Ametropus albrighti is shown to be a junior subjective synonym of A. neavei, n. syn. Ametropus neavei sensu lato demonstrates stable structural characteristics with respect to male genitalia and larval morphology, whereas degree of maculation and therefore abdominal patterning varies both among populations and within populations. Ametropus ammophilus is distinctive, and the two North American species are apparently allopatric based on known distributions, which are reviewed

McCafferty, W. P

English

Valparaiso University

The Great Lakes Entomologist Volume 34 Number 2 - Fall/Winter 2001 Number 2 - Fall/Winter 2001 Article 1 October 2001 Commentary on Ametropus Species (Ephemeroptera: Ametropodidae) in North America W. P. McCafferty Purdue University Follow this and additional works at: https://scholar.valpo.edu/tgle Part of the Entomology Commons Recommended Citation McCafferty, W. P. 2001. "Commentary on Ametropus Species (Ephemeroptera: Ametropodidae) in North America," The Great Lakes Entomologist, vol 34 (2) Available at: https://scholar.valpo.edu/tgle/vol34/iss2/1 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 2001 THE GREAT LAKES ENTOMOLOGIST COMMENTARY ON AMETROPUS SPECIES (EPHEMEROPTERA: AMETROPODIDAE) IN NORTH AMERICA w. P. McCaffertyl ABSTRACT Ametropus albrighti is shown to be a junior subjective synonym ofA. neavei, n. syn. Ametropus neavei sensu lato demonstrates stable structural character­istics with respect to male genitalia and larval morphology, whereas degree of maculation and therefore abdominal patterning varies both among populations and within populations. Ametropus ammophilus is distinctive, and the two North American species are apparently allopatric based on known distribu­tions, which are reviewed. The Holarctic mayfly genus Ametropus Albarda was originally discov­ered in Europe (Albarda 1878) and is now known from Asia and North America as well. Bengtsson (1913) placed this distinctive genus in a separate higher taxon Ametropodidae (originally spelled Ametropidae) where it has resided ei­ther at the family or subfamily level ever since. McCafferty (1981) referred to these mayflies as the sand minnow mayflies because of the peculiar adapta­tions of the larvae for life in sand substrates of rivers (e.g., Soluk and Craig 1988), and workers in g neral have regarded the group's relationships to be with the pisciform mayflies (e.g., Edmunds et al.1976, McCafferty 1991). McDunnough (1928) first discovered the genus in North America. His report was based on a single female adult from Alberta that he described as Ametropus neavei McDunnough. Traver (1935) described a second species, A. albrighti Traver, based on a series oflarvae taken from the San ,Juan River in extreme northwest New Mexico. Edmunds (1954), Woodbury and Argyle (1963), and Pearson et al. (1968) all referred to additional larval populations of A. albrighti from the Green River in northeastern Utah. Newell (1970) listedA. neavei from Montana but provided no substant ating locale data (presumably he saw a female adult that matched McDunnough's [1928] description). Allen and Edmunds (1956) listed an unnamed species of Ametropus from Oregon, and later Allen and Edmunds (1976) described that species asA. ammophilus Allen and Edmunds based on male and female adults and larvae from Oregon, Washington, and we tern Montana. At the same time, Allen and Edmunds (1976) described the male adult and female subimago of A. albrighti, and pro­vided additional records of that species from northwestern Utah, northeastern Colorado, and southwestern Wyoming. Allen and Edmunds (1976) also pro­vided descriptions of the male adult and larva of A. neavei for the first time, along with a record of the species from Saskatchewan. Allen and Edmunds (1976) concluded thatA. neauei was restricted to Alberta and Saskatchewan; thatA. albrighti was restrictcd to the Colorado River Drain­age System in Colorado, New Mexico, Utah, and Wyoming; and that A. ammophilus was restricted to Oregon, Washington, and areas east to western Montana. Subsequent