PROC. ENTOMOL. SOC. WASH. 111(1), 2009, pp. 57-67

TRICORYTHODES MINUTUS TRAVER, A NEW SYNONYM OF TRICORYTHODES EXPLICATUS EATON (EPHEMEROPTERA: )

DAVIDE. BAUMGARDNER

Department of Entomology, Texas A&M University, College Station, Texas 77843-2475, U.S.A (e-mail: [email protected])

Abstract.-Based upon a study of adults, larvae, and reared adults from throughout North and Central America, Tricorythodes minutus Traver, 1935 is shown to be a junior synonym of Tricorythodes explicatus (Eaton, 1892) (n. syn.). Previous characters postulated to separate the two species are shown to be variable and unreliable. This study has clearly revealed the presence of a single, widespread and morphologically variable species distributed throughout much of western, central, and northeastern North America and northern Central America. Key Words: Ephemeroptera, Leptohyphidae, Tricorythodes explicatus, Tricor­ ythodes minutus,

Among Nearctic Tricorythodes, the justify the synonymy of T. minutus with exact identities of T. explicatus (Eaton) T. explicatus. and T. minutus Traver have remained vague and uncertain. Although Kilgore MATERIALS AND METHODS and Allen (1973) and Allen and Murvosh Collections (and their acronyms) ( 1987) described the larval stages of these housing materials used in this study two species and contrasted their larval include: The Natural History Museum, characters, much taxonomic confusion London (BMNH), C.P. Gillette Muse­ still remained. The possibility that these um of Diversity, Colorado two species might be synonymous has States University, Ft. Collins (CSU) and been mentioned in the literature several Texas A&M University, College Station times (Lugo-Ortiz and McCafferty 1995, (T AMU). In material examined sum­ McCafferty et al. 1997, Baumgardner maries, larval collections are abbreviat­ and Bowles 2005). After an extensive ed by the capital letter "L," preceded by study of numerous specimens from the number of specimens examined. throughout North and Central America Collectors are identified by the follow­ (which included larvae and reared ing initials: DEB (David E. Baumgard­ adults), it is apparent that both are the ner); BCK (Boris C. Kondratieff). Legs, same species. The purpose of this paper gills, and mouthparts of over 40 speci­ is to trace the historical taxonomy of the mens were removed and mounted on two species (and two associated species slides for detailed study. Figures were synonymized with T. minutus), and drawn using a camera lucida. Global positioning measures are given in longi­ * Accepted by David R. Smith tude/latitude coordinates as degrees, 58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON minutes, seconds. Seta] descriptive ter­ labeled as such in The Natural History minology follows Baumgardner and Museum. Ulmer (1920) established the Avila (2006). genus Tricorythodes and designated Tri­ corythus explicatus as the type species. RES ULTS AND DISCUSSION Banks (1903) documented the species for Tricorythodes explicatus was described the first time in the United States from by Eaton (1892), as Tricorythus explica­ Arizona, and McDunnough (1931) listed tus, based upon a series of adults it as occurring in Texas and Wyoming in collected by Herbert Knowles Morrison the United States and Saskatchewan and (1854-1885) from "N. Sonora, Mexico." Alberta in Canada. Traver (1935) ques­ However, it appears highly likely that the tioned the identification of the specimens type series was collected from the Gra­ from Canada, while Allen and Murvosh ham Mountains north of Fort Huachuca (1987) considered them to be T minutus. in extreme southern Arizona (Selander Spieth (194 I) indicated that the species and Valurie 1962). Morrison was a occurred as far north as Yellowstone professional collector who collected in Park in Wyoming. Allen and Murvosh Arizona in I 880 and 1882 (Essig 1931 ). (I 987) described the larval stage of T Horn (1886) notes that "The collection explicatus based upon specimens collect­ made by Morrison teaches very little. ed from what they believed was the type How far he went below the boundary I locality of the species in Sonora, Mexico. do not know, although my series was They also included a taxonomic key to purchased as from Arizona purely, and separate T explicatus from T minutus. the same series is quoted by the authors Tricorythodes minutus was described of the "Biologia" as from northern by Traver (1935) from adults from the Sonora." It appears that perhaps Morri­ Provo River in Utah. It was distin­ son, as a commercial collector, labeled guished from other known Tricorythodes his specimens from extreme southern adults on the basis of coloration (paler Arizona as being from northern Sonora than T stygiatus McDunnough) and (Mexico) so that he could then sell them lack of black maculation on the femora to the Biologia researchers as being from and tibiae (present on T allectus (Need­ their area of interest. To the American ham)) and from T texanus Traver due to scientists, he apparently gave them the its much darker coloration and more correct Arizona locality. Despite this shallow excavation of the apical margin apparent error in labeling, Essig ( 1931) of the styliger plate. Kilgore and Allen considered Morrison one of the greatest (1973) described the larval stage of T American entomological explorers, with minutus and listed it as occurring extensive collections made throughout throughout Arizona and New Mexico. the western and southeastern United Among characters they listed for the States. Morrison contracted a fatal case larvae of T minutus were a 3-segmented of dysentery on his last collecting trip to maxillary palp, presence of long margin­ Key West, Florida in 1885 (Essig 1931). al setae on the femora, abdominal terga The original brief description by Ea­ 7-9 with well-developed posterolateral ton (1892) focused upon coloration and projections, and operculate gills pale included two figures, one of the wing and with black markings (Kilgore and Allen one of the forceps (without the penes). I 973: fig. 37). Tricorythodes minutus was Since a holotype was not designated in subsequently reported from throughout the original type series, Kimmins (1934) western North America (Allen and Mur­ designated a specimen from this series as vosh 1987, McCafferty et al. 1993, Lugo­ the lectotype. This specimen is clearly Ortiz and McCafferty 1995, Mccafferty VOLUME 111, NUMBER 1 59

