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Ephemeroptera: Ephemerellidae of Gunnison County, Colorado

Drunella doddsii
Green Drake, Western Green Drake, Great Red Spinner, Ginger Quill

(Needham, 1927)

Updated 24 March 2024
TSN 698494

Locations collected

Meyers Gulch, West Elk Creek, East Elk Creek (Argyle and Edmunds, 1962). Also common in the Coal Creek, Copper Creek, East River, Slate River, Brush Creek and tributaries.

Good Links

On this website:
Drunella Introduction

Other Websites:
Photos, Map, Museums, DNA - Barcode of Life Data System

North American distribution map - flyfishingentomology.com


Allan,JD 1987 Macroinvertebrate drift in a Rocky Mountain stream. Hydrobiologia 144, 261-268.

Allen,RK and Edmunds Jr,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

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.
     Discussed as Ephemerella doddsi. Quote from page 186: "This species was uncommon in the collection. It is particularly well adapted to the torrential habitat occurring in several streams in this drainage, but was only taken from three collection streams. Most of the specimens may have emerged before the intensive July collections."

Balistrieri,LS; Mebane,CA and Schmidt,TS 2020 Time-dependent accumulation of Cd, Co, Cu, Ni, and Zn in mayfly and caddisfly larvae in experimental streams: Metal sensitivity, uptake pathways, and mixture toxicity. Science of the Total Environment, 732. html

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.

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.

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

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
     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."

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.

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
      The authors studied the acute short term tolerance of aquatic insects to low oxygen. They used the 96 hour Median Tolerance Limit. They discussed Drunella doddsii as Ephemerella doddsi. The TLm96 for D. doddsii was 5.2mg/l and 46% oxygen saturation.

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
     Abstract: "The mature larvae of fifteen species of aquatic insects (Diptera, Ephemeroptera, Plecoptera, and Trichoptera) and the scud (Amphipoda) were tested to determine their relative sensitivity to heated waters under laboratory conditions. The temperature at which 50% died after 96 hours (TLm96) was recorded as the lethal temperature. This ranged from 11.7 C for the torrential stream mayfly, Cinygmula par Baton, to 32.6 C for the snipefly, Atherix variegata Walker. "       They discussed Drunella doddsii as Ephemerella doddsi. The TLm96 for D. doddsii was 15.45°C.

Gersich,FM and Brusven,MA 1982 Volcanic ash accumulation and ash-voiding mechanisms of aquatic insects. Journal of the Kansas Entomological Society, 55(2) 290-296.
     Abstract: "Four species of aquatic insects (Hesperoperla pacifica, Rhyacophila acropedes/R. vao, Drunella doddsi and Rhithrogena robusta) were subjected to suspended ash concentration of ca. 2000 mg/1 in laboratory streams for 48 hr. Macro- and microscopic examination revealed substantial ash accumulation on the exo-skeleton; however, acute toxicity was not noted. Ash-impacted aquatic insects placed in a clean water environment voided appreciable amounts of ash within 24 hr. We conclude that the four species studied had high short-term exposure tolerances to ash and that behavioral attributes allowed for the removal of exoskeletal ash deposits once the perturbation ceased."

Gilpin,BR and Brusven,MA 1970 Food habits and ecology of mayflies of the St. Maries River in Idaho. Melanderia 4:19-40. PDF
     Discussed as Ephemerella (Drunella) doddsi.

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

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
     While variously lumping Drunella species from all over the world, they note there was an early fumble on the name of this mayfly. Even though there are many references to this insect as D. doddsi in the scientific and fishing literature, the authors point out the name really has two ii's at the end, i.e Drunella doddsii. See Needham's description below.

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.
     Abstract: "Natural assemblages of stream benthic macroinvertebrates were collected using artificial substrates from a Rocky Mountain stream and exposed for 10 d to a mixture of heavy metals (Cd, Cu, and Zn) in stream microcosms. Metal levels were 0, 1х, 5х, and 10х where х = 1.1, 12, and 110 μg/L Cd, Cu, and Zn, respectively. The 1х treatment was similar to chronic criteria values recommended by the US Environmental Protection Agency (EPA) for each metal and total metal levels measured in water at the Arkansas River, Colorado, a US EPA superfund site. Most ephemeropterans and plecopterans were sensitive to metals; however, some taxa within these groups were metal tolerant. Densities of Baetis tricaudatus (Ephemeroptera:Baetidae), Epeorus longimanus and Rhithrogena hageni (Ephemeroptera:Heptageniidae), and Drunella grandis and D. doddsi (Ephemeroptera:Ephemerellidae) were reduced in the 1х treatment. The response of D. grandis to metals was size-dependent with small larvae being more sensitive than large ones (p = 0.02). Chironomids were generally tolerant to metals. These data show that a metal mixture was extremely toxic to stream macroinvertebrates from a Rocky Mountain stream. Our results were similar to field biomonitoring studies at the Arkansas River and Eagle River, Colorado, that examined the effects of metals on stream macroinvertebrate communities. We suggest that multispecies experiments using indigenous stream organisms be combined with field biomonitoring to rigorously define the biological effects of heavy metals on lotic systems. "

