Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2018-03-12
Page range: 105–127
Abstract views: 197
PDF downloaded: 9

The state of systematics of North American Baetis Leach, 1815 (Ephemeroptera: Baetidae), with recommendations for identification of larvae

Rhithron Associates, Inc. 33 Fort Missoula Road, Missoula, MT 59804, USA.
Division of Science, Indiana University Purdue University Columbus, 4601 Central Avenue, Columbus, IN 47203, USA
Rhithron Associates, Inc. 33 Fort Missoula Road, Missoula, MT 59804, USA.
Ephemeroptera mayfly standard taxonomic effort biomonitoring larval key


The North American species of Baetis Leach (Ephemeroptera: Baetidae) are reviewed. Nearly one-third of species are either unknown or inadequately described in the larval stage, a fact not reflected in most keys or standard taxonomic efforts for bioassessment, which typically recommend species-level identifications of larvae. Furthermore, our new observations indicate that some previously published stage associations should be viewed as only tentative, and molecular evidence suggests that current species taxonomy does not reflect biological species. In order to acknowledge these deficiencies, but at the same time provide a degree of higher taxonomic resolution beyond the genus level, we recommend a scheme for identifications incorporating previously established species groups and the species complexes and species included within them. Species complexes are proposed for instances when there are either multiple species that cannot be differentiated in the larval stage or when multiple lines of evidence indicate more than one actual species is included in a single species concept. Complexes include B. flavistriga complex (B. flavistriga McDunnough + B. phoebus McDunnough + B. rusticans McDunnough), B. intercalaris complex (B. intercalaris McDunnough), B. vernus complex (B. brunneicolor McDunnough + B. vernus Curtis), B. bicaudatus complex (B. bicaudatus Dodds), B. tricaudatus complex (B. tricaudatus Dodds), and B. piscatoris complex (B. piscatoris Traver + B. palisadi Mayo + B. persecutus McDunnough [=B. persecutor McCafferty n. obj. syn]). A new larval identification key incorporating the B. piscatoris complex is provided.



  1. Adler, P.H., Currie, D.C. & Wood, D.M. (2004) The Black Flies (Simuliidae) of North America. Cornell University Press, Ithaca, 960 pp.

    Adler, P.H. & Kim, C.K. (1986) The black flies (Simuliidae, Diptera) of Pennsylvania: Bionomics, taxonomy, and distribution. Pennsylvania State University Agricultural Experiment Station Bulletin, 856, 1–88.

    Allen, R.K. & Murvosh, C.M. (1983) Taxonomy and zoogeography of the mayflies (Ephemeroptera: Insecta) of Baja California. Annals of the Entomological Society of America, 76, 425–433.

    Allen, R.K. & Murvosh, C.M. (1987) New Baetidae from the southwestern United States and northern Mexico (Ephemeroptera: Insecta), with notes. Canadian Entomologist, 119, 1095–1099.

    Banks, N. (1924) Descriptions of new neuropteroid insects. Bulletin of the Museum of Comparative Zoology, 65, 419–455.

    Barbour, M.T., Gerritsen, J., Snyder, B.D. & Stribling, J. (1999) Rapid Bioassessment Protocols for Use in Streams and Wadeable Rivers: Periphyton, Benthic Macroinvertebrates and Fish. 2nd Edition. EPA 841-B-99-002. Washington, D.C., 337 pp. Available from: (accessed 28 February 2018)

    Barnard, K.H. (1932) South African may-flies (Ephemeroptera). Transactions of the Royal Society of South Africa, 20, 201–259.

    Bauernfeind, E. & Soldán, T. (2012) The Mayflies of Europe (Ephemeroptera). Brill, Ollerup, 779 pp.

    Bergman, E.A. & Hilsenhoff, W.L. (1978) Baetis (Ephemeroptera: Baetidae) of Wisconsin. Great Lakes Entomologist, 11, 125–135.

    Bisconti, R., Canestrelli, D., Tenchini, R., Belfiore, C., Buffagni, A. & Nascetti, G. (2016) Cryptic diversity and multiple origins of the widespread mayfly species group Baetis rhodani (Ephemeroptera: Baetidae) on northwestern Mediterranean islands. Ecology and Evolution, 6, 7901–7910.

