalt,RE; Kondratieff,BC; Johnson,KP; Davis,MA; Lee,JJ and Durfee,RS 2021 Phylogenomics of the north American Plecoptera. Systematic Entomology, 46(1), pp.287-305. HTML
     Abstract: "Stoneflies (Insecta: Plecoptera) provide essential ecosystem services and are vital components of aquatic ecological systems worldwide. Despite this importance, a well-supported and fully-resolved phylogeny of the order has remained elusive for over a century. Using transcriptome data from 94 species, we performed maximum likelihood and multispecies coalescent analyses with 1400 orthologous genes. This taxon sample includes representatives of all families, subfamilies and tribes of the North American fauna, providing the most complete molecular phylogenetic study of the North American Plecoptera to date. Analyses recovered high support for the resolution of previously unresolved or contested relationships and the elucidation of a few novel relationships among historically accepted clades. Results included recovering (i) Perlidae as the earliest diverging family of Perloidea, (ii) the clade Nemouridae + Capniidae instead of the traditionally recognized Leuctridae + Capniidae, (iii) Peltoperlidae as sister to four Systellognatha families and (iv) non-monophyly of Chloroperlidae due to placement of the genus Kathroperla Banks. The position of Taeniopterygidae and Leuctridae remain inconclusive, as the placement of these taxa was not consistent between analyses of different data types nor was strong support for their relationships to other stoneflies recovered in a four-cluster likelihood analysis. However, our results for the North American taxa establish a robust foundation for future phylogenetic studies of the Plecoptera world fauna."

Stanford,JA and Gaufin,AR 1974 Hyporheic communities of two Montana rivers. Science 185:700-702. PDF
     Abstract: Collections of stream organisms from a domestic water supply system adjacent to the Tobacco River revealed that a detritus-based community exists in subterranean waters circulating through floodplain gravels at least 4.2 meters below and 50 meters laterally from the river channel. Several stone fly species spend their entire nymphal life cycles in underground habitats of the Tobacco and Flathead rivers.

Stanford,JA and Ward,JV 1985 The effects of regulation on the limnology of the Gunnison River: A North American case history. In: Regulated Rivers. Eds: Lillehammer,A; Saltveit,S Universitetsforlaget As., Oslo, Norway, 467-480.

Stanford,JA and Ward,JV 1988 The hyporheic habitat of river ecosystems. Nature, 335(6185), pp.64-66. PDF
     Abstract: "Contemporary river ecology is based primarily on biogeochemical studies of the river channel and interactions with shoreline vegetation, even though most rivers have extensive floodplain aquifers that are hydraulically connected to the channel. The hyporheic zone, the interstitial habitat penetrated by riverine animals, is characterized as being spatially limited to no more than a few metres, in most cases centimetres, away from the river channel1-9. However, riverine invertebrates were collected in hundreds per sample within a grid of shallow (10 m) wells located on the flood-plain up to 2 km from the channel of the Flathead River, Montana, USA. Preliminary mass transport calculations indicate that nutrients discharged from the hyporheic zone may be crucial to biotic productivity in the river channel. The strength and spatial magnitude of these interactions demonstrate an unexplored dimension in the ecology of gravel-bed rivers."

Stanford,JA and Ward,JV 1989 Serial discontinuities in a Rocky Mountain river. I. Distribution and abundance of Plecoptera. Regulated Rivers: Research and Management 3, 169-175.

