Thraulodes telegraphicus Needham & Murphy 1924

Kluge, Nikita J., 2020, Systematic position of Thraulodes Ulmer 1920 (Ephemeroptera: Leptophlebiidae) and descriptions of new and little-known species, Zootaxa 4756 (1), pp. 1-142 : 34-43

3. Thraulodes telegraphicus Needham & Murphy 1924

( Figs 66–121 )

Thraulodes telegraphicus Needham & Murphy 1924: 41 , figs 99, 102 (♂ and ♀ imago); Traver 1946: 428 (holotype, synony-

my); Traver & Edmunds 1967: 379, figs 40, 60 (♂ and ♀ imago).

Thraulodes plicatus Needham & Murphy 1924: 42 , fig. 100 (♂ and ♀ subimago) (holotype redescribed and synonymy estab-

lished by Traver 1946).

Thraulodes sp. 1, sp. 2, sp. 4, sp. 6: Roback 1966: 145, pl. 13 (larva).

Material examined. PERU: Region Junin, Provincia Satipo: Rio Satipo in Satipo, 5.I.2006, coll. N. Kluge: 1 L-S ♂, 4 larvae; Rio Sotorani near Pichanaki (10°57’S, 74°52’W), 8–10.I.2006, coll. N. Kluge: 1 L-S-I ♀, 1 S-I ♂, 2 S-I ♀, 24 larvae; Puerto Prado (11°09’S, 74°14’W), 10–14.IX.2013, coll. N. Kluge & L. Sheyko: 11 L-S-I ♂, 5 L-S-I ♀, 6 L-S ♀, 10 L/S ♂, 6 L/S ♀, 63 larvae; Rio Shima 5 km E Puerto Prado, (11°07’S, 74°13’W), alt. 350m, 6.I.2006, coll. A. Petrov: 2 S(m); Region Huanuco, Provincia Leoncio Prado, Rio Chunatahua near Cayumba (9°30’S, 75°57’W) and Rio Cantaria near Puente Perez (9°21’S, 75°58’W), 13–25.I.2006, coll., N. Kluge: 20 L-S-I ♂, 2 L-S/I ♂, 7 L- S ♂, 27 L-S-I ♀, 2 L-S/I ♀, 15 L-S ♀, 2 L/S ♂, 2 L/S ♀, 74 larvae; Region Huanuco, Provincia Ambo: river Huertas near Ambo, 28.VIII.2013, coll. N. Kluge: 2 larvae; Huacar (10°9’S, 76°15’W), 1–5.IX.2013: 1 L-S-I ♀, 1 L-S ♀, 32 larvae; Region Loreto, Provincia Ucayali, Aguas Termales, 17 km NNE Contamana (7°12’S, 74°57’W), 26.VII– 4.VIII.2013, coll. N. Kluge & L. Sheyko: 1 S ♂.

Descriptions. Larva. CUTICULAR COLORATION ( Figs 66–67 , 70–82 ). Dorsal side of head, thorax and abdomen with highly variable coloration. Head either medially brown and laterally light ( Fig. 66 ), or mostly brown ( Fig. 67 ); pronotum either mostly light with contrasting brown maculae on these or that margins ( Figs 66 , 70–73 ), or mostly brown with light lateral margins ( Figs 67 , 74–75 ); mesonotum mostly brown, either with several separate blanks, or crossed by wide blanks ( Fig. 77 ); abdominal terga either brown with blanks on lateral parts, or terga I–III and VI–VII brown and terga IV–V and VIII–X contrastingly light ( Figs 66 , 81 ). Femora with brown and light areas, tibiae and tarsi unicolor light ( Figs 77–80 ).

HYPODERMAL COLORATION ( Fig. 68 ). Abdominal terga, besides cuticular coloration (see above) often with paired transverse or round blackish spots corresponding to midway gray spots of imago. Tergalii either light grayish, or light brownish-reddish, or colorless; tracheae often black, sometimes light gray or colorless.

SHAPE AND SETATION. Clypeus widened distally; labrum 1.2–1.5 times wider than clypeus ( Fig. 76 ). Labrum widest at 1/3 length from base; initial fore margin (often turned ventrally) without median emargination, slightly convex, with all 5 denticles wide; anterior transverse setal row regular, as wide as 3 or 5 denticles ( Fig. 88 ). Maxilla with 14–17 pectinate setae in apical-ventral row.

