Plecoptera of Gunnison County, Colorado Annotated Publication List
Updated 7 March 2026
Arranged by the first author's last name. Species names are linked to pages on this website. Scientific words and phrases are usually linked to wikipedia.
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Many older publications are available from the Biodiversity Library. The International Journal of Stonefly Research Illiesia has stopped publishing new papers.
Alexander,KD and 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. PDF
>They studied Claassenia in the Gunnison river near Almont.
Alexander,KD and 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 and 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 and 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 and 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,JD 1975a The distributional ecology and diversity of benthic insects in Cement Creek, Colorado. Ecology 56(5) 1040-1053. PDF
A fascinating aspect of this study is that it was done in the early 1970's when beaver were locally extinct, or at a landscape level, functionally extinct. Some of the sites on Cement Creek used in this study have changed and been colonized by beavers and their ponds since roughly the late 1990s.
Abstract: "Distributional patterns and species diversity of benthic insects in an alpine stream in Gunnison County, Colorado, USA were investigated on several levels of spatial scale, from faunal replacement over 1,000 vertical m to microdistribution within the stony substratum. Ecotones including zonation in terrestrial vegetation and in trout distribution did not appear to affect faunal replacement. Competition among congeners accounted for 7—30% of the cases, while in the majority of species, faunal replacement appeared to be associated with gradual changes in the physical gradient. Trout zonation may affect total numbers of insects, however, as the trout—free headwaters had two to six times higher insect densities. Microdistribution was investigated by measures of species and substratum patterning in a series of microhabitats (0.093 m2) at a series of sites (separated by 75—150 vertical m), and by field colonization experiments with various substratum choices. I hypothesize that increased substratum complexity leads to greater species richness based on several lines of evidence: (1) different species showed different substratum preferences, (2) colonization of mixed substrata generally resulted in greater mean species richness than did colonization of a single substratum type, and (3) both species diversity and substratum complexity were greatest at the within—microhabitat level. However, substratum composition showed little variation along the elevational gradient and did not appear to be a cause of faunal replacement. The several scales of investigation were complementary, as congeners exhibiting sharp mutual exclusion in vertical distribution had similar microhabitat preferences, while other congeners showed less exclusion and differed in microhabitat preferences. Most of species diversity as measured by H′ was found within habitats rather than between habitats while species richness depended equally upon within—habitat variation (owing to rare species) and between—habitat variation (owing to faunal replacement)."
Allan,JD 1975b Faunal replacement: altitudinal zonation in an alpine stream. Verhandlungen der Internationale Vereinigung für Theoretische und Angewandte Limnologie 19:1015-1022. PDF
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: "The feeding ecology of predaceous stoneflies was investigated over 18 mo in Cement Creek, Colorado, USA. Three species were studied at each of three sites over their entire life cycles to elucidate any differences due to species and site. Diet typically changed, over the course of development, from diatoms and chrionomids in the earliest instars, to primarily chironomids, to a broader diet in which mayflies increased in importance relative to chironomids. Stonefly diet was similar in individuals of a given size at a particular site, regardless of species, but differed among size categories and sites within a species. Two factors appeared to determine what prey were eaten. The maximum size of prey ingested increased with size of stonefly, at approximately 10% of predator body mass. No evidence of preference for larger prey was observed within a predator size category, however. Rather, the availability of prey depends upon the abundance of the vulnerable size class, which in turn is affected by the juxtaposition of predator and prey life cycles. The mass of a full foregut was related to mass of stonefly by the power law for each species. By combining these estimates of gut clearance rate, total prey consumption by stoneflies at each site was estimated to range from 2.7 to 3.3 g°m—2°yr—1 (dry mass). While prey availability is notoriously difficult to measure, this appears to be substantial predation pressure."
Allan,JD 1982 The effects of reduction in trout density on the invertebrate community of a mountain stream. Ecology 63(5) 1444-1455. PDF
Abstract: "An experimental reduction in trout density was carried out for 4 yr to determine whether the numbers or species composition of aquatic invertebrates would be affected. In a Colorado stream, the standing crop of trout (mainly Salvelinus fontinalis) prior to this study was 4.86 g/m2, typical of infertile trout streams. Repeated electroshocking of a 1220 m long experimental section kept trout stocks at 10—25% of this initial value during the summers of 1975—1978. Density of invertebrates in the benthos (number of invertebrates per square metre) showed no consistent differences between the trout removal section and upstream and downstream control sections for the majority to taxa examined. Drift density (number of invertebrates per 1000 cubic metres) also failed to show an effect of the trout removal. Some taxa were significantly more abundant at one site than another, but this was attributable to changes along the stream gradient or differences in phenology among the sites rather than experimental reduction of trout. As trout stomach content analysis revealed very intensive grazing of a few taxa aquatic insects, their failure to increase in the trout removal section was unexpected. Statistical analysis showed that because of the variability of the system, a doubling or halving of numbers must occur to be detectable. Within these limits, it is concluded that removal of trout had no significant effect on the prey community, and two explanations are offered. Trout may actually consume only a small fraction of total prey, although this conclusion is limited by inability of current techniques to sample the benthos fully. It is also likely that because streams do not provide areas where fish are predictably absent, the invertebrate community is highly adapted to fish predation and so is not sensitive to manipulations in fish density."
Allan,JD 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 and McClintock,NL 1987a Prey preference of stoneflies: sedentary vs mobile prey. Oikos 49 (3) 323-331.
Allan,JD; Flecker,AS and McClintock,NL 1987b Prey size selection by carnivorous stoneflies. Limnology and Oceanography 32 (4) 864-872. PDF
Abstract: "To investigate prey size preference by carnivorous stoneflies, we conducted choice experiments and directly observed predatory behavior in laboratory streams. Small, medium, and large size classes of Hesperoperla pacifica (Perlidae), small Kogotus modestus, and medium and large Megarcys signata (both of the Perlodidae) were offered an array of sizes of the mayfly Baetis as prey. Small predators displayed strong positive selection for the smallest size class of prey, and preference declined steadily with increasing prey size. Large predators displayed hump-shaped preference curves, exhibiting strongest preference for intermediate size classes of prey. Thus size selectivity was demonstrated and varied with the size class of predator.