    Burks, B.D. (1953) The mayflies, or Ephemeroptera, of Illinois. Bulletin of the Illinois Natural History Survey, 26, 1–216.

    Cordero, R.D., Sánchez-Ramírez, S. & Currie, D.C. (2017) DNA barcoding of aquatic insects reveals unforeseen diversity and recurrent population divergence patterns through broad-scale sampling in northern Canada. Polar Biology, 40, 1687–1695.

    Curtis, J. (1834) Description of some hitherto nondescript British species of mayflies of anglers. The London and Edinburgh Philosophical Magazine and Journal of Science, 3 (4), 120–218.

    Day, W.C. (1954) New species of California mayflies in the genus Baetis (Ephemeroptera). Pan-Pacific Entomologist, 30, 29–34.

    Dodds, G.S. (1923) Mayflies from Colorado. Descriptions of certain species and notes on others. Transactions of the American Entomological Society, 49, 93–114.

    Durfee, R.S. & Kondratieff, B.C. (1993) Description of adults of Baetis magnus (Ephemeroptera: Baetidae). Entomological News, 104, 227–232.

    Durfee, R.S. & Kondratieff, B.C. (1999) Notes on North American Baetis (Ephemeroptera: Baetidae): Baetis moffatti new synonym of B. tricaudatus and range extension for B. bundyae. Entomological News, 110, 177–180.

    Eaton, A.E. (1870) On Some new British species of Ephemeridae. Transactions of the Royal Entomological Society of London, 18, 1–8.

    Eaton, A.E. (1871) A monograph on the Ephemeridae. Transactions of the Entomological Society of London, 1871, 1–164.

    Edmunds, G.F. & Allen, R.K. (1957) A Checklist of the Ephemeroptera of North America North of Mexico. Annals of the Entomological Society of America, 50, 317–324.

    Edmunds Junior, G.F. & Allen, R.K. (1957) A new species of Baetis from Oregon (Ephemeroptera: Baetidae). Journal of the Kansas Entomological Society, 30, 57–58.

    Epler, J.H. (2001) Identification manual for the larval Chironomidae (Diptera) of North and South Carolina. A guide to the taxonomy of the midges of the southeastern United States, including Florida. Special Publication SJ2001-SP13 North Carolina Department of Environmental and Natural Resources , and St. John’s River Water Management District, Palatka, FL, 526 pp.

    Epler, J.H. (2014) Identification Guide to the Larvae of the Tribe Tanytarsini (Diptera: Chironomidae) in Florida. State of Florida Department of Environmental Protection Division of Environmental Assessment and Restoration, Tallahassee, 81 pp. Available from: (accessed 23 February 2018)

    Funk, D.H., Sweeney, B.W. & Jackson, J.K. (2008) A taxonomic reassessment of the Drunella lata (Morgan) species complex (Ephemeroptera:Ephemerellidae) in northeastern North America. Journal of the North American Benthological Society, 27, 647–663.

    Gattolliat, J.-L. & Nieto, C. (2009) The family Baetidae (Insecta: Ephemeroptera): synthesis and future challenges. Aquatic Insects, 31, 41–62.

    Gehara, M., Crawford, A.J., Orrico, V.G.D., Rodríguez, A., Lötters, S., Fouquet, A., Barrientos, L.S., Brusquetti, F., la Riva, I., Ernst, R., Urrutia, G.G., Glaw, F., Guayasamin, J.M., Hölting, M., Jansen, M., Kok, P.J.R., Kwet, A., Lingnau, R., Lyra, M., Moravec, J., Pombal Jr, J.P., Rojas-Runjaic, F.J.M., Schulze, A., Señaris, J.C., Solé, M., Rodrigues, M.T., Twomey, E., Haddad, C.F.B., Vences, M. & Köhler, J. (2014) High levels of diversity uncovered in a widespread nominal taxon: Continental phylogeography of the neotropical tree frog Dendropsophus minutus. PLoS ONE, 9, 1–12.

    Hamilton, H.R. & Clifford, H.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. Archiv für Hydrobiologie Supplementum, 65, 197–234.

    Harker, J.E. (1954) The Ephemeroptera of Eastern Australia. Transactions of the Royal Entomological Society of London, 105, 241–268.

