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Publications about the Ephemeroptera (Mayflies) of Gunnison County, Colorado

Updated 1 March 2024

Check out The Virtual Library from Ephemeroptera Galactica, it has a very complete bibliography for mayflies worldwide and is a great source for downloadable PDF files of many publications. Many of the PDF links on this website are to the Virtual Library.

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Aaraard,K; Solem,JO; Nøst,T and Hannssen,O 1997 The macrobenthos of the pristine steam Skiftesåa, Høylandet, Norway. Hydrobiologia 348: 81-94.

Alexander,RD and Brown Jr,WL 1963 Mating behavior and the origin of insect wings. Occasional papers of the Museum of Zoology University of Michigan 628:1-19.

Alexander,LC; Delion,M; Hawthorne,DJ; Lamp,WO and Funk,DH 2009 Mitochondrial lineages and DNA barcoding of closely related species in the mayfly genus Ephemerella (Ephemeroptera: Ephemerellidae). Journal of the North American Benthological Society, 28(3) 584-595. PDF

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

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

Allan,JD 1978a 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,JD 1978b Trout predation and the size composition of stream drift. Limnology and Oceanography 23 6, 1231-1237.

Allan,JD 1981 Determinants of diet of brook trout (Salvelinus fontinalis) in a mountain stream. Canadian Journal of Fisheries and Aquatic Sciences 38, 184-192. PDF

Allan,JD 1985 The production ecology of Ephemeroptera in a Rocky Mountain stream. Internationale Vereinigung für Theoretische und Angewandte Limnologie Verhandlungen 22, 3233-3237. PDF

Allan,JD 1987 Macroinvertebrate drift in a Rocky Mountain stream. Hydrobiologia 144, 261-268.
     Based at the Rocky Mountain Biological Lab in Gothic, the author studied Cement Creek in Gunnison County during the spring, summer and fall of 1975-1978. He found that drift densities (number of animals per 100 m³) were 10 times higher at night. 24 hour totals approached 2000 animals/100m³ in mid-summer down to 500 animals/m³ in the fall. Quote from the abstract: "Ephemeroptera, especially Baetis, dominated the drift." He studied the effects of stream discharge, benthic density (number animals/m² of bottom samples) and temperature on drift finding that discharge and benthic density significantly affected some taxa and not others, while temperature had no effect on drift. He found his results varied by whether he used drift density (number animals/m³ of water) or drift rate (number animals past a certain point for a specific time). Quote from page 265: "Expressing drift on a per unit volume basis eliminated a significant dependency of drift on flow in 5 or 8 taxa examined. Benthic density typically was the best predictor of 24 hour drift density." More information on these taxa: Baetis bicaudatus, Cinygmula spp., Epeorus longimanus

Allan,JD and Feifarek,BP 1989 Distances travelled by drifting mayfly nymphs: factors influencing return to the substrate. Journal of the North American Benthological Society 8 4, 322-330.

Allan,JD; Flecker,AS and McClintock,NL 1986 Diel epibenthis activity of mayfly nymphs, and its nonconcordance with behavioral drift. Limnology and Oceanography 31 5, 1057-1065.

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

Allan,JD and Russek,E 1985 The quantification of stream drift. Canadian Journal of Fisheries and Aquatic Sciences 42:210-215. PDF
     Although the purpose of many drift studies is to describe quantitatively the abundance of drifting invertebrates and make comparisons between seasons or sites, almost no investigations have employed replicate sampling. We analyzed drift collections from a Rocky Mountain stream in order to investigate the variability of drift sampling. The data were normalized and the variances stabilized for each taxon examined by data transformation. The fourth root transformation was favored for five taxa and the logarithmic transformation for three. Using the 95% confidence limits on 24-k drift density for an abundant mayfly (Baetis bicaudatus), we found that six to seven replicates are required to obtain 95% CL + 58% of the mean. Drift sampling appears to require fewer replicates than benthic sampling for comparable precision. Investigators may fail to replicate drift samples because they elect to sample frequently over 24 hr in order to quantify the diel periodicity of drift. However, when comparison between sites or dates is the principal goal, we recommend that the effort normally put into frequent sampling over 24 h be invested instead in replicated sampling just after dark, whew drift normally is greatest. When we regressed drift from the first night sample against total drift from the remainder of the 24-h period, 60-90% of the variation in the latter was predicted from the single nighttime sample. Thus, little information appears to be lost by this recommended procedure.

Allen,RK 1968 New species and records of Ephemerella (Ephemerella) in western North America (Ephemeroptera: Ephemerellidae). Journal of the Kansas Entomological Society 41(4):557-567 PDF

Allen,RK 1974 Neochoroterpes, a new subgenus of Choroterpes Eaton from North America (Ephemeroptera: Leptophlebiidae). Canadian Entomologist 106:161-168.

Allen,RK 1984 A new classification of the subfamily Ephemerellinae and the description of a new genus. Pan-Pacific Entomologist 60, 245-247.

Allen,RK and Chao,ESM 1978 Mayflies of the Southwest: new species and records of Heptageniidae. Pan-Pacific Entomologist 54, 311-315PDF.

Allen,RK and Chao,ESM 1981 Mayflies of the Southwest: new records and notes of Siphlonuridae (Ephemeroptera: Baetidae). Pan-Pacific Entomologist 57, 449-456. PDF

Allen RK and Edmunds Jr GF. 1959 A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). I. The subgenus Timpanoga. Canadian Entomologist 91:51-58. PDF

Allen,RK and Edmunds,GF 1962 A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). V. The subgenus Drunella in North America. Miscellaneous Publications of the Entomological Society of America 3, 147-179. PDF

Allen,RK and Edmunds Jr,JF 1963 A revision of the genus Ephemerella (Ephemeroptera: Ephemerellidae). VI. The subgenus Serratella in North America. Annals of the Entomological Society of America 56:583-600. PDF

Alvarez,M and Peckarsky,BL 2013 The influence of moss on grazers in high-altitude streams: food, refuge or both? Freshwater Biology, 58(9) 1982-1994. PDF

Argyle,DW and Edmunds Jr,GF 1962 Mayflies (Ephemeroptera) of the Curecanti Reservoir Basins Gunnison River, Colorado. University of Utah Anthropological Papers 59 8, 178-189.
Table 1. Distribution of mayflies in the Curecanti region.
Taxonomy Translation (page 182)
Argyle and Edmunds Name 2005 Name
Siphlonurus occidentalis same
Heptagenia solitaria same
Rhitrogena robusta same
Epeorus albertae same
Epeorus longimanus same
Ephemerella doddsi Drunella doddsii
Ephemerella grandis Drunella grandis grandis
Ephemerella hecuba hecuba Timpanoga hecuba hecuba
Ephemerella inermis same
Ephemerella margarita Attenella margarita
Ephemerella tibialis Serratella tibialis
Tricoythodes minutus same
Paraleptophlebia pallipes Paraleptophlebia memorialis

Allen,RK and Murvosh,CM 1987 Leptophlebiidae of the southwestern United States and northwestern Mexico (Insecta: Ephemeroptera). Great Basin Naturalist 47(2):283-286. PDF


Ball,SL; Hebert,PDN; Burian,SK and Webb,JM 2005 Biological identification of mayflies (Ephemeroptera) using DNA barcodes. Journal of the North American Benthological Society 24 (3) 508-524. PDF

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

Banks,N 1924 Descriptions of new neuropteroid insects. Bulletin of the Museum of Comparative Zoology. 65(12):419-455, pl. 1-4.

Barber-James,HM; Gattolliat,J; Sartori,M and MD Hubbard 2008 Global diversity of mayflies (Ephemeroptera, Insecta) in freshwater. Hydrobiologia 595(1) 339-350. DOI:10.1007/s10750-007-9028-y Abstract and Full Text

Baumgardner,DE 2009 Tricorythus minutus Traver, a new synonym of Tricorythodes explicatus Eaton (Ephemeroptera: Leptohyphidae). Proceedings of the Entomological Society of Washington 111(1):57-67. PDF

Bedarik,AF and Edmunds,GF 1980 Descriptions of larval Heptagenia from the Rocky Mountain region (Ephemeroptera: Heptageniidae). Pan-Pacific Entomologist 56, 51-62.

Bengtsson,S 1908 Berättelse öfver en resa i entomologiskt syfte til mellersta Sverige Sommaren 1907. K. Svenska Vetenskapsakad. Årsbok 6: 237-46.

Bengtsson,S 1930 Kritische Bemerkungen über einige Nordische Ephemeropteren, nebst Beschreibung neuer Larven. Acta Universitatis Lundensis. 2(2) 1-27. PDF

Benton,MJ and Pritchard,G 1990 Mayfly locomotory responses to endoparasitic infection and predator presence: the effects on predator encounter rate. Freshwater Biology 23(2) 363-371 Abstract

Bergman,EA and Hilsenhoff,WL 1978 Parthenogenisis in the mayfly genus Baetis (Ephemeroptera: Baetidae). Annals of the Entomological Society of America 71:167-168.

Behmer,DJ and Hawkins,CP 1986 Effects of overhead canopy on macroinvertebrate production in a Utah stream. Freshwater Biology 16:287-300.

