Systematics and Phylogeny of Ulmeritus-Ulmeritoides ...

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Systematics and Phylogeny of Ulmeritus-Ulmeritoides revisited (Ephemeroptera: Leptophlebiidae) F.F. SALLES1 & E. DOMÍNGUEZ2 1

Depto. de Ciências Agrárias e Biológicas, Universidade Federal do Espírito Santo, CEP 29.933-415, São Mateus, ES, Brazil. E-mail: [email protected] 2 CONICET- Instituto de Biodiversidad Neotropical, Facultad de Ciencias Naturales e IML, Universidad Nacional de Tucumán, Argentina. E-mail:[email protected]

Abstract Based on specimens recently collected from Brazil, two new species of Ulmeritoides, as well as the nymph and female imago of the atypical U. flavopedes are described. A new synonymy is proposed, U. flavopedes (=U. oepa). In order to reevaluate the relationships of the species of the Ulmeritus-Ulmeritoides group based on these new evidences, a cladistic analysis is carried out. The monophyly of Ulmeritus and Ulmeritoides is reconfirmed, and a clear differentiation between the Central and South American species of Ulmeritoides emerges from the analysis. Key words: Mayfly, Neotropics, new species, identification key, cladistics

Introduction The genus Ulmeritus was erected by Traver (1956), and then the same author established three new subgenera (Traver 1959). Domínguez (1991) reviewed the status of the genus. In this paper, he elevated the subgenus Ulmeritoides Traver, 1959 to generic status, and transferred U. flavopedes (Spieth 1943), the only species in the subgenus Pseudulmeritus Traver, 1959 to Ulmeritoides. Domínguez (1995) carried out a cladistic analysis of the Ulmeritus-Ulmeritoides group, and described several new species. The data matrix was composed of 12 species: two out-groups (Meridialaris and Atopophlebia), three Ulmeritus and seven Ulmeritoides. Of the 10 species of Ulmeritus-Ulmeritoides treated, in only 5 the nymphal stage was included. As a result of the analysis, the monophyly of the genera Ulmeritus and Ulmeritoides was reconfirmed, as well as the position of U. flavopedes within Ulmeritoides. Since Domínguez (1995), several new species were described and the nymphal stages of others became known (e.g. Lopes et al. 2003, Ávila & Flowers 2005, Mariano & Froehlich 2007). Besides that, recently collected material from Brazil resulted in the discovery of two new species and the unknown nymph of the atypical species U. flavopedes. For this reason, in addition to the descriptions of the new species and stages, we considered it necessary to carry out a new cladistic analysis, incorporating the new evidence in the matrix to see if the relationships proposed in the 1995 paper remained.

Material and methods Material from the following institutions was studied: Instituto-Fundación Miguel Lillo, Tucumán, Argentina (IFML); Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil (INPA); Coleção Zoológica Norte Capixaba, Universidade Federal do Espírito Santo, São Mateus, Brazil (CZNC); Universidad del Valle, Cali, Colombia (UVC); Universidad de la República, Montevideo, Uruguay (URU); Instituto Nacional de Biodiversidad, Costa Rica (INBio); Florida Agricultural and Mechanical University, Tallahassee, Florida, USA