Femora: Stout setae on anterior surface parallel-sided or widened distally, apically blunt ( Fig. 83 ). Irregular row of hairs near inner margin sparse on fore femur, dense on middle and hind femora.

Fore tibia ( Fig. 84 ): outer hairs form two irregular rows; inner-anterior row of recurved hairs absent; inner-anterior row of stout setae represented by few (2–4) blunt stout setae near tibia base; inner field of stout pointed setae dense (i.e. setae longer than distances between them), consists of pointed bipectinate and smooth setae, situated irregularly (about 3 setae in cross section).

Hind tibia ( Fig. 85 ): outer-anterior row of stout setae consists of two clearly different types of stout setae: numerous short rounded ones and several long spoon-like ones; outer-posterior row of stout setae consists of stout blunt setae of various lengths; hairs located between these rows, numerous and form more than one row (besides row of hairs posteriad of outer-posterior row of stout setae); stout setae of inner-anterior row short, widened distally and truncated.

Claws with 6–10 denticles on rigid portion, with or without minute denticles on articulatory portion.

Tergalii ( Figs 90–93 ): wide; on both lamellae main trachea with many branches directed toward costal and anal margins; dorsal lamella with costal margin most convex in proximal part and anal margin most convex in distal part, gradually narrowed toward apex, with slender apical filament; ventral lamella widest near base, gradually narrowed toward apex, with slender apical filament.

Male genitalia in last larval instar ( Figs 106–109 ): protogonostyli separated one from another by deep emargination. Each protopenes lobe with gonopore-bearing process located near median margin; gonopore opened caudally.

Subimago ( Figs 94–95 , 115 ). CUTICULAR COLORATION. Cuticle mostly colorless. Flagellum of antenna brown. Pronotum colorless. Mesonotum with colorless and brown areas; antelateroparapsidal suture contrastingly colored with dark brown, lateroparapsidal suture and some other areas lighter brown; chromozone of medioscutum light brown, chromozone of submedioscutum contrastingly lighter ( Figs 94–95 ), sometimes colorless. Fore femur mostly colorless, more or less colored by brown on outer margin and on apex; middle and hind femora colorless; on each leg tibia and tarsus light brown. Cuticle of wings colorless, microtrichia light brownish.Abdominal terga mostly colorless, diffusively shed by brown medially; sterna colorless; gonostyli colorless; caudalii light brownish.

TEXTURE. On tarsi of all legs, 1st tarsomere with microtrichia (as tibia), 2nd–5th tarsomeres coved by blunt microlepides; pointed microlepides present near apical margins of 2nd–4th tarsomeres of middle and hind legs (as in Fig. 473 ).

Male imago ( Figs 96–99 , 101–105 , 110–120 ). Described by Needham & Murphy (1924) and Traver & Edmunds (1967).

Head brown. Antenna with scapus ocher with brown macula; pedicellus proximally ocher, distally brown; flagellum brownish with ocher base. Dorsal eyes contiguous medially, dull orange.

Prothorax ocher with pair of dark brown oblique stripes on pleura. Mesonotum and metanotum ocher or/and brown. Lateral sides of pterothorax with brown and ocher areas. Ventral side of pterothorax with mesothoracic epi- sterna, basisternum, furcasternal protuberances and furcasternal median impression either brown with ocher ( Fig. 98 ), or brown.

Cuticle of legs mostly colorless; on fore leg apex of femur and base of tibia with cuticle brown. Legs with following hypodermal pigmentation on whitish-ocher background: Fore femur brown with diffusive spot(s) or interrupted band on proximal half and with multicolored apical band occupying distal 0.4 of femur; apical band brownish-yellowish, proximally bordered by brown, on inner and outer sides with brown longitudinal stripes. Middle and hind femora with multicolored pre-apical band within apical 0.4 of femur; pre-apical band brownish-yellowish, proximally bordered by brown, on inner and outer sides with brown longitudinal stripes; apical 1.0 of femur nonpigmented. Tibia of each leg apically dark brown. In selected individual, relation of fore femur to fore wing length 75:325; proportions femur/tibia/tarsomeres on fore leg 75:107:2:26:20:18:10; on middle leg 70:80:4:4:3:3:9; on hind leg 85:90:4:4:3:3:9.

