Two new species of Cloeodes Traver (Ephemeroptera: Baetidae) from Espírito Santo, Southeastern Brazil

Zootaxa 3058: 1–21 (2011) www.mapress.com / zootaxa/ Copyright © 2011 · Magnolia Press

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

Two new species of Cloeodes Traver (Ephemeroptera: Baetidae) from Espírito Santo, Southeastern Brazil FABIANA CRISTE MASSARIOL1 & FREDERICO FALCÃO SALLES2 1

Programa de Pós-graduação em Biodiversidade Tropical, Universidade Federal do Espírito Santo, CEP 29.933-415, São Mateus, ES, Brazil. E-mail: [email protected] 2 Laboratório de Sistemática e Ecologia de Insetos, Depto de Ciências Agrárias e Biológicas, Universidade Federal do Espírito Santo, CEP 29.933-415, São Mateus, ES, Brazil. E-mail: [email protected]

Abstract In the present work, two new species of Cloeodes Traver are described based on nymphs and adults collected in the State of Espírito Santo, Southeastern Brazil. The main characteristics that distinguish the new species from its congeners are, in C. itajara sp. nov.: a) labrum with dorsal arc of setae composed of 12 setae, b) segment III of labial palp with robust and pectinate setae on inner margin, c) fore femur with apex projected, with 5−6 blunt setae, d) male imago with abdominal terga V−VII with a anterolateral triangular black mark; in C. aymore sp. nov.: a) labrum with dorsal arc of setae composed of 1 + 0 + 3 setae, b) fore femur with apex projected, with 2 blunt setae, c) male imago with abdominal terga IV with kidney-like median brown mark.

Key words: taxonomy, new species, South America, mayfly, macroinvertebrate

Introduction The genus Cloeodes Traver, 1938 (Ephemeroptera: Baetidae) has a widespread pantropical distribution with representatives in North America (Wiersema & Baumgardner 1999), Central (Traver 1938) and South America (Nieto & Richard 2008), Africa (Waltz & McCafferty 1994, Jacobus et al. 2006), Madagascar (Lugo-Ortiz et al. 1999) and Southeast Asia (Soldán & Yang 2003). Despite being one of the genus of the family with a higher number of representatives in South America, Cloeodes is poorly documented in Brazil. In South America the following species has been recorded so far: Cloeodes anduzei Traver 1943; C. auwe Salles & Batista, 2004; C. barituensis Nieto & Richard, 2008; C. binocularis Needham & Murphy 1924; C. espinillo Nieto & Richard, 2008; C. hydation McCafferty & Lugo-Ortiz, 1995; C. incus Waltz & McCafferty 1987; C. irvingi Waltz & McCafferty, 1987; C. jaragua Salles & Lugo-Ortiz, 2003; C. nocturnus Navas 1923; C. opacus Nieto & Richard, 2008; C. penai Morihara & Edmunds, 1980; C. redactus Waltz & McCafferty, 1987; C. turbinopsis Needham & Murphy, 1924. Of theses, only five species have been recorded from Brazil: Cloeodes hydation described from nymphs and adults of Mato Grosso and later recorded from Minas Gerais (McCafferty & Lugo-Ortiz 1995; Salles et al. 2004a); C. irvingi described only from nymphs from Paraguay and later recorded from all states of Southeastern Brazil (Lugo-Ortiz et al. 2002; Salles et al. 2003; Salles et al. 2004a); C. jaragua known from nymphs and adults exclusively from the State of Rio de Janeiro (Salles & Lugo-Ortiz 2003; Gonçalves et al. 2010); C. auwe described based on nymphs from Mato Grosso; and C. opacus, a species described from nymphs, originally from Argentina, and recently recorded from the State of Espírito Santo (Salles et al. 2010). Recently, during a survey of mayflies from the State of Espírito Santo, two new species of Cloeodes were found; the aim of this paper is to describe the new species based on nymphs and adults.

Accepted by M. Hubbard: 9 Sep. 2011; published: 13 Oct. 2011

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Material and methods The State of Espírito Santo is located in Southeastern Brazil (Fig. 1) and has an area of 45597Km2. The relief is characterized as mountainous, with altitudes ranging from sea level to 2897m, whose peak is the Pico da Bandeira, at the Parque Nacional do Caparaó (IPEMA 2005). Field trips were carried out in several areas within the state, especially in environmental protected ones, such as Parque Nacional do Caparaó, Reserva Biológica de Sooretama, Reserva Biológica do Córrego do Veado, and Reserva Biológica Augsuto Ruschi. TABLE 1. Sample sites at Espírito Santo State and at its border with Minas Gerais State (Alto Caparaó city and Espera Feliz city), followed by municipality, locality, geographic coordenates, and altitudes. Point