    Hawkins, C.P., Norris, R.H., Hogue, J.N. & Feminella, J.W. (2000) Development and evaluation of predictive models for measuring the biological integrity of streams. Ecological Applications, 10, 1456–1477.[1456:DAEOPM]2.0.CO;2

    Hilsenhoff, W.L. (1977) Use of Arthropods to evaluate water quality of streams. Wisconsin Department of Natural Resources Technical Bulletin, 100, 1–15.

    Hilsenhoff, W.L. (1987) An improved biotic index of organic stream pollution. Great Lakes Entomologist, 20, 31–19.

    Hubbard, M.D. (1974) Some specific homonyms in mayflies (Ephemeroptera). Journal of the Kansas Entomological Society, 47, 357–359.

    Ide, F.P. (1937) Descriptions of eastern North American species of baetine mayflies with particular reference to the nymphal stages. Canadian Entomologist, 69, 219–231, 235–243.

    Jackson, J.K., Battle, J.M., White, B.P., Pilgrim, E.M., Eric, D., Miller, P.E., Sweeney, B.W. & Stein, E.D. (2014) Cryptic biodiversity in streams: a comparison of macroinvertebrate communities based on morphological and DNA barcode identifications. Freshwater Science, 33, 312–324.

    Jacob, U. (2003) Baetis Leach 1815, sensu stricto oder sensu lato. Ein Beitrag zum Gattungskonzept auf der Grundlage von Artengruppen mit Bestimmungsschlüsseln. Lauterbornia, 47, 59–129.

    Jacobus, L.M. (2010) Taxonomic review of the Caudatella heterocaudata (McDunnough) and C. hystrix (Traver) complexes (Insecta: Ephemeroptera: Ephemerellidae). Psyche, 2010, 1–5.

    Jacobus, L.M., Wiersema, N.A. & Webb, J.M. (2014) Identification of Far Northern and Western North American Mayfly Larvae (Insecta: Ephemeroptera), North of Mexico. In: Unpublished workshop manual, Joint Aquatic Science meeting. Portland, Oregon, pp. 176.

    Kazlauskas, R.S. (1972) Neues uber das System der Eintagsfliegen der Familie Baetidae (Ephemeroptera). In: Proceedings of the 13th International Congress of Entomology, Moscow, 1972, pp. 337–338.

    Kerans, B.L. & Karr, J.R. (1994) A Benthic Index of Biotic Integrity (B-IBI) for rivers of the Tennessee Valley. Ecological Applications, 4, 768–785.

    Klubertanz, T.H. (2016) Mayfly Larvae of Wisonsin. University of Wisconsin-Extension, Madison, WI.

    Kluge, N.J. & Novikova, E.A. (2016) New tribe Labiobaetini tribus n., redefinition of Pseudopannota Waltz & McCafferty 1987 and descriptions of new and little known species from Zambia and Uganda. Zootaxa, 4169 (1), 1–43.

    Koryak, M., Stafford, L.J. & Sykora, J.L. (2003) Urban drainage aquatic invertebrate signature species in Allegheny County, Pennsylvania: Gammarus minus (Crustaces, Amphipoda), and Baetis flavistriga (Insecta, Ephemeroptera). Journal of the Pennsylvania Academy of Science, 76, 86–87.

    Kwon-Chung, K.J., Bennett, J.E., Wickes, B.L., Meyer, W., Cuomo, C.A., Wollenburg, K.R., Bicanic, T.A., Castañeda, E., Chang, Y.C., Chen, J., Cogliati, M., Dromer, F., Ellis, D., Filler, S.G., Fisher, M.C., Harrison, T.S., Holland, S.M., Kohno, S., Kronstad, J.W., Lazera, M., Levitz, S.M., Lionakis, M.S., May, R.C., Ngamskulrongroj, P., Pappas, P.G., Perfect, J.R., Rickerts, V., Sorrell, T.C., Walsh, T.J., Williamson, P.R., Xu, J., Zelazny, A.M. & Casadevall, A. (2017) The case for adopting the “Species Complex” nomenclature for the etiologic agents of Cryptococcosis. mSphere, 2, e00357-16.

    Lager, T.M., Johnson, M.D. & McCafferty, W.P. (1982) The mayflies of northeastern Minnesota (Ephemeroptera). Proceedings of the Entomological Society of Washington, 84, 729–741.