Fore wing with longitudinal and cross veins ocher; costal braces and bases of Sc and R bordered by brown. Costal cross veins proximad of bulla either absent, of few, colorless and very thin, so that visible only under high magnification (as in Fig. 142 ); occasionally proximal part of costal field with several well-visible cross veins. Pterostigmatic cross veins moderately dense, oblique, arched, non-branched ( Fig. 96 ) or occasionally anastomozed. Hind wing with veins colorless, with brown macula on costal brace. Proximal coloration of fore and hind wings appears during larval development and present both in subimago and imago.

Abdominal hypodermal coloration: Tergum I ocher-brown. Terga II–VI mostly light ocher, translucent, with or without brown band on posterior margin, medially more or less tinged with darker ocher; terga III–VI or III–VI with pair of smaller or larger gray midway spots; pleura with gray stigmatic dots. Terga VII–X with orange or ocherbrownish coloration: tergum VII mostly colored with latero-posterior areas whitish, tergum VIII colored anteriorlymedially and whitish posteriorly-laterally, terga XI–X mostly colored; lateral margins of terga VIII and IX with blackish-brown longitudinal stripes. Abdominal sterna I–IX light ocher; posterolateral angles of sterna VII and VIII with pair of blackish-brown spots. Caudalii with segments alternated as following: black; white; white proximally and black distally; white.

Genitalia ( Figs 101–120 ): Styliger ocher. Gonostyli either entirely ocher, or with long segment distally brownish ( Fig. 102 ). Penis either entirely light ocher, or partly with brown cuticular or/and hypodermal coloration. Dorsal extension of styliger small and prominent. Penis lobes long and divergent; each lobe gradually widened apically, without lateral pouch; in other respects shape of penis lobes variable. Apico-lateral area usually forming well-expressed ear, occasionally without ear ( Fig. 119 ). Telopenes in form of spear-like rolls, more or less straight, arising on dorsal side of penes and directed medially or medially-caudally (not inclined dorsally or ventrally), with groove opened dorsally-laterally; length of spear-like rolls varies individually.

Female imago ( Fig. 100 ). Coloration of head, thorax, legs, wings and caudalii as in male. Abdominal terga II–VI ocher with brownish; coloration of other abdominal segments as in male.

Eggs ( Fig. 121 ). Mostly barrel-shaped. Each KTC surrounded by smooth, ring-like cover. Few small protuberances located close to these rings; other chorion without protuberances.

Dimension. Fore wing length (and approximate body length) 7.5–9.5 mm (this study); 10–11 (according to Traver & Edmunds 1967).

Distribution. Peru ( Needham & Murphy 1924; this study).

Synonymy. Th. telegraphicus is one of the most common mayfly species in low altitudes of eastern slopes of Andes (regions Junin and Huanuco of Peru). Originally ( Needham & Murphy 1924), imago of this species was described from the Region of Junin under the name Th. telegraphicus , and subimago—under the name Th. plicatus . Synonymy of these two names was established by Traver (1946), who reexamined the both holotypes.

Larvae were described by Roback (1966) from Region of Huanuco without species names. Probably, among six larval forms distinguished by Roback, the forms reported as Thraulodes sp. 1, Th. sp. 2, Th. sp. 4 and Th. sp. 6 belong to Th. telegraphicus . The forms «2» and «6» have contrastingly light pronotum and contrasting dark and light areas on mesonotum and abdomen (as in Figs 66 , 70–73, 81 ), while the forms «1» and «4» have more uniform brown coloration (as in Figs 67 , 75, 82 ). The forms «1» and «2» have tergalii with tracheal branches by sides of main trachea, while the forms «4» and «6» are said to lack these branches. Among material examined by me, all larvae have side trachea, but in some larvae from Huacar the side trachea are colorless and can be overlooked. Two other larval forms described by Roback, are different from Th. telegraphicus . The form «3» (describes as one larva from Puente Perez) differs from Th. telegraphicus by acute small stout femoral setae ( Roback 1966: fig. 184); its systematic position remains unknown. The form «5» is T. quevedoensis (see below).

Figures

FIGURE 121. Thraulodes telegraphicus, eggs.

FIGURES 66–69. Thraulodes telegraphicus. 66–67, last instar larvae with intensive cuticular coloration and poorly visible hypodermal coloration; 68, younger larva with light cuticle and well visible blackish hypodermal markings on abdomen; 69, claw. Abbreviations: artic.p., articulatory portion; rigid p., rigid portion of claw.