Direct observations of encounters between predators and their prey revealed that percent attacks/ encounters by small predators was strongly biased toward small prey and by large predators was weakly biased toward large prey. Capture success always was greater and handling times shorter with small prey compared to large prey."
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. PDF
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. PDF
Banks,N 1908 Neuropteroid insects - notes and descriptions. Transactions of the American Entomological Society 34:255-267. PDF
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 and Gaufin,AR 1972 The Amphinemura venusta complex of western North America. Contributions of the Science and Natural History Museum 226, 1-16. 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
Bottorff,RL and 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 and Dimick,JJ 1990 Drumming behavior of four western nearctic Isoperla species (Plecoptera:Perlodidae). Annals of the Entomological Society of America 83 (5) 991-997.
Boumans,L and Baumann,RW 2012 Amphinemura palmeni is a valid Holarctic stonefly species (Plecoptera: Nemouridae). Zootaxa, 3537(1), pp.59-75. PDF
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 and 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. PDF
Abstract: "We used a phylogenetically based comparative approach to evaluate the potential for physiological studies to reveal patterns of diversity in traits related to susceptibility to an environmental stressor, the trace metal cadmium (Cd). Physiological traits related to Cd bioaccumulation, compartmentalization, and ultimately susceptibility were measured in 21 aquatic insect species representing the orders Ephemeroptera, Plecoptera, and Trichoptera. We mapped these experimentally derived physiological traits onto a phylogeny and quantified the tendency for related species to be similar (phylogenetic signal). All traits related to Cd bioaccumulation and susceptibility exhibited statistically significant phylogenetic signal, although the signal strength varied among traits. Conventional and phylogenetically based regression models were compared, revealing great variability within orders but consistent, strong differences among insect families. Uptake and elimination rate constants were positively correlated among species, but only when effects of body size and phylogeny were incorporated in the analysis. Together, uptake and elimination rates predicted dramatic Cd bioaccumulation differences among species that agreed with field-based measurements. We discovered a potential tradeoff between the ability to eliminate Cd and the ability to detoxify it across species, particularly mayflies. The best-fit regression models were driven by phylogenetic parameters (especially differences among families) rather than functional traits, suggesting that it may eventually be possible to predict a taxon's physiological performance based on its phylogenetic position, provided adequate physiological information is available for close relatives. There appears to be great potential for evolutionary physiological approaches to augment our understanding of insect responses to environmental stressors in nature."
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
Abstract: "A 46km section of the Dolores River, in southwest Colorado, was studied to determine the relative abundance of invertebrate functional groups over an altitudinal gradient of 500m during three seasons. The Dolores River is a third order steam with an average width of 11m in upper 8 km of the study area. In the lower 38 km of the study area, it is a fourth order stream with an average width of 15 m. Benthic invertebrates were collected with a modified Hess sampler in October 1980, and March and August 1981, from 11 stations on the Dolores River. Despite little change in either stream order, width or apparent food resources in the study area, there were noticeable differences in the relative abundance of functional groups, with shredders most abundant upstream and collectors most abundant in the mid-reaches. The observed trends were highly dependent upon season with shredders abundant at most stations only in spring. This was a result of life history patterns of winter stoneflies, the primary shredders. Collector-gatherers were most abundant at the upper-middle stations in summer, but were less abundant in the other two seasons. In general, the pattern appeared to conform more to the altitudinal shifts in benthic species composition than to stream order or width. This led to shifts in the assigned functional groups without noticeable changes in food resources."
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 and 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 and 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
Abstract: "Three new fossil species of stoneflies (Plecoptera: Nemouridae and Leuctridae) from
Eocene Baltic amber are being described: Zealeuctra cornuta n. sp., Lednia zilli n. sp., and Podmosta attenuata n. sp.. Extant species of these three genera are found in Eastern Asia and in the Nearctic region. It is very probably that the genera must have been widely spread across the northern hemisphere in the Cretaceous period, before Europe was an archipelago in Eocene. The current state of knowledge about the seventeen Plecoptera species of Baltic amber is shortly presented. Due to discovered homonymies, the following nomenclatural corrections are proposed: Leuctra fusca Pictet, 1856 in Leuctra electrofusca Caruso & Wichard, 2010 and Nemoura affinis Berendt, 1856 in Nemoura electroaffinis Caruso & Wichard, 2010."
Caruso,C and Wichard,W 2011 Paleogeographic Distribution of Leuctridae and Nemouridae genera preserved in Baltic amber, with the description of Palaeopsole weiterschani n. gen., n. sp.(Plecoptera). Entomologie heute, 23, pp.69-77.
Abstract: " The description of the new fossil species Palaeopsole weiterschani n. gen., n. sp. adds the genus Palaeopsole to the extant six genera of the families Leuctridae and Nemouridae, which have been found in Eocene Baltic amber. These Eocene fossils are the oldest representatives of their extant genera, so far. But the Eocene European archipelago is neither place nor time in which these genera originated. The origin of these seven genera is unknown, but took very probably place in the northern hemisphere before the Eocene. After Laurasia had finally broken apart at early Jurassic, probably not till then the genera developed and spread out across the warm-temperate and subtropical northern hemisphere. The extant Nearctic genera - Lednia, Megaleuctra, and Zealeuctra, including the transberingian Podmosta, and including Leuctra and Nemoura which are nearly distributed all over the Holarctic region - spread probably across the northern hemisphere via the temporary Beringian and Turgai Strait land bridges at Cretaceous. Towards the end of Eocene the temperature decreased worldwide and forced the Nearctic and Holarctic genera to adapt to temperate and boreal climates, whereas the recent relatives of genus Palaeopsole, probably genus Rhopalopsole, continued and live in the subtropical area of the Oriental region and Southeast Asia."
