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De Oliveira, Laura A., Do Nascimento, Jeane M. C., Couceiro, Sheyla R. M. (2021): A new species of Tricorythopsis Traver, 1958 (Ephemeroptera: Leptohyphidae) from Pará state, Brazil. Zootaxa 5023 (1): 131-141, DOI: https://doi.org/10.11646/zootaxa.5023.1.8

Tricorythopsis similis sp. nov. (Figs 2–5) Diagnosis. Nymph: 1) maxillary palp 1-segmented (Fig. 4E); 2) segment II of labial palpi shorter than segment I and longer than segment III (Fig. 4F); 3) femora and tibiae with margins covered by long and pectinate setae (Figs 5A–D); 4) tarsal claws with 4 to 5 marginal denticles and 4+2 very small, submarginal denticles (Fig. 5E); 5) abdominal terga III–VII with acute, serrated tubercles medially located on posterior margins (Figs 2A–C, 3C); 6) transversal line of operculate gill absent (Fig. 3D). Mature nymph. Length: body, 2.3 mm; antenna, 0.7 mm; mesonotum, 0.8 mm; caudal filaments broken. Body robust, base of the abdomen wider than the apex (Fig. 2A). General coloration yellowish brown, with blackish and purplish marks irregularly distributed. Head. Yellowish brown, with few, blackish marks irregularly distributed. Tubercles absent. Eyes and ocelli blackish (Figs 2A–B; 3A). Antennae: scape and pedicel whitish yellow, flagellomeres grayish translucent on basal half and whitish translucent on distal half. Mouthparts yellowish translucent (Figs 3B; 4A–G). Labrum broad, with shallow anteromedial emargination; with pores distributed on basal 1/4 (Fig. 4A). Hypopharynx as in Figure 4B. Mandible: basal half with purplish marks irregularly distributed; incisors and molars dark brown (Figs 4C–D); basal 3/5 with spaced pores. Maxilla with pectinate setae at the apex; palp 1-segmented, approximately 2.5 times longer than wide, with a terminal seta (Fig. 4E). Labium with spaced pores; glossa not fused, with filiform setae; paraglossa with filiform setae; segment II of labial palp shorter than segment I and longer than segment III (Fig. 4F); lateral margin of submentum with short, filiform setae (Fig. 4F). Thorax. Yellowish brown, with blackish marks irregularly distributed and with purplish marks near medial line (Figs 2A–B; 3A). Pronotum without tubercles; anterolateral corner projected and rounded (Fig. 3A). Mesonotum with a pair of small anterolateral tubercles; with a pair of slightly developed submedial humps near apex of forewing pads (Fig. 3A). Developing wings with dark gray veins; internal margin blackish basally. Legs. General coloration yellowish white, with purplish and grayish marks; dorsal surface of femora with circular, whitish marks irregularly distributed (Figs 5A–D). Coxae projections absent. Femora almost 1.5 times longer than wide. Femora and tibiae with margins covered by long and pectinate setae. Fore femur with a transversal row of long, pectinate setae on submedial region of the dorsal surface. Tarsal claws (Fig. 5E) almost entirely yellowish white, except by apical region yellowish brown; with 4 to 5 marginal denticles and 4+2 very small, submarginal denticles; with apical seta. Abdomen. Terga yellowish brown, with blackish marks; segments VII–IX with a paired sublateral, transversal, blackish strip (Figs 2A–C; 3C). Sterna whitish yellow. Segments III–IX with lateral margins expanded; segments VI–IX with posterolateral projections bordered with thin setae (projection less developed on segment VI) (Figs 2C; 3C). Terga III–VII with acute, serrated tubercles medially located on posterior margins; tubercle slightly shorter on tergum III (Figs 2A–C; 3C). Gills. Operculate gills yellowish brown, with purplish or grayish marks; oval, almost 1.4 times longer than wide; reaching the segment VII (Figs 2A–C; 3C–D). Ventral lamellae without fringed lobes, without ridges. Transversal line of operculate gill absent. Caudal filaments yellowish brown, with short setae at segment joints. Adults. Unknown Etymology. The specific epithet is a Latin word meaning alike. It is in reference to the general similarity of this species and T. rondoniensis. Habitat. Nymphs of Tricorythopsis similis sp. nov. were collected in two slow-flowing rivers (with width varying to 6–12 m) and associated to marginal vegetation in sections exposed to sunlight (Figs 1D–E). Material examined. HOLOTYPE. Mature nymph, Brazil, Pará state, Santarém, Ponte do Juá stream, 02°26’41”S; 54°47’21”W; 06.xii.2019, Santos SE col. (INPA-EPH 028). PARATYPE. Nymph (parts on slide), Brazil, Pará, Mojuí dos Campos, Mojuí dos Caboclos stream, 02°42’03”S; 54°41’01”W, 05.vii.2020, Oliveira LA col. (INPA-EPH 029).