Blinn,DW and Ruiter,DE 2006 Tolerance values of stream caddisflies (Trichoptera) in the lower Colorado river basin, USA. The Southwestern Naturalist 51(3):326-337.

Brinkman,SF and Johnston,WD 2008 Acute toxicity of aqueous copper, cadmium, and zinc to the mayfly Rhithrogena hageni. Archives of Environmental Contamination and Toxicology 54:3, 466-472. PDF

Brinkman,SF and Johnston,WD 2012 Acute toxicity of zinc to several aquatic species native to the Rocky Mountains. Archives of Environmental Contamination and Toxicology, 62(2), 272-281.

Brittain,JE 2008 Mayflies, biodiversity and climate change. International Advances in the Ecology, Zoogeography and Systematics of Mayflies and Stoneflies (128)1-14. PDF
     Abstract: "Mayflies (Ephemeroptera) are an ancient order of insects that are globally distributed in both northern and southern hemispheres and have survived major environmental shifts. Despite the problems associated with selection processes operating in both terrestrial and aquatic environments, mayflies have successfully colonized a wide range of freshwater habitats from the tropics to the arctic, a somewhat greater range than other hemimetabolic aquatic insects such as the Plecoptera and Odonata. While many species of Ephemeroptera require specific environmental cues, others display considerable flexibility in life cycle length and timing in relation to environmental changes. This is particularly apparent in arctic and alpine species. Climate change scenarios predict rapid shifts across many environmental gradients, including temperature and the frequency and magnitude of floods and droughts. Changes in the mayfly fauna are hypothesized in the light of the environmental tolerances, life cycle plasticity and the dispersal mechanisms of present day mayflies. During periods of rapid environmental transition certain species traits will be beneficial. Generalists will do better; specialists with strict environmental limits and poor powers of dispersal may become extinct."

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.

Buchwalter,DB and Luoma,SN 2005 Differences in dissolved cadmium and zinc uptake among stream insects: mechanistic explanations. Environmental Science and Technology 39, 498-504.

Burton,GJ and McRae,TM 1972 Phoretic attachment of Simulium larvae and pupae to mayfly and dragonfly nymphs. Mosquito News 32(3):436-443.

Bushnell,JH; Foster,SQ AND Wahle,BM 1987 Annotated inventory of invertebrate populations of an alpine lake and stream chain in Colorado. Great Basin Naturalist 47, 500-511.


Cain,DJ; Luoma,SN and Wallace,WG 2004 Linking metal bioaccumulation of aquatic insects to their distribution patterns in a mining-impacted river. Environmental Toxicology and Chemistry 23, 1463-1473.

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

Canton,SP; Cline,LD; Short,R and Ward,JV 1984 The macroinvertebrates and fish of a Colorado stream during a period of fluctuating discharge. Freshwater Biology 14(3) 311-316.

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
     Curious about the effects of chronic metal contamination on mayfly (and other taxa) production, the authors compared two clean reference streams to three polluted streams in the mountains near Leadville, Colorado. The clean streams had 36 and 56 µ/L of Zinc respectively while the polluted streams had low (105µg/L) medium (210µg/L) or high (293µg/L) levels of Zinc. They found that secondary production of the Heptageniidae Rhitrogena robusta, Cinygmula sp. and Epeorus longimanus was lower in slightly to moderately conatminated streams. These taxa and many others were absent from seriously polluted streams and thus their productivity approached zero in streams with the highest levels of Zinc pollution. Growth estimates from feeding experiments discovered that the reduction in secondary production was mostly due to the decline of Rhitrogena robusta with increasing levels of Zinc.

Caudill,CC 2002 Metapopulation biology of the mayfly Callibaetis ferrugineus hageni in high elevation beaver ponds. Cornell University Ph.D. Thesis

Caudill,CC 2003 Measuring dispersal in a metapopulation using stable isotope enrichment: high rates of sex-biased dispersal between patches in a mayfly metapopulation. Oikos 101 3, 624-630.

Caudill,CC and Peckarsky,BL. 2003. Lack of appropriate behavioral or developmental responses by mayfly larvae to trout predators. Ecology 84(8):2133-2144. Abstract

Chaika,V; Pikula,K; Vshivkova,T; Zakharenko,A; Reva,G; Drozdov,K; Vardavas,AI; Stivaktakis,PD; Nikolouzakis,TK; Stratidakis,AK and Kokkinakis,MN 2020 The toxic influence and biodegradation of carbon nanofibers in freshwater invertebrates of the families Gammaridae, Ephemerellidae, and Chironomidae. Toxicology Reports, 7, pp.947-954. PDF
     Abstract: "Carbon nanofibers (CNFs) are widely used in consumer products today. In this study, we assessed the effects of CNFs on the digestive system of three freshwater invertebrate species (Gammaridae, Ephemerellidae, and Chironomidae). The aquatic insects Diamesa sp., Drunella cryptomeria, and Gammarus suifunensis were incubated with the CNFs at the concentration of 100 mg/L during the 7-days period. Histological examination of the whole specimens and the longitudinal sections revealed no toxic effects of CNFs. However, a noticeable change in the structure of the CNFs accumulated in the intestines of the aquatic insects was found by Raman spectroscopy. The registered decrease in the relative proportion of amorphous carbon included in the CNF sample was found in the intestines of Diamesa sp. and D. cryptomeria. The registered effect can indicate a biodegradation of amorphous carbon in the digestive tract of these two insect species. In contrast, the decrease of highly structured carbons and the decrease of G-bonds intensity were registered in the digestive tract of G. suifunensis. This observation demonstrates the partial biodegradation of CNFs in the digestive tract of G. suifunensis."

Check,GR 1982 A revision of the North American spcies of Callibaetis (Ephemeroptera: Baetidae). Ph.D. Dissertation, University of Minnesota, USA.

Clark,JL and Clements,WH 2006 The use of in situ and stream microcosm experiments to assess population- and community-level responses to metals. Environ Toxicol Chem. 25(9)2306-2312.

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. Quote from page 632: "In particular, abundance of the mayflies Rhithrogena robusta (Fig. 5b), Cinygmula sp.(Fig. 5c), and Drunella doddsi(Fig. 5d), and the stonefly Sweltsa sp.(Fig. 5e) was significantly lower at medium- and high-metal stations."

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 1982a Aquatic insects as measures of trace element presence in water: Cadmium and Molybdenum. Aquatic Toxicology and Hazard Assessment: Fith Conference, ASTM STP 766, J.G. Pearson, R.B. Foster, and W.E. Bishop, Eds., American Society for Testing and Materials, pgs 316-325.

Colburn,T 1982b Measurement of low levels of molybdenum in the environment by using aquatic insects. 29, 422-428.
     The author measured molybdenum (Mo) levels in several aquatic imsect species in the upper Gunnison Basin.

Corkum,LD 1980 Carnivory in Ephemerella inermis Eaton nymphs (Ephemeroptera: Ephemerellidae). Entomological News 91(5):161-163.

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

Corkum,LD μμand Clifford,HF 1981 Function of caudal filaments and correlated structures in mayfly nymphs, with special reference to Baetis (Ephemeroptera). Quaestiones Entomologicae 17:129-146. PDF

Courtney,LA and Clements,WH 2000 Sensitivity to acidic pH in benthic invertebrate assemblages with different histories of exposure to metals. Journal of the North American Benthological Society 19 (1) 112-127.Abstract

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."

Cowan,CA and Peckarsky,BL 1990 Feeding by a lotic mayfly grazer as quantified by gut fluorescence. Journal of the North American Benthological Society 9(4)368-378. PDF

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

Culp,JM and Scrimgeour,GJ 1993 Size-dependent diel foraging periodicity of a mayfly grazer in streams with and without fish. Oikos 68(2)242-250. PDF


Dahl,J and Peckarsky,BL 2002. Induced morphological defenses in the wild: predator effects on a mayfly, Drunella coloradensis. Ecology 83:1620-1634. PDF

De Jong,GD 2015 Acute toxicity of natural acidic hydrothermal alteration scar runoff and historic frequency of potential exposure to a native mayfly in the Red River, New Mexico. Environmental Earth Sciences, 74(9), pp.6871-6876.

De Jong,GD 2024 Habitat and benthic community correlates of Epeorus longimanus (Ephemeroptera: Heptageniidae) microdistribution in a Colorado, USA, stream riffle. Annals of the Entomological Society of America, 117(1), pp.21-26.

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

Dobrin,M and Giberson,DJ 2004 Life history and production of mayflies, stoneflies, and caddisflies (Ephemeroptera, Plecoptera, and Trichoptera) in a spring-fed stream in Prince Edward Island, Canada: evidence for population asynchrony in spring habitats? Can. J. Zool./Rev. Can. Zool. 81(6): 1083-1095

Dodds,GS 1923 Mayflies from Colorado: descriptions of certain species and notes on others. Transactions of American Entomological Society 69: 93-116. PDF
     Dodds collected and reared mayflies in Boulder Creek in the Front Range of Colorado during the early 1900s. He described several mayfly species found in Gunnison County.