FIGURES 70–85. Thraulodes telegraphicus, larval exuviae (70–82 at the same magnification): 70–75, right half of pronotum and mesonotum; 76, head with labrum and mandibles; 77–78 and 79–80, fore and hind legs of two specimens; 81–82, abdominal terga; 83, anterior side of hind femur; 84, anterior side of fore tibia; 85, anterior side of hind tibia. Abbreviations: i-a.r, inneranterior row of stout setae; o-a.r, outer-anterior row of stout setae; o-p.r, outer-posterior row of stout setae.

FIGURES 86–93. Thraulodes telegraphicus. 86, median longitudinal section of labrum in usual condition; 87, the same, in spread condition; 88, labrum in spread condition in dorsal view (setal rows shown as dotted lines); 89, transverse section of hind tibia; 90–93, tergalii I, III, VI and VII. Abbreviations: anter.r, anterior transverse row of setae; dent., five denticles separated by six sensilla; poster.r, posterior transverse row of setae; i.f, inner field of stout pointed setae; i-a.h, inner-anterior recurved hairs; i-a.r, inner-anterior row of stout setae; o-a.r, outer-anterior row of stout setae; o.h, outer hairs; o-p.r, outer-posterior row of stout setae; p.r, row of posterior pointed pectinate setae.

FIGURES 94–100. Thraulodes telegraphicus. 94–95, right half of subimaginal exuviae of mesonotum of two different specimens; 96, apex of fore wing; 97–98, male imago in lateral and ventral view; 99, abdominal terga and sterna of male imago (black dots are algae under cuticle); 100, female imago. Abbreviations: acz.ms, achromozone of medioscutum; acz.sms, achromozone of submedioscutum; cz.ms, chromozone of medioscutum; cz.sms, chromozone of submedioscutum; mp.s, medioparapsidal suture; ms.cz.s, medioscutal chromozonal suture; sms.cz.s, submedioscutal chromozonal suture.

FIGURES 101–105. Thraulodes telegraphicus. 101–102, genitalia of the same specimen; 103, penis of another specimen; 104– 105, part of penis with telopenis forming spear-like roll.Abbreviations: bl, basal lobe; dp, dorsal projection of styliger; gd, gonoduct; gs1, gs2, gs3, gs4, segments of gonostylus; m.gs, gonostylar muscle; m.lat, lateral penial muscle; m.med, median penial muscle; m.s, styligeral muscle; m-v.r, medio-ventral ridge (recurved flap); sp, seminal pump; tp, spear-like roll (telopenis).

FIGURES 106–111. Thraulodes telegraphicus. 106, protogonostyli of last instar larva (hypoderm of gonostyli and styliger and hidden cuticle of protopenis shown by interrupted lines); 107, enlarged cuticle of protopenis and hypoderm of developing penis inside; 108–109, the same, at later stage of development; 110–111, genital of male imago (110, styliger ventrally; 111, penis dorsally at left and ventrally at right). Abbreviation: bl, basal lobe; dp, dorsal projection of styliger; gd, gonoduct; m.gs, gonostylar muscle; m.med, median penial muscle; m.s, styligeral muscle; m-v.r, medio-ventral ridge (recurved flap); sp, seminal pump; tp, spear-like roll (telopenis).

FIGURES 112–120. Thraulodes telegraphicus. 112, genitalia of male imago in lateral view; 113, apex of penis with spear-like roll in dorsal view; 114, genitalia of the same specimen in ventral view; 115, their subimaginal exuviae at the same magnification; 116, hind wing; 117–120, penis of four different specimens. Abbreviations: dp, dorsal projection of styliger; tp, spear-like roll (telopenis).

FIGURES 464–473. Thraulodes nigrabdominalis sp. n. (holotype). 464, left half of subimaginal exuviae of mesonotum; 465, 466, male imago; 467–469, fore, middle and hind legs of male imago; 470, subimaginal exuviae of fore leg; 471, genitalia of male imago; 472, apex of penis; 473, subimaginal exuviae of tarsus.

FIGURES 133–142. Thraulodes panamensis sp. n. 133, pterostigma; 134–136, male imago in lateral in ventral and dorsal view; 137–138, left half of subimaginal exuviae of mesonotum of two different specimens; 139, female imago; 140–141, genitalia in lateral and ventral view; 142, costal margin of fore wing (arrows show proximal costal cross veins) (133, 134, 136, 137, 141, 142, holotype).