Accepted by L. Jacobus: 29 Oct. 2012; published: 4 Dec 2012

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(FAMU); National Museum of Natural History, Smithsonian Institution, Washington, D.C. USA (NMNH); American Museum of Natural History, New York, USA (AMNH); Illinois Natural History Survey, Illinois, USA (INHS). The usual techniques of preparation and illustration for the group were employed. Examined material: Ulmeritus balteatus: 1 male imago, URUGUAY, Lavalleja Prov., Aguas Blancas, 24/11/ 1963, F. Achaval (URU); 4 male and 6 female subimagos, Picada de Rodriguez, Río Cebollati, 5/I/1957, C. S. Carbonell (URU); 5 nymphs, Arequita, Torrente de Aguas Santa Lucía, 27/I/1957, C. S. Carbonell (URU); 4 female imagos, 6 male and 8 female subimagos, Tacuarembó Prov., Tacuarembó Chico, 20/I/1960, C. S. Carbonell (URU); 15 male and 25 female subimagos, Artigas Prov., Arroyo Tres Cruces (Potrero Sucio), 20/II/1955, C. S. Carbonell (URU). Ulmeritus carbonelli: URUGUAY, Artigas Prov., Sepulturas. Holotype male imago, Alotype female imago, 15 female imagos, 15/I/1952, C. S. Carbonell et al. (URU); Arroyo de la Invernada, 20 male imagos, 4 female imagos, 19/II/1954, C. S. Carbonell (URU); Arroyo Tres Cruces (Potrero Sucio), 8 male subimagos, 3 female subimagos, 20/II/1955, C. S. Carbonell (URU); Salto Prov., Arapey, 4 Km de las termas, 4 female imagos, 3 male subimagos, 2 female subimagos, 12/III/1972, C. S. Carbonell (URU); Paso Yacaré, 8 nymphs, 9/I/1978, L. C. de Zolessi et al. (URU); Belén, 5 nymphs, 4/VI/78, L. C. de Zolessi (URU); C. L. de Vaz, Rio Tacuari, 20 Km SE de Malo, 5 male imagos, 4 female imagos, 12/III/63, C. S. Carbonell (URU). Ulmeritoides acosa: Paratype male imago: COSTA RICA, Guanacaste Prov. Santa Elena Peninsula, Rio Mairena, 2 Km arriba de Cuajiniquil, 27/I/2005, S. Avila, R. Tiffer, R. W. Flowers (IFML); 6 nymphs, Maquenco, Quebrada Maquenco, 392 m, 16/II/2005 same collectors (IFML); 3 nymphs, same locality as holotype, 29/VII/ 2004, S. Avila (IFML). Ulmeritoides chavarriae: Paratypes, male imago and 5 nymphs: COSTA RICA, Guanacaste Prov., Area de conservación Guanacaste, P N Santa Rosa, Sector Santa Elena, Rio Cuajiniquil at road to Playa Potrero Grande, 12/XII/2004, S. Avila, M. M. Chavarria, R. W. Flowers (IFML). Ulmeritoides flavopedes: Holotype male imago: SURINAM, Moengo, 12/IV/1939, D. C. Geijskes col. (AMNH). Paratype: SURINAM, Litani river, Feti Creek, 17/VII/1939, D. C. Geijskes (AMNH). 2 male and 2 female imagoes (reared, with nymphal exuviae), 10 nymphs, BRAZIL, Roraima, Boa Vista, Rio Murupú , Falcão J. N., 12/II/2007 (INPA). Ulmeritoides guanacaste: Holotype male imago: COSTA RICA, Guanacaste, Arroyo # 1; Estación Pitilla, Sendero La Laguna, C. de La Rosa, 12/VI/1989 (INBio); Paratypes 1 male imago, 2 nymphs (same data as holotype) (INBio & FAMU). Ulmeritoides haarupi: 3 male imagos, 11 female imagoes (Holotype, Allotype and Paratypes of U. fidalgoi). ARGENTINA, Misiones, Bompland, Arroyo Martires, E. Domínguez, 26/XI/1986 (IFML). Ulmeritoides huitoto: Holotype male imago: COLOMBIA, Amazonas, Leticia, Caño Km 11, road to Tarapacá. Zuñiga, Domínguez, Molineri (UVC & IFML). Ulmeritoides misionensis: Holotype male imago: ARGENTINA, Misiones, Inta San Vicente, E. Domínguez, 30/XI/1986 (IFML); Paratypes: 1 male imago, 40 nymphs (IFML, FAMU & NMNH). Ulmeritoides uruguayensis: Holotype male imago: URUGUAY, Artigas, Arroyo de la Invernada, C. S. Carbonell, 21/II/1954 (URU). Paratypes: 2 male imagos, 20 female imagos (idem Holotype); 1 male imago, 15 female imagos, Sepulturas, Rio Cuareim, C. S. C. Carbonell, 13/I/1952 (URU). 1 male imago, BRAZIL, Roraima, Alto Alegre, Igarapé da 1ª ponte do ramal do Traiano, Falcão, J.N. 4/II/2007 (INPA). 2 male imagos, BRAZIL, Maranhão, Carolina, Hotel Lajes, 21–22/vii/2010, Boldrini, R. col. (CZNC). Ulmeritoides spinulipenis: Holotype male imago: URUGUAY, Tacuarembó, Tacuarembó chico, C. S. Carbonell, 20/I/1960 (URU). Paratypes: 2 male imagos (idem Holotype) (IFML). Ulmeritoides tifferae: Holotype male imago: COSTA RICA, Guanacaste, Quebrada Alcornoque, Cerro El Hacha, P.N. Guanacaste, C. de La Rosa, 18/VII/1989. 6 male imagos, 18 nymphs (INBio, IFML & FAMU).

Key to the species of the Ulmeritus-Ulmeritoides complex ADULTS 1.

Vein Sc of hind wings 9/10 length of wing; membrane of wings with dark maculae at cross veins (Fig. 2, in Domínguez 1991, p. 164); apex of each penis round, lobe with ventral digitiform projection (Fig. 3, in Domínguez 1991, p. 164)… . . .