et al. 1997). It has also been reported varieties occurring in what would nor­ from north central and northeastern mally be considered the range of T North America (Alba-Tercedor and explicatus or T minutus, and can find Flannagan 1995, Randolph and McCaff­ no reliable characters in the adult stage erty 1998). separating the two species. Two other species of Tricorythodes, T As no characters were found to fa/lax Traver and T fallacina McDun­ separate the two species in the adult, nough, had previously been synonymized the larval stages of each species were with T minutus. Tricorythodes Jal/ax was examined. Although the key of Allen and described by Traver (1935) from adults Murvosh (1987) and characters given for from California, and its overall similarity the larval stage of T minutus by Kilgore to T explicatus was noted by Traver who and Allen (1973) appear to offer many distinguished it by its paler coloration. distinguishing characters, these charac­ Edmunds et al. (1976) synonymized T ters proved unreliable and sometimes Jal/ax with T. minutus because the two incorrectly applied. species completely intergraded along the Firstly, Kilgore and Allen (1973) borders of California, Oregon, and indicated that T minutus possessed a 3- Washington. They considered T. Jal/ax segmented maxillary palp, whereas Allen a "clinal variant" of T. minutus. and Murvosh (1987) indicated that T Tricorythodes fallacina was described explicatus possessed a 2-segmented max­ by McDunnough (1939) from adults illary palp. Examination of numerous from British Columbia, Canada. The larvae from throughout the reported new species was considered a "smaller, range of T minutus showed that all more northerly species" within the T possessed a 2-segmented maxillary palp explicatus complex. McDunnough ( 1939) with a single apical seta (Fig. 1); howev­ considered T explicatus to be a large er, some individuals possessed a weakly dark species occurring in Texas and developed articulation near the base of adjoining regions and T. Jal/ax a smaller, the second segment, giving the appear­ paler western species. Traver (1935) and ance of a 3-segmented maxillary palp McDunnough (1939) noted that all three (Fig. 2). This condition of a weakly species shared morphological characters articulated palp has been demonstrated such as the type and shape of genitalia for other leptohyphid such as and a shallow excavation of the apical Vacupernius packeri (Allen) (Baumgard­ margin of the styliger plate. Edmunds ner 2003) and some South American and Allen (1957) synonymized T falla­ Tricorythodes (Molineri 2002). None of cina with T minutus without comment. the larvae examined possessed a 3- However, this synonymy appears to be segmented maxillary palp; all possessed well justified on the basis of discussion a 2-segmented maxillary palp, with given in Edmunds et al. (1976) discussing about 10% of the larvae having traces the clinal variants of T fa/lax, T. of the weakly developed articulation. fallacina, and T. minutus. Secondly, Kilgore and Allen (1973) Thus, characters used to separate the indicated that T minutus possess well­ adult stages of T explicatus from T developed posterolateral projections on minutus are few and based upon differ­ abdominal terga 7-9, whereas in T. ences in overall size and coloration. I explicatus, terga 7-9 lacked well-devel­ have examined long series of adults from oped posterolateral projections (Allen western North America, Mexico, and and Murvosh 1987). However, examina­ Guatemala which would be attributed to tion of numerous larvae throughout the T. explicatus and T minutus, including reported range of T. explicatus clearly