    Leach, W.E. (1815) Entomology. Brewster’s Edinburgh Encyclopedia, 9, 52–172.

    Lehmkuhl, D.M. (1973) A new species of Baetis (Ephemeroptera) from ponds in the Canadian Arctic, with biological notes. The Canadian Entomologist, 105, 343–346.

    Leonard, J.W. (1950) A new Baetis from Michigan. Annals of the Entomological Society of America, 43, 155–159.

    Lester, G.T., McCafferty, W.P. & Edmondson, M.R. (2002) New mayfly (Ephemeroptera) records from Idaho. Entomological News, 113, 131–136.

    Lucentini, L., Rebora, M., Puletti, M.E., Gigliarelli, L., Fontaneto, D., Gaino, E. & Panara, F. (2011) Geographical and seasonal evidence of cryptic diversity in the Baetis rhodani complex (Ephemeroptera, Baetidae) revealed by means of DNA taxonomy. Hydrobiologia, 673, 215–228.

    Mayo, V.K. (1952) New western Ephemeroptera, III. Pan-Pacific Entomologist, 23, 93–103.

    McCafferty, W.P. (1997) Name adjustments and a new synonym for North American Ephemeroptera species. Entomological News, 108, 318, 320.

    McCafferty, W.P. (2007) Two new synonyms in North American Baetis Leach (Ephemeroptera: Baetidae). Proceedings of the Entomological Society of Washington, 109, 735–736.

    McCafferty, W.P. & Jacobus, L.M. (2017) Mayfly Central. Available from: (accessed 12 July 2017)

    McCafferty, W.P., Jacobus, L.M., Provonsha, A.V. & Wiersema, N.A. (2017) Ephemeroptera. In: Morse, J.C., McCafferty, W.P., Stark, B.P. & Jacobus, L.M. (Eds.), Larvae of the Southeastern USA Mayfly, Stonefly, and Caddisfly Species. Clemson University Public Service Publishing, Clemson, South Carolina, pp. 15–160.

    McCafferty, W.P., Lenat, D.R., Jacobus, L.M. & Meyer, M.D. (2010) The Mayflies (Ephemeroptera) of the Southeastern United States. Transactions of the American Entomological Society, 136, 221–233.

    McCafferty, W.P. & Lugo-Ortiz, C.R. (1996) Los efemerópteros (Ephemeroptera) de América Central. Revista Nicaraguense de Entomologia, 35, 19–28.

    McCafferty, W.P. & Meyer, M.D. (2007) Insecta, Ephemeroptera: transcontinental range extensions in western North America. Check List, 3, 51–54.

    McCafferty, W.P., Meyer, M.D., Randolph, R.P. & Webb, J.M. (2008) Evaluation of mayfly species originally described as Baetis Leach (Ephemeroptera: Baetidae) from California. Proceedings of the Entomological Society of Washington, 110, 577–591.

    McCafferty, W.P. & Pereira, C. (1984) Effects of developmental thermal regimes on two mayfly species and their taxonomic interpretation. Annals of the Entomological Society of America, 77, 69–87.

    McCafferty, W.P. & Randolph, R.P. (1998) Canada mayflies: a faunistic compendium. Proceedings of the Entomological Society of Ontario, 129, 47–97.

    McCafferty, W.P., Randolph, R.P. & Jacobus, L.M. (2012) Mayflies of the Intermountain West. American Entomological Institute, Gainesville, FL, 317 pp.

    McCafferty, W.P. & Silldorff, E.L. (1998) Reared association and equivalency of Baetis adonis and B. caelestis (Ephemeroptera: Baetidae). Entomological News, 109, 261–265.

    McCafferty, W.P. & Waltz, R.D. (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, W.P. & Waltz, R.D. (1990) Revisionary synopsis of the Baetidae (Ephemeroptera) of North and Middle America. Transactions of the American Entomological Society, 116, 769–799.

    McDunnough, J. (1923) New Canadian Ephemeridae with notes. The Canadian Entomologist, 55, 39–50.

    McDunnough, J. (1924) New Ephemeridae from Illinois. The Canadian Entomologist, 56, 7–9.