Stanford,JA and Ward,JV 1993 An ecosystem perspective of alluvial rivers: connectivity and the hyporheic corridor. Journal of the North American Benthological Society, 48-60.
     Abstract: " Floodplains of large alluvial rivers are often expansive and characterized by high volume hyporheic flow through lattice-like substrata, probably formed by glacial outwash or lateral migration of the river channel over long time periods. River water downwells into the floodplain at the upstream end; and, depending on bedrock geomorphology and other factors, groundwater from the unconfined aquifer upwells directly into the channel or into floodplain springbrooks at rates determined by head pressure of the water mass moving through the floodplain hydrologic system. These large scale (km3) hyporheic zones contain speciose food webs, including specialized insects with hypogean and epigean life history stages (amphibionts) and obligate groundwater species (stygobionts). Biogeochemical processes in the hyporheic zone may naturally load groundwaters with bioavailable solutes that appear to exert proximal controls on production and biodiversity of surface benthos and riparian vegetation. The effect is especially evident in floodplain springbrooks. Dynamic convergence of aquifer-riverine components adds physical heterogeneity and functional complexity to floodplain landscapes. Because reaches of aggraded alluvium and attendant ecotonal processes occur serially, like beads on a string, along the river continuum, we propose the concept of a hyporheic corridor in alluvial rivers. We expect predictable zonation of groundwater communities and other aquifer-riverine convergence properties within the corridor from headwaters to river mouth. The landscape-level significance and connectivity of processes along the hyporheic corridor must be better understood if river ecosystems, especially those involving large floodplain components, are to be protected and/or rehabilitated. "

Stanger,JA and Baumann,RW 1993 A revision of the stonefly genus Taenionema (Plecoptera: Taeniopterygidae). Transactions of the American Entomological Society 119 3, 171-229.
     The key for Taenionema, has diagnoses, illustrations of pertinent taxonomic characters and a proposed phylogeny.

Stark,BP and Baumann,RW 2018 Two New Stonefly Species in the Sweltsa coloradensis (Banks) Complex (Plecoptera: Chloroperlidae). Illiesia, 14(02):30-43. https://doi.org/10.25031/2018/14.02 PDF

Stark,BP; Baumann,RW; Kondratieff,BC and Stewart,KW 2013 Larval and egg morphology of Paraperla frontalis (Banks 1902) and P. wilsoni Ricker 1965 (Plecoptera: Chloroperlidae). Illiesia 9(08):101-108. PDF

Stark,BP and Green,S 2011 Eggs of western Nearctic Acroneuriinae (Plecoptera: Perlidae). Illiesia 2011 7(17):157-166. PDF

Stark,BP and Gaufin,AR 1976a The nearctic genera of Perlidae (Plecoptera). Miscellaneous Publications of the Entomological Society of America 10, 1-80.
     Has keys, illustrations, range maps and discussions of biogeography.

Stark,BP and Gaufin,AR 1976b The nearctic species of Acroneuria (Plecoptera: Perlidae). Journal of the Kansas Entomological Society 49 2, 221-253.
     Keys, diagnoses, illustrations.

Stark,BP and Kyzar,JW 2001 Systematics of nearctic Paraleuctra with description of a new genus (Plecoptera: Leuctridae). Tijdschrift voor Entomologie 144 (1)119-135. PDF
     Has a key to nearctic adults of the stonefly family Leuctridae including their newly described genus Pomoleuctra.

Stark,BP; Oblad,BR and Gaufin,AR 1973 An annotated list of the Stoneflies (Plecoptera) of Colorado Part I. Entomological News 84 9, 269-277.
     Very useful historic distribution information, many nomenclature changes have occurred since this was published. Now you may want to review Kondratieff and Baumann, 2002 for additional information.

Stark,BP and Sivec,I 2010 Systematic notes on the genus Claassenia Wu (Plecoptera: Perlidae), with description of a new species. Illiesia 6(24):303-314. PDF

Stark,BP and Szczytko,SW 1982. Egg morphology and phylogeny in Pteronarcyidae (Plecoptera). Annals of the Entomological Society of America 75: 519-529.

Stark,BP and Szczytko,SW 1986 North American stoneflies (Plecsoptera) Systematics, distribution, and taxonomic references. Great Basin Naturalist 46: 383-397.

