Centroptella (Crassolus?) bifida ( Shi & Tong 2019 ), 2020Centroptella Chopralla Waltz & McCafferty 1987

Kluge, Nikita J., 2021, Review of Centroptella Braasch & Soldán 1980 (Ephemeroptera, Baetidae), Zootaxa 5054 (1), pp. 1-144 : 94-141

2.6. Centroptella (Crassolus?) bifida ( Shi & Tong 2019)

( Fig. 421 )

Bungona (Chopralla) bifida Shi & Tong 2019: 578 (larva);

Centroptella bifida: Kluge, Godunko & Svitok 2020: 92 .

Distribution. China.

Comments. Centroptella bifida is known as larvae only. According to the original description, its larvae differ from others as «submarginal setae on labrum are forked apically» ( Shi & Tong 2019: 578, 584, fig. 37). Actually, these setae are bifurcate in some other species as well, particularly in C. ingridae ( Fig. 395 ), which in the same paper by Shi & Tong (2019) was redescribed under the name « Bungona (Chopralla) liebenauae » and told to have «labrum surface with simple submarginal setae» ( Shi & Tong 2019: 584).

Form of right prostheca (pressed to kinetodontium and lacking long process) suggests that C. bifida belongs either to Chopralla , or to Crassolus . Male larvae used for the original description, are not mature enough to have subimaginal gonostyli, so their pose is unknown (Xiaoli Tong, personal communication). Judging by the presence of denticles on posterior margins of abdominal terga II–X and the presence of setal rows (j) on abdominal sterna III–VI ( Shi & Tong 2019: 578), C. bifida probably belongs to Crassolus .

Judging by the picture provided by Xiaoli Tong ( Fig. 421 ), hypodermal coloration of mature male larva is limited by reddish stripes on posterior margins of abdominal terga, that clearly separates this species from such representatives of Crassolus , as C. ludmilae , C. pontica , C. inzingae and C. saxophila , whose males have more extensive hypodermal coloration on abdominal terga.

2.7. Centroptella (Crassolus?) sp. « Nepal »

( Figs 422–428 )

Material examined. Central NEPAL, Helambu Area , 27°55’N 83°30’E, Gohare Khola opposite to village Mahenkai, about 500 m from falling to Melamchi River, 4.III.2007, coll. M. Chertoprud: 1 male larva of penultimate instar (other specimens in Chertoprud’s collection) .

Distribution. Nepal.

Comments. Characters of this larva are reported in the Table 3 and shown on the Figs 422–428 ). Pose of developing subimaginal gonostyli is unknown; judging by the presence of denticles on posterior margins of abdominal terga II–X and the presence of setal rows on abdominal sterna III–VI, this species probably belongs to Crassolus .

3. Subgenus Chopralla Waltz & McCafferty 1987

( Figs 1 , 429–686 )

Genus Chopralla Waltz & McCafferty 1987a: 182 .

Subgenus Chopralla of genus Bungona: Salles, Gattolliat & Sartori 2016: 104 .

Synonym of Centroptella: Kluge, Godunko & Svitok 2020: 88 .

Type species: Centroptella ceylonensis Müller-Liebenau 1983 .

Diagnosis (see Centroptella s. l: Classification and Table 2).

(1) In larva, posterior arm of U-shaped row of long setae on fore- and middle tibiae more transverse than longitudinal and crosses tibia [on hind leg longitudinal—see Centroptella s. l. (2)] ( Figs 560–562 , 662–666 ). The same in Crassolus , in contrast to Centroptella s. str.

(2) Larval claw usually with 2 rows of denticles ( Figs 548–548 ). The same in Crassolus , in contrast to Centroptella s. str.

(3) In larva, at least abdominal terga I–IV without denticles on posterior margins; in various species denticles are present beginning either from tergum V (in C. colorata , C. papuanica sp. n., C. ghatensis sp. n. and C. kangi sp. n.), or from tergum VIII (in C. bintang ), or from tergum IX (in C. ceylonensis , C. pusilla and C. rufostriata sp. n.).

(4) In larva, abdominal sternum VII without regular rows of bifurcate setae (j), so that such rows occur on sterna IV–V only (in contrast to Centroptella s. str. and Crassolus , which have regular rows on sterna IV–VI).

(5) In last larval instar, developing subimaginal gonostyli folded under larval cuticle in the « Nigrobaetis - type » pose, i.e. with 3rd segments directed caudally ( Figs 469 , 529 , 571 , 609 , 627 , 654 ). The same in many other Baetovectata, but in contrast to Centroptella s. str. and Crassolus , whose gonostyli are folded in the peculiar « Crassolus - type » pose.

(6) Gonovectes without halberd-like widenings apically ( Figs 465–468 , 527 , 572 ) (in contrast to Crassolus ).

(7) Sterno-styligeral muscle is retained (in contrast to Crassolus ), being rather weak and often divided into pair of convergent muscles ( Figs 465–466 , 527 , 572 ) (the same in Centroptella s. str.).

Other characters. In larva, anterior margin of mesonotum is sharply curved ( Figs 431 , 499 , 434 , 599 , 633 ), in contrast to straight or shallowly curved in Crassolus and Centroptella s. str. ( Figs 24–25 , 77 , 104 , 128 , 185 , 274 , 338–340 ).

Bifurcate setae on metasternum (i) (see Centroptella s. l.) form a pair of regular rows ( Fig. 517 ) (in contrast to Centroptella s. str. and Crassolus , whose bifurcate setae of metasternum are arranged less regularly).

Both male and female imagines have characteristic coloration: thorax contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; abdomen of female has the same coloration ( Figs 531 , 577 , 519 ); abdomen of male has either the same coloration ( Figs 451 , 520 ), or has no reddish coloration on most of terga ( Fig. 566 ). This coloration differs from Crassolus and Centroptella s. str., whose thorax has brown cuticular coloration both on dorsal and on ventral sides at least in males ( Figs 49 , 93 , 143 , 159–160 , 298 , 379 ).

Distribution. The Oriental Region.

Composition. Centroptella (Chopralla) ceylonensis (= C. similis syn. n.), C. (Chopralla) ghatensis sp. n., C. (Chopralla) rufostriata sp. n., C. (Chopralla) papuanica sp. n., C. (Chopralla) pusilla , C. (Chopralla) kangi sp. n., C. (Chopralla) colorata (= C. fusina syn. n.), C. (Chopralla) bintang .

3.1. Centroptella (Chopralla) ceylonensis Müller-Liebenau 1983

( Figs 429–488 )

Centroptella ceylonensis Müller-Liebenau 1983: 486 (larva); Centroptella similis Müller-Liebenau 1983: 487 (larva), syn. n.

Chopralla ceylonensis Waltz & McCafferty 1987a: 183 ;

Chopralla similis: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) ceylonensis: Salles, Gattolliat & Sartori 2016: 104 , figs 4E, 5G, 6G, 6M;

Bungona (Chopralla) similis: Salles, Gattolliat & Sartori 2016: 104 ;

Bungona (Chopralla) pusilla and Bungona (Centroptella) pusilla: Selvakumar, Kubendran, Chandra Kailash & Sidhu Avtar Kaur 2017: 274 (larva).