L' 60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

I~

.· .. · .· ... . : .• 6 2

Figs. 1-6. Tricorythodes explicatus, mature larva. I, Maxillary palp [Scale bar AJ. 2, Maxillary palp (arrow indicates location of weak suture line) [AJ. 3, Operculate gill (dorsal), "typical form" [BJ. 4, Operculate gill (dorsal), "pale form" [BJ. 5, Maxilla [DJ. 6, Left mandible [CJ. Scale bars (mm): A, D = 0.1; B = 0.5; C=0.2. indicated some variation in the degree of 7-9, in addition to a 2-segmented maxil­ development of the posterolateral projec­ lary palp. These larvae were identical to tions on abdominal terga 7-9, including T. minutus larvae throughout the western both larval instar and gender. For exam­ United States. In general, more mature ple, examination of almost 100 male and larvae, and female larvae, possessed more female larvae from Queretaro, Mexico extensively developed posterolateral pro­ (located in south-central Mexico) pos­ jections on terga 7-9 than male and/or sessed posterolateral projections on terga immature larvae. VOLUME 111, NUMBER 1 61

Thirdly, Allen and Murvosh (1987) characters proposed by Allen and Mur­ distinguished T. explicatus larvae from vosh (1987) to separate the two species in T. minutus larvae based upon leg seta­ the larval stage were artificial or errone­ tion. Their key indicated that T. explica­ ous. Given the lack of any distinct, stable tus possessed a "double row of long, morphological characters to separate the heavy spines" on the inner margin of the species, it is clear that T. minutus is a front and middle tibiae and had "short, synonym of T. explicatus and that a heavy spines on the inner margin of all single, widely distributed and morpho­ tarsi" (Allen and Murvosh 1987: fig. 1), logically variable species is present while T. minutus larvae lacked these throughout western, north-central, and spinelike setae on the inner margin of northeastern North America. the front and middle tibiae and tarsi Perhaps the most distinctive morpho­ (Allen and Murvosh 1987: fig. 3). Both logical feature present on larvae of T. comparisons were incorrect. While lar­ explicatus is the pale, triangular shaped vae within what is considered the range macula on the operculate gill (Fig. 3). of T. explicatus did possess the described This feature will readily distinguish T. leg setation, so did all larvae examined explicatus larvae from all other known from within what is considered the range Tricorythodes larvae in North America. of T. minutus. In addition, all known It is important to note, however, that this species of Tricorythodes larvae in North feature is somewhat variable among America possess setae on the inner individual specimens. As some larvae margin of the tibiae and tarsi. If these reach the final instar, the black maculae tarsi become broken away, or the leg is largely disappear, and the operculate gill slide mounted in such a way as they are becomes mostly pale, with black maculae not visible, the spinelike setae may confined to the base and outer margin appear absent. However, careful exami­ (Fig. 4). This phenomenon has been nation will reveal sockets from which the observed for a number of individuals in setae arise. It appears that either Allen a variety of populations. and Murvosh (1987) were examining Tricorythodes explicatus is a highly larvae with these setae broken away or variable species in terms of its body size perhaps were not examining the legs and adult coloration. During the course under high magnification. In any case, of this study, it was extremely rare to the characters used to separate these two find large series of individuals from a species in the larval stages are clearly single population that clearly match a artificial. description of T. exp/icatus, T. minutus Finally, the figure used by Allen and or T fallax. It was more common to find Murvosh (1987: fig. 2) to diagram the a mix of individual specimens matching operculate gill of T explicatus appears to the various concepts of each of these be the same exact figure used by Kilgore species. The "pale gill" forms were often and Allen (1973: fig. 37) to illustrate the small individuals with very pale adult operculate gill of T minutus. coloration, the reared adults of which Almost all larvae examined from what were similar in coloration to that de­ Allen and Murvosh (1987) considered scribed for T. fallax; however, these the range of T. minutus matched the individuals were typical of the size range description of T. explicatus and not T. of T. minutus. The T. explicatus form minutus. Those larvae examined from (large and very dark in coloration, i.e., central Mexico and Guatemala also browns and blacks) is common in match the description of T. exp/icatus collections from numerous locations and not T. minutus. It appears that the from the extreme southwestern United 1