to the review ofthe genus Ametropus in North America by Allen and Edmunds (1976), however, several additional reports under the vari­ous species names were forthcoming. A etropus albrighti was reported from Saskatchewan by Lehmkuhl (1976a) and by Dosdall and Lehmkuhl (1989). Ametropus ammophilus was reported from northern California by Allen (1977). Ametropus neavei was reported from additional localities in Alberta by Clifford ]Department of Entomology, Purdue University, West Lafayette, IN 47907, pat_mccafferty@entm.purdue.edu 1McCafferty: Commentary on Ametropus Species (Ephemeroptera: AmetropodiPublished by ValpoScholar, 20012 THE GREAT LAKES ENTOMOLOGIST Vol. 34, No.2 and Barton (1979), Barton (1980), and Soluk and Craig (1988); from the upper peninsula of Michigan by Steven and Hilsenhoff (1979); from the Northwest Territories by Cobb et al. (1995); and again from Montana by Waltz et al. (1998). ANALYSIS OF SPECIES Ametropus ammophilus is a distinctive species within Ametropus with respect to its much larger size, cleft penes of its male genitalia, and complex abdominal color pattern. These distinctions are accurately reflected in the spe­cies treatment and key provided by Allen and E munds (1976) and therefore all reports under that name can reasonably be expected to have been applied to the correct species. It is known to range from northern California north through Oregon, Washington and, in the north, east to western Montana, including a first report from Idaho by Lester et al. (2002). It may also occur in Alberta (see below). My study of materials ofAmetropus from north-central and far eastern Mon­tana and re-examination of most of the materials on which Allen and Edmunds (1976) based their review suggested tha reports variously assigned to A. lbrighti orA. neavei, other than those of the types, may not be reliable. Importantly, the key provided by Allen and Edmunds (1976), and probably used by every worker since that time when faced with i entifyingAmetropus larvae and adults, is inac­curate based on the actual concepts of the species. This fact was borne out by the examination of specimens used by Allen and Edmunds (1976) in their study. Cou­plet 2 indicated thatA. neavei had abdominal terga 5-7 with inverted T-shaped markings. That statement actually is referable to A. albrighti based on the de­scriptions and figures provided by Allen and Edmunds (1976) and all prior descrip­tions. In the same sense, the key stat ment that abdominal terga 5-7 have trian­gular shaped markings in A. albrighti would actually have been applicable to A. neavei based on available descriptions at the time. If workers had used the macu­lation characters as stated in the Allen and Edmunds key, in the absence offurther checking of formal descriptions, then identifications certainly could have been, and probably were, confused. Ostensibly, A. neavei and A. albrighti differed in the intensity of th bro\\'I1 coloration of the meso thorax in the adults, in slight size differences, and i the abdominal tergal patterning. No differences in male genitalia and no larval struc­tural differences (generally the important species characters in many Ephemeroptera species) had been identified, and I have not been able to find any. My examination ofmature larval and adult specimens belonging to popu­lations from northern Saskatchewan to New Mexicdemonstrate no reliable size differences that could be correlated wi h the coloration patterns th t Allen and Edmunds assigned toA. albrighti and A. neavei, and certainly the "usually less than 16 mm" (A. albrighti) vs. "usually greater than 17 mm" (A. neavei) could not be applied with any consistency. Ametropus albrighti adults were thought to have dark brown mesonota, and larvae and adults were thought to have inverted T-shaped markings on middle abdominal terga. Ametropus neauei were thought to have light brown mesonota in the adults and triangular markings on middle abdominal terga in larvae and adults. However, whereas some populations I examined appeared to be consistent with respect to thoracic or abdominal characteristics that had been associated either withA. albrighti orA. neauei, a few populations showed a mixture of shape and degree of abdominal markings or mixture of thoracic color intensity. In addition, most adults with a dark brown thorax, as previ­ously associated with A. albrighti, also possessed well maculated abdominal terga (i.e., extensive triangular markings) as previously associated with A. neauei, The variation found on abdominal tergum 6 of three male adults of the same population from the North Saskatchewan River is shown in Figures 1-3. 