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 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
Chen,ZT 2025 Three new stoneflies (Insecta: Plecoptera) from Eocene Baltic amber. European journal of taxonomy, 1026, pp.107-122. PDF
Abstract: "Three new stonefly species are described and illustrated from Eocene Baltic amber found in Lithuania: †Palaeopsole spinosa sp. nov. (family Leuctridae Klapálek, 1905), †Podmosta biloba sp. nov. (family Nemouridae Newman, 1853), and †Isoperla lituana sp. nov. (family Perlodidae Klapálek, 1909). The generic diagnosis of †Palaeopsole Caruso & Wichard, 2011 is also emended. These new fossils enhance the understanding of the extinct Plecoptera fauna in Baltic amber."
Claassen,PW 1923 New species of North American Plecoptera. Canadian Entomologist 55, 257-263,281-292.
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.
Clayton,JA and Westbrook,CJ 2008 The effect of the Grand Ditch on the abundance of benthic invertebrates in the Colorado River, Rocky Mountain National Park. River Research and Applications, 24(7), pp.975-987. PDF
Abstract: "We investigate herein the hypothesis that there is a significant relationship between bed particle mobility and benthic invertebrate abundance in the gravel-bed channel of the upper Colorado River in Rocky Mountain National Park. A large diversion channel called the Grand Ditch normally diverts a significant portion (~50%) of the annual snowmelt runoff from the watershed northward out of the basin. In May 2003, a ~30-m section of the ditch was breached, contributing substantially to the magnitude and duration of discharge in the Colorado River until the ditch breach was repaired in July of that year. As a result, all grain sizes in the river channel were mobilized, which contrasted sharply with the minimal gravel transport experienced during the exceptional drought of the previous year. Benthic macroinvertebrates were collected in the field using a Surber sampler at the same six locations for both years, and the number of individuals of the orders ephemeroptera (mayflies), plecoptera (stoneflies), trichoptera (caddisflies) and diptera (e.g. chironomids) was counted in the laboratory. The total number of individuals was 240% higher in 2003, and the proportion of mayflies in the samples increased from 25% in 2002 to 40% in 2003. In 2003, samples were also taken immediately upstream and downstream of a large flow obstruction in the channel in order to further isolate the relative importance of sediment transport against other variables affecting the stream habitat. Numbers of individuals for all taxa collected (particularly ephemeroptera and plecoptera) were nearly an order of magnitude higher at the upstream site than at the downstream, protected location. These results have important implications for the ecosystem management of streams within Rocky Mountain National Park and elsewhere."
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 and 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.
Clements,WH; Herbst,DB; Hornberger,MI; Mebane,CA and Short,TM 2021 Long-term monitoring reveals convergent patterns of recovery from mining contamination across 4 western US watersheds. Freshwater Science, 40(2), pp.407-426. PDF
Abstract: "Long-term studies of stream ecosystems are essential for assessing restoration success because they allow researchers to quantify recovery trajectories, gauge the relative influence of episodic events, and determine the time required to achieve clean-up objectives. To quantify responses of benthic macroinvertebrate assemblages to stream remediation, we integrated results of 4 long-term (20-29 y) assessments of mining-impacted watersheds that were broadly distributed across the western US (California, Colorado, Idaho, Montana). Using a before-after control-impact (BACI) study design, we observed substantial reductions in metal concentrations and corresponding improvements of benthic assemblages following remediation. Recovery rates were relatively consistent, and streams typically recovered within 10 to 15 y after remediation was initiated (mean = 10.25 y), although episodic events changed trajectories at some sites. Differences in recovery among watersheds were likely determined by a number of factors, including the severity of contamination, effectiveness of remediation, proximity to upstream sources of colonization, and hydrologic variation. We also observed considerable variation in the rate and extent of recovery among assemblage metrics. For example, total abundance and richness recovered rapidly at most sites, but the composition of benthic macroinvertebrate assemblages remained substantially altered compared with reference sites. Using piecewise linear regression, we estimated a threshold response of Ephemeroptera, Plecoptera, and Trichoptera (EPT) species richness at ~1 cumulative criteria unit (CCU), which is the sum of the fractions of chronic water-quality criteria for metals measured, suggesting this value was protective of benthic assemblages. However, EPT richness was reduced by ~20% at 2× this CCU value, indicating that moderate exceedances of water-quality criteria could substantially affect stream biodiversity. Non-metric multidimensional scaling analyses identified common sets of species trait states across the 4 watersheds that were associated with either metal contamination or with recovering and intact reference stream assemblages. Our study illustrates the importance of long-term studies for quantifying responses to stream restoration and the usefulness of BACI designs for demonstrating cause-and-effect relationships between restoration treatments and community recovery. Because these 4 watersheds were among the most severely polluted sites in the western US, our study demonstrates the value of these investments in watershed restoration and the potential for success under the most extreme conditions."
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
Crossman,J; Bradley,C; Windsor,FM and Milner,AM 2023 Water source dynamics influence macroinvertebrate communities across groundwater-fed streams in a glacierized catchment. Hydrobiologia, 850(8), pp.1801-1816. PDF
Abstract: "Groundwater contributions to streamflow significantly influence the structure and function of riverine ecosystems, particularly in glacierized catchments where there are marked differences in water sources and subsurface flow paths. Here, we investigated spatial and temporal variation in relationships between water sources, flow paths, physical and chemical processes, organic matter, microbial biofilms, and macroinvertebrates across groundwater-fed streams in the glacierized Toklat River catchment of Denali National Park, Alaska. Streams fed predominantly by seepage from the valley sides were perennial, whereas streams sustained by glacial meltwater seepage were ephemeral. Differences in environmental conditions between flow regimes appeared to influence spatial and temporal patterns of organic matter, linking to macroinvertebrate community dynamics. Macroinvertebrates in perennial streams were supported by fine particulate organic matter from subsurface flow paths during summer, transitioning to a combination of fine particulate matter and leaf litter in autumn. In comparison, macroinvertebrates inhabiting ephemeral streams, which only flowed during autumn, were supported by leaf litter. Some macroinvertebrate taxa were unaffected by turnover in organic matter, indicating potential plasticity in organic matter resource use. Findings highlight the importance of considering spatial and temporal variation in groundwater-fed streams, considering that projected hydrological changes under a changing climate may have significant implications for these systems."