62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 7-8. Tricorythodes explicatus, lectotype male imago. 7, Penes (dorsal). Scale bar = 0.1 mm. 8, Forewing. Scale bar = 0.5 mm.

States, through the Rocky Mountains, tion of the apical margin of the sty­ and did appear to be the most common liger plate was present in all adult variant. Larval characteristics of the specimens examined (Fig. 7). The lee maxilla, mandibles and the two opercu­ totype and paralectotypes of T. explica­ late gill patterns remain constant in tus were of the T. minutus size with individuals from throughout the range the coloration as in the description of of T. explicatus. The shallow excava- T. explicatus. This was yet another VOLUME 111, NUMBER I 63 indication of the equivalency of these segmented (Fig. 2); some individuals species. with faint annulation giving appearance In summary, characters postulated by of 3-segmented maxillary palp (Fig. 2); Allen and Murvosh (1987), such as the (3) mandibles with numerous, elongate number of segments of the maxillary setae along out margin (Fig. 6); (4) palps, and differences in leg setation and abdominal terga 2-9 with extensive black abdominal posterolateral projections are coloration across dorsum of each ter­ unreliable and largely artificial charac­ gum, contrasting strongly with white to ters. Characters proposed to separate the pale yellow base coloration of each adults of the two species are based upon abdominal segment; segment 10 with size and coloration differences and are little, if any, coloration. not sufficient to diagnose two distinct In the adult stage, T explicatus can be species. Examination of large numbers of distinguished from all other North and individuals throughout the range of the Central American species by the follow­ species has clearly demonstrated a single, ing combination of characters: (1) penes morphologically variable and wide­ of typical Tricorythodes form but with spread species. The known range of T very shallow excavation of apical margin explicatus includes southern Canada, of styliger plate (Fig. 7); (2) forewing western, central, and northeastern North (Fig. 8), veins A and CuP well devel­ America, south to Mexico and Guate­ oped, reaching wing margin; vein CuP mala. The larvae can be found in a vast not strongly curved towards A not variety of aquatic habitats from small, meeting at wing margin; vein 1Cu2 well temporary streams to larger rivers. developed, free, not meeting with vein 1Cu 1• Tricorythodes explicatus (Eaton) Type material examined.-Tricor­ (Figs. 1-8) ythodes minutus Traver. Holotype: male imago (Cornell University). Type Local­ Tricorythus explicatus Eaton 1892: 11. ity: USA: Utah: Provo River, 22.vii. Tricorythodes explicatus: Ulmer 1920: 51 1926. Condition poor, faded, with wings (transferred to Tricorythodes). missing; preserved in ethyl alcohol. Tricorythodes fallax Traver 1935: 634; Paratypes: Same data as holotype; 8 Edmunds et al. 1976: 259 (synonym of male imagos, all in poor condition. T minutus). Tricorythodes explicatus (Eaton). Lec­ Tricorythodes fallacina McDunnough 1939: totype (designated by Kimmins 1934): 53; Edmunds and Allen 1957: 322 (syn­ male imago. Verbatim label data [brack­ onym of T minutus). eted information added]: (1) Type [round Tricorythodes minutus Traver 1935: 636. label with red circle along border]. (2) N. N. syn. Sonora, I Mexico. I Morrison. (3) Brit. Diagnosis.-In the larval stage, T Mus. I 1905-88. (4) B.C.A. Neuropt. I explicatus can be distinguished from all Tricorythus I explicatus, I Eaton. Pinned, other North and Central American in poor condition; thorax and right species by the following combination of forewing are all that remains on pin; 3 characters: (l) operculate gill with prom­ slides which include: (1) left forewing; (2) inent basal ridge and much less promi­ 1 set of legs; (3) approximately 2/3rd of nent external ridge; often with large, abdomen and genitalia. Paralectotypes: 4 pale macula at base of gill (Fig. 3); (2) male imagos, pinned, in excellent condi­ maxilla with numerous elongate setae tion; collection data and all other labels along outer basal margin below maxil­ exactly same as lectotype specimen, lary palp (Fig. 5); maxillary palp 2- except each specimen with a circular 64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON paratype label with faded yellow circle Oak Creek at Bootlegger Camp along border (added by Kimmins 1934). Ground, Hwy. 89A, Oak Creek Canyon Other material examined.