2The Great Lakes Entomologist, Vol. 34, No. 2 [2001], Art. 1https://scholar.valpo.edu/tgle/vol34/iss2/12001 THE GREAT LAKES ENTOMOLOGIST 3 It is strongly suggested from this that the difference between an inverted T and the triangle described by Allen and Edmunds is a matter of degree of macula­tion. The one larva I have from far eastern Montana has tergal patterning that is intermediate between the two extremes (close t Figure 2). Such degrees of maculation are often intraspecifically variable among populations, within popu­lations, or may be influenced by temperature of developmental regime over time (e.g., see McCafferty and Periera 1984). Allen and Edmunds (1976) also stated that abdominal tergum 2 in A. albrighti had an inverted T-shaped marking. In all specimens that would oth­erwise appear to be A. al righti based on most other characteristics, this ter­gum always had a variously developed triangular marking, including the lar­val specimens identified and examined as A. albrighti by Allen and Edmunds (1976). The dark brown medial stripe described for tergum 9 ofA. neavei may or may not be present in individual larvae of the same populations of both of what otherwise would appear to be A. neavei or A. albrighti. Based on all of the above, it is apparent that A. albrighti and A. neavei represent a single North American species that is structurally stable but incon­sistently variable with respect to degree of maculation. Therefore, I here place A. albrighti as a subjective junior synonym of A. neavei, new synonym. This synonymy obviates the potential problems with the as ignments of distribution records commented on above. Th tw now-recognized North Ameri­can species, A. ammophilus and A. neavei, are not only easily characterized from each other, but they appear, to a large degree, to be allopatric. A ammophilus has been reported from the Pacific Coast states and Idaho and Montana, as mentioned previously. Although both North American species possibly occur in the Athabasca River region of Alberta (J. M. Webb, pers. comm.), the only other state or province in which they both have been reported to occur is Montana, where they are clearly disjunct. In Montana, the records associated with A. ammophilus are from Missoula and Lake Counties (Allen and Edmunds 1976), both in the mountainous western part ofthat state, whereas the records associ­ated ·withA. neavei are from Custer (Waltz et al. 1998) and Hill Counties (see Material Examined below), from the eastern and central plains area of the state, respectively. In addition to Alberta and central and eastern Montana, Ametropus neavei is now known from western Colorado, northern Michigan, north­ern New Mexico, the Northwest Territories, Saskatch wan, northeastern Utah, and southwestern Wyoming. 1 2 3 Figures, 1-3, Ametropus neavei male adults, single Saskatchewan population, abdominal tergum 6: Fig.l, individual variant 1; Fig. 2, individual variant 2; Fig. 3, individual variant 3. 3McCafferty: Commentary on Ametropus Species (Ephemeroptera: AmetropodiPublished by ValpoScholar, 20014 THE GREAT LAKES ENTOMOLOGIST Vol. 34, No.2 Distributions of certain other mayflies also encompass ranges from New Mexico to Saskatchewan, for example, Camelobaetidius warreni (see McCafferty and Randolph 2000) and Lachlania saskatchewanensis (see McCafferty et a1. 