Research Article
Research Article
New species and records of Cloeodes Traver, 1938 (Ephemeroptera, Baetidae) from Costa Rica
expand article infoOscar Vásquez-Bolaños, Fabián Sibaja-Araya, Meyer Guevara-Mora
‡ Universidad Nacional, Heredia, Costa Rica
Open Access


The nymph of Cloeodes danta sp. nov. is described from male and female nymphs collected from highland streams in the Caribbean Slope of the Costa Rica Central Volcanic Mountain Range. Adults are unknown. In addition, C. excogitatus and C. redactus are recorded for the first time in the country. Cloeodes danta sp. nov. can be differentiated from all described species by the predominantly brownish coloration on females and a similar coloration on males but with segments VII–IX light yellow and light brown, with no conspicuous marks or patterns; abundant scale-bases throughout most parts of the body; hindwings pads absent; the presence of three spines in the corners of the posterior margin of sternum III, and the posterior margin of tergum III with 28–30 spines on each side of the middle line (spine with a base width up to 0.5× spine length).


diversity, freshwater, mayflies, Neotropics, taxonomy


The genus Cloeodes (Ephemeroptera, Baetidae) was erected by Traver (1938) for a group of species described from Puerto Rico with hind wings absent. Later, the genus was reviewed by Waltz and McCafferty (1987) and established for other species distributed across the Neotropics, Southern Nearctic, Afrotropics, and Oriental region. More recently, through phylogenetic analysis the genus has been depicted as exclusive for the New World within a complex including Bungona spp., Crassolus spp. and Potamocleoen spp. (Salles et al. 2016). Currently, the number of Cloeodes species has doubled in the last ten years with 26 species described for the Neotropics (Salles et al. 2016; Kluge 2017).

Despite the increase in the number of papers related to Cloeodes and the report of 142 mayfly species in Central America and the Caribbean (Cornejo and Flowers 2015; Salles et al. 2016), there are only two Cloeodes species formally reported from the Central American region: C. excogitatus (Waltz & McCafferty, 1987) and C. redactus (Waltz & McCafferty, 1987), registered from Guatemala and Honduras, respectively (McCafferty and Lugo-Ortiz 1996; McCafferty et al. 2004). In Costa Rica, the presence of the genus has been recorded for lowland and midland streams with scarce human intervention and good water quality (McCafferty et al. 1992; Flowers and De la Rosa 2010), but no species have been formally reported until now.

During an ongoing study of the richness and taxonomy of the family Baetidae in Costa Rica, nymphs of a new Cloeodes species and nymphs of C. excogitatus and C. redactus were collected in low order streams. This work provides a complete description of this new species and diagnostic characters for distinguishing C. excogitatus and C. redactus, in order to improve the taxonomic knowledge of the genus for future natural history or ecological studies of mayflies, which have crucial knowledge gaps in the tropics (Balian et al. 2008; Ramírez and Gutiérrez-Fonseca 2014).

Material and methods

Nymphs of these species were collected using a hand net to scrape large stone substrates from different streams in Costa Rica (Fig. 1). Line drawings for the new species description were made from original pictures taken with an AmScope 1803 digital camera adapted to a Premiere (MRP–161) microscope and stereomicroscope Premiere SMZ-05. The type material examined is preserved in 80% ethanol and Euparal permanent slides, and it has been deposited in Museo de Nacional de Costa Rica, San José Province; Laboratorio de Entomología (LEUNA), Escuela de Ciencias Biológicas, Universidad Nacional, Heredia; Museo de Zoología, Universidad de Costa Rica; and Purdue University Entomological Research Collection (PERC), West Lafayette, Indiana, USA.

Figure 1. 

Geographic distribution in Costa Rica of Cloeodes danta sp. nov. (black circle), Cloeodes excogitatus (black triangle) and Cloeodes redactus (black square).


Cloeodes danta Vásquez-Bolaños, Sibaja-Araya & Guevara-Mora, sp. nov.

