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Ephemeroptera: Heptageniidae of Gunnison County, Colorado

Rhithrogena hageni
Western March Browns, Red Quill, Western Black Quill

(Eaton) 1885
Updated 19 Nov 2020
TSN 100583


There are several papers about an ectoparasitic midge larvae called Symbiocladius that refer to this species by one of it's old names Rhithrogena doddsi.

Good Links

On this website:
Introduction to Rhithrogena

Other Websites:
PAN Pesticides database: http://www.pesticideinfo.org/List_AquireAll.jsp?Species=5367&Effect=


Argyle,DW; Edmunds,GF 1962 Mayflies (Ephemeroptera) of the Curecanti Reservoir Basins Gunnison River, Colorado. University of Utah Anthropological Papers 59 8, 178-189.

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
     Abstract; "Heptageniid mayfly nymphs have been suggested as sensitive indicators of metal contamination in streams based on biomonitoring studies, experimentation in situ, and experimentation in microcosm. Laboratory tests were conducted to evaluate the sensitivity of Rhithrogena hageni, a heptageniid mayfly, to waterborne copper, cadmium, and zinc. Tests were conducted with soft water (hardness = 40-50 mg/L) at about 12 degrees C. Toxicity endpoints were survival and moulting (%/day). Median 96 hr lethal concentrations were 0.137, 10.5, and 50.5 mg/L for copper, cadmium and zinc, respectively. The average daily moulting rate of survivors significantly decreased after exposure to these metals in solution."

Buchwalter DB, Cain DJ, Clements WH, Luoma SN. 2007 Using biodynamic models to reconcile differences between laboratory toxicity tests and field biomonitoring with aquatic insects. Environmental Science & Technology. 41(13):4821-8.
     Abstract: "Aquatic insects often dominate lotic ecosystems, yet these organisms are under-represented in trace metal toxicity databases. Furthermore, toxicity data for aquatic insects do not appear to reflect their actual sensitivities to metals in nature, because the concentrations required to elicit toxicity in the laboratory are considerably higher than those found to impact insect communities in the field. New approaches are therefore needed to better understand how and why insects are differentially susceptible to metal exposures. Biodynamic modeling is a powerful tool for understanding interspecific differences in trace metal bioaccumulation. Because bioaccumulation alone does not necessarily correlate with toxicity, we combined biokinetic parameters associated with dissolved cadmium exposures with studies of the subcellular compartmentalization of accumulated Cd. This combination of physiological traits allowed us to make predictions of susceptibility differences to dissolved Cd in three aquatic insect taxa: Ephemerella excrucians, Rhithrogena morrisoni, and Rhyacophila sp. We compared these predictions with long-term field monitoring data and toxicity tests with closely related taxa: Ephemerella infrequens, Rhithrogena hageni, and Rhyacophila brunnea. Kinetic parameters allowed us
to estimate steady-state concentrations, the time required to reach steady state, and the concentrations of Cd projected to be in potentially toxic compartments for different species. Species-specific physiological traits identified using biodynamic models provided a means for better understanding why toxicity assays with insects have failed to provide meaningful estimates for metal concentrations that would be expected to be protective in nature. "

Clark JL, 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.
     Abstract: " We conducted field and stream microcosm experiments to assess population-level (density, size distribution) and community-level (species richness metrics, multivariate analysis of community composition) responses of macroinvertebrates to heavy metals in the Arkansas River, a mining-polluted stream in Colorado, USA. Experiments were conducted in spring and summer to coincide with early and late developmental stages (i.e., instars) of the mayfly Rhithrogena hageni. Results of field experiments showed significant mortality at metal-contaminated sites during summer when mayfly populations were dominated by small, early instars (mean dry wt = 0.13 mg). In contrast, no significant mortality was observed in spring when organisms were larger (mean dry wt = 1.78 mg). Multivariate analyses based on abundance of dominant taxa clearly separated reference and metal-impacted stations in summer experiments but showed little separation in spring. We observed no significant effects of metals on species richness, number of mayfly species, or EPT (species richness of Ephemeroptera, Plecoptera, and Trichoptera) in.either field experiment. Using stream microcosms, we established concentration-response relationships between heavy metals and R. hageni density, species richness, mayfly richness, and EPT. Density of R. hageni was generally more sensitive to metals than measures of species richness, and summer populations of R. hageni were more sensitive to metals than spring populations. Because the presence of large, relatively tolerant individuals in spring coincided with periods of higher metal concentrations, R. hageni was protected from toxic effects in this system. We conclude that phenology and developmental stage are important factors influencing responses of some aquatic macroinvertebrates to metals. Thus, timing bioassessments to coincide with the presence of these sensitive life stages can improve our ability to detect subtle contaminant effects."

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

Corkum LD and 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 1998 Effects of acidic pH on benthic macroinvertebrate communities in stream microcosms. Hydrobiologia, 379(1-3)135-145

Dodds,GS 1923 Mayflies from Colorado: descriptions of certain species and notes on others. Transactions of American Entomological Society 69, 93-116. PDF

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.
     Described as Rhithrogena hageni.

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

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
     Abstract: " Adult mayflies (Ephemeroptera) were sampled using sticky traps at several locations along a Rocky Mountain stream. Eight genera were collected, of which imagos of Baetis spp and Rhithrogena hageni were most common. Baetis was represented almost exclusively by females and flight direction was significantly in the upstream direction. In R. hageni both males and females were included. Females appeared to fly into clearings or downstream, whereas males showed no directionality, or a downstream bias. This difference in adult flight direction may be partly explainable by the greater amount of downstream drift in Baetis relative to R. hageni. Finally, mermithid nematodes were found to be frequent parasites of Baetis but not of R. hageni causing total absence of eggs in infected females."