Stark,BP and Szczytko,SW 1988. Egg morphology and phylogeny in Arcynopterygini (Plecoptera: Perlodidae) Journal of the Kansas Entomological Society 61(2) 143-160.Abstract and First Page

Stark,BP; Stewart,KW 2005 Nymphs of four western nearctic Sweltsa species (Plecoptera: Chloroperlidae). Transactions of American Entomological Society 131 1+2, 189-200.

Stark,BP; Stewart,KW; Szczytko,SW and Baumann,RW 1998. Common Names of Stoneflies (Plecoptera) from the United States and Canada. Ohio Biological Survey Notes 1:1-18. PDF

Stark,BP and Szczytko,SW 1988 Egg morphology and phylogeny in Arcynopterygini (Plecoptera: Perlodidae) Journal of the Kansas Entomological Society 61(2) 143-160.First Page

Stark,BP; Szczytko,SW and Baumann,RW 1986 North American stoneflies (Plecoptera): systematics, distribution, and taxonomic references. Great Basin Naturalist 46, 383-397.
     Another useful general reference.

Stark,BP; Szczytko,SW and Baumann,RW 2010 North American stoneflies (Plecoptera): systematics, distribution, and taxonomic referen ces. Western North American Naturalist, 46(3), 383-397. PDF

Stark,BP; Szczytko,SW and Nelson,CR 1998. American stoneflies: a photographic guide to the Plecoptera. The Caddis Press. Columbus Ohio. 126 pages.

Stark,JD and Banks,JE 2003 Population-level effects of pesticides and other toxicants on arthropods. Annual Review of Entomology 48:505-19.

Stewart,KW 2009 New descriptions of North American Taenionema larvae (Plecoptera: Taeniopterygidae). Illiesia 2009 5(12):128-145. PDF

Stewart,KW 2001 Vibrational communication (drumming) and mate-searching behavior of stoneflies (Plecoptera); evolutionary considerations. In Trends in Research in Ephemeroptera and Plecoptera (pp. 217-225). Springer US.
     Abstract: " A long recognized but little explored mode of intersexual communication in insects is use of low-frequency substrate-borne vibrational signals. Representatives of 10 insect orders are known to have adopted this mode; range of communication, informational content and receiver integration of signals and energy costs are discussed. Arctoperlarian stoneflies represent the epitome of evolution of vibrational communication. Their ancestral signals were monophasic volleys of evenly spaced drumbeats. Derived signals to achieve species-specificity and possibly to enable sexual.selection or some measure of reproductive fitness has involved modification of the ancestral form toward complex signals through: (1) changes in the rhythmic patterning of calls, (2) patterns of ?-? duetting, and/or changes in the method of signal production such as rubbing or tremulation. Proposed paradigms for the evolution of vibrational communication and evolution of signal patterns are presented, with examples of the signals of several arctoperlarian species. The entire mating system of Arctoperlaria is discussed, and searching behavior in relation to vibrational communication is presented for Pteronarcella badia, Claassenia sabulosa, Perlinella drymo and Suwallia sp. "

Stewart,KW; Abbott,JC; Bottorff,RL 1995 Drumming signals of two stonefly species: a newly discovered duet pattern in Plecoptera. Entomological News 106 1, 13-18.

Stewart,KW and Alexander,NH 2008 The nymphs of three Nemouridae species (Plecoptera) from Oregon temporary headwaters streams. Transactions of the North AMerican Entomological Society 134: 173-183.

Stewart,KW; Baumann,RW and Stark,BP 1973 The distribution and past dispersal of southwestern United States Plecoptera. Transactions of the American Entomological Society (1890-), 99(4), 507-546.

Stewart,KW; Bottorff,RL; Knight,AW; Moring,JB 1991 Drumming of four North American euholognathan stonefly species, and a new complex signal pattern in Nemoura spiniloba Jewett (Plecoptera:Nemouridae). Annals of the Entomological Society of America 84 2, 201-206.