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 and 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
Cutler,DR; Edwards Jr,TC; Beard,KH; Cutler,A; Hess,KT; Gibson,J and Lawler,JJ 2007 Random forests for classification in ecology. Ecology, 88(11), pp.2783-2792. PDF
Abstract: "Classification procedures are some of the most widely used statistical methods in ecology. Random forests (RF) is a new and powerful statistical classifier that is well established in other disciplines but is relatively unknown in ecology. Advantages of RF compared to other statistical classifiers include (1) very high classification accuracy; (2) a novel method of determining variable importance; (3) ability to model complex interactions among predictor variables; (4) flexibility to perform several types of statistical data analysis, including regression, classification, survival analysis, and unsupervised learning; and (5) an algorithm for imputing missing values. We compared the accuracies of RF and four other commonly used statistical classifiers using data on invasive plant species presence in Lava Beds National Monument, California, USA, rare lichen species presence in the Pacific Northwest, USA, and nest sites for cavity nesting birds in the Uinta Mountains, Utah, USA. We observed high classification accuracy in all applications as measured by cross-validation and, in the case of the lichen data, by independent test data, when comparing RF to other common classification methods. We also observed that the variables that RF identified as most important for classifying invasive plant species coincided with expectations based on the literature."
Delk,JK; Kilgore,MJ and 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(1), p.13163 PDF
Abstract: "While most global productivity is driven by modern photosynthesis, river ecosystems are supplied by locally fixed and imported carbon that spans a range of ages. Alluvial aquifers of gravel-bedded river floodplains present a conundrum: despite no possibility for photosynthesis in groundwater and extreme paucity of labile organic carbon, they support diverse and abundant large-bodied consumers (stoneflies, Insecta: Plecoptera). Here we show that up to a majority of the biomass carbon composition of these top consumers in four floodplain aquifers of Montana and Washington is methane-derived. The methane carbon ranges in age from modern to up to >50,000 years old and is mostly derived from biogenic sources, although a thermogenic contribution could not be excluded. We document one of the most expansive ecosystems to contain site-wide macroinvertebrate biomass comprised of methane-derived carbon and thereby advance contemporary understanding of basal resources supporting riverine productivity."
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
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
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 and 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 and 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.
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 and 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."
Eichert,A; Sproul,J; Tolman,ER; Birrell,J; Meek,J; Heckenhauer,J; Nelson,CR; Dudchenko,O; Jeong,J; Weisz,D and Aiden,EL 2025 An unusually large genome from an unusually large stonefly: A chromosome-length genome assembly for the giant salmonfly, Pteronarcys californica (Plecoptera: Pteronarcyidae). Journal of Heredity, 116(3), pp.315-323. PDF
Abstract: "Pteronarcys californica is commonly referred to as the giant salmonfly and is the largest species of stonefly (Insecta: Plecoptera) in the western United States. Historically, it was widespread and abundant in western rivers, but populations have experienced a substantial decline in the past few decades, becoming locally extirpated in numerous rivers in Utah, Colorado, and Montana. Although previous research has explored the ecological variables conducive to the survivability of populations of the giant salmonfly, a lack of genomic resources hampers exploration of how genetic variation is spread across extant populations. To accelerate research on this imperiled species, we present a de novo chromosomal-length genome assembly of P. californica generated from PacBio HiFi sequencing and Hi-C chromosome conformation capture. Our assembly includes 14 predicted pseudo chromosomes and 98.8% of Insecta universal core orthologs. At 2.40 gigabases, the P. californica assembly is the largest of available stonefly assemblies, highlighting at least a 9.5-fold variation in assembly size across the order. Repetitive elements account for much of the genome size increase in P. californica relative to other stonefly species, with the content of Class I retroelements alone exceeding the entire assembly size of all but two other species studied. We also observed preliminary suborder-specific trends in genome size that merit testing with more robust taxon sampling."
Elmork,K and 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
Fritz,KM; Kashuba,RO; Pond,GJ; Christensen,JR; Alexander,LC; Washington,BJ; Johnson,BR; Walters,DM; Thoeny,WT and Weaver,PC 2023 Identifying invertebrate indicators for streamflow duration assessments in forested headwater streams. Freshwater Science, 42(3), pp.247-267. PDF
Fuller,RL and Stewart,KW 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 and 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,JS; Beesley,SS and Van Kirk,RW 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
Heinold,B and Pomeranz,J 2011 Colorado River Aquatic Resources Investigations Federal Aid Project F-237R-18 R. Barry Nehring General Professional V Co-Authors. 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."