Figures 2, 3, 4, 5, 6

Material examined

Holotype : mature ♂ nymph slide-mounted in Euparal, Costa Rica, Heredia Province, Central Volcanic Mountain Range, Cerro Chompipe zone, Refugio de Vida Silvestre Cerro Dantas, Río Las Vueltas, 10°5'38.24"N, 84°3'42.57"W, 2000 m above sea level, VI/09/2017, Sibaja-Araya F. and Acuña F. (colls), deposited in Museo de Nacional de Costa Rica, San José Province. Paratypes: 5 mature nymphs same data as holotype, preserved in 95% alcohol (mouthparts, legs, gills, terga and paraprocts in micro-vial) deposited at Laboratorio de Entomología (LEUNA), Escuela de Ciencias Biológicas, Universidad Nacional, Heredia (1♂, 1♀); Museo de Zoología, Universidad de Costa Rica (1♀) and PERC, West Lafayette, Indiana, USA (1♂, 1♀).

Additional material

Nine nymphs, Quebrada Grande, Refugio de Vida Silvestre Cerro Dantas, 10°5'38.24"N, 84°3'42.57"W, about 1 km from the Río Las Vueltas at 2054 m asl, where type material was collected, Heredia Province, Costa Rica V/13/2018, F. Sibaja, D. Romero, M. Guevara, D. Segura, O. Vásquez (colls), deposited at Laboratorio de Entomología (LEUNA) of the Escuela de Ciencias Biológicas, Universidad Nacional.

Figure 2. 

Cloeodes danta sp. nov. Body coloration A male B female.


Mature nymph. 1) Brownish body coloration, without conspicuous marks or patterns (Fig. 2), 2) abundant scale-bases throughout most parts of the body (Figs 5b, 6h, 6i, 6k), 3) Absence of hind wing pads, 4) Tibia with two parallel lines of fine long hairs (Fig. 5b), 5) posterior margin of tergum III with 28–30 spines on each side of the middle line (Fig. 6h), 6) sterna with 26–28 variable size spines on each side of the middle line and three thick, spine-like teeth in each corner of the posterior margin of sternum III (Fig. 6i, 6j), 7) paraproct with about 13–15 spines (Fig. 7k).


(based on last instar male and female nymphs; adults unknown).

Size (Mature nymphs): Body length: 5.3–6 mm males, 5.5–6.2 mm females; antennae 1.5–1.7 mm; cerci 2.4–2.7 mm; terminal filament 2.2–2.5 mm.

Body coloration: Brownish in general (Fig. 2A, 2B), the head light brownish, with clearer area from central ocelli to the border of clypeus and between antennal and labrum bases; ocelli black, with two tiny white symmetrical dots on each side and clear brownish coloration, darker toward ocellus; eyes black, turbinated portion of compound eyes brownish. Fore wing pads brownish, foreleg brown with slightly lighter areas. Abdomen with even brownish coloration on females and males with predominantly brownish color but with segments VII–IX light yellow and light brown, both with no patterns of marks or spots, and upper corners of abdominal segments darkened; terga I–VIII with visible black posterior sigilla in the middle of every segment, terga VII–IX lighter with upper corners darkened; sterna pale yellowish-brown.

Head (Fig. 3A): Capsule longer than wide. Antennae subequal to 1.3× length of head, with scape subequal to 1.3× times length of pedicel (Fig. 3B). Intra-antennal extension of frons prolongs to ocelli.

Figure 3. 

Cloeodes danta sp. nov. A head color pattern and intra-antennal extension detail B scape and pedicel C labrum (left d. v., right v. v.) D detail of labrum asymmetry on anterior margin.

Labrum (Fig. 3C). Subrectangular, broader than long, anterolateral margins rounded; dorsally with anterior margin with about 20 small, double frayed setae. Lateral margin with eight apically frayed setae; arc of anterodorsal setae with four simple setae; intermediate seta tiny; and several small scattered simple setae near posterior margin. Ventrally with small curved fine setae near anterior margin, and seven small simple setae near lateral margin. Anterior margin slightly asymmetric, with the left side of the cleft not extended the same length as the right side (Fig. 3D).

Left mandible (Fig. 4A). Incisors with seven denticles, middle one reduced and others similar in size; prostheca robust, apically with three lobes and four elongated projections. Row of 5 or 6 minute spine-like setae between prostheca and molar region, only visible at high magnification (40×). Ventral surface of molar region with tuft of spines next to thumb as part of molar structure (Fig. 4B).

Figure 4. 

Cloeodes danta sp. nov. A left mandible B detail of molar region, left mandible C right mandible D hypopharynx (left d. v., right v. v.) E maxilla.

