Ephemeroptera: Ephemerellidae of Gunnison County, ColoradoIntroduction to the Mayfly genus Ephemerella
Pale Morning Dun, PMDs, Pale Morning Olive, Pale Morning Spinner, Pale Olive QuillWalsh 1862
Updated Twosday 2/22/2022 :-)
TSN 101233
Provisional Species List
Ephemerella aurivillii
Ephemerella excrucians (inermis)
Ephemerella dorothea infrequens
Ephemerella tibialis
Notes
This genus has many lovely translucent little yellow species. The taxonomy has been changing recently, but no matter, the fishermen recognize the local species of Pale Morning Dun in streams across our planet.
Good Links
On this website:
Ephemerella apopsis
Ephemerella inermis
Introduction to Ephemerellidae
Other Websites:
Photos, Map, Taxon Identifier Numbers - from the Global Biodiversity Information Facility Ephemerella at GBIF
Photos, Map, Museums, DNA - Barcode of Life Data System
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.
References
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
Abstract: "We compared genetic lineages in the mayfly genus Ephemerella (Ephemeroptera: Ephemerellidae) identified from mitochondrial DNA (mtDNA) to current taxonomy in 9 morphological taxa, including 2 geographically widespread species, Ephemerella invaria ( = E. inconstans, E. rotunda, E. floripara) and Ephemerella dorothea ( = E. infrequens). Maximum likelihood and parsimony analyses of the mtDNA sequences placed E. inconstans and E. invaria in a well-supported clade; however, mean Kimura 2-parameter genetic distance between the lineages was high (5.2%) relative to distance within lineages (1.3%). The phylogenetic relationships of synonyms E. rotunda and E. floripara were not resolved, but estimates of mean genetic distance to E. invaria were high for both (8.5% and 11.6%, respectively). Populations of E. dorothea were grouped in 2 well-supported clades (12.9% mean divergence) that did not include the synonym E. infrequens (20.9% mean divergence, based on a single sample). A large genetic distance (18.6%) also was found between eastern and western populations of Ephemerella excrucians. Western samples of Ephemerella aurivillii were so genetically distant from all other lineages (32.2%) that doubt about its congeneric status is raised. mtDNA data have been useful for identifying genetic lineages in Ephemerella, but our results do not support use of cytochrome oxidase I (COI) as a DNA barcode to identify species in this genus because we also found evidence of incomplete mtDNA lineage sorting in this gene. Use of the barcoding gene rediscovered some old taxonomic problems in Ephemerella, a result that emphasizes the importance of completing empirical systematic description of species before using single-character systems for identification."
Allard,M and Moreau,G 1987 Effects of experimental acidification on a lotic macroinvertebrate community. Hydrobiologia, 144, pp.37-49.
Abstract: "A three month experimental acidification was carried out on lotic bottom communities. Experiments were conducted under semi-natural conditions in plasticized wooden channels. Acidified communities (pH 4.0), with or without added aluminum, were compared with a reference community (pH 6.3-6.9). Added aluminum concentrations were respectively 0.2 and 0.4 mg 1-1 in experiments performed in 1982 and 1983. Water chemistry and taxonomic composition of the macroinvertebrate communities were monitored. Under acidified conditions, results were similar, with or without added aluminum. Mean abundances of all groups of organisms were lowered. Mayflies nearly completely disappeared from the acidified channels. The only organism not affected by the acidification was Microtendipes sp. Differences in the organism response were observed: Orthocladiinae (Rheocricotopus, Parametriocnemus, Corynoneura, Thienemanniella, Nanocladius, Cricotopus) and Ephemeroptera (Baetis, Habrophlebia, Habrophlebiodes, Paraleptophlebia, Ephemerella), especially early instars, were very sensitive to low pH, Chironomini and Tanypodinae were much less sensitive, while Tanytarsini were intermediate; Oligochaeta and Nematoda were difficult to classify, their response being different from one year to another. Organisms inhabiting the surface of artificial substrates disappeared very rapidly from the system, while those buried inside had a delayed reaction to acidification. Aluminum which was mainly in the monomeric form was not responsible for community modifications. Direct action of hydrogen ions through a physiological stress seems a more credible explanation. These results, induced by a continuous experimental acidification, suggest that if this small headwater stream undergoes acidification, the resulting invertebrate community will be very simplified, with only resistant species able to cope with the acid conditions."