Municipality

Locality

Coordinates

Altitud

PT 01

Águia Branca

Pedra Torta, propriedade do Sr. Aleônsio

19° 03' 56.0" S, 40° 42' 45.3" W

156m

PT 02

Alto Caparaó

Córrego entre Casa Queimada e Cachoeira da Farofa

20° 27' 45.8'' S, 41° 48' 24.7'' W

2160m

PT 03

Alto Caparaó

Rio São Domingos, Cachoeira da Farofa

20° 28' 19.5'' S , 41° 49' 41.7'' W

1972m

PT 04

Alto Caparaó

Vale Verde

20° 25' 11,6'' S, 41° 50' 45.7'' W

1309m

PT 05

Brejetuba

Cachoeira Bernardes

20° 04' 29.9'' S, 41° 18' 86.9'' W

613m

PT 06

Domingos Martins

Cachoeira Floriano

20° 24' 69.2'' S, 40° 56' 57.0'' W

1044m

PT 07

Domingos Martins

Faz. Batista Belon, Cachoeira do Galo Bravo

20° 25' 32.7'' S, 41° 02' 09.8'' W

889m

PT 08

Espera Feliz

Cachoeira Vale a Pena

20° 32' 19.6'' S, 41° 51' 25.2'' W

1053m

PT 09

Espera Feliz

Rio São Domingos, Cachoeira do Aurélio

20° 28' 57.5'' S, 41° 50' 14.5'' W

1800m

PT 10

Espera Feliz

Rio São Domingos, Macieira

20° 28' 52.6'' S, 41° 49' 44.6'' W

1854m

PT 11

Espera Feliz

Rio São Domingos, Cachoeira Sete Pilões

20° 28' 57.1'' S, 41° 49' 50.4'' W

1850m

PT 12

Fundão

Timbuí, Hotel Fazenda Camping

19° 56' 02.0'' S, 40° 24' 45.0'' W

57m

PT 13

Ibitirama

Rio do Tecnotruta, propriedade "Sonho Meu", 20° 28' 08.8'' S, 41° 43' 22.5'' W Sr. Menário

959m

PT 14

Ibitirama

Rio entre Pedra Roxa e Santa Marta

20° 28' 43.0'' S, 41° 42' 15.8'' W

833m

PT 15

Ibitirama

Rio Pedra Roxa, sede do Ibama

20° 23' 48.1'' S, 41° 44' 08.1'' W

1063m

PT 16

Ibitirama

Rio Pedra Roxa, afluente

20° 23' 48.1'' S, 41° 44' 08.1'' W

1063m

PT 17

Ibitirama

Rio Pedra Roxa, Vertente da Pedra Roxa

20° 24' 20.7'' S, 41° 43' 35.6'' W

997m

PT 18

Itaguaçú

Sobreiro, Cachoeira Christopher

19° 45' 36.5'' S, 40° 55' 94.3'' W

66m

PT 19

Itarana

Barra Jatibocas, Rio Jatibocas

19° 58' 74.0'' S, 40° 52' 91.6'' W

450m

PT 20

Itarana

Barra Jatibocas, Cachoeira da Ponte

19° 57' 84.6'' S, 40° 52' 14.5'' W

777m

PT 21

Iúna

Córrego na estrada

20° 24' 05.5'' S, 41° 43' 45.9'' W

1015m

PT 22

Iúna

Vale Encantado

20° 24' 38,7'' S, 41° 50' 03.6'' W

1976m

PT 23

Santa Teresa

Nova Lombardia, Sítio Capitel de Santo Antônio, Córrego Escavado

19° 52' 31.7" S, 40° 31' 47.3" W

705m

PT 24

Santa Teresa

Nova Lombardia, Sítio Capitel de Santo Antônio, Córrego Grande, Pinguela

19° 52' 16.0" S, 40° 31' 43.3" W

718m

PT 25

Santa Teresa

Nova Lombardia, Sítio Capitel de Santo Antônio, Córrego Grande, Sede

19° 52' 31.0'' S, 40° 31' 49.1'' W

768m

PT 26

Santa Teresa

Nova Lombardia, Sítio Capitel de Santo Antônio, Córrego Grande, Cachoeira Grande

19° 52' 30.8" S, 40° 31' 56.6" W

712m

PT 27

Santa Teresa

Nova Lombardia, Sítio Capitel de Santo Antônio, Poção

19° 52' 30.9" S, 40° 32' 07.4" W

739m

PT 28

Santa Teresa

Reserva Biológica Augusto Ruschi, Cachoeira 19° 53' 20.6" S, 40° 32' 41.5" W do meio

803m

PT 29

Santa Teresa

Santa Bárbara, Cachoeira Strutz

169m

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19° 50' 10.1'' S, 40° 43' 76.7'' W

MASSARIOL & SALLES

FIGURE 1. Map showing geographic location of Espírito Santo State. FIGURES 2−3. Geographic distribution of new species of Cloeodes from Espírito Santo. 2. Cloeodes itajara. 3. C. aymore.

In the examined material item, PT refers to the collection point (Table 1), in parentheses we present the number of individuals studied, followed by the stage examined (N for nymphs and A for adults) and finally, the collection date. The taxonomic descriptions herein were generated from the free program DELTA – Description Language for Taxonomy (Dallwitz 1980; Dallwitz et al. 1993). Pictures were taken using an OPTON Q719K-AC microscope with a TA-0124S digital camera or a Leica (M165C) stereomicroscope with a DFC420 digital camera. Line drawings were prepared with the aid of photographs and the programs Adobe Illustrator® and Adobe Photoshop® according to Coleman (2003, 2006). For making the distribution map of the new species we used the free program DIVA-GIS 5.2.

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The material examined is housed in the following institutions: Invertebrate Collection of the Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Brazil, Invertebrate Collection of the Centro Universitário Norte do Espírito Santo, Universidade Federal do Espírito Santo (CEUNES), Entomological Collection Prof. José Alfredo Pinheiro Dutra, Departamento de Zoologia, Universidade Federal do Rio de Janeiro (DZRJ), and Instituto Miguel Lillo, Tucumán, Argentina (IML).