Stewart,KW; Harper,PP 1996 Plecoptera. In: An Introduction to the Aquatic Insects of North America. 3rd ed. Eds: Merritt,RW; Cummins,KW Kendall/Hunt Publishing Company, Dubuque, Iowa, 217-266.
     The classic North American aquatic insect (including Stoneflies) key and reference.

Stewart,KW; Hassage,RL; Holder,SJ and Oswood,MW 1990 Life cycles of six stonefly species (Plecoptera) in subarctic and arctic Alaska streams. Annals of the Entomological Society of America 83(2)207-214.
     Abstract: Nymphal growth and emergence of adults are described for six species of stoneflies (Plecoptera) found in subarctic and arctic Alaska. The two Nemouridae studied are semivoltine; adults of Zapada haysi (Ricker) are present from May to July and adults of Nemoura arctica Esben-Petersen occur from June to July. The remaining four species are univoltine. Plumiperla diversa (Frison) (Chloroperlidae) has most of its growth occurring during the summer with emergence the following May-September. Taenionema pacificum (Banks) (Taeniopterygidae) completes nymphal growth by the end of January and has an early emergence (April-June). Adults of Arcynopteryx compacta (McLachlan) (Perlodidae) are present from May to August, and growth of nymphs is rapid during summer and fall. Isoperla petersoni Needham ∓ Christenson (Perlodidae) adults are present from June to mid-August, and nymphal growth is interrupted by winter and resumes in the spring; the three other univoltine species studied tend to complete growth before the onset of winter. Growth of these six species is tied to seasonal temperature variation.

Stewart,KW and Maketon,M 1991 Structures used by Nearctic stoneflies (Plecoptera) for drumming, and their relationship to behavioral pattern diversity. Aquatic insects, 13(1), 33-53.
     Abstract: " There has been some co-evolution of male ventral abdominal structure with method of producing vibrational call signals and call pattern diversity in Plecoptera. Male Antarctoperlaria have no specialized ventral abdominal structures and are not known to drum. Male Arctoperlaria without vesicles, lobes, knobs or hammers all produce relatively simple calls by tapping on substrate, or have derived a tremulation method of producing calls. Elongate, moveable vesicles on abdominal segment 9 are homologous structures in ancestral and some extant Euholognatha families, and are primarily associated with ancestral call patterns. Allocapnia and Taeniopteryx, that have secondarily lost vesicles, have retained the ancestral drumming pattern. The lobes, knobs and hammers of Systellognatha, that are characterized with SEM in this paper, are probably non-homologous with vesicles of Euholognatha, and presumably have arisen independently in various families and genera. Systellognathan species with these structures have either retained ancestral, tapping call patterns, or have derived highly specialized grouped, or phased patterns or a rubbing method of signal production. Rubbing has been derived only in association with the knobs of Peltoperlidae or hammers of Perlidae. Complex drumming pattern diversity is unnecessary in females, and as expected, we have found no complex answer patterns, rubbing method of answering or specialized drumming structures in them. "

Stewart,KW and Oswood,MW 2006 The Stoneflies (Plecoptera) of Alaska and Western Canada. In The Caddis Press, Columbus, Ohio. vi. + 325.

Stewart,KW and Ricker,WE 1997 The stoneflies of the Yukon. pgs 201-222 in Danks,HV and Downes,JA (Eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1034 pp.

Stewart,KW and Stark,BP 2002 Nymphs of North American Stonefly Genera. 2nd edition The Caddis Press, Columbus, Ohio. 510 pages.
     The best key for Stonefly nymphs to genus in North America. Excellent illustrations, straightforward identification keys and a broad literature review make this book a neccessity.

Stewart,KW and Stark,BP 2008 Chapter 14: Plecoptera. In: An Introduction to the Aquatic Insects of North America. 4th ed. Eds: Merritt,RW; Cummins,KW; Berg,MB Kendall/Hunt Publishing Company, Dubuque, Iowa, 311-384.