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
     Quote: "All taxa showed variability in the proportion of their diet derived from autochthonous compared to allochthonous sources across streams (Supplementary Table 1). Generally, mean consumer autochthony ranged from around 0.25 to 0.5 proportionally of the diet. Certain taxa, such as the rhyacophilid caddisfly Rhyacophila angelita (mean of dietary proportion derived from the epilithon of 0.39-0.48 for their prey across streams), the simuliid blackfly Simulium sp. (0.24-0.31), the perlodid stonefly Megarcys signata (0.48-0.53) and nemourid stonefly Zapada sp. (0.51-0.53) varied little in their resource use across streams. In contrast, the baetid mayfly Baetis bicaudatis (0.22-0.52), the perlodid stonefly Kogotus modestus (0.28-0.61), and the ephemerellid mayflies Drunella doddsi (0.26-0.55), and Drunella grandis (0.25-0.42) exhibited more variability in their resource use across streams. Full stable isotope results and biplots are available in Supplemental Figures 1 and 2."

Magnum,FA and Winget,RN 1991 Environmental profile of Drunella doddsi (Needham) (Ephemeroptera: Ephemerellidae). Journal of Freshwater Ecology 6 (1) 11-22.
     Abstract: "Data from 813 stream stations in 11 western states have been analyzed to provide a description of the niche width of Drunella (Eatonella) doddsi Needham. The concept of niche width refers to a species' distribution according to environmental conditions, reflecting environmental tolerances. Drunella doddsi was found over a broad range of elevations, and average minimum stream flows. However, its distribution was highly correlated to streams with moderate to high gradients, coarse substrates, dense riparian vegetation, low to moderate levels of alkalinity, very low levels of sulfates, and low specific conductance. This narrow niche width seems to indicate an intolerance of D. doddsi to many types of environmental conditions. Specific habitat selection also requires competitive prevalence over other species that would utilize that habitat."

Maret,TR; Cain,DJ; MacCoy,DE and 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.
     Drunella doddsii, in spite of being in a genus considered sensitive to heavy metals, was found at two of the most contaminated sites in this study.

McCafferty,WP; Durfee,RS and Kondratieff,BC 1993Colorado mayflies (Ephemeroptera): an annotated inventory. Southwestern Naturalist 38 (3) 252-274. PDF
     Quote from page 265: "All reports have been as Ephemerella doddsi except for those of Dodds (1923) and Dodds and Hisaw (1925) from Boulder and Gilpin Co. (South Boulder Cr.). Although Eaton (1884) is often credited with the first Colorado record of this common western mayfly (e.g. Traver, 1935), that particular record is actually from Idaho on the Colorado Territory."

McCafferty,WP and Provonsha, AV The Mayflies of North AmericaSpecies List (Version 8Feb2011)
     Here is the geographic range and synonyms:
Drunella doddsii (Needham), 1927 [CAN:FN,NW;USA:FN,NW,SW]
    * Drunella doddsi (Needham), 1927 (spell.)
    * Ephemerella doddsi Needham, 1927 (comb.,spell.)
    * Ephemerella doddsii Needham, 1927 (orig.)

McCarty,JD; Cross,WF; Albertson,LK; Tumolo,BB and Sklar,LS 2022 Life histories and production of three Rocky Mountain aquatic insects along an elevation-driven temperature gradient. Hydrobiologia, 849(16), pp.3633-3652. PDF
     Abstract: "Although temperature is known to influence individual traits such as growth, body size, and fecundity, few studies have examined how these relationships influence population-level secondary production in natural settings. We quantified life history traits and production of three dominant aquatic insects (Ephemeroptera: Ephemerella infrequens; Drunella doddsii; Trichoptera: Hydropsyche cockerelli) along an elevation-driven thermal gradient in the northern Rockies. We predicted that production would be highest at sites where temperature regimes lead to the largest terminal body size, individual fecundity, and reproductive potential (i.e., eggs per female x abundance of mature nymphs or larvae). In general, we found that temperature had idiosyncratic effects on life history traits of the study taxa, with no consistent effect of temperature on production. Although growth rates were generally highest during the warm months, growth did not consistently covary with temperature among sites along the elevation gradient. Terminal body size also differed among sites and was inconsistently related to mean temperature and reproductive potential. One of three taxa, D. doddsii, showed patterns entirely consistent with predictions, including smallest body size, reproductive potential, and secondary production at the warmest, low-elevation sites. Our findings suggest that connections among temperature regime, life history characteristics, and secondary production may not be straightforward, and are likely influenced by characteristics unmeasured in our study, including factors that influence survivorship throughout the larval phase, as well as adult mating or oviposition success. Such additional information will enrich our understanding of thermal effects on aquatic insects, and may contribute to predicting how these taxa may respond to ongoing changes in climate."