Hotaling,S 2017 Genetic Perspectives on Biodiversity in Rocky Mountain Alpine Streams. PhD Thesis, University of Kentucky. 243 pages. PDF
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
Abstract: "Evolutionary biologists have long sought to disentangle phylogenetic relationships among taxa spanning the tree of life, an increasingly important task as anthropogenic influences accelerate population declines and species extinctions, particularly in insects. Phylogenetic analyses are commonly used to identify unique evolutionary lineages, to clarify taxonomic designations of the focal taxa, and to inform conservation decisions. Advances in DNA sequencing techniques have increasingly facilitated the ability of researchers to apply genomic methods to phylogenetic analyses, even for non-model organisms. Stoneflies are non-model insects that are important bioindicators of the quality of freshwater habitats and landscape disturbance as they spend the immature stages of their life cycles in fresh water, and the adult stages in terrestrial environments. Phylogenetic relationships within the stonefly genus Suwallia (Insecta: Plecoptera: Chloroperlidae) are poorly understood, and have never been assessed using molecular data. We used DNA sequence data from genome-wide ultraconserved element loci to generate the first molecular phylogeny for the group and assess its monophyly. We found that Palearctic and Nearctic Suwallia do not form reciprocally monophyletic clades, and that a biogeographic history including dispersal, vicariance, and founder event speciation via jump dispersal best explains the geographic distribution of this group. Our results also strongly suggest that Neaviperla forcipata (Neave, 1929) is nested within Suwallia, and the concept of the genus Suwallia should be revised to include it. Thus, we formally propose a new taxonomic combination wherein Neaviperla forcipata (Neave, 1929) is reclassified as Suwallia forcipata (Neave, 1929). Moreover, some Suwallia species (e.g., S. amoenacolens, S. kerzhneri, S. marginata, S. pallidula, and S. starki) exhibit pronounced cryptic diversity that is worthy of further investigation. These findings provide a first glimpse into the evolutionary history of Suwallia, improve our understanding of stonefly diversity in the tribe Suwallini, and highlight areas where additional research is needed."
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 and 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.
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."
This is an 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.
Kotalik,CJ; Clements,WH and Cadmus,P 2017 Effects of magnesium chloride road deicer on montane stream benthic communities. Hydrobiologia, 799(1), pp.193-202. PDF
Abstract: "Montane streams often intercept and run parallel to roads and highways where road deicer is seasonally applied for snow and ice removal. This research used stream mesocosms to evaluate the effects of MgCl2 road deicer to a Rocky Mountain stream benthic community in Colorado, USA. Measured responses included macroinvertebrate drift, community composition metrics, and macroinvertebrate biomass after a 10-day exposure. Natural benthic communities were exposed to concentrations of liquid MgCl2 road deicer that bracketed the U.S. Environmental Protection Agency (U.S. EPA) surface water chronic chloride ‘aquatic life criteria’ (230 mg Cl—/l). Results showed no effects on macroinvertebrate drift, but significant reductions in abundance, taxa richness, and community biomass. Specifically, stonefly (Plecoptera) and mayfly (Ephemeroptera) abundance decreased at Cl— concentrations below the U.S. EPA chronic chloride water quality standard, and at concentrations substantially lower than those generated from traditional laboratory toxicity tests. However, caddisflies (Trichoptera), midges (Chironomidae) and other dipterans were tolerant to all MgCl2 treatments. We conclude that MgCl2 road deicer has the potential to impair montane stream benthic communities at relatively low ionic concentrations, and regulatory agencies should manage for and establish regionally appropriate application rates for this stressor."
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
Matsuda,K; Buckingham,SD; Kleier,D; Rauh,JJ; Grauso,M and Sattelle,DB 2001 Neonicotinoids: insecticides acting on insect nicotinic acetylcholine receptors. Trends in pharmacological sciences, 22(11), pp.573-580.
Abstract: "Imidacloprid is increasingly used worldwide as an insecticide. It is an agonist at nicotinic acetylcholine receptors (nAChRs) and shows selective toxicity for insects over vertebrates. Recent studies using binding assays, molecular biology and electrophysiology suggest that both α- and non-α-subunits of nAChRs contribute to interactions of these receptors with imidacloprid. Electrostatic interactions of the nitroimine group and bridgehead nitrogen in imidacloprid with particular nAChR amino acid residues are likely to have key roles in determining the selective toxicity of imidacloprid. Chemical calculation of atomic charges of the insecticide molecule and a site-directed mutagenesis study support this hypothesis."
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.
Meyerhoff,RD 1991 Post-eruption recovery and secondary production of grazing insects in two streams near Mt. St. Helens. phd Thesis, Oregon State University. 217 pages. PDF
Abstract: "The eruption of Mt. St. Helens provided the opportunity to study secondary production of grazing insects in the context of disturbance. Two stream sites were chosen that significantly differed in how their watersheds were impacted by the eruption. Clearwater Creek was catastrophically disturbed (physical alteration of habitat, loss of riparian vegetation, and abundant ashfall); Elk Creek was disturbed only by heavy ashfall. Secondary production of the insect community was estimated for 1985 and 1986. The relative importance of disturbance history and between-site habitat differences in determining secondary production was assessed by placing results in the context of ten years (1980-1989) of summer data from both streams. Few insects were found in Clearwater Creek four months after the eruption. In contrast, the community at Elk Creek was diverse and dominated by long-lived taxa. From 1980 to 1989 at Clearwater Creek there was a gradual shift from dominance by Chironomidae to an increasingly diverse community with an abundance of mayflies, stoneflies, and caddisflies. Disturbance from ash scour during the 1980-81 winter reset the benthos of Elk Creek. However, the recovery process in that stream after 1982 was rapid. The annual production of insects in 1985 was 9.7 g dry wt/m2 at Elk Creek and 8.6 g/m2 at Clearwater Creek. In 1986, production was higher at Clearwater Creek (27.4 g/m2) than at Elk Creek (16.3 g/m2). Aufwuchs grazers were the most important insects (ca. 75% of total insect production) of both streams in both years. After differences in assimilation efficiencies of food types were considered, it was found that 84.3% of grazer production at Clearwater Creek depended on algae as compared to 74.3% at Elk Creek. The disturbance history of each site was an important factor determining the insect community structure and the importance of grazers at each site. Consequently, secondary production in 1985-1986 was influenced to a large degree by the 1980 eruption. However, between-site differences in basin dimensions, substrate size, and riparian vegetation likely controlled the productive capacity of each stream."