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. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritus… 2 Vein Sc of hind wings 8/10 or less length of wing (Figs. 6, 17); wings without color pattern, except in costal area; penis lobes without digitiform projections (Figs. 8, 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides… 4 2(1). Cross veins of hind wings few (around 20); maculae only around subcostal cross veins. . . . . . . . . . Ulmeritus saopaulensis 2’. Cross veins of hind wings numerous (33 to 50); maculae around all cross veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3(2’). Maculae of fore wings forming bands (Fig. 199A, Domínguez et al. 2006 p.530); ventral prolongation of penes short, approximately as long as wide (Fig. 199C, Domínguez et al. 2006 p.530) . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritus balteatus 3’. Maculae of fore wings not forming bands (Fig. 199E, Domínguez et al. 2006 p.530); ventral prolongation of penes long, approximately twice as long as wide (Fig. 199G, Domínguez et al. 2006 p.530) . . . . . . . . . . . . . . . . . . Ulmeritus carbonelli 4(1’). Fore wings with costal and subcostal areas tinged with brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4’. Fore wings with costal and subcostal areas hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 5(4). Penis lobes with apex rounded and with a ventral keel (Figs. 5–8, Avila & Flowers 2005, p. 5) . . . . . . . . . . . . . . . . . . . . . 6 5’. Penis lobes with apex straight, ventral keel absent (Figs. 8, 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 6(5). Ventral keel of penis lobe located near apex (Figs. 9F–G, Domínguez 1995, p. 29) . . . . . . . . . . . . . . . . Ulmeritoides tifferae 6’. Ventral keel of penis lobe located on median length (Figs. 5–8, Avila & Flowers 2005, p. 5) . . . . . . . . . . . . . . . . . . . . . . . 7 7(6’). Abdomen translucent yellowish brown, lacking pattern of pale spots or dark bands . . . . . . . . . . . . Ulmeritoides chavarriae 7’. Abdominal terga orange-brown with pale basal spots and dark brown apical transverse spots. . . . . . . . . Ulmeritoides acosa 8(5’). Basal broad region of forceps almost ¼ length of segment I (Fig. 196P, Domínguez et al. 2006 p.521) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides huitoto 8’. Basal broad region of forceps about 1/6 length of segment I (Figs. 7, 18) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9(8’). Penes with apex ending in an inner acute pointed projection (Figs. 8F–G, Domínguez 1995, p. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides misionensis 9’. Inner projection of penes rounded, outer corner with a small ventral spine (Fig. 7C, Domínguez 1995, p. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ulmeritoides haarupi 10(4’). Apex of penis lobes rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10’. Apex of penis lobes not rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11(10). Veins C, Sc and R1 of forewings yellowish; each penis lobe cylindrical and with a lateral groove (Fig. 4F, Domínguez 1995, p. 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides flavopedes 11’. Veins C, Sc and R1 of forewings yellowish, blackish at stigmatic area; each penis lobe flattened and with a ventral keel (Fig. 12G, Domínguez 1995, p. 32). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides guanacaste 12(10’). Penes lobes with several small apical spines (Fig. 6D, Domínguez 1995, p. 23) . . . . . . . . . . . . . . Ulmeritoides spinulipenis 12’. Penis lobes with inner apical margin ending in an acute projection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13(12’). Abdominal color pattern with conspicuous lighter rounded marks . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides uruguayensis 13’. Abdominal color pattern never as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14(13’). Bulla not surrounded with black mark (Fig. 5a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides araponga sp. nov. 14’. Bulla surrounded with black mark (Fig. 16a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides nigribullae sp. nov. 1’.

NYMPHS 1. 1’. 2. 2’. 3(2’). 3’. 4(1’). 4’. 5(4). 5’. 6(5’). 6’. 7(6’). 7’. 8(4’). 8’. 9(8’). 9’. 10(9).

Row of long dorsal setae of labrum divided, without setae medially (Fig. 7, in Domínguez 1991, p. 165); anteromedian emargination of labrum with 5 flattened denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritus… 2 Row of long dorsal setae of labrum entire, forming a single sinusoidal row (Fig. 18, in Domínguez 1991, p. 166); anteromedian emargination of labrum with 5 denticles, medial one sometimes very large . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides… 4 Segment III of maxillary palpi shorter than segment II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritus saopaulensis Segments II and III of maxillary palpi subequal in length (Fig. 9, in Domínguez 1991, p. 165) . . . . . . . . . . . . . . . . . . . . . . . . . 3 Abdominal gills grayish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritus balteatus Abdominal gills grayish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritus carbonelli Medial denticle of anteromedial emargination of labrum subequal to others. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Medial denticle of anteromedial emargination of labrum much larger than others (Fig. 9b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Exterior surface of femora II and III with numerous short acute spines; femora II with a median black spot (Fig. 13, Avila & Flowers 2005, p. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides tifferae Exterior surface of femora II and III devoid of spines except for a row along the dorsal edge; femora II without a median spot (Fig. 12, Avila & Flowers 2005, p. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Abdominal terga with large pale median and/or submedian spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides guanacaste Pale spots on abdominal terga, if present, small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Abdominal terga with dark brown posterior bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides acosa Abdominal terga with posterior bands weak or missing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides chavarriae Posterolateral projections present on abdominal segments VI to IX . . . . . . . . . . . . . . . . . . . . . . . . . . .Ulmeritoides misionensis Posterolateral projections present on abdominal segments II to IX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Tusk on inner margin of maxillae hardly developed (Fig. 11b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Tusk on inner margin of maxillae well developed (Fig. 22b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Lines of pectinate setae on ventral surface of tibia III forming one main line (as in Fig. 28) . . . . . . . . Ulmeritoides flavopedes

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10’. Lines of pectinate setae on ventral surface of tibia III forming almost two lines (Fig. 27). . . . . Ulmeritoides araponga sp. nov. 11(9’). Spines on dorsum of femora II and III numerous (more than 55); lines of pectinate setae on ventral surface of tibia III forming almost two lines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides uruguayensis 11’. Spines on dorsum of femora II and III few (less than 45); lines of pectinate setae on ventral surface of tibia III forming one main line (Fig. 28) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmeritoides nigribullae sp. nov.