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. PDF

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.

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

Durfee,R and Kondratieff,BC 1993 Description of adults of Baetis magnus (Ephemeroptera: Baetidae). Entomological News 104 (5) 227-232.
    The authors examined B. magnus from the Gunnison River near Gunnison among other places. They discuss the details of telling this species apart from Baetis tricaudatus. They note variation in the number and placement of several wing veins across several geographical locations.

Durfee,RS and Kondratieff,BC 1994 New additions to the inventory of Colorado mayflies (Ephemeroptera). Entomological News 105(4):222-227. PDF
     Abstract: " Four mayfly species, Beatis dardanus, Paracloedes minutes, Homoeoneuria alleni, and Rhithrogenia pellucida are reported from Colorado for the first time. Homoeoneuria and Paracloedes represent new generic records for the state. The male terminalia of R. pellicuda from Colorado and a paratype from Minnesota are illustrated and compared, and characters are given to separate this species from the four other Rhithrogena species in Colorado. Previous tentative records for two additional species, Baetis virile and Leptophlebia nebulosa are confirmed. Biogeographic affinities for several species in the North Platte River system are discussed. A total of 43 genera and 101 species of mayflies are now known from Colorado.

Durfee,RS and Kondratieff,BC 1995 Description of adults of Baetis notos (Ephemeroptera:Baetiidae). Entomological News 106 2, 71-74.

Durfee,RS and Kondratieff,BC 1999 Notes on North American Baetis (Ephemeroptera: Baetidae):Baetis moffatti new synonym of B. tricaudatus and range extension for B. bundyae. Entomological News 110 3, 177-180.


Eaton,AE 1869 On Centroptilum, a new genus of the Ephemeridae. Entomologist's Monthly Magazine 6:131-132.

Eaton,AE 1881 An announcement of new genera of the Ephemeridae. Entomologist's Monthly Magazine 17:191-197.
     The Reverend Alfred Edwin Eaton describes the mayfly genus Choroterpes in this paper.

Eaton,AE 1883-1888. A revisional monograph of recent Ephemeridae or mayflies. Transactions of the Linnean Society of London, Second Series, Zoology 3:1-352, 65 pl.

Eaton,AE 1892 Fam. Ephemeridae. Biologica Centrali-Americana 38:1-16, 1 pl.

Edmunds Jr,GF 1945 Ovoviviparous mayflies of the genus Callibaetis (Ephemeroptera: Baetidae). Entomological News 56:169-171.

Edmunds Jr,GF 1948 A new genus of mayflies from western North America (Leptophlebiinae). Proceedings of the Biological Society of Washington 61:141-148.

Edmunds Jr,GF 1952a Studies on the Ephemeroptera Part I. The phylogeny and classification of the Ephemeroptera, with a study of flight mechanics and evolution of their wings. PhD Thesis, University of Massachusetts. 103 pages.

Edmunds Jr,GF. 1952b Studies on the Ephemeroptera Part II. The taxonomy and biology of the mayflies in Utah. PhD Thesis, University of Massachusetts. 399 pages.

Edmunds Jr,GF 1962 The food habits of the nymphs of the mayfly Siphlonurus occidentalis. Proceedings of the Utah Academy of Science, Arts and Letters 37, 73-74.

Edmunds Jr, GF 1995 Habitat differences between northern and southern populations of mayflies of the western United States. Pages 171-176 in Corkum LD; Ciborowski JJH. Current Directions in Research on Ephemeroptera. Canadian Scholars' Press, Inc. Toronto.

Edmunds Jr,GF and Allen,RK 1964 The Rocky Mountain species of Epeorus (Iron) Eaton (Ephemeroptera: Heptageniidae. Journal of the Kansas Entomological Society 37 4, 275-288. PDF

Edmunds Jr,GF and McCafferty,WP 1988 The mayfly subimago. Annual review of entomology, 33(1)509-527. PDF

Edmunds Jr,GF and Musser,GG 1960 The mayfly fauna of the Green River in the Flaming Gorge Reservoir Basin, Wyoming and Utah. University of Utah Anthropological Papers. 48:111-123.
     Introduction: " The construction of Flaming Gorge dam on the Green River in extreme north-eastern Utah will inundate large areas of lotic water and change the habitat to standing water. Several tributary streams will also be inundated on their lower protions but their fauna, which is largely montane, will be maintained in their exposed upper reaches. The mayfly fauna of the Green River proper is one that is so unique and diverse that its loss by inundation is very regrettable.
The construction of a dam in the gorge will affect the fauna profoundly in two ways. The long, narrow deep lake will be unsuitable for most of the present river fauna. Below the dam, the river will be much cooler than at present as the cold water is released from the bottom of the thermally stratified lake. This will unquestionably materially alter the fauna for a number of miles downstream. It is probable that all of the most interesting and rare elements of the mayfly fauna of the river will become extinct in this section of the river.
The first recorded mayfly collectionsfrom the Green River, where it cuts a gorge through the east end of the Uinta Mountains in Utah, were made by O.A. Peterson in 1908 at "CampDouglas", the campsite of the dinasaur quarry then being worked by personnel of Carnegie Museum. Adults of Traverella albertana and Anepeorus rusticus were reported as collected; no additional specimens of the latter genus have been collected there or at any other locality in the western United States."

Edmunds Jr,GF; Tennessen,KJ 1996 Ephemeroptera. In: An Introduction to the Aquatic Insects of North America. 3rd ed. Eds: Merritt,RW; Cummins,KW Kendall/Hunt Publishing Company, Dubuque, Iowa, 126-163.

Edmunds Jr,GF; Jensen,SL and Berner,L 1976 The Mayflies of North and Central America. University of Minnesota Press, Minneapolis, Minnesota. 330 pages.
     Has a key to adult mayflies, however the taxonomy has changed since then, you may need to consult www.itis.gov frequently. Merritt, Cummins and Berg is better for mayfly larvae.

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.

Encalada,AC and Peckarsky,BL 2006. Selective oviposition of the mayfly Baetis bicaudatus. Oecologia 148:526-537. Abstract


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

Flecker,AS and Allan,JD 1988 Flight direction in some Rocky Mountain mayflies (Ephemeroptera), with observations of parasitism. Aquatic Insects 10(1):33-42. PDF

Flecker,AS; Allan,JD and McClintock,NL 1988 Male body size and mating sucess in swarms of the mayfly Epeorus longimanus. Holarctic Ecology 11 4, 280-285. PDF

Flowers,RW 1980 A review of the Nearctic Heptagenia (Heptageniidae, Ephemeroptera). Pages 93-102 in Flannagan JF; Marshall KE (eds.), 1980, Advances in Ephemeroptera Biology. Plenum Press, New York. PDF

Flowers,RW 1980 Two new genera of Nearctic Heptageniidae (Ephemeroptera). Florida Entomologist 63:296-307.
     Splits the new genera Leucrocuta and Nixe from Heptagenia.

Fuller,RL; Roelofs,JL and Fry,TJ. 1986 The importance of algae to stream invertebrates. Journal of the North American Benthological Society 5(4)290-296. PDF


Gaufin,AR; Clubb,R and Newell,R 1974 Studies on the tolerance of aquatic insects to low oxygen concentrations. Great Basin Naturalist 34:45-59. PDF
     Abstract: "Acute, short-term (96-hour) tests were conducted to, determine the relative sensitivity of low oxygen concentrations to 20 species of aquatic insects. In addition, the longer-term effects of low oxygen levels on the survival, molting, growth, and emergence of 21 species were studied. This paper encompasses work conducted at the University of Montana Biological Station from 1968 to 1970 and at the University of Utah from 1966 to 1972. An evaluation of the average minimum dissolved oxygen requirements of the different groups of aquatic insects tested indicates that the mayflies are the most sensitive, that the stoneflies are next, and that the caddis flies, freshwater shrimp, true flies, and damselflies follow, in that order. While two species of mayfly could tolerate as low a dissolved-oxygen concentration as 3.3 mg/1 for 10 days, a level of 4.6 mg/1 was required for 50-percent survival at 30 days. Fifty percent of the true flies and damselflies tested were able to survive at levels ranging from 2.2 to 2.8 mg/1 for periods ranging from 20 to 92 days."

Gaufin,AR and Hern,S 1971 Laboratory studies on tolerance of aquatic insects to heated waters. Journal of the Kansas Entomological Society 44:240-245. PDF

Gerhardt,A; Bisthoven,LJ de. and Soares,AMVM 2005 Effects of acid mine drainage and acidity on the activity of Choroterpes picteti (Ephemeroptera: Leptophlebiidae). Archives of Environmental Contamination and Toxicology 48:450-458.

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

Gilpin,BR and Brusven,MA 1970 Food habits and ecology of mayflies of the St. Maries River in Idaho. Melanderia 4:19-40. PDF

Glozier,NE; Culp,JM; Scrimgeour,GJ; Halliwell,DB 2000 Comparison of gut fluorescence and gut dry mass techniques for determining feeding periodicity in lotic mayflies. Journal of the North American Benthological Society 19(1):169-175. 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.