Material examined. SRI LANKA: border of Uva and Central Provinces, tributary of river Uma near Randenigala dam, 13–14.I.2011, coll. N. Kluge & L. Sheyko: 5 larvae ; Uva Province, Badulla District, Haputale , 15–23.I.2011, coll. N. Kluge & L. Sheyko: 1 L-S-I ♂ ; Uva Province, Badulla District, Lemastota Oya , 20–21.I.2011, coll. N. Kluge & L. Sheyko: 2 larvae ; Sabaragamuwa Province, river Seetha Sangula in Dalhausie near Sri Pada ( Adam’s Peak ), 24–26.I.2011, coll. N. Kluge & L. Sheyko: 2 S-I ♂, 22 larvae ; the same place and collectors, 1–10.II.2020: 5 L-S-I ♂, 11 L-S-I ♀, 1 S-I ♀, 31 larvae; river Battulu Oya, Fishing Hut (S of Sri Pada = Adam’s Peak ), 8–9.II.2020: L/S ♀, 1 larva ; Central Province, Nuwara Eliya District, river Mahaveli Ganga in Ginigathhena (17 km N Hatton), 31.I–3.II.2011, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 9 S-I ♂, 6 I ♂, 5 S-I ♀, 3 I ♀, 1 S ♀, 3 larvae; Belihuloya , 15– 26.I.2020, coll. N. Kluge & L. Sheylo: 1 L-S-I ♀ , 1 I ♀, 6 larvae; Wellawaya , 22.I.2020, coll. N. Kluge & L. Sheyko: 2 larvae . INDIA: State Karnataka, Udupi District, river Seethanadhi-hole near Someswar , 11.I–1.II.2013, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 3 L-S-I ♀, 1 L-S ♀, 9 larvae. State Kerala, Kottayam District, Erumeli , 22.I.2016, coll. N. Kluge & L. Sheyko: 3 L-S-I ♀ , 3 L; Ernakulam District, Malayatoor , 14–15.II.2016, coll. N. Kluge & L. Sheyko: 10 L-S-I ♂ , 1 L-S ♂, 1 L-S-I ♀, 6 larvae.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 430 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 431 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs. 434–436 ; as in Figs 554–559 ). Abdominal terga with contrasting brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 429 ; Müller-Liebenau 1983: figs 4, 6); medio-anterior and medioposterior sigilla lighter than background (as in Fig 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Figs 429, 437–438 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Figs 456–457, 460 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base (as in Fig. 507 ; Müller-Liebenau 1983: fig.1a). Mandibles ( Figs 439–448 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 432 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them (in some individuals differently on left and right sides, as in Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 434–436 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1983: figs 1i–l). Claw with two rows of 1–6 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1983: figs 1m –n).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles (in single female from Malayatoor terga VII–VIII with irregular small denticles); tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 449 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles (as in Fig. 501 ), with or without 1 pair (rarely 2 pairs) of smaller denticles by sides of them ( Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially ( Fig. 450 ), in male without denticles between protogonostyli ( Fig. 469 ). Paraprocts with thin and colorless antero-median apodeme, with large pointed denticles on posterior margin (as in Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 475–488 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment) located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Figs 437–438 ).

SCALES: Certain areas of pronotum and mesonotum with small, colorless scales. Femora, tibiae and tarsi with colorless, short, oval scales. Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 449 ; Müller-Liebenau 1983: figs 5, 7). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 450 ).

STOUT SETAE: Outer side of femur with regular row of 4–15 long, clavate setae and 2 subapical setae of same form ( Figs 434–436 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Müller-Liebenau 1983: figs 1i–l) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 469 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 453 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 452 ). Legs mostly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and cerci light brownish, sterna nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 451 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 454–455 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in Fig. 392 ). Abdominal terga dark ochre with extensive reddish-brown hypodermal markings ( Figs 456–463 ); abdominal sterna ochre, some sterna with pair of reddish-brown longitudinal stripes sublaterally ( Figs 456–458 ; lateral tracheal trunks and some other abdominal tracheae bordered with blackish. Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Figs 465–468 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment either short, or elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Figs 470–472 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 473–474 ). Oval; chorion with isoedric net-like relief, with very sparsely and irregularly arranged minute pores. Sperm guide small and deep.

Dimension. Fore wing length of male and female 3.2–5 mm.

Synonymy of C. ceylonensis and C. similis . Müller-Liebenau (1983) described Centroptella ceylonensis and C. similis bases on larvae only and stated that «The main difference between both species is the lack of hind wing pads in C. similis sp. n. (in both sexes)». In reality, presence and size of larval vestiges of hind protoptera varies individually in this species (while hind wings are always completely absent in winged stages); among examined larvae of C. ceylonensis from one and the same population (Delhausie near Sri Pada), some individuals have wellexpressed vestiges of hind protoptera, some have very small convexities instead of them, some have not vestiges of hind protoptera at all, and some have vestige of hind protopteron on one side and no such vestige on another side (as in Figs 510–511 ).

Besides this character, Müller-Liebenau (1983) noted: «Also the size ratio of turbinate eyes and the lateral eyes appears somewhat different in the male nymphs of both species». Actually, the drawings of male larval heads ( Müller-Liebenau 1983: figs 1p and 2a) do not allow to distinguish these species.

Müller-Liebenau (1983) also noted that scales on abdominal terga are «in general a little more elongate in C. ceylonensis sp. n. ». Actually, the scales significantly vary in size and proportions on terga of one and the same individual, that is visible from the photos in the original description ( Müller-Liebenau 1983: figs 5, 7).

According to the original description of C. similis , its cuticular coloration of larval abdomen somewhat differs from that of C. ceylonensis . Actually, the coloration of abdominal cuticle varies individually in greater degree than it is shown on the pictures in the original descriptions of C. ceylonensis and C. similis . Thus, the name C. similis should be regarded as a junior synonym of C. ceylonensis .

Distribution. Southern India (Western Ghats) and Sri Lanka.

Variability. Hypodermal coloration of abdomen significantly varies individually within each population ( Figs 456–463 , 470–472 ). The examined specimens from Sri Lanka are larger (fore wing length 4.5–5 mm) then the examined specimens from India, whose fore wing length is 3.5–4 mm. However, examination of many imagines of both sexes individually associated with their larval and subimaginal exuviae allowed to conclude that populations from Sri Lanka and India belong to one and the same species.

Species determination. In Sri Lanka Centroptella (Chopralla) ceylonensis is the single species of Chopralla and cannot be confused with other species. In southern India C. (Ch.) ceylonensis can be confused with the closely related species C. (Ch.) ghatensis sp. n., whose larvae differ only by presence of denticles on abdominal terga V–VIII and winged stages are indistinguishable from C. (Ch.) ceylonensis .

Selvakumar et al. (2017) determined larvae of Chopralla from southern India as C. pusilla . Resolution of the total photo ( Selvakumar et al. 2017: fig. 6) allows to see a pair of large denticles on lateral angles of the posterior projection of tenth abdominal tergum, that differs C. ceylonensis ( Fig. 433 ) from C. pusilla ( Fig. 630 ); other characters of the larvae reported by Selvakumar et al. (2017) (i.e. coloration, mandibular structure, arrangement of denticles on abdominal terga) also agree with the characteristics of C. ceylonensis . The photo indicated as «Foreleg» ( Selvakumar et al. 2017: fig. 10) actually belongs to the hind leg.

3.2. Centroptella (Chopralla) ghatensis sp. n.

( Figs 489–533 )

Etymology. Distributional area of this species is located in the mountains of Western Ghats.

Material examined. Holotype: L-S-I ♂ {number [XXI](1)2013}: INDIA, State Tamilnadu, Tirunelveli District, Courtallam , Chittar river near Peraruvi (= Main Falls ), 5.II.2013, coll. N. Kluge & L. Sheyko. Paratypes: the same locality and collectors, 3–7.II.2013: 6 L-S-I ♂ , 3 L-S-I ♀, 45 larvae; Theni District , Suruli, 24–25.I.2016, coll. N. Kluge & L. Sheyko: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 498 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 499 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless ( Fig. 514 ). Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 502–504 ; as in Figs 554–559 ). Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 500 ); medio-anterior and medioposterior sigilla lighter than background (as in Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than on Fig. 500 Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Fig. 496 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Fig. 497 ).