Mebane,CA; Schmidt,TS; Miller,JL and Balistrieri,LS 2020 Bioaccumulation and toxicity of cadmium, copper, nickel, and zinc and their mixtures to aquatic insect communities. Environmental toxicology and chemistry, 39(4) 812-833. PDF

Mihuc,TB and Minshall, GW 1995 Trophic generalists vs. trophic specialists: implications for food web dynamics in post-fire streams. Ecology, 76(8) 2361-2372.
     Abstract: "The trophic ecology of 11 benthic macroinvertebrate taxa found in Cache Creek, Yellowstone National Park (YNP) was studied to determine if burned organic matter is an important resource and how resource utilization patterns may be altered in post-fire streams. Laboratory food quality experiments were conducted to determine the growth response of each species when grown on several resource types: burned organic matter, periphyton, unburned coarse particulate material (CPM), and unburned fine particulate material (FPM). The central hypothesis of this research was that benthic macroinvertebrates cannot use burned organic matter as a resource. A secondary hypothesis was that some benthic macroinvertebrates are facultative in trophic function, with the ability to use both allochthonous and autochthonous resources for growth. Of the 11 taxa studied, only one (Paraleptophlebia heteronea) could grow on burned organic matter as a resource, indicating that post-fire food webs probably do not exhibit major shifts in resource utilization to burned material. Two species were generalist detrivores (P. heteronea, Ameletus cooki) able to use both natural FPM and CPM resources. Two species were specialist detritivores (Oligophlebodes sigma, Ephemerella infrequens) growing only on unburned CPM resources, and two (Cinygmula mimus, Epeorus albertae) were specialist herbivores utilizing only periphyton. Five species were generalist herbivore-detritivores (Baetis bicaudatus, Drunella doddsi, D. coloradensis, D. spinifera, Zapada columbiana), exhibiting growth on both detritus and periphyton resources. Based on the experimental results, trophic generalists are common food web components in Yellowstone streams. Two of the most abundant benthic macroinvertebrates during post-fire recovery, B. bicaudatus and Z. columbiana, were trophic generalists, indicating that some generalists may be disturbance adapted. In this study, published functional feeding group classification did not indicate obligate resource utilization (growth on only one resource type) for most taxa studied. Comparison of a food web for Cache Creek based on functional feeding group classification and one based on the results of this study indicates that the inclusion of generalists in the web results in a more realistic approximation of food web relationships such as the link-species scaling law. Our results suggest that future research should include spatial and temporal aspects of resource switching and generalist resource utilization by individual lotic primary consumers."

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

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

Prusha,BA; 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.
     Abstract: " Relationships among landscape attributes, instream dissolved organic C (DOC) concentrations, and metal bioaccumulation in Arctopsyche grandis and Drunella doddsi were investigated in the Upper Arkansas River Basin, Colorado. A Geographic Information System (GIS) was used to calculate landscape attributes in 16 watersheds where DOC concentrations were measured from May to August 2001. Metal concentrations in Arctopsyche and Drunella were related to physicochemical characteristics measured in these streams and in the Arkansas River. Results of multiple linear regression showed that % forested area explained 47% and 59% of the variation in maximum and mean DOC concentrations, respectively. Maximum DOC was negatively associated with the concentration of Zn (R2 = 0.25) and Cd (R2 = 0.39) in Arctopsyche. DOC concentration did not describe metal concentrations in Drunella, which accumulated significantly more Zn, Cd, and Cu than Arctopsyche. The higher metal concentrations and the absence of a DOC effect on metal uptake in Drunella most likely resulted from dietary exposure to metal-enriched detritus and periphyton. Our results indicate that % forested area within a watershed can be used to describe DOC concentrations, which in turn influence metal bioaccumulation in Arctopsyche. To our knowledge, this study is the first to quantify the relationship between landscape attributes and DOC, and to demonstrate the influence of DOC on metal bioaccumulation in benthic macroinvertebrates in the field."

Prusha,BA; Kashian,DR and Clements,WH 2004 Effects of total organic carbon and ultraviolet-b radiation on metal accumulation in Arctopsyche grandis and Drunella doddsi from the Upper Arkansas River Basin (Colorado). Presented at the NABS Annual meeting, Athens, Georgia, 2004 in Ecotoxicology http://www.benthos.org/database/allnabstracts.cfm/db/Athens2004abstracts/id/30

Rader,RB and Ward,JV 1987 Resource utilization, overlap and temporal dynamics in a guild of mountain stream insects. Freshwater Biology, 18(3), pp.521-528. PDF

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

The United States Geological Survey (USGS) National Water Quality Assessment Data Warehouse (NAWQA) shows this species is present in Gunnison County. Data as of 1Sep2005

Winget,RN 1993 Habitat partitioning among three species of Ephemerelloidea. Journal of Freshwater Ecology 8 (3) 227-234. PDF

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