Hagen,HA 1861 Synopsis of the Neuroptera of North America with a list of South American species. Smithsonian Miscellaneous Collections 4, 1-344.

Hamilton,H and Clifford,F 1983 The seasonal food habits of mayfly (Ephemeroptera) nymphs from three Alberta, Canada, streams, with special reference to absolute volume and size of particles ingested. Arch. Hydrobiol., Suppl, 65(2/3), 197-234. PDF

Havird,JC; Shah,AA and Chicco,AJ 2020 Powerhouses in the cold: mitochondrial function during thermal acclimation in montane mayflies. Philosophical Transactions of the Royal Society B, 375(1790), p.20190181. PDF

Hawkins,CP 1985 Food habits of species of ephemerellid mayflies (Ephemeroptera: Insecta) in streams of Oregon. American Midland Naturalist 113(2) 343-352. PDF

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

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

Herrmann,J and Andersson,KG 1986 Aluminum impact on respiration of lotic mayflies at low pH. Water, Air and Soil Pollution 30, 703-709.

Hill,WR and Knight, AW 1987 Experimental analysis of the grazing interaction between a mayfly and stream algae. Ecology (68)6:1955-1965.Abstract

Hughes,JM; Mather,PB; Hillyer,MJ; Cleary,C and Peckarsky,B 2003 Genetic structure in a montane mayfly Baetis bicaudatus (Ephemeroptera: Baetidae), from the Rocky Mountains, Colorado. Freshwater Biology 48, 2149-2162.


Irving,EC; Baird,DJ and Culp,JM 2003 Ecotoxicological responses of the mayfly Baetis tricaudatus to dietary and waterborne cadmium: implications for toxicity testing. Environmental Toxicology and Chemistry 22, 1058-1064.


Jacobsen,RE 1995 Symbiotic associations between Chironomidae (Diptera) and Ephemeroptera. Pages 317-332 in Corkum LD; Ciborowski JJH. Current Directions in Research on Ephemeroptera. Canadian Scholars' Press, Inc. Toronto. PDF

Jacobus,LM 2019 Ephemeroptera of Canada. ZooKeys, (819)211-225. HTML

Jacobus,LM and Fleek,ED 2010 Insecta, Ephemeroptera, Ephemerellidae, Attenella margarita (Needham, 1927): Southeastern range extension to North Carolina, USA. Check List 6(2):311-313. PDF

Jacobus,LM and McCafferty,WP 2003 Revisionary contributions to North American Ephemerella and Serratella (Ephemeroptera: Ephemerellidae). Journal of the New York Entomological Society 111:174-193. PDF
     This paper reorganized the taxonomy map for the genus Ephemerella.

Jacobus,LM and McCafferty,WP 2004 Revisionary contributions to the genus Drunella (Ephemeroptera: Ephemerellidae). Journal of the New York Entomological Society 112:127-147. PDF

Jacobus,LM and McCafferty,WP 2006 A new species of Acentrella Bengtsson (Ephemeroptera: Baetidae) from Great Smoky Mountains National Park, USA. Aquatic Insects 28:101-111. PDF
     They describe a new species and provide a key to North American Acentrella species.

Jacobus,LM and McCafferty,WP 2008 Revision of Ephemerellidae genera (Ephemeroptera). Transactions of the American Entomological Society 134: 185-274. PDF

Jensen,PD 2006 Ecological Impact of Selenium and Mercury on two Insect Food Chains. PhD Dissertation, University of California, Riverside, CA, USA.

Jensen,SL 1966 The Mayflies of Idaho (Ephemeroptera). M.S. Thesis, University of Utah, Utah. 364 p.

Jensen,SL and Edmunds Jr,GF 1973 Some phylogenetic relationships within the Heptageniidae. Pages 82-87 in W. L. Peters & J. G. Peters, eds., Proceedings of the First International Conference on Ephemeroptera, E. J. Brill, Leiden.

Johnson,SC 1978 Larvae of Ephemerella inermis and E. infrequens (Ephemeroptera: Ephemerellidae). Pan-Pacific Entomologist 54, 19-25.
     The larvae in our area are difficult to distinguish using the drawings in this paper. Our animals are in between. Further study of mature specimens may work, but don't try to identify young specimens unless you have other information indicating which species it is.


Kerans,BL; Peckarsky BL and Anderson,C 1995 Estimates of mayfly mortality: is stonefly predation a significant source?. Oikos 74(2):315-323. PDF

s Kjӕrstad,G; Webb,JM and Ekrem,T, 2012 A review of the Ephemeroptera of Finnmark–DNA barcodes identify Holarctic relations. Norwegian Journal of Entomology, 59(2), pp.182-195. PDF

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

Kiffney,PM and 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."

Kiffney,PM and Clements,WH 1996 Size-dependent response of macroinvertebrates to metals in experimental streams. Environmental Toxicology and Chemistry 15(8)1352-1356.

Kilgore,JL and Allen,RK 1973 Mayflies of the Southwest: new species, descriptions, and records (Ephemeroptera). Annals of the Entomological Society of America 66(2):321-332. PDF
     Describes the larvae of Choroterpes inornata, Paraleptophlebia memorialis and Tricorythodes minutus among other things. The dorsal view illustrations of these nymphs are wonderful :-)

Kjer,KM 2004 Aligned 18S and insect phylogeny. Systematic biology, 53(3), pp.506-514. PDF
     Abstract: "The nuclear small subunit rRNA (18S) has played a dominant role in the estimation of relationships among insect orders from molecular data. In previous studies, 18S sequences have been aligned by unadjusted automated approaches (computer alignments that are not manually readjusted), most recently with direct optimization (simultaneous alignment and tree building using a program called ”POY”). Parsimony has been the principal optimality criterion. Given the problems associated with the alignment of rRNA, and the recent availability of the doublet model for the analysis of covarying sites using Bayesian MCMC analysis, a different approach is called for in the analysis of these data. In this paper, nucleotide sequence data from the 18S small subunit rRNA gene of insects are aligned manually with reference to secondary structure, and analyzed under Bayesian phylogenetic methods with both GTR+I+G and doublet models in MrBayes. A credible phylogeny of Insecta is recovered that is independent of the morphological data and (unlike many other analyses of 18S in insects) not contradictory to traditional ideas of insect ordinal relationships based on morphology. Hexapoda, including Collembola, are monophyletic. Paraneoptera are the sister taxon to a monophyletic Holometabola but weakly supported. Ephemeroptera are supported as the sister taxon of Neoptera, and this result is interpreted with respect to the evolution of direct sperm transfer and the evolution of flight. Many other relationships are well-supported but several taxa remain problematic, e.g., there is virtually no support for relationships among orthopteroid orders. A website is made available that provides aligned 18S data in formats that include structural symbols and Nexus formats."

Kluge,NJ 2022 Taxonomic significance of microlepides on subimaginal tarsi of Ephemeroptera. Zootaxa, 5159(2), pp.151-186.
     Abstract: "In each mayfly individual, the surface of the subimaginal cuticle significantly differs both from imaginal and larval cuticles, being different on different parts of the body. Most of the subimaginal body, including femora and tibiae, is covered with microtrichia. Tarsal segments are either also covered with microtrichia, or their microtrichia are transformed into microlepides of various shapes. The most usual forms of microlepides are the pointed and the blunt ones. Arrangements of microtrichia, pointed microlepides, blunt microlepides and other forms of microlepides on certain tarsomeres represent good taxonomic characters, which in some cases allow characterization of high level taxa, and in some cases distinguish closely related species. Arrangement of microlepides and microtrichia on subimaginal tarsi of examined mayfly species is given. The following new synonyms are proposed: Afroptilum boettgeri (Kopelke 1980) = Xyrodromeus africanus Lugo-Ortiz McCafferty 1997, syn. n.; Labiobaetis vinosus (Barnard 1932) = L. tenuicrinitus (Kopelke 1980), syn. n.; Baetis parvulus Crass 1947 = Baetis permultus Kopelke 1980, syn. n.; Epeorus gilliesi Braasch 1981 = Epeorus petersi Sivaruban et al. 2013, syn. n.; Dicercomyzon femorale Demoulin 1954 = Dicercomyzon costale Kimmins 1957, syn. n."

Knopp,M and Cormier,R 1997 Mayflies: An anglers study of trout water Ephemeroptera. Lyons Press, Guilford, CT. 366 pages. Buy from Amazon.com
     This book has rather inaccurate illustrations of some larvae. For example the illustration of Caenis amica has the large gill covers drawn on a Baetis-like body. However, once you know what species you're looking at, this book has helpful information on flyfishing and fly patterns for imitating the various life stages of many mayflies.