Results Cloeodes itajara, sp. nov. (Figs. 2, 4−6, 11−22, 35−39, 46, 48, 50−52) Diagnoses. Nymphs. 1) General coloration of abdominal terga yellowish brown washed with brown to dark brown with a lighter medial longitudinal stripe (Fig. 4); 2) Labrum with dorsal arc of setae composed of 12 setae (Fig. 11a); 3) Segment III of labial palp obliquely truncate (Figs. 16a, b); 4) Segment III of labial palp with robust and pectinate setae on inner margin (Fig. 16b); 5) Fore femur with apex projected, with 5−6 blunt setae (Fig. 18a); 6) Gill opaque; trachea dark greyish, inner and outer margins brown and with a light brown stripe parallel to main trunk (Figs. 6, 21a); 7) Caudal filaments with posterior margin of segments with short spines on each segments. Adults. 1) Abdominal terga V−VII with a anterolateral triangular black mark (Figs. 38, 52); 2) Marginal intercalaries absent between Sc-R1 and CuP-A (Fig. 46a); 3) Hind wing present; costal process hooked, located on basal third (Fig. 46b); 4) Hind wing with 2 complete longitudinal veins and 1 incomplete (Fig. 46b); 5) Segment III of forceps elongated (Fig. 48); 6) Posterior margin of subgenital plate rounded (Fig. 48). Description. Nymph Length. Body: 7.5–9.0 mm; cerci: 3.0–3.5 mm; terminal filament: 3.0–3.5 mm; antenna: 2.3 mm. Head (Fig. 4). Coloration: light brown with light yellow narrow median longitudinal stripe on all length of vertex. Compound eyes and lateral ocellus surrounded yellowish. Frons with area between antennal sockets yellowish. Turbinate portion of male compound eyes reddish brown. Antenna light brown; short, 1.3×the length of head capsule. Labrum (Figs. 11a, 11b). Rectangular, broader than long; length about 0.6× maximum width; dorsal surface flat; distal margin with medial emargination and small process. Dorsally with few short, fine, simple setae scattered over surface; dorsal arc of setae composed of 12 long spine-like setae, 5 restricted to medial group; lateral margin bare. Ventrally with submarginal row of setae decreasing in length toward medial region, composed of lateral setae pectinate; anterolateral setae bifid, pectinate; medial setae bifurcated at middle (Fig. 11b); ventral surface with 7 short, blunt setae near lateral and anterolateral margin. Left mandible (Fig. 12). Inner and outer set of incisors respectively with apparently 3 + 4 denticles. Prostheca robust, apically denticulate and with comb-shape structure at apex. Margin between prostheca and mola slightly convex, with spine-like setae on entire length; tuft of setae absent; tuft of spine-like setae at base of mola present; subtriangular process wide, above level of area between prostheca and mola; denticles of mola not constricted; tuft of setae at apex of mola absent. Lateral margins almost straight; bare. Basal half with few short, fine, simple setae scattered over dorsal surface. Right mandible (Fig. 13). Inner and outer set of incisors respectively with apparently 3 + 3 denticles. Prostheca slender, bifurcated at middle, inner lobe long, outer short and both pectinate. Margin between prostheca and mola straight, without setae; tuft of setae absent; tuft of spine-like setae at base of mola present; tuft of setae at apex of mola present, reduced to a bifid setae. Lateral margins almost straight; bare. Basal half with few short, fine, simple setae scattered over dorsal surface. Hypopharynx (Fig. 14). Lingua subequal in length to superlingua; apex with anteromedial lobe rounded; with short, fine, simple setae; distal half not expanded. Superlingua not expanded; fine, simple setae scattered over lateral and distal margin; basal half of lateral margin with short, spine-like setae. Maxilla (Fig. 15). Crown of galea-lacinia with 4 denticles, inner denticle opposed to outer denticles; inner dorsal row of setae with 2 denti-setae, pectinate and bifurcated at base. Medial protuberance of galea with 1 short, spine-like setae + 7 long setae. Maxillary palp subequal in length of galea-lacinia; two segmented; setae on maxillary palp few short, fine and simple setae, micropore scattered over surface and one spine-like setae on the apex of segment II; palp segment II 1.4× length of segment I; apex of last segment constricted.

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MASSARIOL & SALLES

FIGURES 4−6. Cloeodes itajara, nymph. 4. General habitus. 5. Dorsal view, smooth antero-sublateral area of terga. 6. Gill coloration.

Labium (Figs. 16a, 16b). Glossa basally broad, narrowing apically and subequal in length to paraglossa; inner margin with 20 long, spine-like setae increasing in length apically; outer margin with 12–13 long, spine-like setae increasing in length distally; ventral surface with one row of 8 simple setae near inner margin. Paraglossa sub-rectangular or straight, curved only at apex; apex with 2 rows of spine-like setae; outer margin with row of 13 long, spine-like setae; dorsally with a curved row of 6 long setae near inner margin; ventrally with a curved row of 12 spine-like setae near inner margin. Labial palp with segment I 0.8× length of segments II and III combined; segment I covered with few short, simple setae and micropore near to outer margin; segment II without distomedial protuberance; inner margin bare; outer margin with few, short, simple setae; dorsally with row of 7 spine-like, simple setae; segment III obliquely truncate; length 1.1× width; covered with robust, simple setae on distal margin, robust and pectinate setae on inner margin (Fig. 16b); long, spine-like setae scattered over ventral surface, but forming a row near outer margin. Thorax (Fig. 4). Yellowish brown with brownish marks. Fore wing pads yellowish brown. Hind wing pads present. Fore leg (Fig. 17). Coxa brown. Trocanter light brown. Femur yellowish washed with brown, except at base and apex of anterior surface. Tibia light brown. Tarsus brown. Ratio of fore leg 1.3:0.8:0.8:0.2 mm. Fore femur. Length about 3.5× maximum width; dorsally with a row of 11 blunt setae (in lateral view they look like spine-like setae); length of setae about 0.1× maximum width of femur; apex projected; with 5–6 blunt setae (Fig. 18a); ventrally with row of short, spine-like setae restricted to posterior lobe; anterior surface with abundant micropore (restrict to anterior lobe), scale-bases and scales. Tibia. Dorsally with a row of abundant, fine, simple setae; ventrally with row of 10 short, spine-like setae, with a lanceolate setae subapically; anterior surface with abundant scale-base and micropore; tibio-patelar suture present. Subtending bristle present (Fig. 19). NEW SPECIES OF CLOEODES FROM BRAZIL

