Cloeodes redactus Waltz & McCafferty 1987

Kluge, Nikita J., 2017, Contribution to the knowledge of Cloeodes Traver 1938 (Ephemeroptera, Baetidae), Zootaxa 4319 (1), pp. 91-127 : 104-114

Cloeodes redactus Waltz & McCafferty 1987

( Figs 44–91, 46 )

“Genus Poss. Cloeodes ”: Roback 1966: 133, Pl. 3: Figs 31–36 (larva);

Cloeodes (Cloeodes) redactus Waltz & McCafferty 1987b: 204 (larva) (non C. redactus sensu Massariol et al. 2013 );? Cloeodes barituensis: Falcao, Salles & Hamada 2011 : Fig. 89 (larva) (non C. barituensis Nieto & Richard 2008 ).

Material examined. PERU: Region Ucayali, Prov. Padre Abad, Aguaytia, Pampa Yurac , La Choza , 9°04'S, 75°31'W, 23.II.2006, coll. N. Kluge: 2 L/S ♀ ; Region Loreto, Prov. Ucayali: 17 km NNE Contamana, Aguas Termales , 7°11'S, 74°57'W, 26.VII–4.VIII.2013, coll. N. Kluge & L. Sheyko: 5 L-S-I ♂, 1 L-S/I ♂, 7 L-S-I ♀, 8 larvae ; ibid., Pampa Hermosa, Quebrada Huanganaico , 7°12'S, 75°22'W 19–20.VIII.2013, coll. N. Kluge & L. Sheyko: 3 L-S-I ♂, 7 larvae .

Region Loreto, Prov. Maynas, Quebrada El Sabalo (right tributary of Rio Itaya) between Puente Itaya (57 km from Iquitos ) and San Joaquin (at Rio Amazon ), 4°16'S, 73°27'W, 1–16.II.2006, coll. N. Kluge: 1 L-S ♂, 1 L-S-I ♀, 2 L-S ♀, 7 larvae.

Region Junin, Prov. Satipo, Puerto Prado, Rio Meritori , 11°09'S, 74°16'W, 9–15.IX.2013, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂, 1 L-S/I ♂, 2 L-S-I ♀, 2 L-S ♀, 3 L/S ♂.

Descriptions. Larva. CUTICULAR COLORATION ( Figs 58–62 , 69–73 ). Head with colorless and brown areas. Terga and pleura of thorax mainly brown with composite maculation, sterna colorless; in immature larva cuticle of fore protoptera colorless ( Figs 58–59 ), before molt cuticle of protoptera becomes brown ( Fig 72 ). Legs light with or without diffusive darker macula at middle of femur, knee articulation and base of tarsus ( Figs 60–62 ). Most of abdominal terga brown with lighter areas; pigmentation variable, but pigmentation of tergum II always less extensive than pigmentation of tergum III (Figs 5 8–59, 69–71); terga IV and VIII usually lighter than others, often at most part non-pigmented ( Fig. 70 ), sometimes entirely non-pigmented ( Fig. 71 ), rarely as dark as others ( Fig.

69); tergum IX more or less pigmented; tergum X well-pigmented. Abdominal sterna light, sometimes with brown lateral markings; paraprocts brown, as remainder segment X. Caudalii lighter, with brown band near apex ( Fig. 73 ).

HYPODERMAL COLORATION. Black transverse stripes on posterior margins of abdominal terga I–III and V–VII often present ( Figs 58–59 ), sometimes absent (as in imago). Female larva just before molt to subimago with lateral triangular brown maculae on abdominal sterna (as in imago—Fig. 86).