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 and Edmunds,GF 1965 A revision of the genus Ephemerella (Ephemeroptera, Ephemerellidae) VIII. The subgenus Ephemerella in North America. In A Revision of the Genus Ephemerella (Ephemeroptera, Ephemerellidae) VIII. The Subgenus Ephemerella in North America (Vol. 4, p. 244). Entomological Society of America. PDF
Abstract: "The 25 species of the genus Ephemerella Walsh in North America formerly placed in the subgenera Ephemerella, s. s. (=invaria-group auct.), and Chitonophora (Bengtsson) (=needhami-group auct.) are placed in the subgenus Ephemerella, s. s. Characters do not allow grouping of the nymphs and adults in concordant groups that correspond with the 2 subgenera previously recognized. Accounts are given for the nymph and male adult of each species including synonymies, synoptic descriptions, and distribution records and maps. Illustrated keys are given for the nymphs and male adults. E. euterpe Traver is regarded as a synonym of E. maculata Traver. E. lacustris is described from imagoes and nymphs from Wyoming, and E. verruca from Oregon, E. simila from West Virginia, E. rossi from Tennessee, and E. crenula and E. hispida from North Carolina and Tennessee are described from the nymphal stage only."
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
Abstract: "Conceptual and quantitative models were developed to assess time-dependent processes in four sequential experimental stream studies that determined abundances of natural communities of mayfly and caddisfly larvae dosed with single metals (Cd, Co, Cu, Ni, Zn) or multiple metals (Cd + Zn, Co + Cu, Cu + Ni, Cu + Zn, Ni + Zn, Cd + Cu + Zn, Co + Cu + Ni, Cu + Ni + Zn). Metal mixtures contained environmentally relevant metal ratios found in mine drainage. Free metal ion concentrations, accumulation of metals by periphyton, and metal uptake by four families of aquatic insect larvae were either measured (Brachycentridae) or predicted (Ephemerellidae, Heptageniidae, Hydropsychidae) using equilibrium and biodynamic models. Toxicity functions, which included metal accumulations by larvae and metal potencies, were linked to abundances of the insect families. Model results indicated that mayflies accumulated more metal than caddisflies and the relative importance of metal uptake by larvae via dissolved or dietary pathways highly depended on metal uptake rate constants for each insect family and concentrations of metals in food and water. For solution compositions in the experimental streams, accumulations of Cd, Cu, and Zn in larvae occurred primarily through dietary uptake, whereas uptake of dissolved metal was more important for Co and Ni accumulations. Cd, Cu, and Ni were major contributors to toxicity in metal mixtures and for metal ratios examined. Our conceptual approach and quantitative results should aid in designing laboratory experiments and field studies that evaluate metal uptake pathways and metal mixture toxicity to aquatic biota."
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.
Edmunds Jr,GF and McCafferty,WP 1988 The mayfly subimago. Annual review of entomology, 33(1)509-527. PDF
Quote from page 522: "The speed with which subimagos were able to right themselves after being placed on their side with wings contacting water varied immensely among subimagos of different genera observed (G. F. Edmunds,Jr. and T. J. Fink, unpublished). The results could be based on differences in hydrofuge capacity, differences in behavioral response among mayflies, or both. Siphlonurids (Siphlonurus, Ameletus, Parameletus) were slow to respond, and most of the individuals, especially of Siphlonurus, were entrapped on the surface. Heptageniids (Stenacron, Nixe, Heptagenia) righted themselves faster than the siphlonurids and almost always escaped successfully. Leptophlebiids (Paraleptophlebia) and ephemerellids (Ephemerella, Drunella) responded rapidly and escaped so fast that sometimes it was impossible to see if they righted themselves before taking flight"
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.
Working in wadeable streams on the Front Range of Colorado, they found a cryptic species of Ephemerella alongside E. dorothea and infrequens.
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.