SHAPE: Labrum either equally wide at base and middle ( Fig. 507 ), or slightly wider at base. Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Figs 505–506 ). Prostheca of left mandible with 3 blunt processes and 2 pointed processes ( Fig. 505 ). Prostheca of right mandible directed medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes ( Fig. 506 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 508 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them; in some individuals differently on left and right sides ( Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 502–504 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [as in Figs 560–563 ; see Centroptella s. l. (2)]. Claw with two rows of 3–6 denticles in each [(as in Figs 547–548 ; see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 518 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles ( Fig. 501 ), with or without 1 or 2 pairs of smaller denticles by sides of them (as in Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 519 ). Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially (as in Fig. 450 ), in male without denticles between protogonostyli ( Fig. 529 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 501 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 489–495 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment), located on every 4th or every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 496 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 513 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga elongate, oval, with longitudinal striation ( Fig. 518 ). Scales on abdominal sterna, metanotum, thoracic pleura and leg coxa shorter and partly colorless ( Fig. 519 ).

STOUT SETAE: Outer side of femur with regular row of 6–17 long, clavate setae and 2 subapical setae of same form ( Figs 502–504 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets ( Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia,; on hind leg longitudinal (as in Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 529 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 524 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 525 ). Legs mostly colorless or light brownish, with brown articulations. Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 520 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely dark orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 521–533 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 522–523 ; as in Fig. 392 ). Abdominal terga mostly reddish- brown with lighter ochre areas; abdominal sterna ochre; lateral tracheal trunks and some other abdominal tracheae bordered with blackish ( Fig. 526 ). Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Fig. 527 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Fig. 530 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 532–533 ). Oval; chorion with isoedric net-like relief, without pores, Sperm guide small, occupying one cell of net-like relief.

Dimension. Fore wing length 4.8 mm (male); 5.4 mm (female).

Distribution. Western Ghats ( India).

Comparison. Larva and imago of the south Indian species Centroptella (Chopralla) ghatensis sp. n. are very similar to C. (Chopralla) ceylonensis , which inhabit both southern India and Sri Lanka. Larva of C. ghatensis sp. n. differs from C. ceylonensis in having well developed denticles on abdominal terga V–VIII. In India, C. ghatensis sp. n. and C. ceylonensis were collected in different localities, never being found together.

3.3. Centroptella (Chopralla) rufostriata sp. n.

( Figs 1 , 534–590 )

Etymology. Allusion to hypodermal coloration of abdmen, which consists of reddish cross tripes stripes on posterior margins of terga ( Figs 539 , 566–567 ).

Material examined. Holotype: L-S-I ♂ {specimen [XXXVII](14)2009}: INDONESIA, Lombok , Senaru, 23.IX.2009, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 19–26.IX.2009: 11 L-S-I ♂ , 10 L-S-I ♀, 3 L-S ♂, 3 L-S-I ♀, 45 larvae; Java, Bogor, Mount Sulak, Ciapus , altitude 624 m, 24.II.2008, coll. V. Ivanov: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; frons either colorless, or be bordered with brown (i.e. with brown arc connecting paired ocelli) and/or with median brown spot just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 536 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 534 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless; legs with coxae partly colorless, partly brown; trochanteres colorless ( Fig. 537 ). Fore femur with long brown macula on posterior side and brown outer side; other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 551–559 . Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Figs 535, 537–538 ); medio-anterior and medioposterior sigilla lighter than background ( Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than in Figs 535–538 . Caudalii light brownish ( Fig. 541 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and on each abdominal tergum ( Fig. 539 ).

SHAPE: Labrum subrectangular, either equally wide at base and near apex, or slightly wider at base ( Fig. 544 ). Mandibles ( Figs 542–543 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 549 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with extremely small vestiges of hind protoptera (28 exuviae examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 551–553 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 560–563 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–5 denticles in each ( Figs 547–548 ) [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles; tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 564 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–4 moderately long denticles ( Fig. 550 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 565 ), in male without denticles between protogonostyli ( Fig. 571 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 584–590 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 541 ).

SCALES: Scales on pronotum, mesonotum and protoptera short (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 564 ). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 565 ).

STOUT SETAE: Outer side of femur with regular row of 6–11 long, clavate setae and 2 subapical setae of same form ( Figs 551–553 ). Inner-anterior side of each femur with small (10–15 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 560–563 ). Outer-apical seta of tibia small, blunt and elongate ( Figs 560–563 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (j) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows ( Figs 562–563 ) [see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 ( Fig. 571 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 668 ). Thoracic sterna and pleura with brown and colorless areas; legs light brownish, nearly colorless ( Figs 669–670 ). Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago, see below.

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 566 ). Head ochre with reddish- brown. Turbinate eyes relatively high and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Legs either entirely pale ochre ( Fig. 566 ), or tinged with reddish: femora with reddish apically, middle and hind tibiae with longitudinal reddish stripe (as in female— Figs 578–580 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in female— Figs 579–580 ). Abdominal terga I–VI whitish or light ochre, terga VII–VIII reddish, terga IX–X ochre; each tergum I–VIII with narrow, contrasting, reddish-brown stripe on posterior margin; sterna whitish or ochre ( Fig. 567 ). Cerci unicolor pale ochre.

Genitalia ( Fig. 572 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with inner margin of 1st segment forming blunt angle; 2nd segment thickest at middle; 3rd segment elongate, not thickened apically. Penial bridge medially with sclerotized, conic projection ( Fig. 574 ). Gonovectes shallowly bent S-like, i.e. with apices bent cranially; membrane adjacent to their apices forming unpaired sack ( Fig. 572 ).

Imago, female ( Figs 575–580 ). Coloration of head, thorax, legs, wings, cerci and abdominal sterna as in male. Abdominal terga contrastingly darker than sterna: each tergum II–VIII extensively colored with reddish and ocher, with narrow reddish-brown stripe on posterior margin. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 308–309 ).

Egg ( Figs 581–583 ). Oval; chorion with small, shallow cavities forming isoedric net-like relief; minute pores, if present, surround cells of this net. Sperm guide wide, occupying several cell of net-like relief.

Dimension. Fore wing length 4.0– 4.5 mm (male).

Distribution. Sunda Islands: Java and Lombok ( Indonesia).

Comparison. Larva of Cenroptella (Chopralla) rufostriata sp. n. resembles C. (Ch.) ceylonensis and C. (Ch.) pusilla in having a row of denticles on posterior margin of abdominal tergum IX only and in lacking denticles on median part of abdominal tergum X. It can be distinguished from these species by denticles on the trapezoid projection of tergum X ( Fig. 550 ), which are larger than in C. pusilla , but smaller and more numerous (usually 3 on each side) than in C. ceylonensis (which usually has one large denticle on each side).

Male imago of C. rufostriata sp. n. differs from C. ceylonensis by absence of extensive reddish coloration on abdominal terga I–VI and by triangular median projection of penial bridge ( Fig. 574 ). In contrast to other species, male imago of C. rufostriata sp. n. has unusual unpaired membranous sack connected with apices of gonovectes ( Fig. 572 ).

3.4. Centroptella (Chopralla) papuanica sp. n.

( Figs 591–625 )

Material examined. Holotype: L-S-I ♀ {specimen [IV](4)B}: INDONESIA, Papua, Jayapura, Waena , 13.VIII.2012, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 8–13.VIII.2012, coll. N. Kluge & L. Sheyko: 1 L-S-I ♀ , 2 L/S ♂, 1 larva.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; genae brown; dorsal side of head brown. Pronotum and mesonotum with diffusive darker and lighter brown areas ( Fig. 599 ). Thoracic pleura and metanotum brown with lighter areas; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. All femora on outer side mostly colorless, with diffusive longitudinal darkening at midlength nearer outer margin; fore and middle femora with outer side brown and with large brown macula on posterior side; hind femur with outer side colorless, with brown longitudinal macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 611–613 ). Abdominal terga with diffusive darker and lighter brown areas; each tergum II–III and V–VI with pair of transverse blanks located posteriad-laterad of medioposterior sigilla and with smaller, more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light, with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown; medio-anterior and medioposterior sigilla lighter than background ( Fig. 598 ). Caudalii light brownish ( Fig. 617 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and each abdominal tergum (as in winged stages— Fig. 620 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base ( Fig. 604 ). Mandibles: incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially-distally, pressed to kinetodontium, curved and dentate apically, without long branch; median margin of right mandible proximad of prostheca with or without short seta-like processes ( Figs 607–608 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 611–613 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 614–616 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–4 denticles in each [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with long triangular denticles; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 606 ). Posterior projection of tergum X with small, irregular denticles on median part, laterally with paired groups of 2–5 long denticles ( Fig. 610 ). Sterna I–IV without denticles; sterna V–VIII with regular, pointed, triangular denticles ( Fig. 602 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 603 ), in male without denticles between protogonostyli ( Fig. 609 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 610 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 591–597 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 617 ).