Right mandible (Fig. 4C). Incisors with seven denticles, middle one reduced and others similar in size; prostheca with broad base, bifid, inner projection longer, and both parts frayed; 3–4 tiny, simple setae between prostheca and molar region only visible at high magnification (40×); tuft of spines next to molar region.

Hypopharynx (Fig. 4D). Lingua rounded with no apical lobes, slightly longer and broader than superlinguae, both apically covered with short fine hairs on dorsal and ventral surface.

Maxillae (Fig. 4E). Palpi slightly shorter or as long as galea-lacinia, two segmented; segment I slender in mid part; segment II 1.4× length of segment I; both segments with several simple short setae. Crown of galea-lacinia without thick distal dentisetae, with numerous setae on the inner-dorsal and inner ventral rows, and longer and slender towards the biting edge; medial region of galea-lacinia with one short seta and 5 or 6 long, simple setae.

Labium (Fig. 5A). Glossa and paraglossa similar in length, basally broad and apically narrow; with the base of the glossa reaching more than half of the paraglossa; glossae dorsally with 7–8 setae next to the inner margin and some scattered setae, ventrally with 12 or 13 setae on the inner margin and 9 or 10 on the outer margin, increasing in length apically in both cases. Paraglossae curve inwards, apex subtriangular, dorsally with 7 or 8 setae on the inner margin and 17 or 18 setae on outer margin. Palpi with segment I broad and 0.8× length of segment II and III combined, numerous micropores and simple, tiny setae on both dorsal and ventral surfaces; segment II with row of 6 or 7 setae on the dorsal surface; segment III suboval, dorsally with several short simple setae on apex and 20–25 simple setae of different size on the ventral surface, and the inner margin subequal to the outer margin.

Figure 5. 

Cloeodes danta sp. nov. A labium (left d. v., right v. v.) B foreleg C forefemur apex detail D gill III E gill III apex detail.

Thorax: Hindwing pads absent; foreleg with abundant scale-bases and scattered micropores (Fig. 5B). Femur length about 4× maximum with; dorsal edge with 10 or 11 short, simple setae, ventrally bare, apex rounded with no evident projections, and two concave and apically rounded setae. Tibia with two lines of fine hairs arranged parallel to each other along the tibia, subtending bristle elongated and rounded (Fig. 5B, C). Tarsi dorsal edge bare, ventrally with 10 or 11 minute spines, increasing in size distally; tarsal claw length 0.5× length of tarsi. Mid and hind legs are similar to the foreleg.

Abdomen: Gills (Fig. 6A–G). Present on segments I–VII, colorless, oval, margins with curved fine setae (Fig. 6D, 6E); light-brown tracheae extending from main trunk, branching to margins. Gill I smaller than segment II; gill IV length about 1.3× length of segment III; gills on segment VII not extending beyond apex of segment X (Fig. 2A, B).

Terga. Tergum I with no spines on posterior margin; tergum III with 28–30 spines on posterior margin of each side of the middle line, spine bases almost same width as height (Fig. 6J).

Sterna (Fig. 6I). Posterior margin of sternum III with group of 26–28 variable size spines on each side of middle line, spines bases width about one-third of height; group of fine hairs arranged in slightly curved line on each side of sterna and three thick spine-like teeth on each corner of posterior margin of sternum III (Fig. 6H).

Figure 6. 

Cloeodes danta sp. nov. A gill I B gill II C gill III D gill IV E gill V F gill VI G gill VII H abdominal tergum III I abdominal sternum III J detail of sternum spine-like teeth K paraproct.

Paraprocts (Fig. 6K). Lateral margin with 13–15 spines, different in size and arranged in irregular line, with some overlapping; dorsal surface with numerous micropores and scale-bases; posterolateral extension with about 11 or 12 marginal spines, and several scale-bases and micropores.




The name of this species honors the Danta (Tapirus bairdii) (Mammalia: Tapiridae), a common species in the Cerro Chompipe cloud forest zone, whose three-toed back feet resemble the sternal spine-like teeth described as a diagnostic character of C. danta sp. nov.


Costa Rican, Caribbean slope basin, first order streams, above 2000 m asl.


