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1 June 2013 Two-Winged Cloeodes in Brazil: New Species, Stage Description, and Key to South American Species
Fabiana Criste Massariol, Lucas Ramos Costa Lima, Ulisses Dos Santos Pinheiro, Luciano Lopes Quieroz, Leandro Gonçalves Oliveira, Frederico Falcão Salles
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The present work, based on material from northern, central-western, and northeastern Brazil, contributes to the knowledge of the two-winged Cloeodes Traver (Ephemeroptera: Baetidae) in South America. Two new species, C. maracatu, sp. nov. and C. spaceki, sp. nov., are described, the former based on nymphs and reared adults and the latter only on nymphs; the male and female imago of C. auwe and the female imago of C. redactus are described. Based on these findings, an updated key for South American nymphs and male adults of the two-winged Cloeodes is provided.


The genus Cloeodes Traver (Ephemeroptera: Baetidae) is represented by approximately 35 species distributed in the Neotropical, Nearctic, Afrotropical, and Oriental regions (Traver 1938; Waltz and McCafferty 1994; Lugo-Ortiz et al. 1999; Soldán and Yang 2003; Jacobus et al. 2006). The genus is most diverse in the Neotropics, where 22 species have been described so far, including 20 from South America and six from the Antilles, Central America, and southern North America (Waltz and McCafferty 1987a, b; Kluge 1991; Hofman, Sartori, and Tomas 1999; Wiersema and Baumgardner 2000; Nieto and Richard 2008; Nieto and Emmerich 2011; Salles 2011).

All species of Cloeodes outside South America are two-winged (i.e., their hind wings or hind wings pads are absent), but this rule does not apply to South America. Of the 20 species known prior to this study, only six were two-winged: C. anduzei Traver, C. auwe Salles and Batista, C. barituensis Nieto and Richard, C. binocularis Needham and Murphy, C. redactus Waltz and McCaffety, and C. turbinops Needham and Murphy. Cloeodes anduzei, C. binocularis, and C. turbinops were described based only on male imagines, C. barituensis was described based on all stages, and C. auwe and C. redactus are known exclusively at the nymphal stage.

The present study, based on material from several localities in Brazil, contributes to the knowledge of the two-winged representatives of Cloeodes: two new species are described, one based on nymphs from central Brazil and the other based on nymphs and imagines from the Northeastern Region; the male and female imago of C. auwe and the female imago of C. redactus are described; furthermore, new diagnostic characteristics are presented for nymphs of both species. An updated taxonomic key for the South American species of this group is also proposed.

Material and Methods

Taxonomic descriptions and/or diagnoses presented herein were generated from a DELTA (Dallwitz 1980; Dallwitz et al. 1993) database of South American Baetidae genera and species under development (e.g., Salles 2010; Massariol and Salles 2011). The free program DIVA-GIS 5.2 ( was used to make the distribution map of the species.

Photographs were taken using an OPTON Q719K-AC microscope with a TA-0124S digital camera, or a Leica M165C ( stereomicroscope with a DFC420 digital camera. In the latter case, a series of stacked images were processed with the program Leica Application Suite version 3.4.1 to produce final images with enhanced depth of field. Line drawings were made with the aid of a camera lucida, and/or photographs were prepared according to Coleman (2003, 2006).

The material examined is deposited in the following institutions: Invertebrate Collection of the Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Brazil; Coleção Zoológica Norte Capixaba (CZNC), Universidade Federal do Espírito Santo, São Mateus, Brazil; Coleção Entomológica Prof. José Alfredo Pinheiro Dutra (DZRJ), Departamento de Zoologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; and Instituto Miguel Lillo (IML), Tucumán, Argentina.


Cloeodes auwe Salles and Batista, 2004
(Figure 1, 2, 812)

  • Cloeodes auwe Salles and Batista, in Salles et al. 2004: pp. 5; Domínguez et al. 2006: pp. 148.

  • Diagnoses

    Nymphs (Salles at al. 2004; Domínguez et al. 2006, adapted): 1) Antenna about 2.0× the length of head capsule; 2) Labrum with dorsal arc of setae composed of 1 (medial) + 0 (inbetween) + 2 (outer) long, spine-like setae; 3) Segment III of labial palp truncated; 4) Fore-femur with apex without projection, with two blunt setae; 5) Tarsal claw 0.5 to 0.8× length of tarsi; 6) General coloration of abdomen yellowish-white with light brown to dark brown irregular marks, segment I with a circular clear mark at anteromedial region; 7) Spines on posterior margin of tergum I present; 8) Paraproct with eight marginal spines; 9) Caudal filaments with posterior margin of segments with short spines on each segment and long spines on every four segments of cerci and terminal filament.

  • Adults. Male imago: 1) Turbinate eyes with inner margins divergent, touching each other posteriorly (Figure 8, 9); 2) Marginal intercalaries absent between Sc-R2 and CuP-A (Figure 11); 3) Abdominal terga VI with large macula on anterolateral regions (Figure 8); 4) Segment II of forceps with basal constriction (Figure 12); 5) Segment III of forceps elongated (Figure 12); 6) Posterior margin of subgenital plate slightly convex (Figure 12). Female imago: 1) General coloration yellowish-white with abdominal marks similar to male imago.

  • Descriptions

    Male imago

    Length. B