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

Jensen,SL 1966 The Mayflies of Idaho (Ephemeroptera). M.S. Thesis, University of Utah, Utah. 364 p.
     Quote from page 192: "Eaton (1885) [1883-88]) rejected the name Heptagenia brunnea Hagen as a secondary homonym and renamed and redescribed Hagen's sepcimens from California. Traver (1935B) and Edmunds (1952b) provide descriptions of the adults and Edmunds (1952b) of the mature nymphs (as R. doddsi McDunnough).
Taxonomy and Biology: Rhithrogena hageni is closely related to R. futilis McDunnough and R. morrisoni (Banks), the three species representing the Brunnea group in Idaho. Only minute and difficult characters will distinguish the male imagos and mature nymphs of these species.
Nymphs of this species have been collected most commonly in larger streams and rivers, usually above elevations of 5000 feet. They are usually found in the swiftest portions of the current, and in a wide range of summer daytime water temperatures from 44° to 62°F. Adults have been collected from mid-June to August.
Distribution: Rhithrogena hageni is a boreal western North American species previously unreported from Idaho."

Kiffney,PM; Clements,WH 1994 Effects of heavy metals on a macroinvertebrate assemblage from a Rocky Mountain stream in experimental microcosms. Journal of the North American Benthological Society 13 4, 511-523.

Kiffney,PM; Clements,WH 1996 Size-dependent response of macroinvertebrates to metals in experimental streams. Environmental Toxicology and Chemistry 15(8)1352-1356.
     Abstract: "Our previous research has shown that the effects of metals on stream benthic invertebrate populations and communities can vary within and between locations. With this in mind, we examined whether invertebrate body size could explain some of the variation in metal sensitivity within a species. Benthic macroinvertebrates from a pristine Rocky Mountain foothills' stream were collected using artificial substrates and exposed to a mixture of Cd, Cu, and Zn in stream microcosms for 10 d at their respective Colorado chronic criterion levels (4.0, 5.0, and 50 mu g/L). The effects of metals on the ephemeropterans Baetis tricaudatus (Baetidae), Ephemerella infrequens (Ephemerellidae), and Rhithrogena hageni (Heptageniidae) and the plecopteran Pteronarcella badia (Pteronarcyidae) were size dependent, as there was an inverse relationship between body size and survivorship. These results may have important implications for setting water-quality criteria for metals and For using benthic invertebrates in biological assessments. "

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

McCafferty,WP; Durfee,RS; Kondratieff,BC 1993 Colorado mayflies (Ephemeroptera): an annotated inventory. Southwestern Naturalist 38(3) 252-274. PDF
     Quote from page 262: "It is likely that some of the Rhithrogena spp. reported from the upper Gunnison River Drainage by Argyle and Edmunds (1962) are referable to this species."

McCafferty,WP and Provonsha, AV The Mayflies of North AmericaSpecies List (Version 8Feb2011)
     Here is the geographic range and synonyms:
Rhithrogena hageni Eaton, 1885 [CAN:NW;USA:NW,SW]
    * Heptagenia brunnea Hagen, 1875 (hom.: H. brunnea (Rambur), 1842)
    * Rhithrogena brunnea (Hagen), 1875 (hom.comb.)
    * Rhithrogena doddsi McDunnough, 1926 (syn.)

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.
     They found Rhithrogena hageni primarily in the surface substrate samples. This species does not spend much time in the hyporheic zone.

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

Radford,DS and Hartland-Rowe,R 1971 The life cycles of some stream insects (Ephemeroptera, Plecoptera) in Alberta. The Canadian Entomologist, 103(4) 609-617.
     Names have changed since 1971:
1971 Name 2020 Name
Nemoura besametsa Prostoia besametsa
Epeorus deceptivus Epeorus deceptivus
Epeorus longimanus Epeorus longimanus
Ephemerella coloradensis Drunella coloradensis
Arcynopteryx aurea Perlinodes aurea
Nemoura cinctipes Zapada cinctipes
Nemoura columbiana Zapada columbiana
Nemoura oregonensis Zapada oregonensis
Cinygmula ramaleyi Cinygmula ramaleyi
Ephemerella doddsi Drunella doddsi
Rhithrogena doddsi Rhithrogena hageni
Abstract: " The life histories of Nemoura besametsa, Epeorus deceptivus, Epeorus longimanus, and Ephemerella coloradensis are described as "fast seasonal" types and Arcynopteryx aurea, Nemoura cinctipes, Nemoura columbiana, Nemoura oregonensis, Cinygmula ramaleyi, Ephemerella doddsi, and Rhithrogena doddsi as "slow seasonal" types according to Hynes´ (1961) classification. All of the species are univoltine with the exception of N. cinctipes which may be bivoltine. There seems to be a correlation between life cycles and food availability. A means of ecological separation in the four Nemoura species is elucidated. Stream temperature was found to influence growth rates."

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

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

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

Wiens AP; Rosenberg DM; 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
     Discussed as Rhithrogena doddsi.

Brown,WS 2004 Mayflies (Ephemeroptera) of Gunnison County, Colorado, USA