Abstract: "The study of population variability and the reexamination of type material of certain Ephemerellidae species has revealed changes in status and new synonyms. Ephemerella dorothea Needham, 1908 [=E. infrequens McDunnough, 1924, new synonym, =E. mollitia Seemann, 1927, new synonym] is divided into two subspecies: E. d. dorothea, new status, and E. d. infrequens new status [=E. mollitia, new synonym]. New synonyms were discovered for the following: Ephemerella excrucians Walsh, 1862 [=E. inermis Eaton, 1884, new synonym, =E. argo Burks, 1947, new synonym, =E. crenula Allen and Edmunds, 1965, new synonym, =E. lacustris Allen and Edmunds, 1965, new synonym, =E. rossi Allen and Edmunds, 1965, new synonym, =E. rama Allen, 1968, new synonym], E. invaria Walker, 1853 [=E. rotunda Morgan, 1911, new synonym, =E. vernalis Banks, 1914, new synonym, =E. feminina Needham, 1924, new synonym, =E. fratercula McDunnough, 1925, new synonym, =E. inconstans Traver, 1932, new synonym, =E. choctawhatchee Berner, 1946, new synonym, =E. simila Allen and Edmunds, 1965, new synonym, =E. floripara McCafferty, 1985, new synonym], E. mucronata (Bengtsson), 1909 [=E. moffatae Allen, 1977, new synonym], Serratella micheneri (Traver), 1934 [=E. altana Allen, 1968, new synonym], S. serrata (Morgan), 1911 [=S. sordida (McDunnough), 1925, new synonym, =S. carolina (Berner and Allen), 1961, new synonym, =S. spiculosa (Berner and Allen), 1961, new synonym], and S. tibialis."
Jacobus, LM and McCafferty, WP 2008 Revision of Ephemerellidae genera (Ephemeroptera). Transactions of the American Entomological Society 134: 185-274. PDF
Abstract: "The study of population variability and the reexamination of type material of certain Ephemerellidae species has revealed changes in status and new synonyms. Ephemerella dorothea Needham, 1908 [=E. infrequens McDunnough, 1924, new synonym, =E. mollitia Seemann, 1927, new synonym] is divided into two subspecies: E. d. dorothea, new status, and E. d. infrequens new status [=E. mollitia, new synonym]. New synonyms were discovered for the following: Ephemerella excrucians Walsh, 1862 [=E. inermis Eaton, 1884, new synonym, =E. argo Burks, 1947, new synonym, =E. crenula Allen and Edmunds, 1965, new synonym, =E. lacustris Allen and Edmunds, 1965, new synonym, =E. rossi Allen and Edmunds, 1965, new synonym, =E. rama Allen, 1968, new synonym], E. invaria Walker, 1853 [=E. rotunda Morgan, 1911, new synonym, =E. vernalis Banks, 1914, new synonym, =E. feminina Needham, 1924, new synonym, =E. fratercula McDunnough, 1925, new synonym, =E. inconstans Traver, 1932, new synonym, =E. choctawhatchee Berner, 1946, new synonym, =E. simila Allen and Edmunds, 1965, new synonym, =E. floripara McCafferty, 1985, new synonym], E. mucronata (Bengtsson), 1909 [=E. moffatae Allen, 1977, new synonym], Serratella micheneri (Traver), 1934 [=E. altana Allen, 1968, new synonym], S. serrata (Morgan), 1911 [=S. sordida (McDunnough), 1925, new synonym, =S. carolina (Berner and Allen), 1961, new synonym, =S. spiculosa (Berner and Allen), 1961, new synonym], and S. tibialis."
Ogden,TH; Osborne,JT; Jacobus,LM and Whiting,MF 2009 Combined molecular and morphological phylogeny of Ephemerellinae (Ephemerellidae: Ephemeroptera), with remarks about classification. Zootaxa, 1991(1), pp.28-42. PDF
Abstract: "This study represents the first combined molecular and morphological analysis for the mayfly family Ephemerellidae (Ephemeroptera), with a focus on the relationships of genera and species groups of the subfamily Ephemerellinae. The phylogeny was constructed based on DNA sequence data from 3 nuclear (18S rDNA, 28S rDNA, histone H3) and 2 mitochondrial (12S rDNA, 16S rDNA) genes, and 23 morphological characters. Taxon sampling for Ephemerellidae included exemplars from all 25 extant genus groups and additional representatives from those genera with the highest diversity. Ephemerellidae appears to consist of three major clades. Ephemerella, the largest genus of Ephemerellidae, and Serratella were not supported as monophyletic, and each had representatives in two of the three major clades. However, the genera Drunella and Cincticostella were supported as monophyletic. Lineages strongly supported as monophyletic include a grouping of the Timpanoginae genera Timpanoga, Dannella, Dentatella and Eurylophella, and groupings of the Ephemerellinae genera Torleya, Hyrtanella and Crinitella and the genera Kangella, Uracanthella and Teloganopsis. The placement of the Timpanoginae genus Attenella fell within Ephemerellinae, based on molecular and combined data, but it grouped with other Timpanoginae based on morphological data alone. Further study and analysis of Ephemerellidae morphology is needed, and classification should be revised, if it is to reflect phylogenetic relationships."