Ulmeritoides araponga sp. nov. Holotype Male Imago (in alcohol, one pair of wings and genitalia on slides). Length: body, 8.5–8.7 mm; fore wings, 9.5–9.7 mm; hind wings, 1.7–1.8 mm. General coloration brownish, thorax and abdomen washed with black (Figs. 1, 2). Head: light brown, washed with black. Upper portion of eyes light brown, lower portion blackish. Ocelli white, heavily washed with brown laterally, blackish basally. Antennae light brown. Thorax: pronotum light brown, tinged with black on lateral and posterior margins; meso and metanotum light orange-brown, margins and carinae darker; pleura yellowish-brown, margins of sclerites darker; sterna orange-brown, washed with black. Wings (Figs. 5–6): membrane of fore wings (Fig. 5a) hyaline, wing base brown; main longitudinal veins brownish, intercalaries yellowish, cross-veins hyaline; 8 cross-veins basal to bulla. Membrane of hind wings (Figs. 5b, 6) hyaline, base brown; longitudinal and cross-veins hyaline, except C basal to costal projection, brownish. Legs: leg I brown except apex of femur, subapical portion of tibia and claws washed with black and apex of tibia and tarsi lighter; legs II and III yellowish-brown, tibiae lighter, subapical portion of tibia and tarsi of leg II darker; tarsi of leg III washed with gray; claws washed with black. Abdomen (Figs. 1, 2): terga brownish, washed with black, lighter posteriorly; sterna lighter. Genitalia (Figs. 7, 8): styliger plate, penes and base of forceps segment I brownish, distal part of forceps segment I and segments II and III yellowish, lighter towards apex. Internal apex of penes heavily washed with black. Apex of penis lobes straight, ending in rather acute apical and mesal projections. [Caudal filaments broken off and lost]. Mature nymph (in alcohol) (Fig. 29). Body length, 6.8 mm. General coloration: brown to orange-brown with black markings. Head: brown, slightly washed with black, with whitish areas bellow median ocellus and between lateral ocelli and compound eyes, vertex with yellowish irregular marks. Ocelli whitish with inner margins black. Eyes of male with upper portions dark orange-brown, lower portion black. Eyes of female black. Antennae: scape and pedicel grayish-brown, flagellum translucent grayish-brown, paler toward apex. Mouthparts: labrum, mandibles, stipes, paraglossae, and first segment of maxillary and labial palpi brownish, remaining parts paler. Medial denticle on anteromedian emargination of labrum much larger than others (Figs. 9a, b). Basal 2/3 of outer margin of mandibles with few setae, longer on median 1/3, apical 1/3 with setae denser (Fig. 10). Tusk on inner apical margin of maxillae small (Figs. 11a, b). Labium as in Fig. 12. Thorax: nota brown with blackish markings. Anterior half of mesonotum with a yellowish-brown longitudinal band, pleura and sterna grayish-brown. Legs (Figs. 13–15): yellowish-brown to yellowish, coxae washed with black. Leg I with femora with median subapical areas darker, with a blackish spot, apex of tibiae with a blackish band. Leg II with central yellowish and subapical blackish spots, tibiae and tarsi yellowish. Leg III with femora with a median yellowish spot, tibiae and tarsi yellowish. Dorsal surface of all femora with numerous short, pointed spines (Figs. 14, 15). Lines of pectinate setae on ventral surface of tibia III forming almost two lines (Fig. 27). Claws light brown. Abdomen: terga orangebrown, darker posteriorly, lateral margins, including posterolateral projections, yellowish-brown. Sterna yellowishbrown, last segments darker. Gills gray, tracheae and fimbriae gray-violet. Caudal filaments yellowish-brown, alternating segment unions darker. Material: HOLOTYPE: 1 male imago, BRAZIL, Minas Gerais, Araponga, Camping Vale das Luas, S 20° 39' 40.4'' / W 42° 27' 0.06'', alt. 1240 m., 17/iv/2004, F.F. Salles & C. N. Francischetti col. (INPA) PARATYPES: 1 male imago and 3 nymphs, same data as holotype (IFML); 10 nymphs, BRAZIL, Espírito Santo, Parque Nacional do Caparaó, Afluente do Rio Pedra Roxa em frente à sede do IBAMA, S 20° 23' 48.1'' / W 41° 44' 08.1'', alt. 1063 m., 20/iv/2008, F.F. Salles leg. (CZNC); 5 nymphs, Espírito Santo, Parque Estadual da Pedra Azul, Cabeceira do Jucu, S 20° 25' 48.1'' / W 41° 01' 03.8'', alt. 1194 m., 17/x/2012, F.F. Salles & E. Raimundi, col. (CZNC) Biology: Nymphs of the new species were found in packs of leaf litter in the bottom of three streams, both located in well-preserved areas of Atlantic Forest at the Serra da Mantiqueira, Southeastern Brazil, always above 1000 m.s.a.l. Etymology. araponga, name of the city where the holotype was collected.

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FIGURES 1–4. Ulmeritoides spp. nov., male imagos. 1. U. araponga (dorsal view); 2. U. araponga (lateral view); 3. U. nigribullae (dorsal view); 4. U. nigribullae (lateral view).

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FIGURES 5–8. Ulmeritoides araponga sp. nov., male imago. 5a. Fore wing; 5b. Hind wing; 6. Hind wing (enlarged); 7. Genitalia (ventral view); 8. Penis lobe (detail).

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FIGURES 9–15. Ulmeritoides araponga sp. nov., nymph. 9a. Labrum; 9b. Detail of anterior margin of labrum; 10. Mandible; 11a. Maxilla; 11b; Detail of maxillary tusk; 12. Labium, left, ventral view; right, dorsal view; 13. Fore leg; 14. Middle leg; 15. Hind leg.

Discussion. Ulmeritoides araponga sp. nov. can be separated from the other species of the genus by the following combination of characters. In the imago: 1) fore wings hyaline, base brown and without dark spots in bullae in Sc and R1; 2) abdominal color pattern as in Figs. 1–2; 3) no cross-veins basal to bulla in fore wings (Fig. 5a); 4) penis as in Figs. 7–8 . In the nymph: 1) medial denticle on anteromedian emargination of labrum much larger than others (Fig. 9a, b); 2) tusk on inner margin of maxillae hardly developed (Fig. 11a, b); 3) dorsum of femora II–III with numerous, acute spines (Figs. 14, 15); 4) femora II with small median black spot (Fig. 14); lines of pectinate setae on ventral surface of tibia III forming almost two lines (Fig. 27).