Tricorythopsis similis sp. nov. (Figs 2���5) Diagnosis. Nymph: 1) maxillary palp 1-segmented (Fig. 4E); 2) segment II of labial palpi shorter than segment I and longer than segment III (Fig. 4F); 3) femora and tibiae with margins covered by long and pectinate setae (Figs 5A���D); 4) tarsal claws with 4 to 5 marginal denticles and 4+2 very small, submarginal denticles (Fig. 5E); 5) abdominal terga III���VII with acute, serrated tubercles medially located on posterior margins (Figs 2A���C, 3C); 6) transversal line of operculate gill absent (Fig. 3D). Mature nymph. Length: body, 2.3 mm; antenna, 0.7 mm; mesonotum, 0.8 mm; caudal filaments broken. Body robust, base of the abdomen wider than the apex (Fig. 2A). General coloration yellowish brown, with blackish and purplish marks irregularly distributed. Head. Yellowish brown, with few, blackish marks irregularly distributed. Tubercles absent. Eyes and ocelli blackish (Figs 2A���B; 3A). Antennae: scape and pedicel whitish yellow, flagellomeres grayish translucent on basal half and whitish translucent on distal half. Mouthparts yellowish translucent (Figs 3B; 4A���G). Labrum broad, with shallow anteromedial emargination; with pores distributed on basal 1/4 (Fig. 4A). Hypopharynx as in Figure 4B. Mandible: basal half with purplish marks irregularly distributed; incisors and molars dark brown (Figs 4C���D); basal 3/5 with spaced pores. Maxilla with pectinate setae at the apex; palp 1-segmented, approximately 2.5 times longer than wide, with a terminal seta (Fig. 4E). Labium with spaced pores; glossa not fused, with filiform setae; paraglossa with filiform setae; segment II of labial palp shorter than segment I and longer than segment III (Fig. 4F); lateral margin of submentum with short, filiform setae (Fig. 4F). Thorax. Yellowish brown, with blackish marks irregularly distributed and with purplish marks near medial line (Figs 2A���B; 3A). Pronotum without tubercles; anterolateral corner projected and rounded (Fig. 3A). Mesonotum with a pair of small anterolateral tubercles; with a pair of slightly developed submedial humps near apex of forewing pads (Fig. 3A). Developing wings with dark gray veins; internal margin blackish basally. Legs. General coloration yellowish white, with purplish and grayish marks; dorsal surface of femora with circular, whitish marks irregularly distributed (Figs 5A���D). Coxae projections absent. Femora almost 1.5 times longer than wide. Femora and tibiae with margins covered by long and pectinate setae. Fore femur with a transversal row of long, pectinate setae on submedial region of the dorsal surface. Tarsal claws (Fig. 5E) almost entirely yellowish white, except by apical region yellowish brown; with 4 to 5 marginal denticles and 4+2 very small, submarginal denticles; with apical seta. Abdomen. Terga yellowish brown, with blackish marks; segments VII���IX with a paired sublateral, transversal, blackish strip (Figs 2A���C; 3C). Sterna whitish yellow. Segments III���IX with lateral margins expanded; segments VI���IX with posterolateral projections bordered with thin setae (projection less developed on segment VI) (Figs 2C; 3C). Terga III���VII with acute, serrated tubercles medially located on posterior margins; tubercle slightly shorter on tergum III (Figs 2A���C; 3C). Gills. Operculate gills yellowish brown, with purplish or grayish marks; oval, almost 1.4 times longer than wide; reaching the segment VII (Figs 2A���C; 3C���D). Ventral lamellae without fringed lobes, without ridges. Transversal line of operculate gill absent. Caudal filaments yellowish brown, with short setae at segment joints. Adults. Unknown Etymology. The specific