Mihuc,TB; Mihuc,JR 1995 Trophic ecology of five shredders in a Rocky Mountain stream. Journal of Freshwater Ecology 10 (3) 209-216. PDF
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
Morton,SG; Schmidt,TS and Poff,NL 2022 Lack of evidence for indirect effects from stonefly predators on primary production under future climate warming scenarios. Ecoscience, 29(4), pp.283-291. PDF
Abstract: "Consumptive and non-consumptive interactions of predators and prey can have strong direct and indirect effects on primary producers, such as stream algae. Increasing water temperatures may alter these interactions and thus influence productivity in streams. For each of 3 temperature treatments ('ambient', +2°C and +4°C), we measured the amount of algal biomass removed by grazing mayflies from 91 mesocosms after a 24-hour test period under 3 grazing treatments: lethal predators, non-lethal predators, and no predators. At all temperatures, grazers reduced algal biomass (p < 0.01), and the presence of lethal predators effectively dampened mayfly consumption of algae (p < 0.01). However, differences in algal biomass between lethal and non-lethal predator treatments were not significant, indicating that predators had no indirect behaviorally mediated effects on grazer consumption. Grazer removal of algal biomass marginally increased with increasing temperature (p = 0.051). We analyzed video data for changes in grazer foraging and drift behavior. Mayflies increased drift in the presence of lethal predators (p < 0.01) but not non-lethal predators, and no behavioral changes were seen with temperature increases. Mesocosms can help elucidate possible future shifts in trophic interactions due to climate warming. Yet, we found no evidence of indirect stonefly predator effects on grazing mayflies under these warming scenarios."
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 1982 The importance of sampling and sorting techniques in the elucidation of the life cycle of Zapada columbiana (Nemouridae: Plecoptera). Canadian Journal of Zoology, 60(12), pp.3394-3399.
Abstract: "The life cycle of Zapada columbiana (Claassen) was shown to be of 2 years duration at three sites in Alberta, Canada, where the species was previously said to be univoltine. The erroneous conclusion of previous investigators was attributed primarily to the use of nets with too large a mesh size. However other factors such as sorting without magnification, inability to recognize young Z. columbiana, and erroneous assumptions regarding egg incubation times may also have contributed. It is suggested that the temperature regimes of the streams in the Alberta Rocky Mountains are such that Zapada columbiana cannot complete its life cycle in 1 year."
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: "The eggs of eight species of Plecoptera from Alberta were reared at constant temperatures between 2 and 25°C. All species' eggs hatched at 2°C but none hatched at 25°°C. The relationships between temperature and number of days required for development and hatching were fitted to power equations. Slope values (b) ranged from -1.31 for Skwala parallela to 0.03 for Amphinemura banksi; the remainder fell between -0.94 and -0.68. All of these values, except for that for S. parallela, imply that stonefly eggs develop most efficiently at the lowest temperature within the favorable range or develop with equal efficiency throughout the favorable range."
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. PDF
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.
Newell,RL; Baumann,RW and Stanford,JA 2008 Stoneflies of Glacier National Park and Flathead River basin, Montana. International Advances in the ecology, zoogeography, and systematics of mayflies and stoneflies. University of California Publications in Entomology, Berkeley and Los Angeles, pp.173-186.
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. PDF
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. "
Poff,NL; Olden,JD; Viera,NKM; Finn,DS; Simmons,MP; Kondratieff,BC 2006 Functional trait niches of American lotic insects: traits-based ecological applications in light of phylogenetic relationships. Journal of the North American Benthological Society 25 (4) 730-755. PDF
Abstract: "The use of species traits to characterize the functional composition of benthic invertebrate communities has become well established in the ecological literature. This approach holds much potential for predicting changes of both species and species assemblages along environmental gradients in terms of traits that are sensitive to local environmental conditions. Further, in the burgeoning field of biomonitoring, a functional approach provides a predictive basis for understanding community-level responses along gradients of environmental alteration caused by humans. Despite much progress in recent years, the full potential of the functional traits-based approach is currently limited by several factors, both conceptual and methodological. Most notably, we lack adequate understanding of how individual traits are intercorrelated and how this lack of independence among traits reflects phylogenetic (evolutionary) constraint. A better understanding is needed if we are to make the transition from a largely univariate approach that considers single-trait responses along single environmental gradients to a multivariate one that more realistically accounts for the responses of many traits across multiple environmental gradients characteristic of most human-dominated landscapes. Our primary objective in this paper is to explore the issue of inter-trait correlations for lotic insects and to identify opportunities and challenges for advancing the theory and application of traits-based approaches in stream community ecology. We created a new database on species-trait composition of North American lotic insects. Using published accounts and expert opinion, we collected information on 20 species traits (in 59 trait states) that fell into 4 broad categories: life-history, morphological, mobility, and ecological. First, we demonstrate the importance of considering how the linkage of specific trait states within a taxon is critical to developing a more-robust traits-based community ecology. Second, we examine the statistical correlations among traits and trait states for the 311 taxa to identify trait syndromes and specify which traits provide unique (uncorrelated) information that can be used to guide trait selection in ecological studies. Third, we examine the evolutionary associations among traits by mapping trait states onto a phylogentic tree derived from morphological and molecular analyses and classifications from the literature. We examine the evolutionary lability of individual traits by assessing the extent to which they are unconstrained by phylogenic relationships across the taxa. By focusing on the lability of traits within lotic genera of Ephemeroptera, Plecoptera, and Trichoptera, taxa often used as water-quality indicators, we show how a traits-based approach can allow a priori expectations of the differential response of these taxa to specific environmental gradients. We conclude with some ideas about how specific trait linkages, statistical correlations among traits, and evolutionary lability of traits can be used in combination with a mechanistic understanding of trait response along environmental gradients to select robust traits useful for a more predictive community ecology. We indicate how these new insights can direct the research in statistical modeling that is necessary to achieve the full potential of models that can predict how multiple traits will respond along multiple environmental gradients."