SCALES: Scales on occipit, median parts of pronotum and mesonotum very small (with sockets 3–4 mkm and length 4–5 mkm) ( Fig. 600 ). Lateral parts of pronotum, lateral parts of mesonotum and protoptera nearly lack scales. Thoracic pleura and median part of metanotum with scales in opercula-bearing sockets, with longitudinal striation ( Fig. 601 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales in operculae-bearing sockets (sockets 5–6 mkm). Scales on abdominal terga elongate and pointed, with longitudinal striation ( Fig. 606 ). Scales on abdominal sterna either similar ( Fig. 603 ), or partly colorless ( Fig. 602 ).

STOUT SETAE: Outer side of femur with regular row of 5–8 long, clavate setae and 2 subapical setae of same form ( Figs 611–613 ). Inner-anterior side of each femur with small (about 10 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 614–616 ). Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia ( Figs 614–615 ); on hind leg longitudinal ( Fig. 616 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 609 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture lighter ( Fig. 622 ). Thoracic sterna and pleura with brown and colorless areas ( Fig. 621 ). Legs light brownish, nearly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and sterna brown, cerci colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimago developing under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); abdomen light with posterior margins of some terga narrowly bordered with reddish-brown ( Fig. 623 ); median projection of penial bridge shallow and truncate; sterno-styligeral muscle present ( Fig. 609 ) [see Chopralla (7)].

Imago, female ( Figs 618–620 ). Head ochre. Thorax and abdomen contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; pronotum and first 4 intersegmental membranes of abdominal terga with pair of transverse bright red stripes; posterior margins of terga narrowly bordered with reddish-brown. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, femur of fore leg slightly darker apically; tibiae of all legs ochre, slightly darkened in proximal part ( Fig. 618 ). Patella-tibial suture present on middle and hind legs, absent on forelegs ( Fig. 618 ) [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 691–692). Cerci unicolor pale ochre.

Egg ( Figs 624–625 ). Oval; chorion with net-like relief stretched longitudinally. Micropile without sperm guide.

Dimension. Fore wing length of female 4.3 mm.

Distribution. New Guinea.

Comparison. Centroptella (Chopralla) papuanica sp. n. differs from other species of Chopralla in structure of right prostheca, which is pressed to kinetodontium and lacks long branches ( Fig. 608 ). The same structure of right prostheca occurs in all Crassolus , from which Centroptella (Chopralla) papuanica sp. n. differs by the characters listed in Table 2.

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

2.7. Centroptella (Crassolus?) sp. « Nepal »

( Figs 422–428 )

Material examined. Central NEPAL, Helambu Area , 27°55’N 83°30’E, Gohare Khola opposite to village Mahenkai, about 500 m from falling to Melamchi River, 4.III.2007, coll. M. Chertoprud: 1 male larva of penultimate instar (other specimens in Chertoprud’s collection) .

Distribution. Nepal.

Comments. Characters of this larva are reported in the Table 3 and shown on the Figs 422–428 ). Pose of developing subimaginal gonostyli is unknown; judging by the presence of denticles on posterior margins of abdominal terga II–X and the presence of setal rows on abdominal sterna III–VI, this species probably belongs to Crassolus .

3. Subgenus Chopralla Waltz & McCafferty 1987

( Figs 1 , 429–686 )

Genus Chopralla Waltz & McCafferty 1987a: 182 .

Subgenus Chopralla of genus Bungona: Salles, Gattolliat & Sartori 2016: 104 .

Synonym of Centroptella: Kluge, Godunko & Svitok 2020: 88 .

Type species: Centroptella ceylonensis Müller-Liebenau 1983 .

Diagnosis (see Centroptella s. l: Classification and Table 2).

(1) In larva, posterior arm of U-shaped row of long setae on fore- and middle tibiae more transverse than longitudinal and crosses tibia [on hind leg longitudinal—see Centroptella s. l. (2)] ( Figs 560–562 , 662–666 ). The same in Crassolus , in contrast to Centroptella s. str.

(2) Larval claw usually with 2 rows of denticles ( Figs 548–548 ). The same in Crassolus , in contrast to Centroptella s. str.

(3) In larva, at least abdominal terga I–IV without denticles on posterior margins; in various species denticles are present beginning either from tergum V (in C. colorata , C. papuanica sp. n., C. ghatensis sp. n. and C. kangi sp. n.), or from tergum VIII (in C. bintang ), or from tergum IX (in C. ceylonensis , C. pusilla and C. rufostriata sp. n.).

(4) In larva, abdominal sternum VII without regular rows of bifurcate setae (j), so that such rows occur on sterna IV–V only (in contrast to Centroptella s. str. and Crassolus , which have regular rows on sterna IV–VI).

(5) In last larval instar, developing subimaginal gonostyli folded under larval cuticle in the « Nigrobaetis - type » pose, i.e. with 3rd segments directed caudally ( Figs 469 , 529 , 571 , 609 , 627 , 654 ). The same in many other Baetovectata, but in contrast to Centroptella s. str. and Crassolus , whose gonostyli are folded in the peculiar « Crassolus - type » pose.

(6) Gonovectes without halberd-like widenings apically ( Figs 465–468 , 527 , 572 ) (in contrast to Crassolus ).

(7) Sterno-styligeral muscle is retained (in contrast to Crassolus ), being rather weak and often divided into pair of convergent muscles ( Figs 465–466 , 527 , 572 ) (the same in Centroptella s. str.).

Other characters. In larva, anterior margin of mesonotum is sharply curved ( Figs 431 , 499 , 434 , 599 , 633 ), in contrast to straight or shallowly curved in Crassolus and Centroptella s. str. ( Figs 24–25 , 77 , 104 , 128 , 185 , 274 , 338–340 ).

Bifurcate setae on metasternum (i) (see Centroptella s. l.) form a pair of regular rows ( Fig. 517 ) (in contrast to Centroptella s. str. and Crassolus , whose bifurcate setae of metasternum are arranged less regularly).

Both male and female imagines have characteristic coloration: thorax contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; abdomen of female has the same coloration ( Figs 531 , 577 , 519 ); abdomen of male has either the same coloration ( Figs 451 , 520 ), or has no reddish coloration on most of terga ( Fig. 566 ). This coloration differs from Crassolus and Centroptella s. str., whose thorax has brown cuticular coloration both on dorsal and on ventral sides at least in males ( Figs 49 , 93 , 143 , 159–160 , 298 , 379 ).

Distribution. The Oriental Region.

Composition. Centroptella (Chopralla) ceylonensis (= C. similis syn. n.), C. (Chopralla) ghatensis sp. n., C. (Chopralla) rufostriata sp. n., C. (Chopralla) papuanica sp. n., C. (Chopralla) pusilla , C. (Chopralla) kangi sp. n., C. (Chopralla) colorata (= C. fusina syn. n.), C. (Chopralla) bintang .