1997). Such patterns are explicable in view of h storical biogeography that shows connections between th Pleistocene Saskatchewan, Columbia, and Colo­rado drainage systems (see Lehmkuhl 1976b). It remains to be seen ifA. neavei still exists in New Mexico or if has been recently extirpated as a result ofim­poundments on the San Juan River (see McCafferty et a1. 1997). It would also be important to determine if populations still exist in the Green River in Utah below Flaming Gorge Dam. The somewhat rare western psammophilous spe­cies Analetris eximia Edmunds of the family Acanthametropodidae has been taken at the same locales as A. neavei in Wyoming (at the Black's Fork River [Edmunds and Koss 1972]), the same habitat in Saskatchewan (in the Saskatchewan River [Lehmkuhl 1976b]), the same locale in Alberta (in the Athabasca River [Barton 19801), and from the sa e locale in Montana (reported here for the first time from the Milk River, see material examined below). MATERIAL EXAMINED All of the following materials are deposited in the Purdue Entomological Research Collection, West Lafayette, Indi na. Ametropus ammophilus: Mon­tana, Lake Co., Swan R., 4 mi S wan River City, VlI-12-1965 (one larva, one male adultllarval exuviae reared); Oregon, Jackson Co., Rogue R. at j nction St. Hwys. 230 & 62, VlII-24-1954, Edmunds and Allen (one larval exuvia paratype), Deschutes Co., small stream at Lapine, IV-18-1964, M. O. Roberts (three larval paratypes); Washington, Whatcom Co., Nooksack R., nr. Lynden, V-10-1967, K. E. Vander Mey (male adult holotype, two female subimago paratypes, one male adult paratype, four male subimago paratypes, four larval paratypes, nine larval exuviae), V-26-1967 (one male adultllarval exuviae reared). Ametropus neavei: Colorado, Moffat Co., Deerlodge Park, Dinosaur National Monument, 5600 ft., VIII-1-1962, D. Q. Anderson (one larva); Mon­tana, Custer Co., Powder R., XI-1l-1976, G. Romero (one larva), Hill Co., Milk R. above St. John's Br., X-23-1999, D. L. Gustafson (five larvae) (with numer­ous co-occurring larvae ofAnaletris eximia); New Mexico, Farmington, V-1-1935, P. N. Albright (one larva); S skatchewan, North Saskatchewan R., Borden, Hwy 5, V-19-1974, Vl-4-1974, R. Demaray (three female adultsllarval exuviae reared, 16 larvae, three male adultsllarval exuviae reared); Utah, Uintah Co., Green R. at Jensen, Vl-5-1963, Edmunds, Jensen and Peters (two larvae, one male subimagollarval exuviae reared); Wyoming, Sweetwater Co., Black's Fork River at 1-80 west of Green River City, VII-6-1968, R. & D. Koss (five larvae, one male adultllarval exuviae reared, one female adultllarval exuviae reared), VlII-2-1969, A. V. Provonsha (25 larvae), Black's Fork R., 1-80 6 mi E Little America, S. L. Jensen and A. V. Provonsha (three larvae), Green River, Vl-4­1959, Smith and Musser (one larva). ACKNOWLEDGMENTS I thank Luke Jacobus (West Lafayette, IN) for illustrations and review of the manuscript and Pat Randolph (West Lafayette, IN) for his review of the manuscript. Research was supported in part by NSF Grant DEB 9901577. The paper has been given Purdue ARP Journal No. 16450. LITERATURE CITED Albarda, H. 1878. Descriptions of three new European Ephemeridae. Entomologists Monthly Magazine 15:128-130. Allen, R. K. 1977. New records of mayflies (Ephemeroptera) from California and Mexico. Pan-Pacific Entomologist 53:210. 4The Great Lakes Entomologist, Vol. 34, No. 2 [2001], Art. 1https://scholar.valpo.edu/tgle/vol34/iss2/12001 THE GREAT LAKES ENTOMOLOGIST 5 Allen, R. K. and G. F. Edmunds, Jr. 1956. A list of mayflies of Oregon. Proceedings of the utah Academy of Sciences, Arts and Letters 33:85-87. Allen, R. K. and G. F. Edmunds, Jr. 1976. A revision of the genus Ametropus in North America (Ephemeroptera: Ametropodidae).• Journal of the Kansas Entomological Society 49:625-635. Barton, D. R. 1980. Observations on the life histories and biology of Ephemeroptera and Plecoptera in northeastern Alberta. Aquatic Insects 2:97-111. Bengtsson, S. 1913. Undersokningar ofver aggen hos Ephemeriderna. Entomologisk Tidskrift 34:271-320. Clifford, H. F. and D. R. Barton. 