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Tarsus. Dorsally with abundant, long, simple setae; ventrally with a row of 15 spine-like setae and one long lanceolate setae near the apex; tarsal claw bare. Mid and hind legs. Similar to fore leg, except for: hind femur dorsally with a row of 9 blunt setae (in lateral view look spine-like setae); subapical projections of femur less developed on mid leg and practically absent on hind leg (Figs. 18a, b, c). Abdomen (Fig. 4). Yellowish brown washed with brown to dark brown with one lighter medial longitudinal stripe. Segments VIII and IX eventually lighter; segments II−VIII with yellowish smooth antero-sublateral area (Fig. 5). Sterna yellowish. Terga. Surface with abundant scale-bases and micropore; posterior margin with regular spines, 2.3× as long as wide (Fig. 20); spines present in posterior margin of segments: II−X. Sterna. Spines present in posterior margin of segments: V − IX. Gill (Figs. 6, 21a, b). Opaque; trachea dark greyish, inner and outer margins brown and with a light brown stripe parallel to main trunk. Margin with broad spines and short, fine, simple setae (Fig. 21b). Tracheae extending from main trunk to inner and outer margins. Gill I about 0.5 length of segment II; oval. Gill IV as long as length of segments V to half VI combined; oval. Gill VII about 1.5 length of segment VIII; oblong and obliquely truncate. Paraproct (Fig. 22). With 19–21 marginal spines; surface with abundant scale-bases and micropores; posterolateral extension with blunt marginal spines and scale-bases scattered on surface. Caudal filaments. Brown with a yellowish band subdistally. Posterior margin of segments with short spines on each segment. Inner margin of cercus and inner and outer margin of terminal filament with tufts of long, flat setae. Male imago Length. Body: 7.0 mm; fore wing: 8.0 mm; hind wing: 1.2 mm; antenna: 1.2 mm; tibia I: 2.1 mm; tibia II: 1.6 mm; tibia III: 1.4 mm; caudal filament: broken. Head (Figs. 35−37, 50−52). Coloration yellowish white. Antenna yellowish white. Turbinate portion of compound eyes orangishdorsally (yellowish in alcohol), stalk yellowish brown. Dorsal portion of turbinate eyes oblong; length 1.8× width; stalk height 0.4× width of dorsal portion; inner margins parallel, close to eacher other. Thorax (Figs. 37, 39, 50−52) Pronotum light brown (yellowish in alcohol) and mesonotum light brown (yellowish white in alcohol), except anterolateral scutal costa and area between posterior scutal protuberance whitish, metanotum brown. Anteronotal protuberance rounded. Metascutellar protuberance posteriorly pointed (Fig. 39). Legs. Whitish with orange band on apex of femur and black mark on apex of tibia and tarsomeres. (yellowish white in alcohol). Leg I: tibia 1.6× length of femur; tarsus 1.8× length of femur; and with 4 segments decreasing on length apically. Leg II: tibia 1.4× length of femur; tarsus 0.2× length of femur. Leg III: tibia 1.3× length of femur; tarsus 0.3× length of femur. Fore wing (Fig. 46a). Hyaline, except between C and R1 opaque; longitudinal and cross veins light yellowish brown. Stigmatic area with 3 cross veins touching or almost touching subcostal vein and 2 veins not touching subcostal vein; marginal intercalary veins paired, except between veins R1-R2, ICu2-CuP single and between veins Sc-R1, CuP-A absent; length of each intercalary vein between IMA and IMA2 0.6× distance between adjacent longitudinal veins; length of fore wing about 2.6× width. Hind wing present (Fig. 46b). Hyaline; veins light yellowish brown; with 2 complete longitudinal veins and 1 incomplete, not reaching apex of hind wing. Costal process hooked; located on basal third. Abdomen (Figs. 35, 36, 38, 50−52). Terga. Segments I−VI white, VII−X orangish ( yellowish white in alcohol); segments I−VI with a dark brown narrow line on posterior margin; segments V−VII with anterolateral triangular black mark (Figs. 51, 52); segments I−V with lateral orange band. Tracheation not pigmented. Sterna. Segments I−VI white, VII−X yellowish white. Genitalia (Fig. 48). Coloration yellowish white. Forceps segment I sub-rectangular; 0.3× length of segment II; distance between base of forceps 1.3× distance between lateral margins of forceps. Forceps segment II medially with constriction. Forceps segment III elongated, 2.0× as long as wide; 0.1× length of segment II. Posterior margin of subgenital plate rounded. Female imago. Unknown. Etymology. The specific epithet is a combination derived from the tupi-guarani ita (=stone) and jara (=lord), "Lord of the stones".