3.1. Centroptella (Chopralla) ceylonensis Müller-Liebenau 1983

( Figs 429–488 )

Centroptella ceylonensis Müller-Liebenau 1983: 486 (larva); Centroptella similis Müller-Liebenau 1983: 487 (larva), syn. n.

Chopralla ceylonensis Waltz & McCafferty 1987a: 183 ;

Chopralla similis: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) ceylonensis: Salles, Gattolliat & Sartori 2016: 104 , figs 4E, 5G, 6G, 6M;

Bungona (Chopralla) similis: Salles, Gattolliat & Sartori 2016: 104 ;

Bungona (Chopralla) pusilla and Bungona (Centroptella) pusilla: Selvakumar, Kubendran, Chandra Kailash & Sidhu Avtar Kaur 2017: 274 (larva).

Material examined. SRI LANKA: border of Uva and Central Provinces, tributary of river Uma near Randenigala dam, 13–14.I.2011, coll. N. Kluge & L. Sheyko: 5 larvae ; Uva Province, Badulla District, Haputale , 15–23.I.2011, coll. N. Kluge & L. Sheyko: 1 L-S-I ♂ ; Uva Province, Badulla District, Lemastota Oya , 20–21.I.2011, coll. N. Kluge & L. Sheyko: 2 larvae ; Sabaragamuwa Province, river Seetha Sangula in Dalhausie near Sri Pada ( Adam’s Peak ), 24–26.I.2011, coll. N. Kluge & L. Sheyko: 2 S-I ♂, 22 larvae ; the same place and collectors, 1–10.II.2020: 5 L-S-I ♂, 11 L-S-I ♀, 1 S-I ♀, 31 larvae; river Battulu Oya, Fishing Hut (S of Sri Pada = Adam’s Peak ), 8–9.II.2020: L/S ♀, 1 larva ; Central Province, Nuwara Eliya District, river Mahaveli Ganga in Ginigathhena (17 km N Hatton), 31.I–3.II.2011, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 9 S-I ♂, 6 I ♂, 5 S-I ♀, 3 I ♀, 1 S ♀, 3 larvae; Belihuloya , 15– 26.I.2020, coll. N. Kluge & L. Sheylo: 1 L-S-I ♀ , 1 I ♀, 6 larvae; Wellawaya , 22.I.2020, coll. N. Kluge & L. Sheyko: 2 larvae . INDIA: State Karnataka, Udupi District, river Seethanadhi-hole near Someswar , 11.I–1.II.2013, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 3 L-S-I ♀, 1 L-S ♀, 9 larvae. State Kerala, Kottayam District, Erumeli , 22.I.2016, coll. N. Kluge & L. Sheyko: 3 L-S-I ♀ , 3 L; Ernakulam District, Malayatoor , 14–15.II.2016, coll. N. Kluge & L. Sheyko: 10 L-S-I ♂ , 1 L-S ♂, 1 L-S-I ♀, 6 larvae.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 430 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 431 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs. 434–436 ; as in Figs 554–559 ). Abdominal terga with contrasting brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 429 ; Müller-Liebenau 1983: figs 4, 6); medio-anterior and medioposterior sigilla lighter than background (as in Fig 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Figs 429, 437–438 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Figs 456–457, 460 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base (as in Fig. 507 ; Müller-Liebenau 1983: fig.1a). Mandibles ( Figs 439–448 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 432 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them (in some individuals differently on left and right sides, as in Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 434–436 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1983: figs 1i–l). Claw with two rows of 1–6 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1983: figs 1m –n).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles (in single female from Malayatoor terga VII–VIII with irregular small denticles); tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 449 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles (as in Fig. 501 ), with or without 1 pair (rarely 2 pairs) of smaller denticles by sides of them ( Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially ( Fig. 450 ), in male without denticles between protogonostyli ( Fig. 469 ). Paraprocts with thin and colorless antero-median apodeme, with large pointed denticles on posterior margin (as in Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 475–488 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment) located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Figs 437–438 ).

SCALES: Certain areas of pronotum and mesonotum with small, colorless scales. Femora, tibiae and tarsi with colorless, short, oval scales. Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 449 ; Müller-Liebenau 1983: figs 5, 7). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 450 ).

STOUT SETAE: Outer side of femur with regular row of 4–15 long, clavate setae and 2 subapical setae of same form ( Figs 434–436 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Müller-Liebenau 1983: figs 1i–l) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 469 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 453 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 452 ). Legs mostly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and cerci light brownish, sterna nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 451 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 454–455 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in Fig. 392 ). Abdominal terga dark ochre with extensive reddish-brown hypodermal markings ( Figs 456–463 ); abdominal sterna ochre, some sterna with pair of reddish-brown longitudinal stripes sublaterally ( Figs 456–458 ; lateral tracheal trunks and some other abdominal tracheae bordered with blackish. Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Figs 465–468 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment either short, or elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Figs 470–472 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 473–474 ). Oval; chorion with isoedric net-like relief, with very sparsely and irregularly arranged minute pores. Sperm guide small and deep.

Dimension. Fore wing length of male and female 3.2–5 mm.

Synonymy of C. ceylonensis and C. similis . Müller-Liebenau (1983) described Centroptella ceylonensis and C. similis bases on larvae only and stated that «The main difference between both species is the lack of hind wing pads in C. similis sp. n. (in both sexes)». In reality, presence and size of larval vestiges of hind protoptera varies individually in this species (while hind wings are always completely absent in winged stages); among examined larvae of C. ceylonensis from one and the same population (Delhausie near Sri Pada), some individuals have wellexpressed vestiges of hind protoptera, some have very small convexities instead of them, some have not vestiges of hind protoptera at all, and some have vestige of hind protopteron on one side and no such vestige on another side (as in Figs 510–511 ).

Besides this character, Müller-Liebenau (1983) noted: «Also the size ratio of turbinate eyes and the lateral eyes appears somewhat different in the male nymphs of both species». Actually, the drawings of male larval heads ( Müller-Liebenau 1983: figs 1p and 2a) do not allow to distinguish these species.

Müller-Liebenau (1983) also noted that scales on abdominal terga are «in general a little more elongate in C. ceylonensis sp. n. ». Actually, the scales significantly vary in size and proportions on terga of one and the same individual, that is visible from the photos in the original description ( Müller-Liebenau 1983: figs 5, 7).

According to the original description of C. similis , its cuticular coloration of larval abdomen somewhat differs from that of C. ceylonensis . Actually, the coloration of abdominal cuticle varies individually in greater degree than it is shown on the pictures in the original descriptions of C. ceylonensis and C. similis . Thus, the name C. similis should be regarded as a junior synonym of C. ceylonensis .

Distribution. Southern India (Western Ghats) and Sri Lanka.

Variability. Hypodermal coloration of abdomen significantly varies individually within each population ( Figs 456–463 , 470–472 ). The examined specimens from Sri Lanka are larger (fore wing length 4.5–5 mm) then the examined specimens from India, whose fore wing length is 3.5–4 mm. However, examination of many imagines of both sexes individually associated with their larval and subimaginal exuviae allowed to conclude that populations from Sri Lanka and India belong to one and the same species.

Species determination. In Sri Lanka Centroptella (Chopralla) ceylonensis is the single species of Chopralla and cannot be confused with other species. In southern India C. (Ch.) ceylonensis can be confused with the closely related species C. (Ch.) ghatensis sp. n., whose larvae differ only by presence of denticles on abdominal terga V–VIII and winged stages are indistinguishable from C. (Ch.) ceylonensis .

Selvakumar et al. (2017) determined larvae of Chopralla from southern India as C. pusilla . Resolution of the total photo ( Selvakumar et al. 2017: fig. 6) allows to see a pair of large denticles on lateral angles of the posterior projection of tenth abdominal tergum, that differs C. ceylonensis ( Fig. 433 ) from C. pusilla ( Fig. 630 ); other characters of the larvae reported by Selvakumar et al. (2017) (i.e. coloration, mandibular structure, arrangement of denticles on abdominal terga) also agree with the characteristics of C. ceylonensis . The photo indicated as «Foreleg» ( Selvakumar et al. 2017: fig. 10) actually belongs to the hind leg.