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Ulmeritoides nigribullae sp. nov. Holotype Male Imago (in alcohol, one pair of wings and genitalia on slides). Length: body, 5.6–5.8 mm; fore wings, 6.3–6.6 mm; hind wings, 2.25 mm. General coloration orange-brownish, abdomen washed with black (Figs. 3, 4). Head (Figs. 3, 4): yellowish-brown, washed with black. Upper portion of eyes light brown, lower portion blackish. Ocelli white, surrounded with dark brown. Antennae brown. Thorax (Figs. 3, 4): pronotum pale grey, tinged with white and dark brown medially, and with black on lateral margins; mesonotum pale yellowish, with whitish markings, margins and carinae brown; metanotum pale gray washed with brown; pleura orange-brown washed with black; sterna orange-brown. Wings (Figs. 16, 17): membrane of fore wings (Fig. 16) hyaline, wing base brown; longitudinal veins yellowish, cross-veins hyaline; 11 cross-veins basal to bulla, brownish spots on bullae of veins Sc and R1. Membrane of hind wings (Figs. 16b, 17) hyaline, base brown; longitudinal and crossveins hyaline. Legs: leg I brown except apex of tibia, whitish, and tarsi, lighter; femur of leg I with median black spot; legs II and III whitish, tarsi slightly tinged with brown; femur of leg II with median orange-brown spot and subapical orange-brown band; tibia of leg II with subapical brownish-black band; femur of leg III with basal and subapical orange-brown washed with black band; claws washed with brown. Abdomen (Figs. 3, 4): terga orangebrown terga I–II completely washed with black; medial black line on terga I–IX, with anterolateral black marks on terga III–IX, as in Figs. 3, 4; sterna brown, washed with black. Genitalia (Figs. 18, 19): styliger plate, penes, base and distolateral margin of forceps segment I light brown, remnants of forceps segment I and segments II and III yellowish. Apex of penis lobes straight, ending in rather acute apical and mesal projections. [Caudal filaments broken off and lost]. Mature nymph (in alcohol). Body length, 5.5 – 6.4 mm. General coloration: brown to dark orange-brown with black markings. Head: brown, slightly washed with black, with yellowish areas below median ocellus, whitish between lateral ocelli and compound eyes, vertex with yellowish irregular marks. Ocelli whitish with inner margins black. Eyes of male with upper portions dark orange-brown, lower portion blackish. Eyes of female blackish. Antennae: scape and pedicel light grayish-brown, flagellum translucent grayish-brown, paler toward apex. Mouthparts: labrum, mandibles, stipe, paraglossae, and first segment of maxillary and labial palpi brownish, remaining parts paler. Medial denticle on anteromedian emargination of labrum much larger than others (Fig. 20a, b). Basal 2/3 of outer margin of mandibles with few long setae (Fig. 21). Tusk on inner apical margin of maxillae large (Figs. 22a, b). Labium as in Fig. 23. Thorax: nota pale yellowish-brown with blackish and yellowish markings. Pleura and sterna brownish washed with black. Legs (Figs. 24–26). Leg I: femora brown, except base whitish and black median spot; tibia and tarsi dark brown, except apex of tibia and base and apex of tarsi whitish. Leg II: femora grayish, whitish at base with a median darker spot; tibia grayish, with subbasal and apical whitish bands, and subapical brownish marking; tarsi grayish, with subbasal brownish band. Leg III: femora brownishgray, with subbasal, subapical and median black markings; tibia and tarsi similar to leg II, except for lack of subapical blackish marking on apex of tibia. Dorsal surface of all femora with few short, pointed spines (Figs. 25, 26). Lines of pectinate setae on ventral surface of tibia III forming one main line (Fig. 28). Claws grayish, light brown toward apex. Abdomen: terga dark orange-brown, with blackish markings, posterolateral projections of segments VII and VIII lighter. Sterna brown washed with black. Gills gray, tracheae and fimbriae gray-violet. Caudal filaments yellowish-brown, alternating segment unions darker. Material: HOLOTYPE: 1 male imago, BRAZIL, Minas Gerais, Parque Estadual do Rio Doce, Lagoa Carioca, 29/x/2005, F.F. Salles & C. N. Francischetti col. (INPA). PARATYPES: 3 male imagos, 3 nymphs, same data as holotype (1 male imago and 1 nymph at IFML, remaining in CZNC). Biology. Nymphs of the new species were found in packs of leaf litter in the bottom of Lagoa Carioca, a lake located in a well-preserved area of Atlantic Forest in Southeastern Brazil. Etymology. nigribullae, after the black markings on the bullae. Variations. In some nymphs the black bands on tibia I are more extended, covering almost all the segments, while in other the two bands are well defined. Discussion. Ulmeritoides nigribullae sp. nov. can be separated from the other species of the genus by the following combination of characters. In the imago: 1) fore wings hyaline, base brown and with dark spots in bullae in Sc and R1 (Fig. 16a); 2) abdominal color pattern as in Figs. 3, 4; 3) 8–11 cross-veins basal to bulla in fore wings (Fig. 16a); 4) penis as in Figs. 18, 19. In the nymph: 1) medial denticle on anteromedian emargination of labrum much larger than others (Fig.20a, b); 2) tusk on inner margin of maxillae well developed (Figs. 22a, b); 3) dorsum of femora II–III with few short, acute spines (Figs. 25, 26); 4) femora II with medial darker spot (Fig. 25); 5) lines of pectinate setae on ventral surface of tibia III forming one main line (Fig. 28).