epithet is a Latin word meaning alike. It is in reference to the general similarity of this species and T. rondoniensis. Habitat. Nymphs of Tricorythopsis similis sp. nov. were collected in two slow-flowing rivers (with width varying to 6���12 m) and associated to marginal vegetation in sections exposed to sunlight (Figs 1D���E). Material examined. HOLOTYPE. Mature nymph, Brazil, Par�� state, Santar��m, Ponte do Ju�� stream, 02��26���41���S; 54��47���21���W; 06.xii.2019, Santos SE col. (INPA-EPH 028). PARATYPE. Nymph (parts on slide), Brazil, Par��, Moju�� dos Campos, Moju�� dos Caboclos stream, 02��42���03���S; 54��41���01���W, 05.vii.2020, Oliveira LA col. (INPA-EPH 029)., Published as part of De Oliveira, Laura A., Do Nascimento, Jeane M. C. & Couceiro, Sheyla R. M., 2021, A new species of Tricorythopsis Traver, 1958 (Ephemeroptera: Leptohyphidae) from Par�� state, Brazil, pp. 131-141 in Zootaxa 5023 (1) on pages 132-135, DOI: 10.11646/zootaxa.5023.1.8, http://zenodo.org/record/5225581

Leptohyphes cornutus Allen Leptohyphes cornutus Allen 1967: 357; Molineri 2003: 55; Dias et al. (2007 a): 215; Dom��nguez et al. (2006): 269. Material examined. Holotype slides (FAMU) from Brazil, rio Irany (?), iv. 1963, F. Plaumann (Irani river is located near Nova Teutonia in Santa Catarina state). Additional material: 1 nymph (CZNC), Brazil, SANTA CATARINA, Passos Maia, Rio Chapecozinho, S 26 �� 46 ' 1.66 " / W 51 �� 49 ' 49.79 ", 1031 m.a.s.l., 13 /iii/ 2009, F. Salles, E. Raimundi cols.; 2 immature nymphs from Argentina, Misiones, Uruz�� stream, S 25 ���� 51 29 / W 54 �� 10 10, 322m, 17.i. 2004, E. Dominguez col (IBN); 2 nymphs (1 nearly mature) and 1 nymphal exuviae (reared male) same locality (IBN), 23���25 /XI/ 1998, Dom��nguez et al. cols.; 2 immature nymphs (IBN) same locality, 7���11.xii. 1999, C. Molineri; and 2 immature nymphs (IBN) from Misiones, Parque Provincial Esmeralda, S 26 �� 38 47.8 / W 53 �� 59 54.9, 478 m, 23.xi. 2009, E. Dominguez & C. Nieto cols. Diagnosis (Table 1). i) paired tubercles on head (2 pairs), pronotum (two pairs, one very small), and mesonotum (two pairs), abdominal terga without paired tubercles nor remnants of them (in some nymphs a posteromedial small hump is present in terga VII���VIII), and abdominal terga IV���VIII present a pair of sublateral blunt spines; ii) fore femur length/maximum width, 1.4 (very immature) to 1.8 (mature); iii) fore margin of middle and hind femora without spines, hind margin (in mature or nearly mature nymphs) with 4 (middle femur) to 20 (hind femur) rounded and stout spine-like setae on elevated sockets, hind femur with outer apex strongly projected (not so strongly in mature nymphs) bearing 1 or 2 distal spines; iv) tarsal claws denticulation 4 (immature) to 11 (mature) marginal denticles and 1 subdistal submarginal denticle; v) middle coxa with acute projection on dorsum (in mature nymphs a smaller projection is also present in hind coxa); vi) hind wing pads present in females; vii) gill formula 3 / 10 / 8 / 8 / 5, gill V without ventral extension on dorsal lamella. Distribution. Argentina (Misiones), Brazil (Santa Catarina, Goi��s, Minas Gerais (Araponga, Sao Paulo). Discussion. Molineri (2003) incorrectly listed a small nymph (slide IBN 608 CM) as L. cornutus, this nymph was collected from Brazil, Rio de Janeiro, 950 m, Nova Friburgo, municipal water supply, 20.iv. 1977, C.M. & O.S. Flint cols. (IBN). This very immature specimen presents the character states now associated with L. airuoca sp. nov., and thus it is listed and diagnosed under this species. As a consequence L. cornutus is no longer recorded from Rio de Janeiro state as listed in