3.2. Centroptella (Chopralla) ghatensis sp. n.

( Figs 489–533 )

Etymology. Distributional area of this species is located in the mountains of Western Ghats.

Material examined. Holotype: L-S-I ♂ {number [XXI](1)2013}: INDIA, State Tamilnadu, Tirunelveli District, Courtallam , Chittar river near Peraruvi (= Main Falls ), 5.II.2013, coll. N. Kluge & L. Sheyko. Paratypes: the same locality and collectors, 3–7.II.2013: 6 L-S-I ♂ , 3 L-S-I ♀, 45 larvae; Theni District , Suruli, 24–25.I.2016, coll. N. Kluge & L. Sheyko: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 498 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 499 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless ( Fig. 514 ). Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 502–504 ; as in Figs 554–559 ). Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 500 ); medio-anterior and medioposterior sigilla lighter than background (as in Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than on Fig. 500 Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Fig. 496 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Fig. 497 ).

SHAPE: Labrum either equally wide at base and middle ( Fig. 507 ), or slightly wider at base. Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Figs 505–506 ). Prostheca of left mandible with 3 blunt processes and 2 pointed processes ( Fig. 505 ). Prostheca of right mandible directed medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes ( Fig. 506 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 508 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them; in some individuals differently on left and right sides ( Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 502–504 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [as in Figs 560–563 ; see Centroptella s. l. (2)]. Claw with two rows of 3–6 denticles in each [(as in Figs 547–548 ; see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 518 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles ( Fig. 501 ), with or without 1 or 2 pairs of smaller denticles by sides of them (as in Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 519 ). Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially (as in Fig. 450 ), in male without denticles between protogonostyli ( Fig. 529 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 501 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 489–495 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment), located on every 4th or every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 496 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 513 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga elongate, oval, with longitudinal striation ( Fig. 518 ). Scales on abdominal sterna, metanotum, thoracic pleura and leg coxa shorter and partly colorless ( Fig. 519 ).

STOUT SETAE: Outer side of femur with regular row of 6–17 long, clavate setae and 2 subapical setae of same form ( Figs 502–504 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets ( Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia,; on hind leg longitudinal (as in Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 529 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 524 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 525 ). Legs mostly colorless or light brownish, with brown articulations. Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 520 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely dark orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 521–533 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 522–523 ; as in Fig. 392 ). Abdominal terga mostly reddish- brown with lighter ochre areas; abdominal sterna ochre; lateral tracheal trunks and some other abdominal tracheae bordered with blackish ( Fig. 526 ). Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Fig. 527 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Fig. 530 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 532–533 ). Oval; chorion with isoedric net-like relief, without pores, Sperm guide small, occupying one cell of net-like relief.

Dimension. Fore wing length 4.8 mm (male); 5.4 mm (female).

Distribution. Western Ghats ( India).

Comparison. Larva and imago of the south Indian species Centroptella (Chopralla) ghatensis sp. n. are very similar to C. (Chopralla) ceylonensis , which inhabit both southern India and Sri Lanka. Larva of C. ghatensis sp. n. differs from C. ceylonensis in having well developed denticles on abdominal terga V–VIII. In India, C. ghatensis sp. n. and C. ceylonensis were collected in different localities, never being found together.

3.3. Centroptella (Chopralla) rufostriata sp. n.

( Figs 1 , 534–590 )

Etymology. Allusion to hypodermal coloration of abdmen, which consists of reddish cross tripes stripes on posterior margins of terga ( Figs 539 , 566–567 ).

Material examined. Holotype: L-S-I ♂ {specimen [XXXVII](14)2009}: INDONESIA, Lombok , Senaru, 23.IX.2009, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 19–26.IX.2009: 11 L-S-I ♂ , 10 L-S-I ♀, 3 L-S ♂, 3 L-S-I ♀, 45 larvae; Java, Bogor, Mount Sulak, Ciapus , altitude 624 m, 24.II.2008, coll. V. Ivanov: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; frons either colorless, or be bordered with brown (i.e. with brown arc connecting paired ocelli) and/or with median brown spot just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 536 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 534 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless; legs with coxae partly colorless, partly brown; trochanteres colorless ( Fig. 537 ). Fore femur with long brown macula on posterior side and brown outer side; other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 551–559 . Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Figs 535, 537–538 ); medio-anterior and medioposterior sigilla lighter than background ( Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than in Figs 535–538 . Caudalii light brownish ( Fig. 541 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and on each abdominal tergum ( Fig. 539 ).

SHAPE: Labrum subrectangular, either equally wide at base and near apex, or slightly wider at base ( Fig. 544 ). Mandibles ( Figs 542–543 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 549 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with extremely small vestiges of hind protoptera (28 exuviae examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 551–553 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 560–563 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–5 denticles in each ( Figs 547–548 ) [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles; tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 564 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–4 moderately long denticles ( Fig. 550 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 565 ), in male without denticles between protogonostyli ( Fig. 571 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 584–590 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 541 ).

SCALES: Scales on pronotum, mesonotum and protoptera short (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 564 ). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 565 ).

STOUT SETAE: Outer side of femur with regular row of 6–11 long, clavate setae and 2 subapical setae of same form ( Figs 551–553 ). Inner-anterior side of each femur with small (10–15 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 560–563 ). Outer-apical seta of tibia small, blunt and elongate ( Figs 560–563 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (j) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows ( Figs 562–563 ) [see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 ( Fig. 571 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 668 ). Thoracic sterna and pleura with brown and colorless areas; legs light brownish, nearly colorless ( Figs 669–670 ). Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago, see below.

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 566 ). Head ochre with reddish- brown. Turbinate eyes relatively high and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Legs either entirely pale ochre ( Fig. 566 ), or tinged with reddish: femora with reddish apically, middle and hind tibiae with longitudinal reddish stripe (as in female— Figs 578–580 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in female— Figs 579–580 ). Abdominal terga I–VI whitish or light ochre, terga VII–VIII reddish, terga IX–X ochre; each tergum I–VIII with narrow, contrasting, reddish-brown stripe on posterior margin; sterna whitish or ochre ( Fig. 567 ). Cerci unicolor pale ochre.

Genitalia ( Fig. 572 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with inner margin of 1st segment forming blunt angle; 2nd segment thickest at middle; 3rd segment elongate, not thickened apically. Penial bridge medially with sclerotized, conic projection ( Fig. 574 ). Gonovectes shallowly bent S-like, i.e. with apices bent cranially; membrane adjacent to their apices forming unpaired sack ( Fig. 572 ).

Imago, female ( Figs 575–580 ). Coloration of head, thorax, legs, wings, cerci and abdominal sterna as in male. Abdominal terga contrastingly darker than sterna: each tergum II–VIII extensively colored with reddish and ocher, with narrow reddish-brown stripe on posterior margin. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 308–309 ).

Egg ( Figs 581–583 ). Oval; chorion with small, shallow cavities forming isoedric net-like relief; minute pores, if present, surround cells of this net. Sperm guide wide, occupying several cell of net-like relief.

Dimension. Fore wing length 4.0– 4.5 mm (male).

Distribution. Sunda Islands: Java and Lombok ( Indonesia).

Comparison. Larva of Cenroptella (Chopralla) rufostriata sp. n. resembles C. (Ch.) ceylonensis and C. (Ch.) pusilla in having a row of denticles on posterior margin of abdominal tergum IX only and in lacking denticles on median part of abdominal tergum X. It can be distinguished from these species by denticles on the trapezoid projection of tergum X ( Fig. 550 ), which are larger than in C. pusilla , but smaller and more numerous (usually 3 on each side) than in C. ceylonensis (which usually has one large denticle on each side).