Stewart,KW and Stark,BP 2011 Further descriptions of western North American Podmosta larvae and their separation from Ostrocerca larvae (Plecoptera: Nemouridae). Illiesia 7(10):104-117. PDF

Stewart,KW and Szczytko,SW 1983 Drift of Ephemeroptera and Plecoptera in two Colorado rivers. Freshwater Invertebrate Biology. 2(3)117-131. PDF

Stewart,KW; Szczytko,SW and Stark,BP 1982 Drumming behavior of four species of North American Pteronarcyidae (Plecoptera): dialects in Colorado and Alaska Pteronarcella badia. Annals Entomological Society of America 75:530-533.

Stewart,KW and Zeigler,DD 1984 Drumming behavior of twelve North American stonefly (Plecoptera) species: First descriptions in Peltoperlidae, Taeniopterygidae and Chloroperlidae. Aquatic Insects. 6(1) 49 - 61. Abstract

Stewart,KW and Zeigler,DD 1984 The use of larval morphology and drumming in Plecoptera systematics, and further studies of drumming behavior. Annals of Limnology, 20 (1-2) 105-114.

Steyskal, GC 1976 Notes on the nomenclature and taxonomic growth of the Plecoptera. pp 408-410. In: A report on the fifth international symposium on Plecoptera. RW Baumann, ed. Proc. Biol. Soc. Washington. 88:399-428.

Stoaks,RD and Kondratieff,BC 2014 The aquatic macroinvertebrates of a first order Colorado, USA Front Range stream: what could the biodiversity have been before irrigated agriculture?. Journal of the Kansas Entomological Society, 87(1), pp.47-65. PDF

Surdick,RF 1985 Nearctic Genera of Chloroperlinae (Plecoptera: Chloroperlidae). University of Illinois Press, Urbana, IL. 146 pages.
An early key for Chloroperlids, now use Stewart and Stark, 2002.

Surdick,RF 1995a New western nearctic Sweltsa (Plecoptera: Chloroperlidae). Proceedings of the Entomological Society of Washington 97 1, 161-177.

Szczytko,SW and Kondratieff,BC 2015 A review of the Eastern Nearctic Isoperlinae (Plecoptera: Perlodidae) with the description of twenty-two new species. 1, 1-289. PDF

Szczytko,SW and Stewart,KW 1979a The genus Isoperla (Plecoptera) of western North America; holomorphology and systematics, and a new stonefly genus Cascadoperla. Memoirs of the American Entomological Society 32, 1-120.
     The key for Isoperla. Also has diagnoses and many illustrations

Szczytko,SW and Stewart,KW 1979b Three new species of nearctic Isoperla (Plecoptera). Great Basin Naturalist 36, 211-220.

Szczytko,SW and Stewart,KW 1979c Drumming behavior of four Western Nearctic Isoperla (Plecoptera) species. Annals of the Entomological Society of America 72(6)781-786.


Taylor,BW; Anderson,CR and Peckarsky,BL 1998 Effects of size at metamorphosis on stonefly fecundity, longevity, and reproductive success. Oecologia 114, 494-502. Abstract
     Studying Megarcys in streams around the Rocky Mountain Biological Lab in Gothic, Colorado, they found that M. signata was protandrous (males emerge first) and females were about twice the body mass of males in both streams studied. M. signata emerged earlier and larger from a trout stream (East River) than the stoneflies from a smaller fishless tributary (Benthette Brook). Temperature did not affect size at metamorphosis. Many females from Benthette Brook were brachypterous (short-winged), while East River Megarcys females has longer wings. In 1992 East River stoneflies emerged from mid June to mid July with the peak emergence in late June. Benthette brook Megarcys emerged from early July to late August, peaking in Late July. Adults emerged at dawn between 0600 and 0800 hours until later in the season when they emerged at night (after 2200 hours). Adults became more active after dark. Manipulative experiments determined that male body mass did not affect reproductive success. Sometimes male drumming attracted females who mated immediately upon finding the male. Other times the first male to stumble on the female mated with her. Mating usually occurred within the first three days after emergence. Copulation generally lasted all night. Females who mated multiple times had lower total lifetime fecundity than females who mated once. Multiple matings reduced the lifespan of males and females. There was no evidence of parthenogenesis in unmated females. Neither sex fed on sugar water except rarely and dissections of field collected adults showed atrophied digestive tracts.