Habitat preferences in C. danta sp. nov. were observed in relation to elevation. Individuals were found at 2054 m asl in cold waters of two cloud forest streams (Fig. 7); other non-Andean species have been reported between 159–1800 m asl (Nieto and Richard 2008; Gutiérrez and Reinoso-Flórez 2010; Massariol and Salles 2011; Massariol et al. 2013; De Paul et al. 2013; Gutiérrez and Dias 2015). Also, the new species was collected in a pristine forest region, which may be similar to other Cloeodes species in Brazil that seem to prefer remnant forest areas (Salles et al. 2015). In addition, C. danta sp. nov. is common in riffles, on igneous boulders that are covered with periphyton; this is typical for this genus, part of the grazer functional feeding group (Baptista et al. 2006).

Figure 7. 

Habitat of Cloeodes danta sp. nov. at Río Las Vueltas.

Cloeodes excogitatus (Waltz & McCafferty, 1987)

Cloeodes excogitatus Waltz & McCafferty, 1987: 200; McCafferty and Lugo-Ortiz 1996: 23; Wiersema and Baumgardner 2000: 61; McCafferty et al. 2004: 207.

Material examined

Paratype , Cloeodes excogitatus, R.W. Koss and R. Baumann, 1 ♂ / Collected 12/V/1968, Arizona, Oak Creek Canyon, slide-mounted in Euparal (Abs. Alc.) by R.B. Waltz VI/1983, det. by Waltz and McCafferty 1984. (Paratype), PERC (0,012,327). Cloeodes excogitatus, DE Baumgardner, 1 male/ 1 female / Collected 21/V/2004, Arizona, Geenlee CO. San Francisco R. at FS Road, 212, ca 1 mi. N. Clifton. 33°04'30"N, 109°18'04"W, Elev. 3700 ft, (DB04-21), det. M. Meyer 2005. 10 mature nymphs, Río Torito, La Fuente, Santa Teresita, Turrialba, Cartago, Costa Rica, 9°59'13.04"N, 83°40'44.95"W, 1063 m asl, II/28/2017. Romero D. and Sibaja-Araya F, (colls). Material in 95% alcohol, deposited at Laboratorio de Entomología (LEUNA), Escuela de Ciencias Biológicas, Universidad Nacional, Heredia, Costa Rica.


Guatemala, Mexico, USA (Waltz and McCafferty 1987; McCafferty and Lugo-Ortiz 1996; Wiersema and Baumgardner 2000; McCafferty et al. 2004), and Costa Rica.

Cloeodes redactus (Waltz & McCafferty, 1987)

Cloeodes redactus Waltz & McCafferty, 1987: 204; McCafferty and Lugo-Ortiz 1996: 23; Nieto and Emmerich 2011: 58; Massariol et al. 2013: 11; Forero-Céspedes et al. 2016: 462; Kluge 2017: 104.

Material examined

Five mature nymphs, Río Claro tributary, Río Claro, Golfito, Puntarenas, Costa Rica. 8°41'13.05"N, 83°02'48.78"W, 79 m asl, XII/25/2018. Coll. Sibaja-Araya F. (coll.). Material in 95% alcohol deposited at Laboratorio de Entomología, Escuela de Ciencias Biológicas (LEUNA), Universidad Nacional, Heredia, Costa Rica.


Colombia, Honduras, Peru (Waltz and McCafferty 1987; McCafferty and Lugo-Ortiz 1996; Massariol et al. 2013; Forero-Céspedes et al. 2016; Kluge 2017), and Costa Rica.

Figure 8. 

Cloeodes excogitatus: A abdomen coloration B intra-anternal extension (scale:1000 pixels : 2 mm) C spines on tergum III.

Figure 9. 

Cloeodes redactus: A abdomen coloration B intra-anternal extension C spines on tergum III.


Cloeodes danta sp. nov. shares morphological affinities with the following species: C. caraibensis Hofmann et al. 1999, C. excogitatus, C. redactus, and C. maculipes Traver, 1938, including, absence of hind wing pads, segment III of labial palp rounded, maxillary palp shorter than galea-lacinia, and the general shape of other structures like labrum, labium and the apex of the femur that were recognized by Hofmann et al. (1999) and Nieto and Richard (2008). Based on the material and literature reviewed, we consider that Central American and Caribbean species can be separated in two groups according to the depth of the central cleft on the right mandible incisives: in C. excogitatus and C. redactus the cleft breaks down below the level of mandible first inner incisor apex (Waltz and McCafferty 1987: fig. 27; Kluge 2017: fig. 66), while in C. danta sp. nov., C. caraibensis and C. maculipes the cleft breaks down at the level of the first inner incisor apex (Waltz and McCafferty 1987: figs 3, 4 ; Hofmann et al. 1999: fig. 37; Fig. 4C).