Male imago of C. rufostriata sp. n. differs from C. ceylonensis by absence of extensive reddish coloration on abdominal terga I–VI and by triangular median projection of penial bridge ( Fig. 574 ). In contrast to other species, male imago of C. rufostriata sp. n. has unusual unpaired membranous sack connected with apices of gonovectes ( Fig. 572 ).

3.4. Centroptella (Chopralla) papuanica sp. n.

( Figs 591–625 )

Material examined. Holotype: L-S-I ♀ {specimen [IV](4)B}: INDONESIA, Papua, Jayapura, Waena , 13.VIII.2012, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 8–13.VIII.2012, coll. N. Kluge & L. Sheyko: 1 L-S-I ♀ , 2 L/S ♂, 1 larva.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; genae brown; dorsal side of head brown. Pronotum and mesonotum with diffusive darker and lighter brown areas ( Fig. 599 ). Thoracic pleura and metanotum brown with lighter areas; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. All femora on outer side mostly colorless, with diffusive longitudinal darkening at midlength nearer outer margin; fore and middle femora with outer side brown and with large brown macula on posterior side; hind femur with outer side colorless, with brown longitudinal macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 611–613 ). Abdominal terga with diffusive darker and lighter brown areas; each tergum II–III and V–VI with pair of transverse blanks located posteriad-laterad of medioposterior sigilla and with smaller, more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light, with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown; medio-anterior and medioposterior sigilla lighter than background ( Fig. 598 ). Caudalii light brownish ( Fig. 617 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and each abdominal tergum (as in winged stages— Fig. 620 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base ( Fig. 604 ). Mandibles: incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially-distally, pressed to kinetodontium, curved and dentate apically, without long branch; median margin of right mandible proximad of prostheca with or without short seta-like processes ( Figs 607–608 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 611–613 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 614–616 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–4 denticles in each [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with long triangular denticles; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 606 ). Posterior projection of tergum X with small, irregular denticles on median part, laterally with paired groups of 2–5 long denticles ( Fig. 610 ). Sterna I–IV without denticles; sterna V–VIII with regular, pointed, triangular denticles ( Fig. 602 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 603 ), in male without denticles between protogonostyli ( Fig. 609 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 610 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 591–597 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 617 ).

SCALES: Scales on occipit, median parts of pronotum and mesonotum very small (with sockets 3–4 mkm and length 4–5 mkm) ( Fig. 600 ). Lateral parts of pronotum, lateral parts of mesonotum and protoptera nearly lack scales. Thoracic pleura and median part of metanotum with scales in opercula-bearing sockets, with longitudinal striation ( Fig. 601 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales in operculae-bearing sockets (sockets 5–6 mkm). Scales on abdominal terga elongate and pointed, with longitudinal striation ( Fig. 606 ). Scales on abdominal sterna either similar ( Fig. 603 ), or partly colorless ( Fig. 602 ).

STOUT SETAE: Outer side of femur with regular row of 5–8 long, clavate setae and 2 subapical setae of same form ( Figs 611–613 ). Inner-anterior side of each femur with small (about 10 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 614–616 ). Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia ( Figs 614–615 ); on hind leg longitudinal ( Fig. 616 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 609 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture lighter ( Fig. 622 ). Thoracic sterna and pleura with brown and colorless areas ( Fig. 621 ). Legs light brownish, nearly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and sterna brown, cerci colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimago developing under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); abdomen light with posterior margins of some terga narrowly bordered with reddish-brown ( Fig. 623 ); median projection of penial bridge shallow and truncate; sterno-styligeral muscle present ( Fig. 609 ) [see Chopralla (7)].

Imago, female ( Figs 618–620 ). Head ochre. Thorax and abdomen contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; pronotum and first 4 intersegmental membranes of abdominal terga with pair of transverse bright red stripes; posterior margins of terga narrowly bordered with reddish-brown. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, femur of fore leg slightly darker apically; tibiae of all legs ochre, slightly darkened in proximal part ( Fig. 618 ). Patella-tibial suture present on middle and hind legs, absent on forelegs ( Fig. 618 ) [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 691–692). Cerci unicolor pale ochre.

Egg ( Figs 624–625 ). Oval; chorion with net-like relief stretched longitudinally. Micropile without sperm guide.

Dimension. Fore wing length of female 4.3 mm.

Distribution. New Guinea.

Comparison. Centroptella (Chopralla) papuanica sp. n. differs from other species of Chopralla in structure of right prostheca, which is pressed to kinetodontium and lacks long branches ( Fig. 608 ). The same structure of right prostheca occurs in all Crassolus , from which Centroptella (Chopralla) papuanica sp. n. differs by the characters listed in Table 2.

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3. Subgenus Chopralla Waltz & McCafferty 1987

( Figs 1 , 429–686 )

Genus Chopralla Waltz & McCafferty 1987a: 182 .

Subgenus Chopralla of genus Bungona: Salles, Gattolliat & Sartori 2016: 104 .

Synonym of Centroptella: Kluge, Godunko & Svitok 2020: 88 .

Type species: Centroptella ceylonensis Müller-Liebenau 1983 .

Diagnosis (see Centroptella s. l: Classification and Table 2).

(1) In larva, posterior arm of U-shaped row of long setae on fore- and middle tibiae more transverse than longitudinal and crosses tibia [on hind leg longitudinal—see Centroptella s. l. (2)] ( Figs 560–562 , 662–666 ). The same in Crassolus , in contrast to Centroptella s. str.

(2) Larval claw usually with 2 rows of denticles ( Figs 548–548 ). The same in Crassolus , in contrast to Centroptella s. str.

(3) In larva, at least abdominal terga I–IV without denticles on posterior margins; in various species denticles are present beginning either from tergum V (in C. colorata , C. papuanica sp. n., C. ghatensis sp. n. and C. kangi sp. n.), or from tergum VIII (in C. bintang ), or from tergum IX (in C. ceylonensis , C. pusilla and C. rufostriata sp. n.).

(4) In larva, abdominal sternum VII without regular rows of bifurcate setae (j), so that such rows occur on sterna IV–V only (in contrast to Centroptella s. str. and Crassolus , which have regular rows on sterna IV–VI).

(5) In last larval instar, developing subimaginal gonostyli folded under larval cuticle in the « Nigrobaetis - type » pose, i.e. with 3rd segments directed caudally ( Figs 469 , 529 , 571 , 609 , 627 , 654 ). The same in many other Baetovectata, but in contrast to Centroptella s. str. and Crassolus , whose gonostyli are folded in the peculiar « Crassolus - type » pose.

(6) Gonovectes without halberd-like widenings apically ( Figs 465–468 , 527 , 572 ) (in contrast to Crassolus ).

(7) Sterno-styligeral muscle is retained (in contrast to Crassolus ), being rather weak and often divided into pair of convergent muscles ( Figs 465–466 , 527 , 572 ) (the same in Centroptella s. str.).

Other characters. In larva, anterior margin of mesonotum is sharply curved ( Figs 431 , 499 , 434 , 599 , 633 ), in contrast to straight or shallowly curved in Crassolus and Centroptella s. str. ( Figs 24–25 , 77 , 104 , 128 , 185 , 274 , 338–340 ).

Bifurcate setae on metasternum (i) (see Centroptella s. l.) form a pair of regular rows ( Fig. 517 ) (in contrast to Centroptella s. str. and Crassolus , whose bifurcate setae of metasternum are arranged less regularly).

Both male and female imagines have characteristic coloration: thorax contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; abdomen of female has the same coloration ( Figs 531 , 577 , 519 ); abdomen of male has either the same coloration ( Figs 451 , 520 ), or has no reddish coloration on most of terga ( Fig. 566 ). This coloration differs from Crassolus and Centroptella s. str., whose thorax has brown cuticular coloration both on dorsal and on ventral sides at least in males ( Figs 49 , 93 , 143 , 159–160 , 298 , 379 ).