    McDunnough, J. (1925) New Canadian Ephemeridae with notes. III. The Canadian Entomologist, 57, 168–176, 185–192.

    McDunnough, J. (1921) Two new Canadian mayflies (Ephemeridae). The Canadian Entomologist, 53, 117–120.

    McDunnough, J. (1925) The Ephemeroptera of Covey Hill, Que. Transaction of the Royal Society of Canada, 19, 207–223.

    McDunnough, J. (1936) A new Arctic baetid (Ephemeroptera). The Canadian Entomologist, 68, 33–34.

    McDunnough, J. (1938) New species of North American Ephemeroptera with critical notes. Canadian Entomologist, 70, 23–34.

    McDunnough, J. (1939) New British Columbia Ephemeroptera. The Canadian Entomologist, 71, 49–54.

    Meyer, M.D. & McCafferty, W.P. (2007a) Mayflies (Ephemeroptera) of the far western United States. Part 2. Oregon. Transactions of the American Entomological Society, 133, 65–114.[65:MEOTFW]2.0.CO;2

    Meyer, M.D. & McCafferty, W.P. (2007b) Mayflies (Ephemeroptera) of the far western United States. Part I. Washington. Transactions of the American Entomological Society, 133, 21–63.[21:MEOTFW]2.0.CO;2

    Meyer, M.D. & McCafferty, W.P. (2008) Mayflies (Ephemeroptera) of the Far Western United States. Part 3. California. Transactions of the American Entomological Society, 134, 337–430.

    Monaghan, M.T., Gattolliat, J.-L., Sartori, M., Elouard, J.-M., James, H., Derleth, P., Glaizot, O., de Moor, F. & Vogler, A.P. (2005) Trans-oceanic and endemic origins of the small minnow mayflies (Ephemeroptera, Baetidae) of Madagascar. Proceedings of the Royal Society B: Biological Sciences, 272, 1829–1836.

    Morihara, D.K. & McCafferty, W.P. (1979) The Baetis larvae of North America (Ephemeroptera: Baetidae). Transactions of the American Entomological Society, 105, 139–221.

    Morinière, J., Hendrich, L., Balke, M., Beermann, A.J., König, T., Hess, M., Koch, S., Müller, R., Leese, F., Hebert, P.D.N., Hausmann, A., Schubart, C.D. & Haszprunar, G. (2017) A DNA barcode library for Germany′s mayflies, stoneflies and caddisflies (Ephemeroptera, Plecoptera and Trichoptera). Molecular Ecology Resources, 17 (6), pp. 1293–1307.

    Morley, S.A. & Karr, J.R. (2002) Assessing and restoring the health of urban streams in the Puget Sound Basin. Conservation Biology, 16, 1498–1509.

    Moulton, S.R.I., Carter, J.L., Grotheer, S.A., Cuffney, T.F. & Short, T.M. (2000) 00–212 U.S. Geological Survey Open-File Report Methods of analysis by the U.S. Geological Survey National Water Quality Laboratory-processing, taxonomy, and quality control of benthic macroinvertebrate samples. Cambridge University Press, Denver, CO, 1535 pp. Available from: (Accessed 12 Mar. 2018)

    Muller-Liebenau, I. (1969) Revision der europäischen Arten der Gattung Baetis Leach, 1815 (Insecta, Ephemeroptera). Gewasser und Abwasser, 49, 1–214.

    Myers, L.W., Jacobus, L.M. & Kondratieff, B.C. (2008) Insecta, Ephemeroptera: new and additional records from New York (U.S.A.). Check List, 4, 415–423.

    Navás, L. (1924) Insectos de la América Central. Brotéria, Série Zoológica, 21, 55–86.

    NC Department of Environmental Quality (2016) Standard Operating Procedures for Collection and Analysis of Benthic Macroinvertebrates. Raleigh, NC. Available from: (accessed 28 February 2018)

    Novikova, E.A. & Kluge, N.J. (1987) Systematics of the genus Baetis (Ephemeroptera: Baetidae) with description of a new species from Middle Asia. Vestnik Zoologii, 4, 8–19.

    Peckarsky, B.L., Hughes, J.M., Mather, P.B., Hillyer, M.J. & Encalada, A.C. (2005) Are populations of mayflies living in adjacent fish and fishless streams genetically differentiated? Freshwater Biology, 50, 42–51.