Komnick,H and Abel,JH Jr. 1971 Location and fine structure of the chloride cells and their porus plates in Callibaetis spec. (Ephemeropteraa, Baetidae). PDF

Komnick,H; Rhees,RW and Abel,JH 1972 The function of ephemerid chloride cells. Histochemical, autoradiographic and physiological studies with radioactive chloride on Callibaetis. Cytobiologie 5:65-82. PDF

Komnick,H and Stockem,W 1973 The porous plates of coniform chloride cells in mayfly larvae: high-resolution analysis and demonstration of solute pathways. Journal of Cell Science 12:665-681. PDF

Koslucher,DG and Minshall,GW 1973 Food habits of some benthic invertebrates in a northern cool-desert stream (Deep Creek, Curlew Valley, Idaho-Utah). Transactions of the American Microscopical Society, 92(3) 441-452. Abstract

Koss RW and Edmunds,GF Jr. 1974 Ephemeroptera eggs and their contribution to phylogenetic studies of the order. Zoological Journal of the Linnean Society 55:267-349, pl. 1-24. PDF


Larkin,JM; HenkMC and Burton,SD 1990 Occurrence of a Thiothrix sp. attached to mayfly larvae and presence of parasitic bacteria in the Thiothrix sp. Applied and Environmental Microbiology 56:357-361. PDF

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

Leach,WE 1815 Entomology. Brewster's Edinburgh Encyclopaedia 9:57-172.
     Describes the genus Baetis for the first time.

Lehmkuhl,DM 1968 Observations on the life histories of four species of Epeorus in western Oregon (Ephemeroptera: Heptageniidae). Pan-Pacific Entomologist 44(2):129-137. PDF

Li,R; Lei,Z; Li,W; Zhang,W and Zhou,C 2021 Comparative mitogenomic analysis of heptageniid mayflies (Insecta: Ephemeroptera): Conserved intergenic spacer and trna gene duplication. Insects, 12(2), p.170.

Liegeois,M; Sartori,M and Schwander,T 2019 Extremely widespread parthenogenesis and a trade-off between alternative forms of reproduction in mayflies (Ephemeroptera). BioRxiv, 841122. PDF

Lugo-Ortiz,CR; McCafferty,WP 1995 Annotated inventory of the mayflies (Ephemeroptera) of Arizona. Entomological News 106(3) 131-140. PDF

Lugo-Ortiz,CR; McCafferty,WP 1996 Contribution to the taxonomy of Callibaetis (Ephemeroptera: Baetidae) in southwestern North America and Middle America. Aquatic Insects 18:1-9.

Lugo-Ortiz,CR; McCafferty,WP 1998 A new North American genus of Baetidae (Ephemeroptera) and key to Baetis complex genera. Entomological News 109 5, 345-353. PDF

Lugo-Ortiz,CR; McCafferty,WP and Waltz,RD 1994 Contribution to the taxonomy of the Panamerican genus Fallceon (Ephemeroptera:Baetidae) Journal of the New York Entomological Society. 102:460-475.


Ma,Z; Li,R; Zhu,B; Zheng,X and Zhou,C 2022 Comparative mitogenome analyses of subgenera and species groups in Epeorus (Ephemeroptera: Heptageniidae). Insects, 13(7), p.599. PDF

Magnum,FA; Winget,RN 1991 Environmental profile of Drunella (Eatonella) doddsi (Needham) (Ephemeroptera: Ephemerellidae). Journal of Freshwater Ecology 6 1, 11-22.
     This is one of a series of papers by scientists in Utah working on stream samples from all over the Western United States.

Magnum,FA; Winget,RN 1993 Environmental profile of Drunella grandis Eaton (Ephemeroptera: Ephemerellidae) in the Western United States. Journal of Freshwater Ecology 8 2, 133-140.

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

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

Maret,TR; Cain,DJ; MacCoy,DE; Short,TM 2003 Response of benthic invertebrate assemblages to metal exposure and bioaccumulation associated with hard-rock mining in northwestern streams, USA. Journal of the North American Benthological Society 22 4, 598-620. Abstract and entire paper

McCafferty,WP 1977 Biosystematics of Dannella and related subgenera of Ephemerella (Ephemeroptera: Ephemerellidae). Annals of the Entomological Society of America, 70(6) 881-889. PDF McCafferty, WP 1983 Aquatic Entomology: The Fishermens Guide and Ecologists Illustrated Guide to Insects and Their Relatives. Jones and Bartlett Publishers, Inc. 480 pages.

McCafferty,WP 1991 Toward a phylogenetic classification of the Ephemeroptera (Insecta): A commentary on systematics. Annals of the Entomological Society of America 84, 343-360.

McCafferty, WP 1992 New larval desciptions and comparisons of North American Chloroterpes (Ephemeroptera: Leptophlebiidae) Great Lakes Entomologist (25)71-78.

McCafferty,WP 1996 The Ephemeroptera species of North America and index to their complete nomenclature. Transactions of American Entomological Society 122 1, 1-54.
     Checklist, very handy for verifying name changes. The website http://www.entm.purdue.edu/entomology/research/mayfly/basis.html continues to update this information.

McCafferty,WP 1996 Emendations to the Callibaetis (Ephemeroptera: Baetidae) of South America. Entomological News 107 4, 230-232.

McCafferty,WP 1997 Name adjustments and a new synonym for North American Ephemeroptera species. Entomological News 108 4, 318, 320.

McCafferty,WP 1998 Ephemeroptera and the Great American Interchange. Journal of the North American Benthological Society 17(1)1-20First page

McCafferty,WP 1998 Additions and corrections to Ephemeroptera species of North America and index to their complete Nomenclature. Entomological News 109 4, 266-268.

McCafferty,WP; Durfee,RS; Kondratieff,BC 1993 Colorado mayflies (Ephemeroptera): an annotated inventory. Southwestern Naturalist 38 (3) 252-274. PDF
     Abstract: "Fourteen families, 41 genera, and 97 species of Ephemeroptera are confirmed for the state of Colorado. Of these, two families, 11 genera (Acerpenna, Apobaetis, Barbaetis, Camelobaetidius, Centroptilum, Heterocloeon, Ironopsis, Lachlania, Macdunnoa, Neochoroterpes, and Pseudiron), and 37 species are reported for the first time. The probability of certain additional genera eventually being found in Colorado is discussed. Previous and new records from 48 of the 63 Colorado counties are provided. The eastern plains region of the state remains relatively poorly known. Colorado species fit one of the following distributional patterns: continental widespread, continental mountain, northern transcontinental, northwestern, Rocky Mountain, southwestern, eastern-midwestern fringe, or western sand/silt riverine. Taxonomic, distributional, and other data that may be pertinent to Colorado Ephemeroptera are given for each species."

McCafferty,WP and Jacobus,LM 2020 Mayfly central. https://www.entm.purdue.edu/mayfly/

McCafferty,WP; MacDonald,JF 1994a New records of Ephemeroptera in Uath, with notes on biogeography. Entomological News 105 4, 217-221.

McCafferty WP; Waltz RD. 1986. Baetis magnus, new species, formal new name for Baetis sp. B of Morihara and McCafferty (Ephemeroptera: Baetidae). Proceedings of the Entomological Society of Washington 88:604.

McCafferty,WP; Waltz,RD 1990 Revisionary synopsis of the Baetidae (Ephemeroptera) of North and Central America. Transactions of American Entomological Society 116, 769-799. PDF
     Abstract: "Considerable nomenclatural revision of the North and Middle American Baetidae has resulted from comprehensive research aimed at formulating a phylogenetic classification. The 17 Nearctic species in Pseudocloeon not previously assigned to Acentrella or Apobaetis are newly placed in Baetis or Barbaetis. Cloeon in the area is restricted to C. cognatum, while two species previously in Cloeon are placed in Centroptilum and nine in Procloeon, a genus considered for the first time in the Nearctic. Definitions of Centroptilum and Procloeon are modified, both now incorporate species with hindwings and species without hindwings, and 19 species are transferred from Centroptilum to Procloeon. Pseudocentroptilum s. auctt. in North America is synonymized with Procloeon. Neocloeon is recognized as a synonym of Centroptilum s. str. and removed from synonymy with Cloeon. Dactylobaetis is placed in synonymy with Camelobaetidius, and all species, including those from South America, are newly combined. Two species of Baetis are transferred to Acerpenna, and two to Fallceon. A checklist includes 154 currently recognized species among 19 genera in the area, and incorporates 21 new species synonyms and 57 new combinations. The names Baetis armillatus and Baetis cinctutus are substituted for Pseudocloeon parvulum and Pseudocloeon cingulatum, respectively, which otherwise attain homonymic status. Regional distributions for each species and abbreviated synonymies since 1976 are annotated to the checklist. Brief discussions of each genus include information on species diagnosis, revisionary bases, status, and needs. A guide to all nomenclatural changes and added taxa since 1976 provides the bibliographic sources of all such information and serves as a ready index to name equivalencies resulting from extensive recent and present revisions."

McCafferty,WP; Wang,T-Q 1994 Phylogenetics and the classification of the Timpanoga complex (Ephemeroptera: Ephemerellidae). Journal of the North American Benthological Society 13 4, 569-579.

McCafferty,WP; Wang,T-Q 2000 Phylogenetic systematics of the major lineages of Pannote mayflies (Ephemeroptera: Pannota). Transactions of American Entomological Society 126 1, 9-101.

McCafferty,WP; Wigle,MJ; Waltz,RD 1994 Systematics and biology of Acentrella turbida (McDunnough) (Ephemeroptera: Baetidae). Pan-Pacific Entomologist 70, 301-308.