Molineri (2003), Dias et al. (2007 a) and Dom��nguez et al. (2006). Dias et al. (2007 a, b) recorded this species in Esp��rito Santo, S��o Paulo and Minas Gerais. We have not studied the material from Sao Paulo, and only partially that of Minas Gerais, thus we only can eliminate the record from Espirito Santo in the distributional list of this species. The nymphs of L. cornutus are readily distinguished from the other species described here by the presence of two pairs of tubercles in mesonotum (1 or 3 pairs in the others), and by the absence of median tubercles in abdominal terga (single or paired tubercles in the others) (Table 1). There is a possibility that some of the new species described here actually are the nymphal form of Leptohyphes mollipes Needham & Murphy (1924) only known from adults from a dubious locality in Brazil. Nevertheless we prefer to describe the new species because the adult stage of the genus presents very few characters of specific value, and thus species known only from adults are of restricted importance., Published as part of Nascimento, Jeane M. C., Molineri, Carlos & Salles, Frederico F., 2014, Redescription of Leptohyphes cornutus Allen, 1967 (Ephemeroptera: Leptohyphidae) and description of three related new species, pp. 397-415 in Zootaxa 3893 (3) on pages 413-414, DOI: 10.11646/zootaxa.3893.3.5, http://zenodo.org/record/250141, {"references":["Allen, R. K. (1967) New Species of New World Leptohyphinae (Ephemeroptera: Tricorythidae). The Canadian Entomologist, 99, 350 - 375. http: // dx. doi. org / 10.4039 / Ent 99350 - 4","Molineri, C. (2003) Revision of the South American species of Leptohyphes Eaton (Ephemeroptera: Leptohyphidae) with a key to the nymphs. Studies on Neotropical Fauna and Environment, 38, 47 - 70. http: // dx. doi. org / 10.1076 / snfe. 38.1.47.14031","Dias, L. G., Molineri, C. & Ferreira, P. S. F. (2007 a) Ephemerelloidea (Insecta: Ephemeroptera) do Brasil. Papeis Avulsos de Zoologia, 47, 213 - 244.","Dominguez, E., Molineri, C., Pescador, M. L., Hubbard, D. M. & Nieto, C. (2006) Ephemeroptera of South America, Aquatic Biodiversity of Latin America. Pensoft Pulblishers, Bulgaria, 646 pp.","Dias, L. G., Salles, F. F., Polegatto, C. M., Silva, R. M. L. & Froehlich, C. G. (2007 b) Ephemerelloidea (Insecta: Ephemeroptera) do Estado de Sao Paulo. Biota Neotropica, 7, 37 - 40. http: // dx. doi. org / 10.1590 / S 1676 - 06032007000300004"]}

Leptohyphes cornutus Allen from Brazil (Santa Catarina) and Misiones, Argentina is re-studied from new material and newly diagnosed as follows: tubercles on head (two pairs), pronotum (two pairs, anterior pair very small), and mesonotum (two pairs), abdominal terga without paired tubercles, nor remnants of them; fore femur length/maximum width, 1.4?1.8; tarsal claws denticulation 4?11+1; hind wing pads present in females; gill formula 3/10/8/8/5. Three new species are described from the nymphal stage: 1) Leptohyphes airuoca sp. nov. from Minas Gerais and Rio de Janeiro (Brazil) characterized by: i) two paired tubercles on head, two pairs on pronotum, and one pair on mesonotum, abdominal terga VI?IX with single medial tubercle projecting from hind margin; ii) fore femur length/maximum width, 1.7?2.0; iii) tarsal claws denticulation 4+0; iv) hind wing pads absent in female; v) gill formula 3/9/9/7/6. 