SHAPE AND SETATION. Frontal suture pointed, rectangular; face relatively long; in female larva eyes brought together (as in Fig. 6 ). Labrum equally wide at base and at middle or little wider at base, with pair of submedian long setae and 2 pairs of sublateral long setae (as in Fig. 12 ; Roback 1966: Fig. 31 ). Prostheca of left mandible with 3 blunt processes and usually 4 (rarely 3 or 5) pointed processes; median margin of left mandible proximad of prostheca straight or slightly convex, smooth ( Figs 63, 65 ). Prostheca of right mandible directed medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca straight or slightly convex, smooth or with serration, without seta-like processes ( Figs 64, 66 ). Stipes of maxilla proximally with 3–4 setae on ventro-medial side. Maxillary canines and distal dentiseta stout; distal dentiseta widened, with apex somewhat hooked toward canines (as in Fig. 8 ; Roback 1966: Fig. 32 ). Maxillary palp about as long as maxilla, 2-segmented ( Roback 1966: Fig. 34 ). Labium with glossae and paraglossae subequal, both narrowed apically (as in Fig. 14 ). Glossa with 6–8 setae in dorso-lateral row; ventrally with irregularly arranged setae in proximal part and 6–9 setae forming ventro-median row. Paraglossa with lateroapical setae forming one regular row and few (2–4) setae just dorsad of it; with 6–7 setae in ventro-median row; with 3–4 setae in dorso-median row. Distal segment of labial palp not widened apically, with median margin subequal to lateral margin (as in Figs 13, 14 ; Roback 1966: Fig. 36 ).

All thoracic terga without protuberances. Metanotum without vestiges of hind protoptera ( Fig. 61 ). Fore legs largest, middle legs smaller and hind legs smallest; on fore leg tarsus much longer than tibia, on middle and hind legs tarsus subequal to tibia; claws not elongated, much less than half of tarsal length ( Table 4; Figs 60–62 ). Femur parallel-sided, with outer margin straight and inner margin slightly convex; apical projection equally short on femora of all leg pairs (as in Fig. 16 ). Outer side of femur with row of 3–6 small spatulate setae; apical projection with 2 such setae (as in Fig. 16 ). Inner and dorsal side of femur with few stout setae, which are several times smaller than setae on dorsal side. Patella-tibial suture and proximal arc of long setae with contiguous sockets equally developed on tibiae of all leg pairs; inner margin of tibia with irregular small stout pointed setae, apically with 2 larger setae (as in Fig. 16 ). Outer-apical seta blunt, reaching apex of tibia ( Fig. 68 ). Dorsal side of each tarsus with long fine setae forming 2 longitudinal rows ( Fig. 68 ). Claw shorter than 1/2 of tarsus length ( Fig. 68 ).

Scales on abdominal terga and sterna numerous, short, semicircular, colorless and delicate. Posterior margin of abdominal tergum I often entirely smooth, rarely with few irregular short denticles; posterior margins of all terga II–X often with regular long pointed denticles, rarely tergum II without denticles or with few irregular short denticles; on tergum IX, portion of posterior margin behind pair of submedian setae, lacks denticles and projected backward (as in Fig. 18 ). Posterior margins of abdominal sterna I–IV smooth; posterior margins of sterna V–VIII with regular pointed denticles, shorter and lighter than denticles on terga. Each sternum II–VI with pair of regular transverse arched rows of long, fine, simple (not furcate) setae with closely contiguous sockets (as in Fig. 20 ). Tergalius I widest at distal half, with costal and anal margins convex; tergalius II widest near middle, with anal margin more convex than costal margin; tergalii III–IV widest at proximal half, with anal margin more convex than costal margin ( Figs 44–57 ). All tergalii with ribs only on proximal parts of costal and anal margins, without middle rib. Margins bearing costal and anal ribs smooth (not serrated); margin free of ribs slightly notched, with small seta in each notch. Paraprocts with anterior median apodeme small, soft and colorless; each paraproct with 7–8 long pointed denticles on posterior margin, with dense scales as on sterna and terga ( Fig. 67 ). Each swimming seta of caudalii sharply divided into dark proximal part and colorless delicate distal part ( Fig. 73 ). Ventral and dorsal side of paracercus and lateral sides of cerci in middle part with one or several long pointed denticles on each 4th segment.

DEVELOPING MALE GENITALS. In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis -type» pose, with 2nd segment bent by convexity medially, and 3rd segment directed caudally ( Figs 86, 89 ).

RESPIRATORY MOVEMENTS. Larva does not make respiratory vibration by the body (unlike C. vibratorius ). Tergalii unable to respiratory movements.

Subimago. CUTICULAR COLORATION. Cuticle with light brown and colorless areas. Head colorless, only antennae slightly tinged with pale brownish. Pronotum with light brown and colorless areas. Mesonotum at most brown, medioparapsidal suture contrastingly colorless, other sutures darkened ( Fig. 74 ). Thoracic pleura with light brown and colorless areas; lateropostnotal crest narrowly colored by dark brown, diffusively bordered by light brown ( Fig. 75 ). Base of wing light brown, other wing cuticle lighter. Legs at most colorless, tarsi slightly tinged with pale brownish; distal margin of each tarsal segment narrowly bordered by brown. Abdominal terga slightly colored with light brownish, microtrichiae brown. Abdominal sterna lighter. Cerci colorless.