McCullough DA; Minshall GW; Cushing CE. 1979 Bioenergetics of a stream "collector" organism, Tricorythodes minutus (Insecta: Ephemeroptera). Limnology and Oceanography 24:45-58.

McDunnough J. 1921 Two new Canadian May-flies (Ephemeridae). Canadian Entomologist 53:117-120.

McDunnough,J 1923 New Canadian Ephemeridae with notes. Canadian Entomologist 55, 39-50.

McDunnough,J 1926 Notes on North American Ephemeroptera with descriptions of new species. Canadian Entomologist. 58:184-196.

McDunnough,J 1928 The Ephemeroptera of Jasper Park, Alta. Canadian Entomologist 60, 8-10.

McDunnough,J 1931 New species of North American Ephemeroptera. Canadian Entomologist 63, 82-93.

McDunnough J. 1933 The nymph of Cinygma integrum and description of a new heptagenine genus. Canadian Entomologist 65:73-76.

McIntosh,AR; Peckarsky,BL; Taylor,BW 2002 The influence of predatory fish on mayfly drift: extrapolating from experiments to nature. Freshwater Biology 47, 1497-1513.

McIntosh,AR; Peckarsky,BL and Taylor,BW 2004. Predator-induced resource heterogeneity in a stream food web. Ecology 85(8) 2279-2290. Abstract

McPeek, MA and Peckarsky,BL. 1998. Life histories and the strengths of species interactions: combining mortality, growth and fecundity effects. Ecology 79(3):867-879. Abstract

Mebane,CA; Dillon,FS and Hennessy,DP 2012 Acute toxicity of cadmium, lead, zinc, and their mixtures to stream-resident fish and invertebrates. Environmental Toxicology and Chemistry, 31(6), 1334-1348. PDF

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. Technical, not for field identifications, you need a microscope for most of the characters used in the dichotomous keys. Not a first bug book or a book for beginners.

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.

Meyer,MD; McCafferty,WP 2001 Hagen's small minnow mayfly (Ephemeroptera: Baetidae) in North America. Entomological News 112 4, 255-263.

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."

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

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

Morihara,DK; McCafferty,WP 1979a The Baetis larvae of North America (Ephemeroptera: Baetidae). Transactions of American Entomological Society 105, 139-221. PDF
Name in this paper 2008 Name
Baetis hageni Diphetor hageni
Baetis insignificans Acentrella insignificans
Baetis amplus Acentrella ampla
Baetis lapponicus Acentrella lapponicus
Baetis pygmaeus Acerpenna pygmaea
Baetis macdunnoughi Acerpenna macdunnoughi
Baetis longipalpus Pseudocloeon longipalpus
Baetis propinguus Pseudocloeon propinguum
Baetis ephippiatus Pseudocloeon ephippiatus
Baetis macani bundyae Baetis bundyae
Baetis quilleri Fallceon quilleri
Baetis sp. A Baetis adonis
Baetis sp. B Baetis magnus
Baetis sp. C Baetis notos

Morihara,DK; McCafferty,WP 1979b Systematics of the propinquus group of Baetis species (Ephemeroptera: Baetidae). Annals of the Entomological Society of America 72, 130-135. PDF


Needham JG. 1905. Ephemeridae. Bulletin of the New York State Museum 86:17-62, pl. 4-12.

Needham JG. 1908 New data concerning May flies and dragon flies of New York. May flies (Ephemeridae). Bulletin of the New York State Museum 1907:188-194. Ephemerella dorothea Description

Needham JG. 1927. The Rocky Mountain species of the mayfly genus Ephemerella. Annals of the Entomological Society of America 20:107-117.

Needham,JG; Traver,JR; Hsu,Y-C 1935 The Biology of Mayflies. Comstock Publishing Company, Inc, Ithaca, New York. 759 pages.

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

Nelson,SM and Roline,RA. 1993 Selection of the mayfly Rhithrogena hageni as an indicator of metal pollution in the Upper Arkansas River. Journal of Freshwater Ecology 8(2):111-119. PDF

Nelson,SM and Roline,RA 1996 Recovery of stream macroinvertebrates community from mine drainage disturbance. Hydrobiologia 339, 73-84.

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
     They studied the Arkansas River above (surface) and 30 cm below the streambed (hyporheic) macroinvertebrate community before and after a water treatment plant was installed to clean up the Leadville Mine Drainage Tunnel effluent in 1992. Having to contend with issues such as the usual annual variation in insect communities they used some fancy math (correspondance analysis) to intepret changes due to cleaner water. They found the water treatment plant cleaned up the surface water the quickest, with the hyporheic zone showing recovery after several years. The hyporheic community fluctuated, with a lower than expected species richness and abundance in 1995 after the water treatment plant was installed. The authors suspect this was because of the large spring runoff in 1995. Substrate (macroinvertebrates prefer more rocks and less sand), discharge, as well as upwelling and downwelling affected community structure too. Metal concentrations in the hyporheic were different from the surface. Quote from page 222: "The continued remobilization of metals into the hyporheic may still be affecting October hyporheic community, even seven years after initiation of water treatment."

Newell,RL and Hossack,BR 2009 Large, wetland-associated mayflies (Ephemeroptera) of Glacier National Park, Montana. Western North American Naturalist, 69(3) 335-342. Abstract and PDF

Newell RL; Minshall GW. 1978 Effect of temperature on the hatching time of Tricorythodes minutus (Ephemeroptera: Tricorythidae). J. Kans. Ent. Soc. 51:504-506.

Newell RT; Minshall GW. 1978. Life history of a multivoltine mayfly, Tricorythodes minutus: an example of the effect of temperature on the life cycle. Annals of the Entomological Society of America 71:876-881.



Peckarsky,BL 1980a Influence of detritus on colonization of stream invertebrates. Canadian Journal of Fisheries and Aquatic Sciences 37, 957-963.

Peckarsky,BL 1980b Predator-prey interactions between stoneflies and mayflies: Behavioral observations. Ecology 61 4, 932-943. PDF

Peckarsky,BL 1983 Biotic interactions or abiotic limitations? A model of lotic community structure. In: Dynamics of Lotic Ecosystems. Eds: Fontaine III,Thomas D; Bartell,Steven M Ann Arbor Science, Ann Arbor, Michigan, 303-323. PDF

Peckarsky,BL 1985 Do predaceous stoneflies and siltation affect the structure of stream insect communities colonizing enclosures? Canadian Journal of Zoology 63, 1519-1530. PDF

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

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

Peckarsky,BL 1988 Why predaceous stoneflies do not aggregate with their prey. Internationale Vereinigung für Theoretische und Angewandte Limnologie Verhandlungen 23, 2135-2140.

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

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

Peckarsky,BL 1991b 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 1996 Alternative predator avoidance syndromes of stream-dwelling mayfly larvae. Ecology 77(6) 1888-1905.Abstract PDF

Peckarsky,BL; Cook,KZ 1981 Effect of Keystone mine effluent on colonization of stream benthos. Environmental Entomology 10, 864-871.

Peckarsky,BL; Cowan,CA 1995 Microhabitat and activity periodicity of predatory stoneflies and their mayfly prey in a western Colorado stream. Oikos 74(3) 513-521. PDF

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

Peckarsky, B.L., C.A. Cowan, M.A. Penton and C. Anderson. 1993 Sublethal consequences of stream-dwelling predatory stoneflies on mayfly growth and fecundity. Ecology 74(6):1836-1846. Abstract PDF

Peckarsky,BL; Dodson,SI 1980a Do stonefly predators influence benthic distributions in streams? Ecology 61 6, 1275-1282. PDF

Peckarsky,BL; Dodson,SI 1980b An experimental analysis of biological factors contributing to stream community structure. Ecology 61 6, 1283-1290. PDF

Peckarsky,BL; Dodson,SI; 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.

Peckarsky,BL, Encalada,AC and McIntosh, AR 2011 Why do vulnerable mayflies thrive in trout streams? American Entomologist 57(3)152-164.

Peckarsky,BL; Fraissinet,PR; Penton,MA; Conklin Jr.,DJ 1990 Freshwater Macroinvertebrates of Northeastern North America. Cornell University, Ithaca, NY. 442 pages.
     While not local, it works pretty well to the genus level, useful as another tool on the shelf when stuck on a bug identification.

Peckarsky,BL; Hughes,JM; Mather,PB; Hillyer,M; Encalada,AC 2005 Are populations of mayflies living in adjacent fish and fishless streams genetically differentiated? Freshwater Biology 50(1), 42-51. doi: 10.1111/j.1365-2427.2004.01292.x Abstract PDF

Peckarsky,BL, Kerans,B; Taylor,BW and McIntosh,AR. 2008 Predator effects on prey population dynamics in open systems. Oecologia.