    Pereira-da-Conceicoa, L.L., Price, B.W., Barber-James, H.M., Barker, N.P., de Moor, F.C. & Villet, M.H. (2012) Cryptic variation in an ecological indicator organism: mitochondrial and nuclear DNA sequence data confirm distinct lineages of Baetis harrisoni Barnard (Ephemeroptera: Baetidae) in southern Africa. BMC Evolutionary Biology, 12, 26.

    Plotnikoff, R.W. & White, J.S. (1996) 96–323 Taxonomic Laboratory Protocol for Stream Macroinvertebrates Collected by the Washington State Department of Ecology. Washington State Department of Ecology, Olympia, WA, 32 pp.

    Plotnikoff, R.W. & Wiseman, C. (2001) 1-03–028 Benthic Macroinvertebrate Biological Monitoring Protocols for Rivers and Streams 2001 Revision. Washington State Department of Ecology, Olympia, WA. Available from: (accessed 28 February 2018)

    Poteat, M.D., Jacobus, L.M. & Buchwalter, D.B. (2015) The importance of retaining a phylogenetic perspective in traits-based community analyses. Freshwater Biology, 60, 1330–1339.

    Randolph, R.P. & McCafferty, W.P. (2005) The mayflies (Ephemeroptera) of Alaska, including a new species of Heptageniidae. Proceedings of the Entomological Society of Washington, 107, 190–199.

    Ratnasingham, S. & Hebert, P.D.N. (2013) A DNA-based registry for all animal species: the barcode index number (BIN) system. PloS one, 8, e66213.

    Richards, A.B. & Rogers, D.C. (2011) List of Freshwater Macroinvertebrate Taxa from California and Adjacent States including Standard Taxonomic Effort Levels. Southwest Association of Freshwater Invertebrate Taxonomists (SAFIT), 2011, 1–284.

    Savolainen, E., Drotz, M.K., Per-Ola, H. & Saura, A. (2007) The Baetis vernus group (Ephemeroptera: Baetidae) of northernmost Europe: an evidently diverse but poorly understood group of mayflies. Entomologica Fennica, 6, 160–167.

    Smith, A. (2014) New York State Department of Environmental Conservation, Division of Water, Standard Operating Procedure: Biological Monitoring of Surface Waters in New York State. Available from: (accessed 28 February 2018)

    Spitzer, B. (2014) Genetic structure in populations of Baetis tricaudatus in the San Bernardino Mountains. Western North American Naturalist, 74, 434–445.

    Sroka, P. (2012) Systematics and phylogeny of the West Palaearctic representatives of subfamily Baetinae (Insecta: Ephemeroptera): combined analysis of mitochondrial DNA sequences and morphology. Aquatic Insects, 34, 23–53.

    Sroka, P., Godunko, R.J., Novikova, E.A. & Kluge, N.J. (2012) Contribution to the knowledge of the subgenus Rhodobaetis Jacob, 2003 (Ephemeroptera: Baetidae: Baetis) from Central Asia. Part 1. Zootaxa, 3311, 42–60.

    Ståhls, G. & Savolainen, E. (2008) MtDNA COI barcodes reveal cryptic diversity in the Baetis vernus group (Ephemeroptera, Baetidae). Molecular Phylogenetics and Evolution, 46, 82–87.

    Stauffer-Olsen, N.J., O’Grady, P.M. & Resh, V.H. (2017) Temporal patterns of genetic diversity in Baetis tricaudatus (Ephemeroptera:Baetidae) from the Russian River, Northern California. Freshwater Science, 36, 351–363.

    Stephens, J.F. (1836) Family III.—Ephemeridae, Leach. Illustrations of British Entomology, Mandibulata, 6, 54–70.

    Stoddard, J.L., Herlihy, A.T., Peck, D. V, Hughes, R.M., Whittier, T.R. & Tarquinio, E. (2008) A process for creating multimetric indices for large-scale aquatic surveys. Journal of the North American Benthological Society, 27, 878–891.

    Sweeney, B.W., Battle, J.M., Jackson, J.K. & Dapkey, T. (2011) Can DNA barcodes of stream macroinvertebrates improve descriptions of community structure and water quality? Journal of the North American Benthological Society, 30, 195–216.