HYPODERMAL COLORATION. As in imago.

TEXTURE. On all legs of both sexes last tarsal segment entirely covered by pointed microlepides; other tarsal segments at most part covered by blunt microlepides, apically by pointed microlepides ( Table 1).

Tarsal spines of subimago and imago. Tarsus of fore leg in male and female without apical spines. Tarsus of middle and hind leg of both sexes with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Table 1).

Imago, male. COLORATION ( Fig. 76–81 ). Head ocher. Turbinate eyes orange or yellow. Thorax ocher with light brown and white markings. Wing membrane colorless, veins pale ocher or colorless, basal part of costal and subcostal veins and part of costal brace brown. Femora of all legs ocher, with brown or reddish subapical band (hypodermal coloration); main trachea can be colored with dark brown; fore femur also with brown macula apically. Tibiae of all legs pale ocher, with or without hypodermal brownish coloration on inner side. Tarsi of all legs pale ocher, on fore leg cuticle of first segment can be tinged with brownish. Claws as light as tarsus. Abdominal segments I–III and V–VI colorless; segment IV either also colorless ( Fig. 77 ), or with tergum tinged with pale yellowish ( Figs 76, 78 ); segments VII–X pale yellowish; terga always without median markings; terga I– VII with contrasting dark paired transverse stripes near posterior margin: stripes on tergum I largest, stripes on terga IV and VII smallest. Color of these stripes varies from black to brown to light reddish; size of stripes varies from wide and connected medially to small and dot-shaped; stripes can be completely absent on terga IV and/or VII, or on terga II–IV and/or VII, or present on tergum I only. Lateral tracheal trunks and bases of sternal tracheae contrastingly colored by black or brown. Cerci entirely colorless, whitish.

STRUCTURE. Turbinate eyes with facetted surface wider than stem ( Figs 76–79 ). Pterostigma with 3–5 complete and incomplete oblique veins. Two marginal intercalaries nearly in each space (except for few posterior ones), length of most intercalaries less than distance between adjacent longitudinal veins (as in Fig. 32 ). Hind wing absent. Middle and hind tarsus long (about 0.7–0.8 of tibia length), its 1st+2nd segment subequal to 3rd, 4th and 5th segments together ( Table 4). Genitals as in Figs 88, 91 : unistyligers cylindrical; median sterno-styligeral muscle slender, consists of separate bunches, which are either convergent V-shaped ( Fig. 88 ), or crossed ( Fig. 91 ).

Imago, female. COLORATION ( Figs 82–85 ). Coloration of head, thorax, wings and legs similar to that of male. Abdomen ocher, terga VI and VII darker; terga with contrasting dark paired transverse stripes as in male; besides this, lateral parts of terga, pleura and lateral parts of sterna with composite brown and/or reddish maculation; particularly, at least each sternum II–VII with pair of contrasting brown lateral triangular maculae widened toward anterior margin of sternum.

STRUCTURE. Eyes relatively large, elevated above head surface ( Fig. 83 ). Wing either with one marginal intercalary in some spaces and no one marginal intercalary in other spaces, or without marginal intercalaries at all; in other respects wing venation as in male ( Figs 82, 85 ).

Egg. Oval, 0.1 mm length. Chorion irregularly crumpled, without relief ( Fig. 146 ).

Dimension. Fore wing length: male and female 4 mm.

Distribution. Peru.

Variability. Among specimens examined, two forms can be distinguished.

The first form («pallidus») includes specimens collected near Puerto Prado (6 male and 4 female imagoes and subimagoes associated with larvae). They have transverse stripes on abdominal terga at most light brown or reddish, poorly expressed or non-expressed ( Fig. 79 ); larvae of all these 10 specimens have prostheca of left mandible with 3 blunt processes and 3 pointed processes ( Fig. 65 ).