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 1949 Some evolutionary trends in Plecoptera. In Proceedings of the Indiana Academy of Science (Vol. 59, pp. 197-209). PDF
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
Riedl,H 2017 Introduced plant affects the structure and function of riparian food webs, Colorado State University Masters Thesis. 81 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; Davis,MA; Johnson,KP; Teslenko,VA; Lee,JJ; Malison,RL; Hwang,JM; Bae,YJ and Myers,LW 2021 A new family of stoneflies (Insecta: Plecoptera), Kathroperlidae, fam. n., with a phylogenomic analysis of the Paraperlinae (Plecoptera: Chloroperlidae). Insect Systematics and Diversity, 5(4), p.1. PDF
Abstract: "Recent molecular analyses of transcriptome data from 94 species across 92 genera of North American Plecoptera identified the genus Kathroperla Banks, 1920 as sister group to Chloroperlidae + Perlodidae. Given that the genus Kathroperla has historically been included as a member of the family Chloroperlidae, this discovery indicated further investigation of the genus and the subfamily Paraperlinae was needed. Both transcriptome and genome sequencing datasets were generated from 32 species of the infraorder Systellognatha, including all described species of the Paraperlinae, to test the phylogenetic placement of these taxa. From these datasets, a large phylogenomic data matrix of 800 orthologous genes was produced, and multiple analyses were conducted, including both concatenated and coalescent analyses. Morphological comparisons were made among all Paraperlinae using light microscopy. All molecular results support a monophyletic Kathroperla, which is supported as sister taxon to the remaining Perloidea by five of six molecular analyses. Postocular head length is determined to be a distinct morphological character of this genus. Combined molecular and morphological evidence support the designation of Kathroperlidae, fam. n., as the seventeenth family of extant Plecoptera."
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.
Abstract: "Samples were taken year-round at eleven sites along the altitudinal profile (2900-1400 m a.s.l.) of the Gunnison River, a 329 km tributary of the Colorado River, to document the distribution of the Plecoptera and to evaluate responses to hypolimnial-release dams in the headwaters and middle reaches. Twenty-two species were present, with the greatest species richness occurring in an unregulated segment upstream of the middle reach dams; average nymphal biomass over the study period (175 organisms, 395 mg dry mass m-2) was also greatest in this segment. Only four species (58 organisms, 48 mg m-2) were present in the tailwaters of the headwater dam and values were greatly reduced (nine species; 35 organisms, 180 mg m-2) below the middle reach dams. The stonefly community recovered ca. 80 km downstream from the last dam (15 species; 244 organisms, 250 mg m-2), apparently in response to natural resetting of environmental conditions corresponding to those above the middle reach dams. At the most downstream site (11) only four species (four organisms, 16 mg m-2) were present. The observed distributional pattern is a classic serial discontinuity in response to hypolimnial stream regulation in a temperate latitude river."
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 and 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; Dewalt,RE and Oswood,MW 1991 Alaskaperla, a new stonefly genus (Plecoptera: Chloroperlidae), and further descriptions of related Chloroperlidae. Annals of the Entomological Society of America, 84(3), pp.239-247.
Abstract: "Chloroperla ovibovis Ricker is removed from Chloroperla and placed in a new monotypic genus Alaskaperla, and first descriptions of its eggs and nymphs are provided. Vein A2 of the forewing is unbranched, unlike all other North American Chloroperlinae genera except Haploperla, and the delicate, paired aedeagal rods separate A. ovibovis from European Chloroperla sensu Zwick (1967), which have complex, blade-like rods, and from Triznaka, which have no rods. The elongate, tongue-like epiproct and aedeagus are distinctive among Chloroperlinae. Females have a broadly rounded subgenital plate, produced over two-thirds of sternum 9. Nymphs have a distinctive head pattern, a double row of barbed bristles on the mandibles, a single row of paired lacinial bristles and are described in conformance to Stewart & Stark (1988). The ovate eggs average 250 by 190 µm, have shallow hexagonal follicular cell impressions and rounded pits over the chorion surface and have greater development of the collar than other described Chloroperlinae. First SEMs of adult characters are presented for the type species of Chloroperla, selected Haploperla, Rasvena, Triznaka, Siphonoperla, and Suwallia species, and the eggs of Triznaka signata."
Stewart,KW and 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 1984 Nymphs of North American Perlodinae genera (Plecoptera: Perlodidae). The Great Basin Naturalist, pp.373-415. PDF
Abstract: "Nymphs of the type or other representative species of the 22 North American Perlodinae genera are comparatively described and illustrated for the first time. The first complete generic key for the subfamily incorporates recent nymph discoveries and revisions in classification. References to all previous nymph descriptions and illustrations and major life cycle and food habits studies are given for the 53 North American species in the subfamily, and a listing of species and their current distributions by states and provinces is provided for each genus. The previously unknown nymph of Chernokrilus misnomus is described and illustrated."
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 we 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. PDF
Taylor,BW; McIntosh,AR and Peckarsky,BL 2001 Sampling stream invertebrates using electroshocking techniques: implications for basic and applied research. Canadian Journal of Fisheries and Aquatic Sciences, 58(3), pp.437-445. PDF
Abstract: "We present a new technique using electrofishing equipment to collect and quantitatively sample stream invertebrates. We used an electrofishing machine with a small anode to produce a localized field of pulsed direct current to induce invertebrate drift. We quickly obtained large numbers of live invertebrates for experiments by passing the anode over the stream bottom upstream of sampling nets. We compared the results of five techniques: (i) electroshocking inside a modified Hess sampler, (ii) repeated electroshocking over a large area to estimate population size by depletion, (iii) traditional Surber, (iv) Hess, and (v) individual stone sampling. Electroshocking techniques provided estimates of invertebrate density comparable with those of traditional sampling techniques. The electroshocking depletion method that sampled a large area provided higher measures of Ephemeroptera, Plecoptera, and Trichoptera richness. Hess and area-restricted electrobug methods had similar density and diversity estimates, whereas the Surber sampler provided low density estimates, especially for mobile taxa. Density estimates from individual stones were inflated, were biased for mayflies, and had low richness. Samples taken with the electroshocking method were processed 40% faster because these samples contained little detritus. Electroshocking techniques can provide accurate estimates of population size and diversity, minimize disturbance to benthic habitats, and reduce processing time."