    Traver, J.R. (1935) Systematics. In: Needham, J.G., Traver, J.R. & Hsu, Y. (Eds.), The Biology of Mayflies with an Account of the North American Species. Comstock, Ithaca, New York, pp. 239–739.

    U.S. Geological Survey (2017) BioData - Aquatic Bioassessment Data for the Nation available on the World Wide Web. Available from:
    LOGICAL+CLASSIFICATION%7CPLANTS&OrigMetadataNode=GCMD&EntryId=USGS_BioData&MetadataView=gwt&MetadataType=0&lbnode=mdlb1 (accessed 28 February 2018)

    Waite, I.R., Herlihy, A.T., Larsen, D.P., Urquhart, N.S. & Klemm, D.J. (2004) The effects of macroinvertebrate taxonomic resolution in large landscape bioassessments: an example from the Mid-Atlantic Highlands, U.S.A. Freshwater Biology, 49, 474–489.

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

    Waltz, R.D., Baumgardner, D.E. & Kennedy, J.H. (1996) An atypical larval color form of Baetis intercalaris (Ephemeroptera: Baetidae) from Pennsylvania and the Kiamichi River basin of southeastern Oklahoma. Entomological News, 107, 83–87.

    Waltz, R.D. & Burian, S.K. (2008) Ephemeroptera. In: Merritt, R.W., Cummins, K.W. & Berg, M.B. (Eds.), An Introduction to the Aquatic Insects of North America. Kendall/Hunt, Dubuque, Iowa, pp. 181–236.

    Waltz, R.D. & McCafferty, W.P. (1987) New genera of Baetidae for some nearctic species previously included in Baetis Leach (Ephemeroptera). Annals of the Entomological Society of America, 80, 667–670.

    Waltz, R.D. & McCafferty, W.P. (1990) Baetis jesmondis McDunnough, a new junior synonym of Baetis tricaudatus Dodds (Ephemeroptera). Proceedings of the Entomological Society of Washington, 92, 138.

    Webb, J.M., Jacobus, L.M., Funk, D.H., Zhou, X., Kondratieff, B.C., Geraci, C.J., DeWalt, R.E., Baird, D.J., Richard, B., Phillips, I. & Hebert, P.D.N. (2012) A DNA Barcode Library for North American Ephemeroptera: Progress and Prospects. PLoS ONE, 7, e38063.

    Webb, J.M. & Suter, P.J. (2010) Revalidation and redescription of Bungona illiesi (Lugo-Ortiz & McCafferty) (Ephemeroptera: Baetidae) from Australia, based on mitochondrial and morphological evidence. Zootaxa, 2481, 37–51.

    Webb, J.M. & Suter, P.J. (2011) Identification of larvae of Australian Baetidae. Museum Victoria Science Reports, 15, 1–24.

    White, B.P., Pilgrim, E.M., Boykin, L.M., Stein, E.D., Raphael, D. & Mazor, R.D. (2014) Comparison of four species-delimitation methods applied to a DNA barcode data set of insect larvae for use in routine bioassessment. Freshwater Science, 33, 338–348.

    Wiersema, N.A., Nelson, C.R. & Kuehnl, K.F. (2004) A new small minnow mayfly (Ephemeroptera: Baetidae) from Utah, U.S.A. Entomological News, 115, 139–145.

    Williams, H.C., Ormerod, S.J. & Bruford, M.W. (2006) Molecular systematics and phylogeography of the cryptic species complex Baetis rhodani (Ephemeroptera, Baetidae). Molecular Phylogenetics and Evolution, 40, 370–382.

    Zhou, X., Adamowicz, S.J., Jacobus, L.M., DeWalt, R.E. & Hebert, P.D.N. (2009) Towards a comprehensive barcode library for arctic life - Ephemeroptera, Plecoptera, and Trichoptera of Churchill, Manitoba, Canada. Frontiers in Zoology, 6, 30.

    Zhou, X., Jacobus, L.M., DeWalt, R.E., Adamowicz, S.J. & Hebert, P.D.N. (2010) Ephemeroptera, Plecoptera, and Trichoptera fauna of Churchill (Manitoba, Canada): insights into biodiversity patterns from DNA barcoding. Journal of the North American Benthological Society, 29, 814–837.