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FIGURES 16–19. Ulmeritoides nigribullae sp. nov., male imago. 16a. Fore wing; 16b. Hind wing; 17. Hind wing (enlarged); 18. Genitalia (ventral view); 19. Penis lobe (detail).

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FIGURES 20–26. Ulmeritoides nigribullae sp. nov., nymph. 20a. Labrum; 20b. Detail of anterior margin of labrum; 21. Mandible; 22a. Maxilla; 22b; Detail of maxillary tusk; 23. Labium, left, ventral view; right, dorsal view; 24. Fore leg; 25. Middle leg; 26. Hind leg.

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FIGURES 27–28. Ulmeritoides spp. nov., nymph, detail of lines of pectinate setae on ventral surface of tibia III. 27. U. araponga; 28. U. nigribullae.

Ulmeritoides flavopedes (Spieth) Thraulodes flavopedes Spieth, 1943: 11. Atalophlebioides flavopedes; Traver, 1946: 426. Ulmeritus (Pseudulmeritus ) flavopedes; Traver, 1959b: 8. Ulmeritoides flavopedes; Domínguez, 1991: 162; Domínguez, 1995a: 19. Ulmeritoides oepa Lopes, Da-Silva & Py-Daniel, 2003: 195; Domínguez et al. 2006: 527. NEW SYNONYMY.

Female Imago (in alcohol). Body length, 5.4 mm; fore wing,5.5 mm; hind wing 1.2 mm. Coloration similar to the male imago (as described in Domínguez, 1995: 19), except head yellowish white, washed with black. Eyes blackish. Antennae yellowish, with scape and pedicel washed with black. Pronotum light brown, with all margins and medial line blackish. Bullae of fore wings tinged with black. Tarsomeres III–IV of leg I, blackish. Femora II and III, with an apical band besides the basal and medial one, basal and apical bands of femora III, darker. Remarks. In the descriptions of the male imago of U. flavopedes the black markings on the bullae were overlooked; this character is evident from the fresh material here studied. Mature nymph (in alcohol). Body length, 7.4–7.5 mm. General coloration: brown to dark orange-brown. Head: brown, slightly washed with black, with yellowish areas bellow median ocellus and between lateral ocelli and compound eyes. Ocelli whitish with inner margins black. Eyes of male with upper portions dark orange-brown, lower portion blackish. Eyes of female blackish. Antennae: scape and pedicel light-brown, flagellum translucent light-brown, paler toward apex. Mouthparts: labrum, mandibles, stipes, paraglossae, and first segment of maxillary and labial palpi brownish, remaining parts paler. Medial denticle on anteromedian emargination of labrum much

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larger than others. Basal 2/3 of outer margin of mandibles with few long setae. Tusk on inner apical margin of maxillae absent. Thorax: nota pale brown with blackish markings. Pleura and sterna brownish washed with black. Legs. Leg I: femora brown, distal 1/3 darker and with weak black median spot; tibia dark brown, except apex lightbrown; tarsi light-brown. Leg II: femora light-brown, with median spot and apex whitish; tibia light-brown, darker at base and with apical whitish markings; tarsi light brown. Leg III: femora light-brown, with median spot and apex blackish; tibia and tarsi similar to leg II. Dorsal surface of all femora with many short, pointed spines. Lines of pectinate setae on ventral surface of tibia III forming one main line (as in Fig. 28). Claws light brown, darker toward apex. Abdomen: terga orange-brown, with blackish markings on lateral margins of all segments and apical half of segment X, posterolateral projections lighter. Sterna light orange-brown. Gills gray, tracheae and fimbriae gray-violet. Caudal filaments light-brown. Material. 2 male and 2 female imagos, with corresponding nymphal exuvia. Brazil, RR. Boa Vista, Rio Murupú (3,02119; -60,7758), 12/II/2007 J.N. Falcao col. (INPA). Discussion. U. oepa was described based exclusively on nymphs from the state of Roraima. After examination of the type material of this species and comparison with the nymphs recently reared of U. flavopedes, from the same state, it became evident that they are conspecific. As U. flavopedes was described before, U. oepa is considered a Junior Synonym. This is the first formal record of U. flavopedes from Brazil. The nymph of U. flavopedes can be separated from the other species of the genus by the following combination of characters. 1) medial denticle on anteromedian emargination of labrum much larger than others; 2) dorsum of femora II–III with numerous short, acute spines; 3) femora II with small median black spot; 4) lines of pectinate setae on ventral surface of tibia III forming one main line (as in Fig. 28).

FIGURE 29. Ulmeritoides araponga, sp. nov., nymphal habitus.

Phylogeny of the Ulmeritus-Ulmeritoides group Domínguez performed a cladistic analysis on the species of the Ulmeritus-Ulmeritoides complex in 1995. In that study, the first analysis was carried out on a matrix composed of adult characters only. As a result, a single tree was