The second form («nigrostriatus») includes specimens collected near Contamana, Pampa Hermosa and Puente Itaya (11 male and 11 female imagoes and subimagoes associated with larvae). They have transverse stripes on imaginal abdominal terga at most well expressed, contrastingly dark brown or black ( Figs 76–78 ); among them, 18 larvae have prostheca of left mandible with 3 blunt processes and 4 pointed processes ( Fig. 63 ); 2 specimens have prostheca of left mandible with 3 blunt processes and 5 pointed processes; 2 specimens (male and female imagoes reared from larvae near Contamana) have prostheca of left mandible with 3 blunt processes and 3 pointed processes.

Among 2 specimens from Aguaytia (mature female larvae with developed subimaginal parts) one has features of the form «pallidus» (small brown hypodermal stripes on abdominal terga and left prostheca with 3+3 processes) and another has features of the from «nigrostriatus» (long black hypodermal stripes on abdominal terga and left prostheca with 3+4 processes).

The form «pallidus» well agrees with the original description of C. redactus based on larvae from Tingo Maria, for which dark hypodermal stripers on larval abdominal terga are not reported, and left prostheca is figured with 3 blunt processes and 3 pointed processes ( Roback 1966: Fig. 33 ).

I could not find other differences between these two forms, and regard them to be conspecific.

Comparison. Female imago, female subimago and female mature larva of C. redactus can be recognized by presence of contrasting brown hypodermal triangular maculae on abdominal sterna. Imago and subimago of both sexes can be recognized by having contrasting dark transverse lines or paired stripes on posterior margins of abdominal terga ( Figs 76–78 , 84 ), but in some individuals these markings are lighter ( Fig. 79 ) or absent. Larva can be distinguished from other species by the characters given in the key (see above).

Discussion. Preliminary description (without formal species name) was based on 6 larvae from three rivers near Tingo Maria in Peru ( Roback 1966). Formal original description was based on 2 of these larvae from the river Cantaria at Puente Perez ( Waltz & McCafferty 1987b).

Larvae described above, well agree with the original description; they were collected in various localities in Peru, and two specimens were collected near Aguaytia, about 50 km from the type locality.

McCafferty & Lugo-Ortiz (1996: 19–28) reported “ C. redactus ” from Honduras without more details.

Nieto and co-authors ( Nieto & Richard 2008; Nieto & Emmerich 2011) keyed larva of “ C. redactus ” as having denticles on posterior margin on abdominal tergum I, but did not report origin of this information. Actually, denticles on abdominal tergum I are absent or vestigial.

Massariol et al. (2013) described larvae and female imago determined by them as “ C. redactus ”. Description of this female imago is based on one female imago reared from larva in Reserva Ducke in Manaus, while description of larvae are based on 20 larvae collected in Igarape do Anta in Barcelos (F.F. Salles, personal communication). Photos of these larvae contradict the original description of C. redactus : in C. redactus “abdomen dark brown with light pattern ... tergites 4 and 8 lighter” ( Roback 1966) ( Waltz & McCafferty 1987b: Fig. 43 ), while on the photos abdominal terga I–IV and VII–VIII nearly lack cuticular pigmentation, and only terga V–VI and IX–X have brown cuticular pigmentation ( Massariol et al. 2013: Figs 5, 6). Female imago ascribed by Massariol et al. (2013) to C. redactus has abdominal terga with red and orange median markings and sterna washed with orange with lighter sigilla ( Massariol et al. 2013: Fig. 31–32 ), while female imagoes reared by me, have no such red and orange coloration, but have contrasting brown lateral spots on abdominal sterna ( Figs 82–86 ). Thus, larvae and female imago described by Massariol et al. (2013) under the name “ Cloeodes redactus ”, do not belong to this species, and possibly represent a new species.

Falcao et al. (2011) reported larvae of C. barituensis Nieto & Richard 2008 from Roraima, Brazil and gave photos of total dorsal view and labium ( Falcao et al. 2011: Figs 89, 90 ). Coloration of this larva markedly differs from described for C. barituensis . Judging by cuticular coloration (mainly brown with lighter blanks and lightest abdominal segments IV and VIII) and hypodermal coloration (dark transverse stripes on posterior margins of abdominal terga) larva from Roraima can belong to C. redactus .

Figures

FIGURES 37 – 57. Cloeodes spp., tergalii. 37 – 43, tergalii I – VII of Cloeodes vibratorius sp. n.; 44 – 57, tergalii I – VII of two individuals of Cloeodes redactus. For all three individuals vertical line corresponds to length of abdominal segment IV.