Peckarsky,BL 1996 Alternative predator avoidance syndromes of stream-dwelling mayfly larvae. Ecology, 77(6), pp.1888-1905. PDF
Abstract: "Experiments were conducted to compare the patterns, mechanisms, and costs of predator avoidance behavior among larvae of five species of mayflies that co-occur with the predatory stoneflies, Megarcys signata and Kogotus modestus in western Colorado streams. Mayfly drift dispersal behavior, use of high vs. low food (periphyton or detritus) patches, microhabitat use, positioning, and activity periodicity were observed in the presence and absence of predators in circular flow-through chambers using natural stream water. Also, distances from predators at which prey initiated escape responses were compared among prey and predator species. Costs of predator avoidance behavior were assessed by measuring short-term (24 h) feeding rates of mayflies in the presence or absence of predatory stoneflies whose mouthparts were immobilized (glued) to prevent feeding. The intensity and associated costs of predator avoidance behavior of mayfly species were consistent with their relative rates of predation by stoneflies. Megarcys consumes overwintering generation Baetis bicaudatus > Epeorus longimanus > Cinygmula = Ephemerella; Kogotus consumes summer generation Baetis > Epeorus deceptivus = Cinygmula; Megarcys eats more mayflies than Kogotus. While Megarcys induced drift by Baetis, Epeorus, and Cinygmula, this disruptive predator avoidance behavior only reduced food intake by Baetis and Epeorus. The morphologically defended mayfly species, Ephemerella, neither showed escape behavior from Megarcys, nor any cost of its antipredatory posturing behavior. Only Baetis responded by drifting from Kogotus. No mayfly species shifted microhabitats or spent less time on high-food patches in the presence of foraging stoneflies. However, predators enhanced the nocturnal periodicity of Baetis drift, which was negligible in the absence of stoneflies as long as food was abundant. Lack of food also caused some microhabitat and periodicity shifts and increased the magnitude of both day and night drift of Baetis. Thus, Baetis took more risks of predation by visual, drift-feeding fish not only in the presence of predatory stoneflies, but also when food was low or they were hungry. All other mayflies were generally nocturnal in their use of rock surfaces, as long as food was abundant. Finally, the distances at which different mayfly species initiated acute escape responses were also consistent with relative rates of predation. This study demonstrates alternative predator avoidance syndromes by mayfly species ranging from an initial investment in constitutive morphological defenses (e.g., Ephemerella) to induced, energetically costly predator avoidance behaviors (e.g., Baetis). Although the costs of Ephemerella's constitutive defense are unknown, experiments show that prey dispersal is the mechanism underlying fecundity costs of induced responses by Baetis to predators, rather than microhabitat shifts to less favorable resources or temporal changes in foraging activity. A conceptual model suggests that contrasting resource acquisition modes may account for the evolution and maintenance of alternative predator avoidance syndromes along a continuum from Baetis (high mobility) to heptageniids (intermediate mobility) to Ephemerella (low mobility). Prey dispersal (swimming) to avoid capture results in reduction of otherwise high fecundity by Baetis, which trades off morphological defense for enhanced ability to acquire resources. Thus, improved foraging efficiency is the selection pressure maintaining the highly mobile life style in Baetis, which increases resource acquisition and fecundity, offsetting the high mortality costs associated with this behavior."
Short,RA; Canton,SP and Ward,JV 1980 Detrital processing and associated macroinvertebrates in a Colorado mountain stream. Ecology, 61(4), 727-732. PDF
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.
Walsh described the genus Ephemerella in this paper.
Williams,MC and Lichtwardt,RW 1999 Two new Harpellales living in Ephemeroptera nymphs in Colorado Rocky Mountain streams. Mycologia, 91(2) 400-404. PDF
Abstract: "Two new species of harpellid gut fungi (Zygomycota: Trichomycetes) are described from the hindguts of mayfly nymphs inhabiting high altitude Rocky Mountain streams: the new genus and species Legeriosimilis tricaudata living in Ameletus sp. (Siphlonuridae), and the new species Glotzia coloradense from Baetis tricaudatus (Baetidae). Legeriomyces aenigmaticus is reported from a new site and a new ephemeropteran host, Ephemerella sp. (Ephemerellidae), and previously unknown zygospores are described."
"It ain't what you don't know that gets you into trouble.
It's what you know for sure that just ain't so."
--Mark Twain
Brown,WS 2009 Ephemeroptera of Gunnison County, Colorado
www.gunnisoninsects.org
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