-GUATE­ (N34°58'09.4"; Wl 11°45'01", elev. MALA: BAJA VERAPEZ: Rio La Es­ 5210 ft.), 24.v.2004, 20 L, 2'? (reared), tancia at Hwy. 17, Salana (Nl5°05'54"; 16 (reared), DEB (TAMU). Graham Co.; W90°18' 18", elev. 3000 ft), 07.i .2007, Gila R. at Ft. Thomas Road, Ft. Thomas 55 L, DEB (T AMU). SANTA ROSA: (N33°02'59"; Wl09°58'01, elev. 2660 ft.), unnamed creek at Hwy. 16, ca. 13.2 Km 27.v.2004, 1 L, DEB (TAMU). Greenlee S. from Hwy. 16/CA I jct, between Co.; San Francisco R. at FS Rd. 212, Km marker 78 & 79 (NI4°1 I '51"; ca. I mi. N Clifton (N33°04'30"; W90°21'38", elev. 2095 ft), 06.i.2007, Wl09°18'03", elev. 3700 ft.), 21.v.2004, 14 L (TAMU). MEXICO: AGUASCA­ > 100 L, DEB (TAMU). Santa Cruz Co.; LIENTES: Sabinolandia, Rio San Pedro, Patagonia State Park, Sonoita Creek, El Saito de los Salados (N21.85; W02.49), 22.iii.1987, 3 L, R. Leschen (TAMU); 06.xii.1997, 2 L (TAMU). MEXICO: Sonita Cr. at Blue Haven Rd., just off stream at 56 km. between Toluca and Hwy. 82, ca I mi. SW Patagonia Zitaeuaro 4 km. N.E. Bosencheue (elev. (N31°30'57"; WI 10°47'35"), 06.vi.2000, 8200 feet), 24. viii.1977, 4 L, RK Allen 2 L, DEB (TAMU); Sonoita Cr., nr. (CAS). QUERETARO: Puerta de Ale­ Patagonia, l 5.iii.1997, I 6, J. Slusark & griax, Arroyo Los Zunigas (N20°20'28"; K. Byrnes (TAMU). Yavapai Co.; Bub­ Wl00°07' 10", elev. 2000 m), 08.vii.2000, bling Springs off Forest Service Rd. 134, I L, WDS (TAMU); I km S. San Miguel ca. I kn N. Page Springs, 22.iv .1993, 4 L, Palmas, Rio Victoria (N21°05' 13"; S.R. Moulton and K.W. Stewart W99°57' 10"), 09.vii.2000, IOO+L, WDS (TAMU); Oak Cr. at Page Springs Rd (TAMU); Bucareli, Rio Estorax (Co. Rd. 50), Page Springs (N34°45'57"; (N21°02'05"; W99°37'03"), l 1.vii.2000, WI 11°53'27", elev. 3455 ft), 25.v.2004, 12 L, WDS (TAMU); I Km. S. Huasqui­ >75 L, DEB (TAMU); Verde R. @ lico, Arroyo Jalpan (N21°09'04"; Hwy. 260, Camp Verde (N34°32'60"; W99°34'42", elev. 1725 m), l 1.vii.2000, WI 11°51 '00", elev. 3140 ft.), 23.v.1999, I L, WDS (TAMU). 46, 8'? (reared), 38 L, DEB (TAMU); UNITED STA TES: ARIZONA: Co­ Verde R. @ Hwy. 260, Camp Verde chise Co.; San Pedro R., Luis Springs, (N34°32'60"; Wlll 0 51'00", elev. 3140 17.iv.2001, 226, D. Rees (TAMU); San ft.), 23.v.1999, 66 (reared), 9'? (reared), Pedro River, Hereford Rd. Bridge, 25 L, DEB (TAMU); Verde R. @ Per­ 26.xi.1999, 146, A.B. Richards (TAMU); kinsville bridge, ca. 16 mi. N. Jerome on San Pedro River, Rt. 80, Saint David, FSR 318, Prescott Nat. Forest, 6.vi.1993, l.v.1995, 21 L, BCK (TAMU); Cochise 2 L, S.R. Moulton and K.D. Alexander Co, San Pedro River, Riparian Nat. Cons. (TAMU); Verde River@ US Hwy. 89A Area, Rt. 90, 30.iv.1995, >1006, BCK bridge, Cottonwood, 4-5.vi.1993, 38 6 (CSU). Coconino Co.; Oak Cr. at Hwy. and '? imagos, S.R. Moulton and K.D. 89A, ca. 5 mi. N. Sedona (N34°55'23"; Alexander (TAMU); West Clear Creek at WI 11°44'01"), 04.vi.2000, 4 L, DEB Clear Creek Campground, ca. I mi. from (TAMU); Oak Cr. at Manzanita Camp Hwy. 260, ca. 4 mi SE Camp Verde Ground (Hwy. 89), Coconino Nat. Forest (N34°30'55"; Wlll 0 45'45", elev. 3270 (N34°56'04.1"; Wlll 0 44'46.0"), 25-26. ft.), 26.v.2004, 11 L, DEB (TAMU); East v.1999, 16 L, DEB (TAMU); Oak Cr. at Verde River, Camp Verde, 29.iv.1995, Pine Flat Camp Ground (Hwy. 89) 2 L, BCK (CSU); Oak Creek, Red Rock (N35°00'52"; Wl 11°44'26"), 03-05.vi. Crossing, 20.i.1988, 1 L, 1 6, 1 '?, BCK 2000, 15 L, 3 '? (reared), DEB (TAMU); (CSU). COLORADO: Arapahoe Co.; VOLUME Ill, NUMBER I 65