2) Leptohyphes cornutillus sp. nov. from Espirito Santo (Brazil), with: i) paired tubercles on head (two pairs), pronotum (two pairs, anterior pair very small), and mesonotum (three pairs), abdominal terga with remnants of paired tubercles on hind margin (blunt and short undulations in dorsal view); ii) fore femur length/maximum width, 2.1?2.3; iii) tarsal claws denticulation 6+1; iv) hind wing pads present in females; v) gill formula 3/11/9/9/6. And 3) Leptohyphes nebulosus sp. nov. from Espirito Santo (Brazil), with: i) two paired tubercles on head, one pair on pronotum, and one pair on mesonotum, abdominal terga VI?VII with paired submedian tubercles on hind margin; ii) fore femur length/maximum width, 1.6; iii) tarsal claws denticulation 5+1; iv) hind wing pads present in females; v) gill formula 3/9/9/9/6. Fil: Nascimento, Jeane M. C.. Instituto Nacional de Pesquisas da Amazônia. Divisão de Curso de Entomologia; Brasil Fil: Molineri, Carlos. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Centro Cientifico Tecnologico Tucuman. Instituto de Biodiversidad Neotropical; Argentina Fil: Salles, Frederico F.. Universidade Federal Do Espirito Santo; Brasil

Leptohyphes cornutillus sp. nov. (Figs. 6���9 and 14) Material examined. HOLOTYPE, male mature nymph, BRAZIL, Esp��rito Santo, Domingos Martins, Megalcinho, 30 /iii/ 2010, S 20 ˚ 15 ��� 0.6 ��� / W 40 ˚ 43 ��� 26.9 ���, 622m, Salles FF leg. (INPA). PARATYPE, 1 female mature nymph (parts on slide), same data as holotype (CZNC). Mature nymph. Length of male: body, 5.0 mm; hind femur, 1.5 mm; caudal filaments, 4.0 mm. Length of female (mm): body, 8.2 mm; hind femur, 2.0 mm; caudal filaments, 4.5 mm. General coloration brownish, almost completely shaded with black (Figs 6, 7). Head (Fig. 7 A) with two pairs of tubercles, a submedian smaller and sublateral longer and pointed; clypeus well developed and subquadrate. Gena (Fig. 7 A) strongly projected. Mouth parts (Fig. 8). Labrum (Fig. 8 A): with an anteromedial marked indentation on fore margin. Labium (Fig. 8 C): submentum with pointed and projected anterolateral corner; glossae shorter than paraglossae; labial palp with basal segment three times the length of segment two, apical segment subequal in length to segment two. Hypopharynx (Fig. 8 D): lingua with short and strong setae. Maxillae (Fig. 8 B): maxillary palp three-segmented with setae at joints, suture on galea and lacinea complete. Mandibles (Fig. 8 E, F): left mandible with fused and strong incisives, both mandibles with a dense row of setae on dorsolateral surface; prostheca reduced (left), or absent (right). Thorax. Pronotum subrectangular with two pairs of sublateral tubercles, a large and pointed tubercle near hind margin and a smaller blunt anterolateral tubercle. Mesonotum with three pairs of tubercles: a short sublateral pair on fore margin, a longer and acute sublateral pair at 1 / 3 from anterior margin and a submedian pair of tubercles at apex of wing pads. Mesonotum with blunt anterolateral projection. Hind wing pad present in both sexes, small. Thoracic sterna paler than terga. Legs: brownish yellow shaded with black, except on apex of tibia and tarsi, tarsi brownish. Fore leg (Fig. 9 A): femur ratio length/maximum width 2.1���2.3; transversal row of seven stout spines at 1 / 3 from base; fore margin bare, hind margin with five stout spines (on elevated sockets) distally to transversal row; apex of femur medially and dorsally projected. Tibia subequal in length to femur with row of setae on inner margin; tarsus half the length of tibia with row of setae on inner margin, dorsum with elevated ridge; tarsal claw (Fig. 9 C) apically curved with six marginal denticles and one subapical submarginal denticle. Middle and hind legs (Figs. 9 B): coxae with short semicircular dorsal projection; hind femur ratio length/maximum width 2.2; with dorsally row of six blunt spines at base, fore margin bare, hind margin with ten stout spines on elevated sockets, dorsal surface