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Distribution (Fig. 2). Brazil - Espírito Santo (Ibitirama), Minas Gerais (Alto Caparaó, Espera Feliz). Material examined. Holotype. Male imago with corresponding nymphal exuvia, Brazil, Espírito Santo, Parque Nacional do Caparaó, Ibitirama, Pedra Roxa River, 20/iv/2008, 20°23'48.1''S/ 41°44'08.1''W, Salles, F.F., Massariol, F.C., Boldrini, R., Lima, M.M. and Britto, P.V.A. col. (INPA). Paratypes. Ten nymphs, Brazil, Minas Gerais, Parque Nacional do Caparaó, Espera Feliz, São Domingos River, Sete Pilões Waterfall, 22/iv/2008, rock, low current, Salles, F.F., Massariol, F.C., Boldrini, R., Lima, M.M. and Britto, P.V.A. col. (six at CEUNES, two at DZRJ, two at IML). Additional material. PT 02: (12N) 23/iv/2008, PT 03: (60N) 23/iv/2008, PT 03: (37N) 25/iii/2009, PT 04: (51N) 23/iii/2009, PT 08: (3N) 27/iii/2009, PT 09: (24N) 22/iv/2008, PT 11: (67N) 22/iv/2008, PT 11: (13N) 25/ iii/2009, PT 13: (32N) 21/iv/2008, PT 15: (12N) 20/iv/2008, PT 16: (19N) 20/iv/2008 (CEUNES). Life cycle association. Rearing. Biology. Most of the nymphs of the new species were found on rocky substrates, such as slab and stones, preferable in areas with low or no current, as in other species of the genus (Salles & Lugo-Ortiz 2003; Salles et al. 2004b). They were also collected in leaf litter and marginal vegetation.

Cloeodes aymore, sp. nov. (Figs. 3, 7−10, 23−34, 40−45, 47, 49, 53−55) Diagnoses. Nymphs. 1) Segment VI of abdominal terga with a anteromedial subtriangular dark mark (Fig. 7); 2) Labrum with dorsal arc of setae composed of 1 + 0 + 3 setae (Fig. 23a); 3) Segment III of labial palp rounded and covered with spine-like setae along margins and ventral surface (Fig. 28); 4) Fore femur with apex projected, with 2 blunt setae (Fig. 30a); 5) Posterior margin of terga with long spines often intercalated by short spines, both slightly outwardly (Fig. 32a) (in some individual, see comments bellow, spines can be regular - Fig. 32b); 6) Caudal filaments with posterior margin of segments with short spines on each segments and long spines on every two segments on cercus and every four segments on terminal filament. Adults. Male Imago. 1) Abdominal terga with segments I, VII−X brown and II−VI translucent whitish; segment IV with kidney-like median brown mark (Fig. 40); 2) Marginal intercalaries absent between Sc-R1 and CuPA (Fig. 47a); 3) Hind wing present; costal process hooked, located on basal third (Fig. 47b); 4) Hind wing with 2 complete longitudinal veins (Fig. 47b); 5) Posterior margin of subgenital plate truncate (Fig. 49). Female imago. 1) Abdominal sterna yellowish with lateral dark brown marks on segments II−VIII in the female imago (Fig. 44); 2) Hind wing present; costal process hooked, located on basal third; 3) Hind wing with 2 complete longitudinal veins. Description. Nymph Length. Body: 5.0–7.5 mm; cerci: 2.5–3.0 mm; terminal filament: 2.5–3.0 mm; antenna: 1.9 mm. Head (Figs. 7, 8, 53). Coloration: yellowish white with a light brown mark between compound eyes and lateral ocellus. Vertex brown with yellowish vermiform marks. Frons with dark brown stripe extending from lateral ocellus and almost reaching the clipeo base. Turbinate portion of male compound eyes orangenish brown. Antenna yellowish; short, 1.6× the length of head capsule. Labrum (Figs. 23a, b). Rectangular, broader than long; length about 0.8× maximum width; dorsal surface flat; distal margin with medial emargination and small process. Dorsally with few short, fine, simple setae scattered over surface and scale-bases; dorsal arc of setae composed of 1 + 0 + 3 long, spine-like setae; lateral margin bare. Ventrally with submarginal row of setae decreasing in length toward medial region, composed of lateral and anterolateral frayed and bifurcated at middle; medial setae frayed and bifurcated at base (Fig. 23b); ventral surface with 4 short, blunt setae near lateral and anterolateral margin. Left mandible (Fig. 24). Inner and outer set of incisors respectively with apparently 3 + 4 denticles. Prostheca robust, apically denticulate and with comb-shape structure at apex. Margin between prostheca and mola straight, without crenulations; tuft of setae absent; tuft of spine-like setae at base of mola present; subtriangular process wide, above level of area between prostheca and mola; denticles of mola not constricted; tuft of setae at apex of mola absent. Lateral margins almost straight; bare. Basal half with scale bases. Right mandible (Fig. 25). Inner and outer set of incisors respectively with apparently 3 + 4 denticles. Prostheca slender, bifurcated at middle, inner lobe long, outer short and both pectinate. Margin between prostheca and mola

