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Ephemeroptera: Ephemerellidae of Gunnison County, Colorado

Ephemerella dorothea infrequens
Pale Morning Dun, PMD

Needham, 1908

Updated 22 January 2019
TSN 698506

Good Links

On this website:
Introduction to Ephemerellidae
Ephemerella Introduction

Other Websites:
Book Pale Morning Dun, a book by Richard Dokey. The link is to a few paragraphs about the artifical fly imitation that his Grandfather used to tie by hand and his experience of the start of the PMD hatch every year.

Photo Adult male - go to the bottom of the page and among many great photos, there is a Pale Morning Dun or Ephemerella dorothea second from the right.

PAN Pesticides database: http://www.pesticideinfo.org/List_AquireAll.jsp?Species=5162&Effect=


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

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 and Technology 41, 4821-4828.

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

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

Johnson,SC 1978 Larvae of Ephemerella inermis and E. infrequens (Ephemeroptera: Ephemerellidae). Pan-Pacific Entomologist 54, 19-25.
     After the Ephemerella revision by Jacobus and McCafferty in 2003, this paper could be titled "Larvae of Ephemerella excrucians and E. dorothea infrequens."

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. "

McCafferty,WP and Provonsha, AV The Mayflies of North America Species List (Version 8Feb2011)
     Here is the geographic range and synonyms:
Ephemerella dorothea infrequens McDunnough, 1924 [CAN:NW;USA:FN,NW,SW]
    * Ephemerella infrequens McDunnough, 1924 (orig.)
    * Ephemerella mollitia Seemann, 1927 (syn.)

McDunnough J. 1924. New North American Ephemeridae. Canadian Entomologist 56:221-226.
     Described as Ephemerella infrequens.

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.

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 below the streambed rocks (hyporheic) before and after a water treatment plant was installed to clean up the Leadville Mine Drainage Tunnel effluent in 1992. At their clean reference site they found Ephemerella pr. infrequens in the surface samples with an average density of 1907 individuals per square meter. The hyporheic samples had a mean density of 881/m².

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

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.

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

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 and Cowan,CA 1995 Microhabitat and activity periodicity of predatory stoneflies and their mayfly prey in a western Colorado stream. Oikos, 513-521. PDF
     Abstract: " Experiments were conducted to determine whether overlap between microhabitat preferences and activity periodicities of four mayfly species and their stonefly predators could explain species-specific differences in predator-prey encounter frequencies. Preferences for rock type (slate or granite), flow microhabitat (high or low), rock surface (top, bottom, upstream or downstream sides), and periodicity of drift and the use of rock tops were measured in a stream-side system of flow-through circular Plexiglas chambers receiving natural stream water and light levels. These parameters were compared among the predatory stoneflies, Megarcys signata or Kogotus modestus, and four species of mayflies that vary in their encounter rates with the stoneflies. Based on predator-prey encounter rates previously observed in similar chambers, we expected greater overlap between Megarcys and Ephemerella infrequens and the overwintering generation of the bivoltine mayfly, Baetis bicaudatus than with Cinygmula sp. Likewise, we expected Kogotus microhabitat use to overlap more strongly with that of summer generation Baetis than with later instars of Cinygmula and Epeorus deceptivus. Results ran counter to our predictions, indicating that microhabitats of the prey species with high predator encounter rates did not overlap more strongly with the stoneflies than did mayflies with low predator encounter rates. Most mayflies and stoneflies preferred the bottom surfaces of granite rocks, and showed few flow preferences. Most were nocturnal in their use of top rock surfaces, in drift and feeding activity periodicity. Therefore, nocturnal activity periodicities of both mayflies and stoneflies confirm that mayflies have not evolved feeding periodicity to avoid encounters with foraging stonefly predators. We conclude therefore, that neither temporal nor spatial microhabitat overlap is a reasonable explanation of differential encounter rates between predatory stoneflies and their mayfly prey. Alternative explanations for differential encounter rates are that more abundant or more mobile mayflies have higher encounter rates with predators, and effective pre-contact predator avoidance responses of other mayflies reduce their encounter rates with stoneflies."

Pennack,RW and Ward,JV 1986 Interstital faunal communities of the hyporheic and adjacent groundwater biotopes of a Colorado mountain stream. Archiv für Hydrobiologie Suppl. 74 3, 356-396.
     They found E. infrequens nymphs in the hyporheic zone of the South Platte river in the Front Range of Colorado at 1863 meters elevation.

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.
     Abstract: Field experiments were conducted in the regulated upper Colorado River to assess drift responses of lotic macroinvertebrates to streamflow manipulations. In each of three seasons, drift was collected in one control and two experimental riffles. On the first day, no flow manipulations occurred. Six hours before sunset on the second day, streamflow was simultaneously reduced and elevated in two experimental riffles with instream diversion structures. Following flow elevation, both mean daily drift density and drift rate generally increased for 13 taxa across all seasons. Flow reductions generally induced elevated drift densities for most taxa, but drift rates declined for some taxa. Patterns of diel drift periodicity were less frequently modified by flow manipulations. Taxa with typical nocturnal peaks in drift activity (Baetis spp., Epeorus longimanus, Triznaka signata) generally maintained this pattern despite some increases in diurnal drift. For a few taxa, modification of diel drift patterns occurred, either as nocturnal decreases following reduced flow (Paraleptophlebia heteronea, Ephemerella infrequens) or as diurnal drift increases in response to either elevated flow (Lepidostoma ormeam, Chironomidae larvae) or reduced flow (Simuliidae). With some exceptions, observed drift responses could be used to suggest active versus passive processes of drift entry.

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

Short,RA; Canton,SP and Ward,JV 1980 Detrital processing and associated macroinvertebrates in a Colorado mountain stream. Ecology, 61(4), 727-732. PDF
     E. infrequens nymphs were found in all 4 of the plant species used to make leaf packs; alder, willow, aspen and pine.

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

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

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
      Quote: " ...Ephemerella dorothea infrequens McDunnough, and Teloganopsis deficiens (Morgan) were all monophyletic with a maximum intraspecific divergence greater than the minimum interspecific divergence. "

Brown,WS 2004 Ephemeroptera of Gunnison County, Colorado, USA