with three stout spines at the middle; dorsomedial projection on apex of femur more pronounced than fore leg. Tibia 1.2 x the length of femur, with dorsal ridge, inner margin with strong setae; hind margin with 2���3 strong setae basally and weaker setae distally. Tarsus 0.4 x the length of femur, inner margin with setae, tarsal claw as in fore leg. Abdomen (Fig. 7 B) with strong spines and weak setae on dorsum. Lateral margins of segments III���VI expanded forming pointed flanges; segment VII and VIII strongly expanded laterally. Terga with a pair of blunt submedian projection on hind margin. Posterolateral spines on segments V���IX (flange on IV is also somewhat pointed resembling a posterolateral spine). Abdominal sternum IX with distal V-shaped indentation (female). Gills: operculate gill on abdominal segment II yellowish translucent almost completely shaded with black on dorsum, except medially and along distal margin, ventrally with strong basal spine and two long and pointed lamellae; other gills whitish almost completely shaded with black. Gill formula 3 / 11 / 9 / 9 / 6. Caudal filaments completely covered with blunt scattered setae, and with a whorl of strong spines at the segment joint after every fourth segment. Adults. unknown. Diagnosis (Table 1). i) paired tubercles on head (2 pairs), pronotum (2 pairs, anterior pair very small), and mesonotum (3 pairs), abdominal terga with remnants of paired tubercles on hind margin (blunt and short undulations in dorsal view) (Figs. 6, 7); ii) fore femur length/maximum width, 2.1���2.3 (Fig. 9 A); iii) fore margin of middle and hind femora without spines, hind margin with 5 rounded and stout spine-like setae on elevated sockets (Figs. 9 B, C); iv) tarsal claws denticulation 6 + 1 (Fig. 9 C); v) middle and hind coxa only with short semicircular projection on dorsum (Figs. 9 B, C); vi) hind wing pads present in females; vii) gill formula 3 / 11 / 9 / 9 / 6, gill V without ventral extension on dorsal lamella. Etymology. From cornutus, given the similarity with L. cornutus, and the diminutive suffix ilus. An allusion to the smaller size of the tubercles of the new species. Distribution. Brazil (Esp��rito Santo). Discussion. L. cornutillus is similar to L. cornutus, but they can be differentiated by the presence of three paired tubercles on mesonotum of the first species (L. cornutus shows two pairs), and by the absence of coxal projection in L. cornutillus (character shared with L. airuoca) (Table 1). The form of mandibles in L. cornutillus, with very short canines (Figs. 8 E, F), somewhat resembles those of L. mandibulus Baumgardner (2007), but this Central American species does not present tubercles on body, among other important differences. As stated by Baumgardner (2007), the short canines could be as a result of wear and tear, but we discard this probability because other mouthparts do not show any mark related with wear and tear (Arens 1990)., Published as part of Nascimento, Jeane M. C., Molineri, Carlos & Salles, Frederico F., 2014, Redescription of Leptohyphes cornutus Allen, 1967 (Ephemeroptera: Leptohyphidae) and description of three related new species, pp. 397-415 in Zootaxa 3893 (3) on pages 402-408, DOI: 10.11646/zootaxa.3893.3.5, http://zenodo.org/record/250141, {"references":["Baumgardner, D. E. (2007) New species of Leptohyphidae (Ephemeroptera) from Costa Rica. Proceedings of the Entomological Society of Washington, 109, 416 - 426.","Arens, W. (1990) Wear and tear of mouthparts: a critical problem in stream animals feeding on epilithic algae. Canadian Journal of Zoology, 68, 1896 - 1914. http: // dx. doi. org / 10.1139 / z 90 - 269"]}