Taylor,BW; Anderson,CR and Peckarsky,BL 1999 Egg diapause and semivoltinism in the Nearctic stonefly Megarcys signata (Plecoptera: Perlodidae). Aquatic Insects 21, 179-185.

Taylor,BW; Anderson,CR and Peckarsky,BL 1999 Delayed egg hatching and semivoltinism in the Nearctic stonefly Megarcys signata (Plecoptera:Perlodidae). Aquatic Insects 21, 179-185. Abstract

Theissinger,K; Feldheim,KA; Seitz,A and Pauls,SU 2009 Isolation and characterization of 11 polymorphic trinucleotide microsatellite markers in the stonefly Arcynopteryx compacta (Plecoptera: Perlodidae) Molecular Ecology Resources 9(1)357-359.

Tikkanen,P; Muotka,T; Huhta,A and Juntunen,A 1997 The roles of active predator choice and prey vulnerability in determining the diet of predatory stonefly (Plecoptera) nymphs. Journal of Animal Ecology 66: 36-48.



VanWieren,BJ; Kondratieff,BC and Stark,BP 2001 A review of the North American species of Megarcys Klapálek (Plecoptera:Perlodidae). Proceedings of the Entomological Society of Washington 103 2, 409-427.
     Reviews all five North American species of Megarcys (M.irregularis, M. signata, M, subtruncata, M. watertoni and M. yosemite). Keys to males, females and eggs, diagnoses of each species, illustrations of male and female genitalia as well as photomicrographs of eggs are provided.

Vieira,NK; Barnes,TR and Mitchell,KA 2011 Effects of wildfire and postfire floods on stonefly detritivores of the Pajarito Plateau, New Mexico. Western North American Naturalist, 71(2) 257-270. PDF

Vieira, Nicole K.M., Poff, N. LeRoy, Carlisle, Daren M., Moulton, Stephen R., II, Koski, Marci L., and Kondratieff, Boris C., 2006, A database of lotic invertebrate traits for North America: U.S. Geological Survey Data Series 187, http://pubs.water.usgs.gov/ds187



Ward,JV, Kondratieff,BC and Zuellig,RE 2002 An Illustrated Guide to the Mountain Stream Insects of Colorado. 2nd ed. University Press of Colorado, Boulder, Colorado. 219 pages.
     General reference for aquatic insects in the mountain running waters of Colorado. Used by entomology classes everywhere in Colorado.

Warnick,SL and Bell,HL 1969 The acute toxicity of some heavy metals to different insects. Journal WPCF 41 2, 280-284.

Webb,DW (Ed.) 1996 Current and Selected Bibliographies on Benthic Biology. North American Benthological Society, Windsor, Ontario, Canada. 96 pages.
     This series of bibliographies helps everyone keep up on the current literature. There are more of these for many other years. They contain mostly all new citations every year with a few missed from previous years included.

Wellnitz,T 2014 Can current velocity mediate trophic cascades in a mountain stream?. Freshwater Biology, 59(11) 2245-2255. PDF

Williams,MC; Lichtwardt,RW 1987 Two new Trichomycete species from Zapada spp. (Stonefly) nymphs with an unusual distribution. Mycologia 79, 473-478.
     Studies a common fungus on Plecoptera nymphs.

Winterbourn,MJ; Crowe,ALM 2001 Flight activity of insects along a mountain stream: is directional flight adaptive? Freshwater Biology 46, 1479-1489.