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.
Usinger,RL ed., 1956 Aquatic insects of California: with keys to North American genera and California species. Univ of California Press.
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,NK; Clements,WH; Guevara,LS and Jacobs,BF 2004 Resistance and resilience of stream insect communities to repeated hydrologic disturbances after a wildfire. Freshwater Biology, 49(10) 1243-1259. 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 1984 Diversity patterns exhibited by the Plecoptera of a Colorado mountain stream. In Annales de limnologie (Vol. 20, No. 1-2, pp. 123-128). EDP Sciences. PDF
Abstract: "Plecoptera collected from 11 locations along the longitudinal profile (first to fifth orders) of a Rocky Mountain stream (St. Vrain Creek) revealed distinctive spatial patterns of species diversity. Overall diversity was low near the stream source (epirhithron), attained maximum values in middle reaches, then declined dramatically concomitant with the transition from rhithron [steeper, alternating riffles or rapids with pools] to potamon [flat, meandering, slower flowing water] conditions. Shannon-Weaver index values ([math]) were < 1.0 at high elevations and at the potamon station, whereas [math] exceeded 3.0 in middle reaches. Species richness based on summer data alone is depressed by nearly 50 % at some locations, but the summer spatial pattern of [math] values is in general concordance with that derived from the entire data set. Species diversity patterns of consecutive summers were relatively stable considering the substantial differences in discharge. To accurately determine Plecoptera diversity patterns, it was necessary to combine data from both fine (240 µm) and coarse mesh (720 µm samples). Although a myriad of interrelated factors determine the diversity of lotic insects at a given site, summer maximum temperature and annual temperature range appear to be major variables controlling the spatial diversity pattern of Plecoptera."
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 helped us keep up on the 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. As websites such as SciLit and Google Scholar came online, NABS stopped publishing this and later changed their name to The Society for Freshwater Science
Wellnitz,T 2014 Can current velocity mediate trophic cascades in a mountain stream?. Freshwater Biology, 59(11) 2245-2255. PDF
Wigglesworth,VB 2012 The principles of insect physiology. Springer Science & Business Media.
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.
Willkommen,J 2008 The morphology of the pterothorax of Ephemeroptera, Odonata and Plecoptera (Insecta) and the homology of wing base sclerites and flight muscles. Stuttgarter Beiträge zur Naturkunde A, Neue Serie, 1, pp.203-300. PDF
Abstract: "The ability to fly was the decisive factor for the evolutionary success of the most diverse group of insects, the Pterygota. Nevertheless, the ground plan of the functionally important wing base has not been sufficiently clarified.
The aim of this study is to homologise the wing base sclerites of Ephemeroptera, usually regarded as sister group of the remaining Pterygota, with that of other basal pterygote lineages and to reconstruct the ground plan of the wing base of Pterygota. The pterothoracic musculature of representatives of the three basal lineages of Pterygota (Ephemeroptera, Odonata and Neoptera) is also described and discussed.
Contrary to previous hypotheses, it is shown that most elements of the neopteran wing base are also present in Ephemeroptera and Odonata. The wing base in the ground plan of Pterygota is presumably composed of three axillary sclerites. The proximal median plate is probably also present in the ground plan of Pterygota. The first axillary is provided with two muscles. The third axillary is equipped with a short muscle that originates from the epimeron. This muscle is interpreted as another ground plan character of Pterygota. In Plecoptera a second muscle inserts at the third axillary sclerite. It originates from the episternum and is most likely an autapomorphic character of Neoptera. The results imply that the wing base of the Plecoptera is close to the pterygote ground plan. It is assumed that the wing base of Ephemeroptera and Odonata is secondarily stiffened. The so-called basalare and its associated muscles in Ephemeroptera and Odonata are probably not homologous to the basalare and respective muscles in Neoptera.
The enlarged subalare and associated muscles, the large dorsal longitudinal muscle, the small metathorax and shortened hind wings in Ephemeroptera suggest that mayflies have a derived flight apparatus in many respects. The Odonata on the other hand show different specialisations, namely a synthorax, large direct flight musculature, and a fusion of second and third axillary with the proximal median plate. Though the wing base in both taxa is secondarily stiffened, the specialisations of Ephemeroptera and Odonata may have evolved independently from each other."
Winterbourn,MJ and Crowe,ALM 2001 Flight activity of insects along a mountain stream: is directional flight adaptive? Freshwater Biology 46, 1479-1489. PDF
Wolfe,JM; Daley,AC; Legg,DA and Edgecombe,GD 2016 Fossil calibrations for the arthropod Tree of Life. Earth-Science Reviews, 160, pp.43-110. PDF
Abstract: "Fossil age data and molecular sequences are increasingly combined to establish a timescale for the Tree of Life. Arthropods, as the most species-rich and morphologically disparate animal phylum, have received substantial attention, particularly with regard to questions such as the timing of habitat shifts (e.g. terrestrialisation), genome evolution (e.g. gene family duplication and functional evolution), origins of novel characters and behaviours (e.g. wings and flight, venom, silk), biogeography, rate of diversification (e.g. Cambrian explosion, insect coevolution with angiosperms, evolution of crab body plans), and the evolution of arthropod microbiomes. We present herein a series of rigorously vetted calibration fossils for arthropod evolutionary history, taking into account recently published guidelines for best practice in fossil calibration. These are restricted to Palaeozoic and Mesozoic fossils, no deeper than ordinal taxonomic level, nonetheless resulting in 80 fossil calibrations for 102 clades. This work is especially timely owing to the rapid growth of molecular sequence data and the fact that many included fossils have been described within the last five years. This contribution provides a resource for systematists and other biologists interested in deep-time questions in arthropod evolution."
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 www.gunnisoninsects.org
"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
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