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

Peckarsky,BL; McIntosh,AR; Caudill,CC; Dahl,J 2002 Swarming and mating behavior of a mayfly Baetis bicaudatus suggest stabilizing selection for male body size. Behavioral Ecology and Sociobiology 51, 530-537. PDF

Peckarsky,BL; McIntosh,AR; Taylor,BW and Dahl,J 2002 Predator chemicals induce changes in mayfly life history traits: a whole stream manipulation. Ecology 83 3, 612-618. PDF

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

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

Peckarsky,BL; Taylor,BW; McIntosh,AR; McPeek,MA and Lytle,DA 2001 Variation in mayfly size at metamorphosis as a developmental response to risk of predation. Ecology 82, 740-757. Abstract

Pennack,RW 1978 Fresh-water Invertebrates of the United States. 2nd ed. Wiley-Interscience, New York, NY. 803 pages.

Pennuto,CM and deNoyelles Jr,F 1993 Behavioral responses of Drunella coloradensis (Ephemeroptera) nymphs to short-term pH reductions. Canadian Journal of Fisheries and Aquatic Sciences 50, 2692-2697.

Perry,SA; Perry,WB and Stanford,JA 1986. Effects of stream regulation on density, growth, and emergence of two mayflies (Ephemeroptera: Ephemerellidae) and a caddisfly (Trichoptera: Hydropsychidae) in two Rocky Mountain rivers (U.S.A.). Canadian Journal of Zoology 64(3):656-666.

Peters,WL 1988. Origins of the North American Ephemeroptera fauna, especially the Leptophlebiidae). Memoirs of the Entomological Society of Canada 144:13-24.

Peters,WL and Edmunds Jr,GF 1961 The mayflies (Ephemeroptera) of the Navajo Reservoir Basin, New Mexico and Colorado. University of Utah Anthropological Papers, No. 55, Upper Colorado Series, No. 5, pp.

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. "

Ploskey,GR and Brown,AV 1980 Downstream drift of the mayfly Baetis flavistriga as a passive phenonmenon. American Midland Naturalist 104:405-409. PDF

Poff,NL and Ward,JV 1988 Use of occupied Glossosoma verdona (Trichoptera: Glossosomatidae) cases by early instars of Baetis spp.(Ephemeroptera: Baetidae) in a Rocky Mountain stream. Entomological news (USA).

Poff,NL and Ward,JV 1991 Drift responses of benthic invertebrates to experimental streamflow variation in a hydrologically stable stream. Canadian Journal of Fisheries and Aquatic Sciences, 48(10): 1926-1936.

Poff,NL; Wellnitz,T; Monroe,JB 2003 Redundancy among three herbivorous insects across an experimental current velocity gradient. Oecologia 134, 262-269. PDF

Pritchard,G; Zloty,J 1994 Life histories of two Ameletus mayflies (Ephemeroptera) in two mountain streams: the influence of temperature, body size, and parasitism. Journal of the North American Benthological Society 13 4, 557-568.

Provonsha,AV 1990 A revision of the genus Caenis in North America (Ephemeroptera: Caenidae). Transactions of American Entomological Society 116, 801-884.

Prusha,BA and Clements,WH 2004 Landscape attributes, dissolved organic C, and metal bioaccumulation in aquatic macroinvertebrates (Arkansas River Basin, Colorado). Journal of the North American Benthological Society 23 2, 327-339.



Raddum,G; Fjellheim,A and Velle,G 1980 Increased growth and distribution of Ephemerella aurivillii (Ephemeroptera) after hydropower regulation of the Aurland catchment in Western Norway. River Research and Applications 24:688-697.

Rader RB. 1997 A functional classification of the drift: traits that influence invertebrate availability to salmonids. Canadian Journal of Fisheries and Aquatic Sciences 54:1211-1234.

Rader RB; Ward JV. 1987 Mayfly production in a Colorado mountain stream: an assessment of methods for synchronous and non-synchronous species. Hydrobiologia 148:145-150.

Rader RB; 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

Rader RB; Ward JV. 1989 Influence of impoundments on mayfly diets, life histories, and production. Journal of the North American Benthological Society 8:64-73.

Rader RB; Ward JV. 1989 The influence of environmental predictability/disturbance characteristics on the structure of a guild of mountain stream insects. Oikos 54:107-116.

Rader RB; Ward JV. 1990 Diel migration and microhabitat distribution of a benthic stream assemblage. Canadian Journal of Fisheries and Aquatic Sciences 47:711-718.

Rader RB; Ward JV. 1990 Mayfly growth and population density in constant and variable temperature regimes. Great Basin Naturalist 50:97-106.

Radford, DS and Hartlan-Rowe,R 1971 The life cycles of some stream insects (Ephemeroptera, Plecoptera) in Alberta. Canadian Entomologist 103: 609-617.

Randolph,RP; McCafferty,WP 1996 First larval descriptions of two species of Paraleptophlebia (Ephemeroptera: Leptophlebiidae). American Midland Naturalist 107 4, 225-229.

Richards C; Minshall GW. 1988. The influence of periphyton abundance on Baetis bicaudatus distribution and colonization in a small stream. Journal of the North American Benthological Society 7(2):77-86.

Riddell DJ, Culp JM; Baird,DJ 2005 Behavioral responses to sublethal cadmium exposure within an experimental aquatic food web. Environmental Toxicology and Chemistry 24: 431-441.

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 identified the macroinvertebrates to genus level and 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.

Rowe,L; Hudson,J and Berrill,M 1988 Hatching success of mayfly eggs at low pH. Canadian Journal of Fisheries and Aquatic Sciences 45:1649-1652.


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."

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

Short,RA 1983 Food habits and dietary overlap among six stream collector species. Freshwater Invertebrate Biology 2:132-138. PDF

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

Sibley,PK; Kaushik, NK and Kreutzweiser,DP 1991 Impact of a pulse application of permethrin on the macroinvertebrate community of a headwater stream Environmental Pollution 70(1)35-55.

Slater,J; Kondratieff,BC 2004 A review of the mayfly genus Cinygmula McDunnough (Ephemeroptera: Heptageniidae) in Colorado. Journal of the Kansas Entomological Society 77 2, 121-126.

Sproul,JS; Houston,DD; Davis,N; Barrington,E; Oh,SY; Evans,RP and Shiozawa,DK 2014 Comparative phylogeography of codistributed aquatic insects in western North America: insights into dispersal and regional patterns of genetic structure. Freshwater Biology, 59(10), pp.2051-2063.

Stanford JA; Gaufin AR. 1974 Hyporheic communities of two Montana rivers. Science 185:700-702.
     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; 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.

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

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

Stitt,RP Rockwell,RW Legg,DE and Lockwood,JA 2006 Evaluation of Cinygmula (Ephemeroptera: Heptageniidae) Drift Behavior as an Indicator of Aqueous Copper Contamination. Journal of the Kentucky Academy of Science 67(2) 102-108. Abstract

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


Taylor,BW; McIntosh,AR and Peckarsky,BL 2002 Reach-scale manipulations show invertebrate grazers depress algal resources in streams. Limnology and Oceanography 47 (3) 893-899.

Torres-Ruiz,M, Wehr,JD; Perrone,AA. 2007 Trophic relations in a stream food web: importance of fatty acids for macroinvertebrate consumers. Journal of the North American Benthological Society 26: 509-522. Abstract

Traver,JR 1935 Two new genera of North American Heptageniidae (Ephemerida). Canadian Entomologist 67:31-38.

Traver,JR 1935 Part II, Systematic. In: The Biology of Mayflies with a Systematic Account of North American Species. Eds: Needham,JG; Traver,JR; Hsu,YC Comstock Publications, Ithaca, New York, 239-739.

Tronstad,LM; Hotaling,S; Giersch,JJ; Wilmot,OJ and Finn,DS 2020 Headwaters fed by subterranean ice: potential climate refugia for mountain stream communities? Western North American Naturalist, 80(3), pp.395-407. PDF


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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

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Vance,SA 1996 The effect of the mermithid parasite Gasteromermis sp. (Nematoda: Mermithidae) on the drift behaviour of its mayfly host, Baetis bicaudatus (Ephemeroptera: Baetidae): a tradeoff between avoiding predators and locating food. Canadian Journal of Zoology 74, 1907-1913.

Vance,SA; Peckarsky,BL 1997 The effect of mermithid parasitism on predation of nymphal Baetis bicaudatus (Ephemeroptera) by invertebrates. Oecologia 110, 147-152.

Vance,SA 1996 Morphological and behavioural sex reversal in mermithid-infected mayflies. Proceedings of the Royal Society of London B 263, 907-912.

Vannote,RL; Sweeney,BW 1980 Geographic analysis of thermal quilibria: a conceptual model for evaluating the effect of natural and modified thermal regimes on aquatic insect communities. The American Naturalist 115 (5), 667-695.


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Walley GS. 1930. Review of Ephemerella nymphs of western North America (Ephemeroptera). Canadian Entomologist 62(1):12-20, pl. 2-3. PDF

Walsh, BD 1862 List of the Pseudoneuroptera of Illinois contained in the cabinet of the writer, with descriptions of over forty new species, with notes on their structural affinities. Proceedings of the Academy of Natural Sciences Philadelphia 14:361-402.

Waltz,RD 1986 Revisionary studies of the mayfly family Baetidae (Ephemeroptera). PhD dissertation, Purdue University 259 pp.

Waltz,RD 1995 Baetis ochris, a new synonym of Baetis flavistriga (Ephemeroptera: Baetidae). Entomological News 106: 75-76.