In regard to C. caraibensis from Lesser Antilles, these species mentioned above have setae on segment III of the labial palp as long as the setae on the glossa and paraglossa (Waltz and McCafferty 1987: fig. 6; Hofmann et al. 1999: fig. 41), while in C. danta these setae are shorter (Fig. 5A). Also, labrum intermediate setae are well developed in C. maculipes (Waltz and McCafferty 1987: fig. 2), while in C. danta sp. nov. they are minute (Fig. 3C); in regard to C. caraibensis this character is absent, and this species also shares some diagnostic characters with C. danta sp. nov. like scale-bases present on most of the body, shape of III segment of labial palp and spines on the corner of the posterior margins of the sterna (Hofmann et al. 1999: figs 36, 41, 50); however, C. danta sp. nov. can be differentiated by no spots or color pattern on the abdomen, right mandible prostheca with two well-developed branches, glossa apically not lobulated, distal portion of gills rounded, minute spines on the terga and sterna margins, and the posterior margin of the paraproct with no bifurcated spines (Figs 2A, 2B, 4C, 4D, 6A–6G, 6K).

Furthermore, C. danta sp. nov. shares similar features with C. excogitatus, such as the number of spines in the terga III, the shape segment III of labial and the abundance of setae on it (20–25), the number of spines in the paraproct, and body size (Waltz and McCafferty 1987, but C. danta sp. nov. can be identified by the different abdominal color pattern, abundant scale-bases throughout most parts of the body, the maxillar palp is equal in length to the galea-lacinia and the absence of a narrow intra-antennal extension. Also, the new species resembles C. redactus in the length of the maxillary palp being about as long as the galea-lacinia, the number of spines in the paraproct and the lack of any projections on the apex of the femur (Waltz and McCafferty 1987; Kluge 2017); however it could be distinguished by the shape and number of spines in terga III, the absence of the colorless spots in some terga, a larger body size, the presence of an intermediate setae on the labrum, and the presence of thick spine-like teeth in the inferior corner of sterna III.

In order to improve the identification of Cloeodes species in the Central American region, we provide a key to distinguish C. danta sp. nov., C. excogitatus and C. redactus. This will be a useful tool for future aquatic research in the region, which has been increasing over the last 20 years due to development of water quality monitoring using aquatic insects (Alonso-Eguía et al. 2014; Carrie et al. 2017; Castillo 2018; Dávila-Recinos et al. 2019; Pérez 2019).

Key to mature nymphs of Cloeodes species in Central America

1 Abdomen with brownish uniform color pattern, without spots or distinctive marks; body covered with scale-bases; intermediate setae minute; cleft on right mandible incisive breaks down at level of first inner incisor apex and thick spines like teeth present in sternal III corners (Figs 2A, 2B, 4C, 6I, 6J); highlands (2054 m asl) C. danta sp. nov
Abdomen with marks or spotted color pattern (Figs 8A, 9A); body not covered with scale-bases cleft on right mandible incisives breaks down below level of mandible first inner incisor apex; intermediate setae absent; no spines on sternal corners of sterna III (Waltz and McCafferty 1987; Kluge 2017); mid or lowlands 2
2 Abdominal coloration on terga I–III and V–VII with 3 pale spots (the middle spot being smaller); intra-antennal extension narrow, labrum arc of anterodorsal setae with 5 simple setae; segment III of labial palp ventrally with 20–25 setae; maxillary palp shorter than galea-lacinia; 30–35 spines on tergum III (Fig. 8B, C; Waltz and McCafferty 1987: figs 25, 36, 40); midlands (1063 m asl) C. excogitatus
Abdominal coloration with terga I–III and V–VII with dark brown transverse stripes on posterior margins; intra-antennal extension not narrowed; labrum arc of anterodorsal setae with 3 simple setae; segment III of the labial palp with 30–35 setae; maxillary palp about same length as galea-lacinia; 15–20 spines on tergum III (Fig. 9B, C; Kluge, 2017: figs 6, 12, 14, 58, 59); lowlands (79 m asl) C. redactus


We thank our team at the Laboratorio de Entomología (LEUNA) (Carlos Esquivel, David Romero, and Francisco Bravo) for their help both in the field and in the lab with their companionship. We also thank the Escuela de Ciencias Biológicas from Universidad Nacional for their financial and logistical support. We are also grateful with Luke Jacobus (Division of Science, Indiana University Purdue University Columbus for the comments in the early draft, and Sudeep Chandra (Aquatic Ecosystems Laboratory University of Nevada Reno) for facilitating a space to examine part of the material.