South Platte R., Mineral Rd., 23.v.1993, 7 L, BCK (CSU); Mancos River, Rt. 25 6', S. Fitzgerald (TAMU). Archuleta 666, 18.vii.1990, 536', BCK (CSU). Rio Co, Piedra River, Rd. 193, l.vii.1996, Blanco Co.; Douglas Creek @ Rt. 64, 266', BCK (CSU). Boulder Co.; South 19.vii.1996, 23 L, 186', 15SJ, BCK & R. Boulder Creek, Baseline Rd., 3.x.1995, Durfee (TAMU). Saguache Co, Indian 166', BCK (TAMU). Chaffee Co, Trout Springs NA, 15.viii.1999, 16', BCK Cr, F.R. 215 and US 285, 4.ix.1993, 56', (CSU). Weld Co, 9.ix.1986, 1 6', D. BCK & R. Durfee (CSU). Costilla Co, Thompson (CSU); Lone Tree Creek, Trinchera Creek, 12 Road, 30. vi.1996, Terry Ranch, 10 mi. NW Carr, 17.vi. 9 L, BCK (CSU). Douglas Co.; Deckers 1989, 4 6', 1 6' , P.A. Opler (CSU). Yuma Resort, BLT, 23.viii.1988, 66', P.A. Opler Co.; S. Fork Republican R., US Hwy. 385 (CU); South Plate, Trumbull, 1O.ix.1993, bridge, 29.iv.1993, 18 L, BCK & R. 126', lOSJ, K. Rogers, (CSU). Garfield Durfee (TAMU); Arikaree River, Bow­ Co, 31.viii.1975, 16', lSJ, D.E. Ruiter man Ranch, 23.vi.1999, 1 L, G. Doyle, (CSU). Gunnison Co.; Gunnison River, (CSU). MISSOURI: Dallas Co., Niangua Riverway Picnic Area, 3 mi. W. of Gun­ R. at Hwy. M, 3.5 mi. E.jct. with Hwy. 38 nison, 31. viii.1991, 22 6', BCK, R. Durfee ( N37°31'1 l"; W92°59'03", elev. 1,120 ft), (TAMU). Jackson Co.; Grizzly Creek, 10.viii.2005, 3 L, DEB (TAMU). MON­ Peterson SWA, off Rt. 14, 28.vii.1991, T ANA: Big Horn Co, Big Hom River, 4 L, R. Durfee & B. Painter (TAMU); 5.ix.1991, 176', D.E. Ruiter (TAMU). North Platte River, Verner SWA, 7 mi. NEBRASKA: Garden Co, Blue Ck, SW of Walden, 22. viii.1991, 42 J, R. Rackett Rd., 22. v.1998, 3 6', 8 SJ, BCK Durfee (CSU). Kit Carson Co, South (TAMU). Keith Co, Otter Ck., St. Hwy. Fork Republican, N of Stratton, 92, 22.v.1998, 11 L, 2SJ, BCK (TAMU). l 7.vii.1986, 4 L, 166', BCK (CSU). Lar­ Sioux Co, Sowbelly Canyon, 6 mi. NE of imer Co.; Buckhorn Creek, about 6 miles Harrison, 12.vi.1989, >3006', BCK west of Horsetooth Reservior, l 2ix. l 986, (CSU). NEW MEXICO: Catron Co.; 46', G.W. Gerlich (TAMU); Head of San Francisco R., Gila National Forest, Spring Creek, Stream, l 1.ix.1986, 4 J, D. ca. 5 mi. S. Glenwood (N33°14' 18"; Brigham (TAMU); Mail Cr., BLT, Wl08°52'47, elev. 4560 ft.), 20.v.2004, 14.x.1988, 36', P.A. Opler (TAMU); 2 6' (reared), > 100 larvae, DEB (TAMU); Poudre River, nr. N. Colorado Nature creek at FS Road 233, ca. 5 mi. NE Re­ Area, 21.ix.1986, 6 L, 39 6', BCK serve, Gila National Forest (N33°43'54"; (TAMU); Ft. Collins, 27. vii.1979, 2 6', Wl08°42'28", elev. 5960 ft.), 20.v.2004, D.E. Ruiter (CSU); Heatheridge & Pros­ 1 L, DEB (TAMU); Middle Fork pect, 27.vii.1979, 66', D.E. Ruiter (CSU); Gila R., Gila National Monument Mail Creek, Fort Collins, l O.xiii.1989, (N33°13'47"; Wl08°15'58", elev. 5720 40 6', P.A. Opler (CSU); Poudre R., ft.), 18.v.2004, 7 L, DEB (TAMU); Tay­ Shields St. Br. Ft. Collins, l 4.ix.1986, lor Creek, above Wall Lake, 02.viii.1993, 1 L, 36', S. Klahn (CSU). Moffat Co.; 12 L, >50 adults, R. Durfee (TAMU). Yampa River, Echo Park, Dinosaur NM, Grant Co.; Gila R., Gila Riparian Re­ 19.vii.1993, 186', 4SJ, 2L, BCK & R. serve, Gila Wilderness, ca. 6.7 mi. NE Durfee (CSU); Yampa River, Tee Pee Cliff (N33°02'39"; Wl08°31'47", elev. Campground, Dinosaur NM, 28. vii.1995, 4650 ft.), 19.v.2004, 6 L, DEB (TAMU). 386' (TAMU); Green River, Echo Park, Lincoln Co.; Rio Ruidoso at Main St. in Dinosaur NM, 24.vii.1992, 2 L, DAW Ruidoso (N33°19'14"; Wl05°42'43", elev. (CSU); Yampa R., Craig, 15 L, (CSU). 4000 ft), 22.v.2000, 5 L, DEB (TAMU). Montezuma Co, Mancos River, Mancos Sandoval Co.; Jemez R. at Vista Linda Canyon, Mesa Verde NF, 10.vii.1998, Camp Sight (Santa Fe Nat Forest),