Zanetell,BA and Peckarsky,B 1996 Stoneflies as ecological engineers-hungry predators reduce fine sediments in stream beds. Freshwater biology, 36(3), pp.569-577.

Ziegler,DD and Stewart,KW 1977 Drumming behavior of eleven Nearctic stonefly (Plecoptera) species Annals of the Entomological Society of America. 70(4)495-505.

Zeigler,DD and Stewart,KW 1985 Age effects of drumming behavior of Pteronarcella badia (Plecoptera) males. Entomological News 96(4) 157-160

Zenger,JT and Baumann,RW 2004 The holarctic winter stonefly genus Isocapnia, with an emphasis on the North American fauna (Plecoptera: Capniidae) Monographs of the Western North American Naturalist 2(1):65-95.Abstract

Zuellig,RE; Heinold,BD; Kondratieff,BC and Ruiter,DE 2012 Diversity and Distribution of Mayflies (Ephemeroptera), Stoneflies (Plecoptera), and Caddisflies (Trichoptera) of the South Platte River Basin, Colorado, Nebraska, and Wyoming, 1873-2010.U.S. Geological Survey Data Series 606, 257 p. PDF - caution 46MB

Zuellig,RE; Kondratieff,BC; Rhodes,HA 2002 Benthos recovery after an episodic sediment release into a Colorado Rocky Mountain river. Western North American Naturalist 62 1, 59-72.

Zwick,P 1973 Insecta: Plecoptera Phylogenetisches System und Katalog. Das Tierreich - Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen., 94, 465 pp.

Zwick,P 1982 The stonefly collection of F. Klapálek in Prague, with notes on the Nemouridae (Plecoptera). Aquatic Insects 4(1)39-48.

Zwick,P 1989 Notes on Plecoptera (18) Skwala americana (Klapálek, 1912), comb. n., the valid name for Skwala parallela (Frison, 1936). Aquatic Insects 11 3, 181-182.

Zwick,P 2000 Phylogenetic system and zoogeography of the Plecoptera. Annual review of entomology, 45(1), pp.709-746.
     Abstract: " Information about the phylogenetic relationships of Plecoptera is summarized. The few characters supporting monophyly of the order are outlined. Several characters of possible significance for the search for the closest relatives of the stoneflies are discussed, but the sister-group of the order remains unknown. Numerous characters supporting the presently recognized phylogenetic system of Plecoptera are presented, alternative classifications are discussed, and suggestions for future studies are made. Notes on zoogeography are appended. The order as such is old (Permian fossils), but phylogenetic relationships and global distribution patterns suggest that evolution of the extant suborders started with the breakup of Pangaea. There is evidence of extensive recent speciation in all parts of the world."

Zwick,P 2006 New family characters of larval Plecoptera, with an analysis of the Chloroperlidae: Paraperlinae. Aquatic Insects, 28:13-22.

Brown,WS 2004 Stoneflies or Plecoptera of Gunnison County, Colorado, USA

"One final paragraph of advice: Do not burn yourself out. Be as I am - a reluctant enthusiast... a part-time crusader, a half-hearted fanatic. Save the other half of yourselves and your lives for pleasure and adventure. It is not enough to fight for the land; it is even more important to enjoy it. While you can. While it is still there. So get out there and mess around with your friends, ramble out yonder and explore the forests, encounter the griz, climb the mountains. Run the rivers, breathe deep of that yet sweet and lucid air, sit quietly for a while and contemplate the precious stillness, that lovely, mysterious and awesome space. Enjoy yourselves, keep your brain in your head and your head firmly attached to your body, the body active and alive, and I promise you this much: I promise you this one sweet victory over our enemies, over those deskbound people with their hearts in a safe deposit box and their eyes hypnotized by desk calculators. I promise you this: you will outlive the bastards." -- Edward Abbey