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slightly convex, without setae; tuft of setae absent; tuft of spine-like setae at base of mola present; tuft of setae at apex of mola present, reduced to a single setae. Lateral margins almost straight; bare. Basal half with scale bases. Hypopharynx (Fig. 26). Lingua subequal in length to superlingua; apex convex; with short, fine, simple setae; distal half not expanded. Superlingua not expanded; fine, simple setae scattered over lateral and distal margin; basal half of lateral margin with short, spine-like setae. Maxilla (Fig. 27). Crown of galea-lacinia with 4 denticles, inner denticle opposed to outer denticles; inner dorsal row of setae with 2 denti-setae, pectinate and bifurcated at base. Medial protuberance of galea with 1 short, spine-like setae + 5 long setae. Maxillary palp subequal in length of galea-lacinia; two segmented; setae on maxillary palp fine, simple setae scattered over surface and micropores on segment I; palp segment II 1.6× length of segment I; apex of last segment smooth and acute. Labium (Fig. 28). Glossa basally broad, narrowing apically and subequal in length to paraglossa; inner margin with 16 long, spine-like setae increasing in length apically; outer margin with 12–13 long, spine-like setae increasing in length distally; ventral surface with one row of 6 simple setae near inner margin. Paraglossa sub-rectangular or straight, curved only at apex; apex with 2 rows of spine-like setae; outer margin with row of 13 long, spine-like setae; dorsally a row of 4 long setae near inner margin; ventrally with a curved row of 9 spine-like setae near inner margin. Labial palp with segment I 1.0× length of segments II and III combined; segment I covered with few short, simple setae and micropore near to outer margin; segment II without distomedial protuberance; inner margin bare; outer margin bare; dorsally with row of 6 spine-like, simple setae; segment III rounded; length 1.1× width; covered with spine-like simple setae along margins and ventral surface. Thorax (Figs. 7, 53). Yellowish washed with light brown. Fore wing pads yellowish. Hind wing pads present. Fore leg (Fig. 29). Coxa yellowish washed with light brown. Trocanter yellowish white. Femur yellowish washed with brown, except basal third yellowish white. Tibia yellowish white except basal and distal margins brown. Tarsus brown. Ratio of fore leg 1.2:0.6:0.8:0.2 mm. Fore femur. Length about 4.5× maximum width; dorsally with a row of 9 blunt setae (in lateral view they look like spine-like setae); length of setae about 0.1× maximum width of femur; apex projected; with 2 blunt setae (Fig. 30a); ventrally with 13 short, spine-like setae; anterior surface with abundant micropore, scale-bases and spine-like setae near the base of ventral margin. Tibia. Dorsally with a row of abundant, fine, simple setae; ventrally with row of 11 short, fine, spine-like setae, apex with pectinate setae; anterior surface with abundant scale-base and micropore; tibio-patelar suture present. Subtending bristle present (Fig. 31). Tarsus. Dorsally bare; ventrally with a row of 15–16 spine-like setae and one long lanceolate setae near the apex; tarsal claw bare. Mid and hind legs. Similar to fore leg, except for: mid and hind femur brown except basal and distal third yellowish; tarsus basally brown and became lighter toward apex. Hind femur dorsally with row of 5 blunt setae (in lateral view they look like spine-like setae). Subapical projections of femur less developed on mid leg and practically absent on hind leg (Figs. 30a, b, c). Abdomen (Figs. 7, 8, 53). Yellowish white washed with brown to dark brown. Segments V, VI, IX and anterior half of X shaded with brown; segment VI with a anteromedial subtriangular dark mark; segments II−VII with smooth, yellowish sublateral area. Sterna yellowish white with segment IX shaded with brown; sublateral brown marks on segments II−VI; lateral margins of segments V and VI brown. Terga. Surface with abundant scale-bases and micropore. Posterior margin with long spines (2.7× as long as wide) often intercalated by short spines (2.2× as long as wide), both slightly outwardly (Fig. 32a). Spines present in posterior margin of segments: III−X. Sterna. Spines present in posterior margin of segments: IV−IX. Gill (Figs. 33a, b). Opaque; trachea dark greyish, inner and outer margins brown. Margin with broad spines and short, fine, simple setae (Fig. 33b). Tracheae extending from main trunk to inner and outer margins. Gill I subequal in length to segment II; oval. Gill IV as long as length of segments V to half VI combined; oval. Gill VII about 1.5 length of segment VIII; oblong and obliquely truncate.

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FIGURES 7−10. Cloeodes aymore, nymph. 7. Dorsal view. 8. Ventral view. 9. Dorsal view (intraspecific variation). 10. Ventral view (intraspecific variation).

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FIGURES 11−16. Cloeodes itajara, nymph. 11. Labrum (a, left d.v., right v.v., b, details of submarginal row of ventral setae). 12. Left mandible (d.v.). 13. Right mandible (d.v.). 14. Hypopharynx (d.v.). 15. Maxilla (v.v.). 16. Labium (a, left d.v., right v.v., b, details of segment III of labial palp).

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FIGURES 17−22. Cloeodes itajara, nymph. 17. Fore leg, anterior surface. 18. Projections of femur apex (a, fore femur, b, mid femur, c, hind femur). 19. Subtending bristle, anterior surface. 20. Posterior margin of terga IV. 21. Gill IV (a, general appearance, b, details of gill margin). 22. Paraproct.

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FIGURES 23−28. Cloeodes aymore, nymph. 23. Labrum (a, left d.v., right v.v., b, details of submarginal row of ventral setae). 24. Left mandible (d.v.). 25. Right mandible (d.v.). 26. Hypopharynx (d.v.). 27. Maxilla (v.v.). 28. Labium (a, left d.v., right v.v.).

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FIGURES 29−34. Cloeodes aymore, nymph. 29. Fore leg, anterior surface. 30. Projections of femur apex (a, fore femur, b, mid femur, c, hind femur). 31. Subtending bristle, anterior surface. 32. Posterior margin of terga IV. 33. Gill IV (a, general appearance, b, details of gill margin). 34. Paraproct (d.v.).

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FIGURES 35−39. Cloeodes itajara, male imago. 35. Dorsal view. 36. Lateral view. 37. Head and thorax, d.v. 38. Abdominal terga V−VII, showing the black, triangular, anterolateral marks. 39. Thorax, lateral view, showing the metascutellar protuberance.

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FIGURES 40−45. Cloeodes aymore, male and female imago. 40. Dorsal view, male. 41. Dorsal view, female. 42. Lateral view, male. 43. Head, lateral view, male. 44. Lateral view, female. 45. Genitalia, v.v., male.