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Hwy. 4, ca. 5 mi. N. Jemez Puebloa 118, ca. 1 mi. W. Ft. Davis (N30°36' 17"; (N35°43'02"; Wl06°43' 17, elev. 5865 ft.), Wl03°53'51, elev. 4943 ft), 14.v.2002, 12.vii.2004, >75 L, 5 6 (reared), 5 1006 imagos, R. Durfee (CSU). Wl04°03'49.9), 24.viii.1996, 4 L, 16, SOUTH DAKOTA: Lawrence Co, Box­ DEB (TAMU). Reeves Co.; unnamed elder Creek, Boxelder Forks Camp­ ck., Hwy 17, Balmorhea (N30°59'13""; ground (N44°1 l '57''; Wl03°32'05"), Wl03°44'35, elev. 3341 ft), 04.v.2002, 12.vii.1997, 96, Baumann and BCK 11 L, DEB (TAMU). Val Verde Co.; (TAMU); Redwater River, S. of Belle Dolan Falls Preserve, Dolan Creek; The Fourche (N44°35'36"; Wl03°52' 10"), Nature Conservancy (N29°53'41", 13.vii.1997, 106, Baumann & BCK Wl00°59'11"), 08.xi.1998, 36, DEB (T AMU). Fall River Co, Fall River, (TAMU). UT AH: Uintah Co, Green Hot Springs, 5.ii.1995, 3 L, 1 6, 1 Mayfly fauna of New Mexico. Allen, R. K. and C. M. Murvosh. 1987. Mayflies Great Basin Naturalist 57: 283-314. (Ephemeroptera: Tricorythidae) of the South­ McDunnough, J. 1931. New North American western United States and Northern Mexico. Caeninae with notes (Ephemeroptera). Cana­ Annals of the Entomological Society of Amer­ dian Entomologist 63: 254--268. ica 80: 35-40. Banks, N. 1903. Neuropteroid from Ar­ ---. 1939. New British Columbian Ephemer­ izona. Proceedings of the Entomological Soci­ optera. Canadian Entomologist 71: 49-54. ety of Washington 5: 237-245. Molineri, C. 2002. Cladistic analysis of the South Baumgardner, D. E. 2003. New synonyms and stage American species of Tricorythodes (Ephemer­ description for three species of Leptohyphidae optera: Leptohyphidae) with the description of (Ephemeroptera). Proceedings of the Entomo­ new species and stages. Aquatic Insects 24: logical Society of Washington I 05: 203-208. 273-308. Baumgardner, D. E. and S. Avila. 2006. Cabecar Randolph, R. P. and W. P. McCafferty. 1998. serratus, a new genus and species of leptohy­ Diversity and distribution of the mayflies phid mayfly from Central America, and (Ephemeroptera) of Illinois, Indiana, Ken­ description of the imaginal stages of Tricor­ tucky, Michigan, Ohio, and Wisconsin. Ohio ythodes sordidus Allen (Ephemeroptera: Lep­ Biological Survey Bulletin (new series) 13 (I), tohyphidae). Zootaxa 1187: 47-59. 188 pp. Baumgardner, D. E. and D. E. Bowles. 2005. Selander, R. B. and P. Vaurie. 1962. A gazetteer to Preliminary survey of the mayflies (Ephemer­ accompany the "Insecta" volumes of the optera) and caddisflies (Trichoptera) of Big "Biologia Centralia-Americana". American Bend Ranch State Park and Big Bend National Museum Novitates 2099: 1-70. Park. Journal of Science 5: 28. Spieth, H. T. 1941. The North American Ephemer­ Eaton, A. E. 1892. Fam. Ephemeridae, pp. 1-16, opteran types of the Rev. A. E. Eaton. Annals plate I. In Godman, F. D. and 0. Salvin, eds. of the Entomological Society of America 34: Biologica Centrali-Americana 38. 87-98. Edmunds, G. F. and R. K. Allen. 1957. A checklist Traver, J. R. 1935. Part II. Systematics, of the Ephemeroptera of North America north pp. 239-739. In Needham, J. G., J. R. Traver, of Mexico. Annals of the Entomological and Y. C. Hsu, eds. The Biology of Mayflies Society of America 50: 317-324. With a Systematic Account of North American Edmunds, G. F., S. L. Jensen, and L. Berner, L. Species. Comstock Publishing Company, 1976. The Mayflies of North and Central Ithaca, New York. America. University of Minnesota Press, Min­ Ulmer, G. 1920. Neue Ephemeropteren. Archiv fiir neapolis. x + 330 pp. Naturgeschichte (A) 85(11): 1-80.