Waltz,RD; Burian,SK 2008 Ephemeroptera. 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, 181-236.

Waltz,RD and McCafferty,WP 1987a New genera of Baetidae for some Nearctic species previously in Baetis Leach (Ephemeroptera). Annals of the Entomological Society of America 80:667-670.

Waltz,RD and McCafferty,WP 1987b Systematics of Pseudocloeon, Acentrella, Baetiella, and Liebebiella, new genus (Ephemeroptera: Baetidae). Journal of the New York Entomological Society 95:553-568.

Waltz,RD and McCafferty,WP 1997 New generic synonymies in Baetidae (Ephemeroptera). Entomological News 108 2, 134-140.
     This paper synonmizes a number of Baetidae genera revised by Ephemeroptera researchers in Taiwan and Asia.

Wang,T-Q and McCafferty,WP 1995 Relationships of Arthropleidae, Heptageniidae, and Pseudironidae (Ephemeroptera: Heptagenioidea). Entomological News 106 5, 251-256.

Wang,TQ and McCafferty,wp 1996 New diagnostic characters for the mayfly family Baetidae (Ephemeroptera). Entomological News 107 2, 207-212. PDF

Wang,T-Q; McCafferty,WP and Bae,YJ 1997 Sister relationship of the Neoephemeridae and Caenidae (Emphemeroptera: Pannota). Entomological News 108 1, 52-56.

Ward,JV 1986 Altitudinal zonation in a Rocky Mountain stream. Arch. Hydrobiol., Suppl. 74:133-199.

Ward,JV and Berner,L 1980 Abundance and altitudinal distribution of Ephemeroptera in a Rocky Mountain stream. In Advances in Ephemeroptera biology (pp. 169-177). Springer US. PDF

Ward,JV and Stanford,JA 1990 Ephemeroptera of the Gunnison River, Colorado, USA. In: Mayflies and Stoneflies. Ed: Campbell,IC Kluwer Academic Publishers, 215-220. PDF

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.
     This is the bible of aquatic insect ID for the mountains of Colorado. Be careful when you collect insects from streams at lower elevations, you will run into species not covered by this valuable book.

Webb,JM; Jacobus,LM; Funk,DH; Zhou,X; Kondratieff,BC; Geraci,CJ; DeWalt,RE Baird,DJ Richard,B Philips,I and Hebert,PDN 2012 A DNA barcode library for North American Ephemeroptera: Progress and prospects. PloS One 7(5): e38063 HTML

Webb JM and McCafferty,WP 2006 Contribution to the taxonomy of Eastern North American Epeorus Eaton (Ephemeroptera: Heptageniidae). Zootaxa 1128:57-64.

Webb,JM and McCafferty,WP 2008 Heptageniidae of the world. Part II. Key to the genera. Canadian Journal of Arthropod Identification, 7(10.37551). PDF
     Abstract: "Keys and diagnoses illustrated with line drawings and colour photographs for the identification of larvae and adult males of the genera of Heptageniidae of the world and female adults of North American Heptageniidae genera are provided. Siberionurus McCafferty is recognized as a junior objective synonym of Ecdyogymnurus Kluge. Epeiron Demoulin is shown to be congeneric with Rhithrogena Eaton. All subgenera that have been proposed for Rhithrogena, Compsoneuria Eaton, and Epeorus Eaton are treated as junior synonyms (Rhithrogena = Himalogena Kluge, N.SYN.; = Sibirigena Kluge, N.SYN.; = Tumungula Zhou & Peters, N.SYN.; Epeorus = Alpiron Braasch, N.SYN.; = Albertiron Kluge, N.SYN.; = Belovius Tshernova; = Caucasiron Kluge, N.SYN.; = Iron Eaton; = Ironopsis Traver; = Proepeorus Kluge, N.SYN.; Compsoneuria = Siamoneuria Braasch, N.SYN.)"

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

Wellnitz,TA; Poff,NL; Cosyleón,G and Steury,B 2001 Current velocity and spatial scale as determinants of the distribution and abundance of two rheophilic herbivorous insects. Landscape Ecology, 16(2), 111-120. PDF

Wellnitz,T and Rader,RB 2003 Mechanisms influencing community composition and succession in mountain stream periphyton; interactions between scouring history, grazing and irradiance. Journal of the North American Benthological Society 22 4, 528-541. Abstract and full text

White,MM and Licthwardt,RW 2004 Fungal symbionts (Harpellales) in Norwegian aquatic insect larvae. Mycologia, 96(4):891-910.

Wiens AP; Rosenberg DM and Evans KW 1975 Symbiocladius equitans (Diptera: Chironomidae), an ectoparasite of Ephemeroptera in the Martin River, Northwest Territories, Canada. Entomologica Germanica 2(2):113-120 PDF

Wiersema,NA and McCafferty,WP 2000 Generic revision of the North and Central American Leptohyphidae (Ephemeroptera: Pannota). Transactions of American Entomological Society 126 3+4, 337-371.
     Discusses the subfamilies Tricorythodinae and Leptohyphinae. Has keys and illustrations to distinguish larvae and adult genera of the Leptohyphidae family.

Wiersema,NA and McCafferty,WP 2004 New specific synonyms and records of North American Centroptilum and Procloeon (Ephemeroptera: Baetidae). Entomological News 115(3):121-128. PDF

Wiersema,NA; Nelson,CR and Kuehnl,KF 2004 A new small minnow mayfly (Ephemeroptera: Baetidae) from Utah, USA. Entomological News 115 3, 139-145.

Wilcox,AC; Peckarsky,BL; Taylor,BW; and Encalada,AC 2008 Hydraulic and geomorphic effects on mayfly drift in high-gradient streams at moderate discharges. Ecohydrology, 1(2), 176-186.

Williams,MC and Lichtwardt,RW 1999. Two new Harpellales living in Ephemeroptera nymphs in Colorado Rocky Mountain streams. Mycologia 91:400-404.

Winget,RN 1993 Habitat partitioning among three species of Ephemerelloidea. Journal of Freshwater Ecology 8 3, 227-234.
     Abstract: "Niche volumes according to several physical and water quality parameters are shown for Drunella grandis, Drunella doddsi and Tricorythodes minutus. Generalist strategies have been adopted by D. grandis for physical habitat characteristics and by T. minutus for water quality characteristics. Drunella doddsi, a specialist, has the smallest physical habitat and water quality niche dimensions of the three mayfly species. Even though each has its own unique niche, there is some overlap between them for most niche dimensions studied. These overlaps are probably present because density independent seasonal limiting factors in western United States streams are as strong or stronger selectors than many density dependent factors. Also, each species has its own unique niche "hypervolume" that changes from station to station and from time to time. "

Winget,RN and Mangum,FA 1991 Environmental profile of Traver (Ephemeroptera: Tricorythidae) in the Western United States. Journal of Freshwater Ecology 6 3, 335-344.

Winget,RN and Mangum,FA 1996 Environmental profile of Drunella coloradensis Dodds (Ephemeroptera: Ephemerellidae) in the Western United States. Journal of Freshwater Ecology 11 2, 225-232.

Wipfli,MS; Hudson,J and Caouette,J 1998 Influence of salmon carcasses on stream productivity: response of biofilm and benthic macroinvertebrates in southeastern Alaska, U.S.A. Can. J. Fish. Aquat. Sci. 55(6): 1503-1511 Abstract




Zhou,X; Adamowicz,SJ; Jacobus,LM; DeWalt,RE and Hebert,PD 2009 Towards a comprehensive barcode library for arctic life-Ephemeroptera, Plecoptera, and Trichoptera of Churchill, Manitoba, Canada. Frontiers in zoology, 6(1), p.30. HTML

Zitnan,D; Zitnanová,I; Spalovská,I; Takác,P; Park,Y and Adams,ME 2003 Conservation of ecdysis-triggering hormone signalling in insects. The Journal of Experimental Biology 206, 1275-1289. Full Text
     They studied Epeorus sp. and Heptagenia sp. among other insects.

Zloty,J 1996 A revision of nearctic Ameletus mayflies based on adult males, with descriptions of seven new species (Ephemeroptera: Ameletidae). Canadian Entomologist 128 (2) 293-346. Abstract PDF

Zloty,JS 1996 Systematics of Nearctic Ameletus mayflies (Ephemeroptera: Ameletidae). Ph.D. dissertation, University of Calgary, Alberta.

Zloty,J and Pritchard,G 1997 Larvae and adults of Ameletus mayflies (Ephemeroptera: Ameletidae) from Alberta. Canadian entomologist, 129(2), 251-290. PDF

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; Kashian,DR; Brooks,ML; Kiffney,PM and Clements,WH 2008 The influence of metal exposure history and ultraviolet-B radiation on benthic communities in Colorado Rocky Mountain streams. Journal of the North American Benthological Society, 27(1), 120-134. PDF

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

Brown, Wendy S. 2004 Extended reading for those obsessed with the Mayflies of central Colorado

"Ever drifting down the stream--
Lingering in the golden gleam--
Life, what is it but a dream?"

           Lewis Carroll