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FIGURES 46−49. Wings and genitalia of new species of Cloeodes. 46. Wings of C. itajara (a, fore wing, b, hind wing). 47. Wings of C. aymore (a, fore wing, b, hind wing). 48. Male genitalia of C. itajara. 49. Male genitalia of C. aymore.

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FIGURES 50−52. Cloeodes itajara, male imago. 50. Dorsal view. 51. Head and thorax, d.v. 52. Lateral view.

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FIGURES 53−55. Cloeodes aymore. 53. Nymph, d.v. 54. Male imago, l.v. 55. Female imago, l.v.

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Paraproct (Fig. 34). With 11–12 marginal spines; surface with abundant scale-bases and micropores; posterolateral extension with blunt marginal spines. Caudal filaments. Yellowish white with one brown band subdistally and one brown band distally (Fig. 53). Posterior margin of segments with short spines on each segment, and long spines on every two segments on cercus and every four segments on terminal filament. Inner margin of cercus and inner and outer margin of terminal filament with tufts of long, flat setae. Male imago Length. Body: 5.0 mm; fore wing: 5.5 mm; hind wing: 1.1 mm; tibia II: 1.1 mm; tibia III: 0.9 mm; caudal filament: broken. Head (Figs. 40, 42, 43, 54). Coloration brown, except yellowish region between ocellus. Turbinate portion of compound eyes yellow dorsally, stalk orangish. Dorsal portion of turbinate eyes oblong; length 1.5× width; stalk height 0.7× width of dorsal portion; inner margins parallel, close to each other. Thorax (Figs. 40, 42, 54) dark brown. Anteronotal protuberance rounded. Metascutellar protuberance posteriorly pointed. Legs. Whitish except margins brown. Apex of tarsomeres brown. Leg II: tibia 1.2× length of femur; tarsus 0.3× length of femur. Leg III: tibia 1.1× length of femur; tarsus 0.3× length of femur. Fore wing (Fig. 47a). Hyaline, except between C and R1 opaque; longitudinal and cross veins yellowish brown. Stigmatic area with 2 cross veins touching or almost touching subcostal vein and 2 veins not touching subcostal vein; marginal intercalary veins paired, except between veins R1-R2, ICu2-CuP single and between veins Sc-R1, CuP-A absent; length of each intercalary vein between IMA and IMA2 0.7 and 0.4× distance between adjacent longitudinal veins; length of fore wing about 2.7× width. Hind wing present (Fig. 47b). Hyaline; veins yellowish brown; with 2 complete longitudinal veins. Costal process hooked; located on basal third. Abdomen (Figs. 40, 42, 54). Terga. Segments I, VII−X brown; II−VI translucent whitish; membrane between II−VII dark brown; segment IV with kidney-like median dark brown mark; 2/3 of segment VI medially with brown mark; segments II−VI with brown mark laterally. Tracheation dark brown. Sterna. Segments I, VII−IX brown; segments II−VI translucent whitish. Genitalia. (Figs. 45, 49) Segment I brown, II and III brown and lighter toward apex. Forceps segment I subrectangular; 0.3× length of segment II; distance between base of forceps 1.0× distance between lateral margins of forceps. Forceps segment II medially with constriction. Forceps segment III elongated, 1.6× as long as wide; 0.1× length of segment II. Posterior margin of subgenital plate truncate. Female Imago Length. Body: 4.5 mm; fore wing: 6.0 mm; hind wing: 1.0 mm; tibia II: 1.1 mm; tibia III: 0.9 mm; caudal filament: broken. Head (Figs. 41, 44, 55). Coloration yellowish white. Compound eyes black. Thorax (Figs. 41, 44, 55) Pronotum yellowish white with brown marks. Mesonotum brown, anteronotal protuberance and medioscutum washed with grey; scutellum whitish and posterior scutal protuberance washed with grey. Metanotum brown. Anteronotal protuberance rounded. Metascutellar protuberance posteriorly pointed. Legs. Whitish with orange band on apex of femur and with brown mark on apex of tibia and tarsomeres. Leg II: tibia 1.1× length of femur; tarsus 0.3× length of femur. Leg III: tibia 1.0× length of femur; tarsus 0.2× length of femur. Fore wing. Hyaline, except between C and R1 opaque; longitudinal and cross veins dark yellowish white. Stigmatic area with 2 cross veins touching or almost touching subcostal vein and 2 veins not touching subcostal vein; marginal intercalary veins paired, except between veins R1-R2, ICu2-CuP single and between veins Sc-R1, CuP-A absent; length of each intercalary vein between IMA and IMA2 0.4× and 0.2× distance between adjacent longitudinal veins; length of fore wing about 2.8× width. Hind wing present. Hyaline; veins dark yellowish white; with 2 complete longitudinal veins. Costal process hooked; located on basal third. Abdomen (Figs. 41, 44, 55). Terga yellowish washed with brown, except median region of segments II−IX translucent; unpigmented area wider toward base of abdomen. Segments IV−VI similar to male imago. AnterolatNEW SPECIES OF CLOEODES FROM BRAZIL