FIGURES 58 – 62. Cloeodes redactus. 58 – 59, immature last instar female larva; 60 – 62, larval exuviae: 60, fore leg; 61, middle leg and half of metanotum; 62, hind leg.

FIGURES 63 – 68. Cloeodes redactus, larva. 63 – 64, left and right mandibles with developing mandibles of next instar inside (form « nigrostriatus »); 65 – 66, left and right mandibles (form « pallidus »); 67, paraprocts; 68, tarsus of hind leg, view from outer side.

FIGURES 69 – 75. Cloeodes redactus. 69 – 73, larval exuviae: 69 – 71, abdomen; 72, right half of pronotum and mesonotum; 73, caudalii; 74 – 75, subimaginal exuviae: 74, half of mesonotum; 75, part of meso- and metapleuron with postsubalar sclerite, lateropostnotal crest and part of middle leg. Abbreviations: I – X, abdominal segments I – X.

FIGURES 76 – 81. Cloeodes redactus, male imagoes. 76 – 78, form « nigrostriatus »; 79, form « pallidus » (with partly shed subimaginal exuviae); 80, fore leg; 81, middle leg.

FIGURES 82 – 85. Cloeodes redactus, female imago, form « nigrostriatus ».

FIGURES 86 – 91. Cloeodes redactus, male genitals. 86 – 88, form « nigrostriatus »; 89 – 91, form « pallidus ». 86 and 89, subimaginal gonostyli developing under larval cuticle; 87 and 90, exuviae of subimaginal genitals; 88 and 91, genitals of male imago (gonovectes shown by interrupted line; at right half gonovectal and gonostylar muscles shown by interrupted lines; median styligeral muscle shown by interrupted lines, areas of anterior attachment of median paraproctal muscles shown by dotted lines). Abbreviations: 1, 2, 3, segments of gonostylus; m. IX-X, areas of anterior attachment of median paraproctal muscles (i. e., muscles going from sternum IX to common base of paraprocts); m. sg, median sterno-styligeral muscle; usg, unistyliger.

FIGURES 26 – 31. Cloeodes vibratorius sp. n. 26 – 27, female imago; 28 – 31, male imago: 28, femur of fore leg; 29, femur of hind leg; 30, abdominal terga and sterna spread on slide; 31, lateral view. Abbreviations: I – X, abdominal segments I – X.

FIGURES 32 – 36. Cloeodes vibratorius sp. n. 32, fore wing of male; 33, subimaginal gonostyli developing under larval cuticle; 34, exuviae of subimaginal genitals; 35, genitals of male imago (gonovectes shown by interrupted line; at right half gonovectal and gonostylar muscles shown by interrupted lines; median styligeral muscle shown by interrupted lines, areas of anterior attachment of median paraproctal muscles shown by dotted lines); 36, gonostylus (32, 34 – 35, holotype). Abbreviations: 1, 2, 3, segments of gonostylus; m. IX-X, areas of anterior attachment of median paraproctal muscles (i. e., muscles going from sternum IX to common base of paraprocts); m. sg, median sterno-styligeral muscle; usg, unistyliger.

FIGURES 6 – 12. Cloeodes vibratorius sp. n., larva. 6, head of female larva; 7, maxilla; 8, apex of maxilla; 9 – 11, left and right mandibles; 12, labrum (apical setae not shown) (10, 12, holotype). Abbreviations: ds 1, ds 2, ds 3, dentisetae.

FIGURES 13 – 17. Cloeodes vibratorius sp. n. 13, labial palp; 14, labium (left half in ventral view with muscles shown by interrupted lines; right half in dorsal view); 15, paraprocts; 16, tibia of fore leg, anterior view (seta and suture on opposite side shown by interrupted lines; only two setae of proximal arched row are shown); 17, tarsus of hind leg, view from outer side (13, holotype).

FIGURES 18 – 25. Cloeodes vibratorius sp. n. 18 – 23, larval exuviae: 18, tergum IX; 19, paraprocts; 20, portion of sternum IV with setal row; 21, fore leg; 22, middle leg and half of metanotum; 23, hind leg; 24 – 25, subimaginal exuviae: 24, left half of mesonotum; 25, postsubalar sclerite and lateropostnotal crest (21 – 23, holotype).

FIGURES 145 – 146. Eggs. 145, Cloeodes vibratorius sp. n.; 146, Cloeodes redactus.