1979. Ohservations on the biology ofAmetropus neavei (Ephemeroptera: Ametropodidae) from a large river in northern Alberta, Canada. Canadian Entomologist 111 :855-858. Cobb, D. G., J. F. Flannagan, P. M. L. Flannagan and R. D. Wickstrom. 1995. Collections of Ephemeroptera from Kluane National Park, Yukon Territory, and Nahanni Na­tional Park, Northwest Territories, pp. 177-184. In: L. D. Corkum and J. J. H. Ciborowski (eds.), Cu rent directions in research on Ephemeroptera. Canadian Schol­ars' Press. Toronto, 478 pp. Dosdall, L. M. and D. M. Lehmkuhl. 1989. Drift of aquatic insects following methoxychlor treatment ofthe Saskatchewan River system. Canadian Entomologist 121:1077-1096. Edmunds, G. F ... Jr. 1954. The mayflies of Utah. Proceedings of the Utah Academy of Sciences, Arts and Letters 31:64-66. Edmunds, G. F., Jr. and R. W. Koss. 1972. A review of the Acanthametropodinae with a description of a new genus (Ephemeroptera: Siphlonuridae). Pan-Pacific Ento­mologist 48:136-144. Edmunds, G. F., Jr., S. L. Jensen and L. Berner. 1976. Mayflies of North and Central America. University of Minnesota Press. Minneapolis, 330 pp.. Lehmkuhl, D. M. 1976a. Mayflies. Blue Jay 34:70-81. Lehmkuhl, D. M. 1976b. Additions to the taxonomy, zoogeography, and biology ofAnaletr s eximia (Acanthametropodinae: Siphlonuridae: Ephemeroptera). Canadian Entomolo­gist 108:199-207. Lester, G. T., W. P. McCafferty, and M. R. Edmondson. 2002.1\ ew mayfly (Ephemeroptera) records from Idaho. EntomoL News 113:131-136. McCafferty, W. P. 1981. Aquatic entomology, J nes and Bartlett Publishers. Boston, 448 pp. McCafferty, W. P. 1991. Toward a phylogenetic classification of the Ephemeroptera (Insecta): a commentary on systematics. Annals of the Entomological Society of America 84:343-360. McCafferty, W. P. and C. Pereira. 1984. Effects of developmental th rmal regimes on two mayfly species and their taxonomic in erpretat on. Annals of the Entomological Society of America 77:69-87. McCafferty, W. P. and R. P. Randolph. 2000, Further contributions to the spatulate clawed Baetidae (Ephemeroptera). Entomological News 111 :259-264. McCafferty, W. P., C. R Lugo-Ortiz and G. Z .• Jacobi.1997. Mayfly fauna of New Mexico. The Great Basin Naturalist 57:283-314. McDunnough, J. 1928. The Ephemeroptera of ,Jasper Park, Alta. CaIladian Entomolo­gist 60:8-10. Newell, R. L. 1970. Checklist of some aquatic insects from Montana, Proceedings of the Montana Academy of Sciences. 30: 45-56. Pearson, W. D., R. H. Kramer andD. R. };'ranklin.1968. Macroinvertebrates in the Green River below Flaming Gorge Dam, 1964-1965 and 1967. Proceedings of th Utah Academy of Sciences, Arts and Letters. 45:148-167. 5McCafferty: Commentary on Ametropus Species (Ephemeroptera: AmetropodiPublished by ValpoScholar, 20016 THE GREAT LAKES ENTOMOLOGIST Vol. 34, No.2 Soluk, D. A. and D. A. Craig. 1988. Vortex feeding from pits in the sand: a unique method of suspension feeding used by a stream invertebrate. Limnology and Oceanography 33:638-645. Steven, J. C. and W. L. Hilsenhoff. 1979. Ametropus neavei (Ephemeroptera: Ametropodidae) in the Upper Peninsula of Michigan. The Great Lakes Entomolo­gist 12:226. Traver, J. R. 1935. Part II: Systematic, pp. 237-739. In: J. G. Needham, J. R. Traver and Y-C. Hsu (eds.), The biology of mayflies. Comstock Publishing Company. Ithaca, New York, 759 pp. Waltz. R. D., G. F. Edmunds, Jr. and G. Lester. 1998. New distributions for R'aptoheptagenia cruentata and Ametropus neavei (Ephem roptera: Heptageniidae, Ametropodidae). Entomological News 109:213-214. Woodbury, A. M. and D. W. Argyle. 1963. Ecology and limn gy of Green River. Univer­sity of Utah Division of Biological Sciences Miscellaneous Papers 1:7-48. 6The Great Lakes Entomologist, Vol. 34, No. 2 [2001], Art. 1https://scholar.valpo.edu/tgle/vol34/iss2/1

Commentary on \u3ci\u3eAmetropus\u3c/i\u3e Species (Ephemeroptera: Ametropodidae) in North America

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Commentary on \u3ci\u3eAmetropus\u3c/i\u3e Species (Ephemeroptera: Ametropodidae) in North America

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