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obtained, separating two monophyletic groups: one formed by the three Ulmeritus species and the other by the seven Ulmeritoides species (with the then recently transferred U. flavopedes). In a second analysis, the known nymphs of five out of the ten treated species were added, without any change neither in the topology of the resulting cladogram nor in the adult character distribution. This analysis (Analysis 0 from here on) gave support to the earlier systematic arrangement proposed by Domínguez (1991), separating the two monophyletic groups (Ulmeritus and Ulmeritoides). At that moment, the nymph of U. flavopedes (Spieth), that was located in a separate subgenus by Traver (1959) due to the unique genital characteristics, remained unknown. Since the 1995 analysis, several new species have been described and the nymphs of U. flavopedes and U. uruguayensis were reared to imago. For this reason we considered a new analysis should be performed on a new matrix with the new evidence, to test the previous results. This was especially important in the case of the recently reared U. flavopedes, to test if its atypical genitalia was correlated with nymphal characters that could justify its separation in a different genus. Cladistic analysis. For the cladistic analysis a matrix of 28 characters (Appendix I) was compiled, containing 13 adult and 15 nymphal external morphological characters. In this study were included the same taxa treated by Domínguez (1995), except for the addition of five new species and the nymphal characters of U. flavopedes and U. uruguayensis. In a first analysis (Analysis 1 from here on) all known taxa were included, and in a second analysis, U. saopaulensis, U. spinulipenis, U. haarupi and U. huitoto known only from adults were excluded to test the influence of the nymphal missing characters (Analysis 2 from here on). Most of the characters were treated in the same way as in Domínguez (1995), except for a few that needed to be recoded due to new evidence, and are discussed later. Binary characters were coded as 0 and 1. Multistate characters were assigned different numbers, and treated in two different ways: additive or non additive (see list of characters). The program WinClada (Nixon, 2002) was used to compile the matrix and the program TNT (Goloboff et al. 2008) was used to analyze it. The data were analyzed under “implied weight” (k=3) and an exhaustive search was performed (with the command “implicit enumeration”). To establish group support, Bremer supports (absolute and relative) were calculated (Bremer, 1988, 1994). Ten thousand suboptimal trees up to 10 units of fit below the optimal trees were kept in memory using “traditional search” option. Characters and coding. Only Characters treated differently from Domínguez (1995) will be discussed here. Character numbers in bold correspond to Domínguez (1995), character numbers in italics correspond to new numeration here: Character 2 (Attachment of vein ICu2 of fore wings) and character 10 (Basal swelling on segment I of forceps) were removed, because the variability presented by the new species turned them uninformative. Character 4 (Number of cross veins basad to bulla), coded as 0/1 is considered here multistate and additive (character 3), because was possible to separate intervals. Character 5 [new]: The location of the coloration of C and Sc was added. Characters 11–12 (ventral projections and apex of penis lobes) were combined and coded as four different characters: 10–13, three binary, and multistate, additive. These new combination allowed a better delimitation of the differences observed. Character 16 (Shape of tusk on inner apical margin of maxillae) changed from 3-state character to a 4-state character (character 17). In both cases was considered as additive. Character 24 (Lines of pectinate spines on ventral surface of tibia III) changed from a 3-state character to a 4-state character (character 25), and considered as non additive. Character 28 [new]: Number of abdominal segments with posterolateral projections.

Results and discussion In analysis 1 (all taxa included) eleven trees were obtained with a score of 3.842. These trees were summarized in a strict consensus tree (Fig. 30). In analysis 2, three trees were obtained, with a score of 3.671. The topology of this consensus tree (Fig. 31) was equivalent to the consensus of analysis 1 (Fig. 30), with a minor change within the South American Ulmeritoides, affecting the relationships of U.misionensis, mainly due to changes in distribution of character 28.

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FIGURE 30. Consensus tree of complete matrix. Numbers above branches correspond to absolute/relative Bremer Support. Black circles= apomorphies, empty circles= homoplasies. Numbers above circles are character numbers, numbers below circles are character states. “*” indicates species known only from adult stages, and removed for second analysis.

In the consensus tree of the whole matrix (Analysis 1, Fig. 30), two main groups are recovered, confirming the monophyly of the genera Ulmeritus and Ulmeritoides, with both groups presenting several synapomorphies (see figure 31). These two groups were also present in Domínguez (1995) analysis. In analysis 1 however, some differences within the two genera when compared with the previous study (analysis 0) were found. In Ulmeritus the resolution among its species is lost, due to the recodification of the character referred to ventral projections of penis lobes. Within Ulmeritoides two groups are recovered. One group is composed by the four Central American species (U. acosa, U. chavarriae, (U.tifferae, U. guanacaste), with Ulmeritoides tifferae as the sister group of U. guanacaste (as in Domínguez, 1995), and no resolution for the other two. The synapomorphy of this clade is the presence of ventral keel on penis (character 11(1)). The other group includes the eight South American species, and

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the synapomorphies are the apex of penis lobes rather straight, inner corner forming an acute angle (character 12(1)) and the presence of denticles on anteromedian emargination of labrum, with the median one larger (character 14(2)) with only two clades formed: one by U. spinulipenis and U. haarupi and the other by U. huitoto and U. araponga. Nevertheless, we consider these two groups rather unstable, because three of the four species involved are only known by the adult stage (see data matrix and asterisks in cladogram). The main difference in Ulmeritoides with the cladogram from Analysis 0 is that U. haarupi (in that paper called U. fidalgoi) was grouped with the Central American species (then only represented by U. tifferae and U. guanacaste); in Analysis 1 is within the South American clade. We consider that the new position of U. haarupi is more logical, due to its geographical distribution. The removal of the species known only by adults (represented in the consensus of Analysis 2, Fig. 31) did not change significantly the relationships of the species, nor the character distribution (only affected character 28). This does not mean that the inclusion of the unknown nymphal characters would not contribute to improve the resolution of the tree, rather the opposite. In conclusion, in the present analysis the monophyly of Ulmeritus and Ulmeritoides is reconfirmed, and a clear differentiation between the Central and South American groups emerge from the analysis. There is still uncertainty among the relationships of the South American species of Ulmeritoides mainly due to the lack of information concerning the nymphs of several of these species.