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eral corner of segments II−VII with circular unpigmented area. Tracheation dark brown. Sterna yellowish with sublateral dark brown subtriangular marks on segments II−VIII. Etymology. The specific epithet is the foreign name for a tribe of South American Indians of eastern Brazil (including the State of Espírito Santo), also known as botocudos. Distribution (Fig. 3). Brazil - Espírito Santo (Águia Branca, Brejetuba, Domingos Martins, Fundão, Ibitirama, Itaguaçú, Itarana, Iúna, Santa Teresa), Minas Gerais (Alto Caparaó, Espera Feliz). Material examined. Holotype. Male imago with corresponding nymphal exuviae, Brazil, Espírito Santo, Parque Nacional do Caparaó, Ibitirama, Pedra Roxa River, 20/iv/2008, 20°23'48.1''S/ 41°44'08.1''W, Salles, F.F., Massariol, F.C., Boldrini, R., Lima, M.M. and Britto, P.V.A. col (INPA). Paratypes. Five nymphs, Brazil, Espírito Santo, Parque Nacional do Caparaó, Ibitirama, Tecnotruta River, propriedade "Sonho meu", 21/iv/2008, rock, moderate current, Salles, F.F., Massariol, F.C., Boldrini, R., Lima, M.M. and Britto, P.V.A. col. (three at CEUNES, two at DZRJ). Five nymphs, same locality, data and collectors, marginal vegetation (three at CEUNES, two at IML). Additional material. PT 01: (26N) 02/i/2009, PT 01: (18N) 23/ii/2009, PT 03: (7N) 23/iv/2008, PT 03: (5N) 26/ iii/2009, PT 04: (4N) 23/iii/2009, PT 05: (1N) 17/ii/2011, PT 06: (73N) 18/ii/2011, PT 07: (310N) 26/iii/2011, PT 07: (4A) 26/iii/2011, PT 08: (26N) 27/iii/2009, PT 09: (7N) 22/iv/2008, PT 10: (1N) 23/iv/2008, PT 11: (6N) 25/ iii/2009, PT 12: (12N) 02/xii/2010, PT 12: (16N) 03/xii/21010, PT 13: (25N) 21/iv/2008, PT 14: (50N) 21/iv/ 2008, PT 15: (11A) 20/iv/2008, PT 15: (8N) 20/iv/2008, PT 16: (11N) 20/iv/2008, PT 17: (2N) 21/iv/2008, PT 18: (9N) 07/ii/2011, PT 19: (6N) 10/ii/2011, PT 20: (7N) 10/ii/2011, PT 21: (61N) 21/iv/2008, PT 22: (2N) 24/iii/ 2009, PT 23: (1N) 16/xi/2007, PT 23: (6N) 19/i/2008, PT 23: (2N) 26/x/2008, PT 24: (1N) 26/x/2008, PT 24: (2N) 18/ii/2009, PT 25: (1N) 19/i/2008, PT 25: (2N) 25/x/2008, PT 26: (14N) 20/i/2008, PT 26: (5N) 25/x/2008, PT 26: (2N) 19/ii/2009, PT 27: (6N) 26/x/2008, PT 28: (7N) 20/ii/2009, PT 29: (6N) 08/ii/2011 (CEUNES). Life cycle association. Rearing. Biology. As in C. itajara and several other species (see above), most of the nymphs of the new species were found in rocky substrates, such as slab and stones, preferable in areas with low or no current. However, a significant proportion of nymphs were also collected in sand in moderate flow, besides marginal vegetation and gravel.

Discussion The new species herein described, C. itajara and C. aymore, share with C. jaragua from Brazil, and C. opacus and C. espinillo from Argentina, the presence of an apical projection in their femora (more developed in the fore femur, smaller in the mid and barely discernible in the hind one). Among them, three species are readily distinguished: Cloeodes itajara can be distinguished because of a series of unique features, such as: gill with a light brown stripe parallel to main trunk; fore femur with apex projected, with 5−6 blunt setae; robust setae on inner margin of segment III of labial palp pectinate; C. jaragua is easily recognizable based on its abdominal color pattern, with segments I−VII or I−VIII blackish, contrasting with segments IX and X whitish; while C. espinillo is the only species of this group in which the dorsal setae of femora are pointed, instead of blunt. Cloeodes opacus and C. aymore, on the other hand, are more similar. The main differences between them are: a) abdominal color pattern – in C. opacus tergites are predominantly dark brown, except for a medial yellow marking on segments I−IX (Fig. 43 of Nieto and Richard 2008), while in C. aymore the coloration of abdominal tergites is more contrasting, with segments I−IV and VII−VIII lighter (yellowish white), and remaining segments darker (brown to dark brown) (Figs. 7 and 9); and b) apical projection on fore femora – in C. opacus, although present (Figs 47 and 48 of Nieto and Richard 2008), it is poorly developed when compared to C. aymore (Figs. 29 and 30a). Noteworthy is also a morphological variation found in nymphs of C. aymore. Some of the studied individuals are darker (Figs. 9, 10), sternites V and VI, as well as paraprocts are entirely pigmented (Fig. 10), and the filaments are predominantly dark with a subapical light band. Besides that, all characteristics are identical, both in nymphs and adults. Individuals of both types were found very often in the same sample, and because of these reasons we are treating them as an intraspecific variation of C. aymore.

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Acknowledgements We would like to express our gratitude to the staff from Reserva Biológica de Sooretama, Reserva Biológica Augusto Ruschi, and Parque Nacional do Caparaó for logistic suport; ICMBio (Instituto Chico Mendes de Conservação da Biodiversidade) and IBAMA (Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis) for collection permission (number 11239-1, 16719-1, 12777-1); FAPES (Fundação de Apoio à Ciência e Tecnologia do Espírito Santo, process number 3632763/07, 45429502/09), CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico, processes number 313117/2009-8, 470731/2009-5) for financial support, and UFES (Universidade Federal do Espírito Santo) for providing scholarship to FCM. We also would like to thank Dr. Carolina Nieto (INSUE-Tucuman) for providing type material of Cloeodes described by her.

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