This work is part of the research project “Estudio preliminar de la ecología y diversidad del orden Ephemeroptera en Costa Rica”, SIA 0230–13, and Laboratorio de Entomología (LEUNA), SIA 0484-18, Escuela de Ciencias Biológicas


  • Alonso-Eguía P, Mora JM, Campbell B, Springer M (2014) Diversidad, conservación y uso de los macroinvertebrados dulceacuícolas de México, Centroamérica, Colombia, Cuba y Puerto Rico. Instituto Mexicano de Tecnología del Agua.
  • Baptista DF, Buss DF, Dias LG, Nessimian JL, Da Silva ER, Neto DM, Andrade LR (2006) Functional feeding groups of Brazilian Ephemeroptera nymphs: ultrastructure of mouthparts. Annales de Limnologie-International Journal of Limnology 42(2): 87–96.
  • Carrie R, Dobson M, Barlow J (2017) Challenges using extrapolated family-level macroinvertebrate metrics in moderately disturbed tropical streams: a case-study from Belize. Hydrobiologia 794(1): 257–271.
  • Castillo FS (2018) Evaluación de la calidad ambiental del Humedal Refugio de Vida Silvestre Sistema Lagunar de Tísma, Masaya, Nicaragua. Thesis. Universidad Nacional Agraria. Managua, Nicaragua.
  • Dávila-Recinos G, Ortiz JR, Reyes-Morales F (2019) Efecto del microhábitat sobre la abundancia y riqueza específica de los macroinvertebrados bentónicos en dos ríos tropicales de montaña, Guatemala. Ciencia, Tecnología y Salud 6(1): 7–21.
  • De Paul MA, Nieto C, De Roman LE (2013) Cloeodes incus (Waltz and McCafferty) (Ephemeroptera: Baetidae): primer registro de Argentina, características hidrológicas y ambientales. Revista de la Sociedad Entomológica Argentina 72(3–4): 227–230.
  • Gutiérrez Y, Dias LG (2015) Ephemeroptera (Insecta) de Caldas-Colombia, claves taxonómicas para los géneros y notas sobre su distribución. Papéis Avulsos de Zoologia 55(2): 13–46.
  • Hofmann C, Sartori M, Thomas A (1999) Les Ephéméroptères (Ephemeroptera) de la Guadeloupe (petites Antilles françaises). Mémoires de la Société Vaudoise des Sciences Naturelles 20(1): 20–70.
  • Massariol FC, Lima RC, Pinheiro UDS, Quieroz LL, Oliveira LG, Salles FF, Takiya D (2013) Two-winged Cloeodes in Brazil: New species, stage descrip-tion, and key to South American species. Journal of Insect Science 13(60): 1–20.
  • Ramirez A, Gutiérrez-Fonseca PE (2014) Studies on Latin American freshwater macroinvertebrates: recent advances and future directions. Revista de Biología Tropical 62: 9–20.
  • Salles FF, Batista JD, Cabette HRS (2004) Baetidae (Insecta: Ephemeroptera) de Nova Xavantina, Mato Grosso, Brasil: novos registros e descrição de uma nova espécie de Cloeodes Traver. Biota Neotropica 4(2): 1–8.
  • Salles FF, Massariol FC, Angeli KB, Lima MM, Gattolliat JL, Sartori MS (2015) Revealing the diversity of Cloeodes Traver, 1938 (Ephemeroptera: Baetidae) in the Neotropics: description of eleven new species from Brazilian mountain ranges. Zootaxa 4020(1): 1–50.
  • Salles FF, Gattolliat JL, Sartori M (2016) Phylogenetic analyses of Cloeodes Traver and related genera (Ephemeroptera: Baetidae). Systematic Entomology 41(1): 93–111.
  • Traver JR (1938) Mayflies of Puerto Rico. The Journal of Agriculture of the University of Puerto Rico 22: 5–42.
  • Waltz RD, McCafferty WP (1987) Revision of the genus Cloeodes Traver (Ephemeroptera: Baetidae). Annals of the Entomological Society of America 80(2): 191–207.