FIGURE 31. Consensus tree of matrix including species with nymphal and adult stages. Numbers above branches correspond to absolute/relative Bremer Support. Black circles= apomorphies, empty circles= homoplasies. Numbers above circles are character numbers, numbers below circles are character states.

Acknowledgments We would like to express our gratitude to: C. Francischetti and E. Raimundi, for field assistence; J. N. Falcão and R. Boldrini, for providing pictures of the type of U. oepa and material of U. uruguayensis, respectively; N. SYSTEMATICS OF ULMERITUS-ULMERITOIDES

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Hamada, for allowing the use of photograph equipments provided by CAPES/Pro-equipamentos; and to Instituto Estadual de Florestas (IEF/MG) for providing facilities in the field work at Parque Estadual do Rio Doce. C. Molineri made constructive comments on an early version of this manuscript. This work was partially funded by FAPES (Fundação de Amparo a Pesquisa do Estado do Espírito Santo), CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) and CONICET- PIP 1484 (Consejo Nacional de Investigaciones Científicas y Técnicas).

References Ávila, A.S. & Flowers, R.W. (2005) New species and records of Ulmeritoides (Ephemeroptera: Leptophlebiidae) from Costa Rica. Zootaxa, 1010, 1–14. Bremer, K. (1988) The limit of amino-acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42: 795–803. Bremer, K. (1994) Branch support and tree stability. Cladistics, 10, 295–304. Domínguez, E. (1991) The status of the genus Ulmeritus (Ephemeroptera: Leptophlebiidae: Atalophlebiinae) and related taxa. In: Alba-Tercedor, J. & Sánchez-Ortega, A. (Eds.) Overview and strategies of Ephemeroptera and Plecoptera. Sandhill Crane Press, Gainesville, Florida, pp 157–167. Domínguez, E. (1995) Cladistic analysis of the Ulmeritus-Ulmeritoides group (Ephemeroptera, Leptophlebiidae), with descriptions of five new species of Ulmeritoides. Journal of the New York Entomological Society, 103(1), 15–38. Lopes, M.J.N., E.R. Da-Silva & Py-Daniel, V. (2003) A new species of Ulmeritoides from Brazil (Ephemeroptera: Leptophlebiidae). Revista de Biologia Tropical, 51, 195–200. Mariano, R. & Froehlich, C.G. (2007) Description of the nymph of Ulmeritoides uruguayensis (Traver) (Ephemeroptera: Leptophlebiidae). Zootaxa, 1642, 61–64. Spieth, H.T. (1943) Taxonomic studies on the Ephemeroptera. III. Some interesting Ephemerids from Surinam and other Neotropical localities. American Museum Novitates, 1244, 1–13. Traver, J.R. (1956) A new genus of Neotropical mayflies (Ephemeroptera, Leptophlebiidae). Proceedings of the Entomological Society of Washington, 58(1),1–13. Traver, J.R. (1959) Uruguayan mayflies. Family Leptophlebiidae: Part I. Revista de la Sociedad Uruguaya de Entomologia, 3, 1–13.

APPENDIX I. Characters used in Cladistic analysis. Adults 1. Fore wing with slanting cross vein above MA fork (< with MA approximately 45º: 0 (absent); 1 (present). 2. Number of cross veins in fore wing: 0 (more than 110); 1 (less than 100). 3. Fore wing cross veins basad to bulla: 0 (absent); 1 (1–5); 2 (7–15). [additive]. 4. Spots around cross veins in fore wing: 0 (absent); 1 (present, not forming bands); 2 (present, forming bands). [additive]. 5. Coloration of C and Sc area in fore wing: 0 (restricted to base); 1 (completely tinged). 6. Coloration of longitudinal veins posterior to and in relation with C, Sc and R1 in fore wing: 0 (same color); 1 (lighter). 7. Location of apex of hind wing costal projection: 0 (in basal ½); 1 (beyond basal ½). 8. Length of Sc of hind wing: 0 (> 0.8 of wing length); 1 (< 0.8 of wing length). 9. Number of cross veins in hind wing: 0 (25 or more); 1 (less than 20). 10. Ventral digitiform projections on penis lobes: 0 (absent); 1 (present). 11. Ventral keel on penis lobes: 0 (absent); 1 (present). 12. Shape of apex of penis lobes: 0 (rounded); 1 (rather straight, inner corner forming an acute angle). 13. Spine on apex of penis lobes: 0 (absent); 1 (present, one); 2 (present, several). [additive]. Nymphs 14. Large denticles on anteromedian emargination of labrum: 0 (absent); 1 (present, subequal); 2 (present, median one larger). [additive]. 15. Dorsal row of setae on labrum: 0 (apical); 1 (medial); 2 (basal). [additive]. 16. Dorsal row of setae on labrum: 0 (entire, straight); 1 (entire, sinusoidal); 2 (divided). [additive]. 17. Tusk on inner apical margin of maxillae: 0 (absent); 1 (present, small); 2 (present, medium); 3 (present, big). [additive]. 18. Maxillae palpifer size: 0 (normal); 1 (enlarged). 19. Maxillae ratio of segment III/segment II of palpi: 0 (0.9). 20. Shape of outer margin of mandible: 0 (evenly curved); 1 (strongly curved). 21. Row of setae at base of outer incisor: 0 (no); 1 (yes). 22. Dorsal row of long spines on labial palpi: 0 (on segment III only); 1 (on segment II and III). 23. Row of long setae on labial paraglossae: 0 (absent); 1 (present, apical); 2 (present, medial). [nonadditive].

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24. Spines on margin of labial glossae: 0 (numerous (>10), small); 1 (few (