Distribution. The Oriental Region.

Composition. Centroptella (Chopralla) ceylonensis (= C. similis syn. n.), C. (Chopralla) ghatensis sp. n., C. (Chopralla) rufostriata sp. n., C. (Chopralla) papuanica sp. n., C. (Chopralla) pusilla , C. (Chopralla) kangi sp. n., C. (Chopralla) colorata (= C. fusina syn. n.), C. (Chopralla) bintang .

3.1. Centroptella (Chopralla) ceylonensis Müller-Liebenau 1983

( Figs 429–488 )

Centroptella ceylonensis Müller-Liebenau 1983: 486 (larva); Centroptella similis Müller-Liebenau 1983: 487 (larva), syn. n.

Chopralla ceylonensis Waltz & McCafferty 1987a: 183 ;

Chopralla similis: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) ceylonensis: Salles, Gattolliat & Sartori 2016: 104 , figs 4E, 5G, 6G, 6M;

Bungona (Chopralla) similis: Salles, Gattolliat & Sartori 2016: 104 ;

Bungona (Chopralla) pusilla and Bungona (Centroptella) pusilla: Selvakumar, Kubendran, Chandra Kailash & Sidhu Avtar Kaur 2017: 274 (larva).

Material examined. SRI LANKA: border of Uva and Central Provinces, tributary of river Uma near Randenigala dam, 13–14.I.2011, coll. N. Kluge & L. Sheyko: 5 larvae ; Uva Province, Badulla District, Haputale , 15–23.I.2011, coll. N. Kluge & L. Sheyko: 1 L-S-I ♂ ; Uva Province, Badulla District, Lemastota Oya , 20–21.I.2011, coll. N. Kluge & L. Sheyko: 2 larvae ; Sabaragamuwa Province, river Seetha Sangula in Dalhausie near Sri Pada ( Adam’s Peak ), 24–26.I.2011, coll. N. Kluge & L. Sheyko: 2 S-I ♂, 22 larvae ; the same place and collectors, 1–10.II.2020: 5 L-S-I ♂, 11 L-S-I ♀, 1 S-I ♀, 31 larvae; river Battulu Oya, Fishing Hut (S of Sri Pada = Adam’s Peak ), 8–9.II.2020: L/S ♀, 1 larva ; Central Province, Nuwara Eliya District, river Mahaveli Ganga in Ginigathhena (17 km N Hatton), 31.I–3.II.2011, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 9 S-I ♂, 6 I ♂, 5 S-I ♀, 3 I ♀, 1 S ♀, 3 larvae; Belihuloya , 15– 26.I.2020, coll. N. Kluge & L. Sheylo: 1 L-S-I ♀ , 1 I ♀, 6 larvae; Wellawaya , 22.I.2020, coll. N. Kluge & L. Sheyko: 2 larvae . INDIA: State Karnataka, Udupi District, river Seethanadhi-hole near Someswar , 11.I–1.II.2013, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 3 L-S-I ♀, 1 L-S ♀, 9 larvae. State Kerala, Kottayam District, Erumeli , 22.I.2016, coll. N. Kluge & L. Sheyko: 3 L-S-I ♀ , 3 L; Ernakulam District, Malayatoor , 14–15.II.2016, coll. N. Kluge & L. Sheyko: 10 L-S-I ♂ , 1 L-S ♂, 1 L-S-I ♀, 6 larvae.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 430 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 431 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs. 434–436 ; as in Figs 554–559 ). Abdominal terga with contrasting brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 429 ; Müller-Liebenau 1983: figs 4, 6); medio-anterior and medioposterior sigilla lighter than background (as in Fig 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Figs 429, 437–438 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Figs 456–457, 460 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base (as in Fig. 507 ; Müller-Liebenau 1983: fig.1a). Mandibles ( Figs 439–448 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 432 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them (in some individuals differently on left and right sides, as in Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 434–436 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1983: figs 1i–l). Claw with two rows of 1–6 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1983: figs 1m –n).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles (in single female from Malayatoor terga VII–VIII with irregular small denticles); tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 449 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles (as in Fig. 501 ), with or without 1 pair (rarely 2 pairs) of smaller denticles by sides of them ( Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially ( Fig. 450 ), in male without denticles between protogonostyli ( Fig. 469 ). Paraprocts with thin and colorless antero-median apodeme, with large pointed denticles on posterior margin (as in Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 475–488 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment) located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Figs 437–438 ).

SCALES: Certain areas of pronotum and mesonotum with small, colorless scales. Femora, tibiae and tarsi with colorless, short, oval scales. Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 449 ; Müller-Liebenau 1983: figs 5, 7). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 450 ).

STOUT SETAE: Outer side of femur with regular row of 4–15 long, clavate setae and 2 subapical setae of same form ( Figs 434–436 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Müller-Liebenau 1983: figs 1i–l) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 469 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 453 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 452 ). Legs mostly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and cerci light brownish, sterna nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 451 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 454–455 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in Fig. 392 ). Abdominal terga dark ochre with extensive reddish-brown hypodermal markings ( Figs 456–463 ); abdominal sterna ochre, some sterna with pair of reddish-brown longitudinal stripes sublaterally ( Figs 456–458 ; lateral tracheal trunks and some other abdominal tracheae bordered with blackish. Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Figs 465–468 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment either short, or elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Figs 470–472 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 473–474 ). Oval; chorion with isoedric net-like relief, with very sparsely and irregularly arranged minute pores. Sperm guide small and deep.

Dimension. Fore wing length of male and female 3.2–5 mm.

Synonymy of C. ceylonensis and C. similis . Müller-Liebenau (1983) described Centroptella ceylonensis and C. similis bases on larvae only and stated that «The main difference between both species is the lack of hind wing pads in C. similis sp. n. (in both sexes)». In reality, presence and size of larval vestiges of hind protoptera varies individually in this species (while hind wings are always completely absent in winged stages); among examined larvae of C. ceylonensis from one and the same population (Delhausie near Sri Pada), some individuals have wellexpressed vestiges of hind protoptera, some have very small convexities instead of them, some have not vestiges of hind protoptera at all, and some have vestige of hind protopteron on one side and no such vestige on another side (as in Figs 510–511 ).

Besides this character, Müller-Liebenau (1983) noted: «Also the size ratio of turbinate eyes and the lateral eyes appears somewhat different in the male nymphs of both species». Actually, the drawings of male larval heads ( Müller-Liebenau 1983: figs 1p and 2a) do not allow to distinguish these species.

Müller-Liebenau (1983) also noted that scales on abdominal terga are «in general a little more elongate in C. ceylonensis sp. n. ». Actually, the scales significantly vary in size and proportions on terga of one and the same individual, that is visible from the photos in the original description ( Müller-Liebenau 1983: figs 5, 7).

According to the original description of C. similis , its cuticular coloration of larval abdomen somewhat differs from that of C. ceylonensis . Actually, the coloration of abdominal cuticle varies individually in greater degree than it is shown on the pictures in the original descriptions of C. ceylonensis and C. similis . Thus, the name C. similis should be regarded as a junior synonym of C. ceylonensis .

Distribution. Southern India (Western Ghats) and Sri Lanka.

Variability. Hypodermal coloration of abdomen significantly varies individually within each population ( Figs 456–463 , 470–472 ). The examined specimens from Sri Lanka are larger (fore wing length 4.5–5 mm) then the examined specimens from India, whose fore wing length is 3.5–4 mm. However, examination of many imagines of both sexes individually associated with their larval and subimaginal exuviae allowed to conclude that populations from Sri Lanka and India belong to one and the same species.

Species determination. In Sri Lanka Centroptella (Chopralla) ceylonensis is the single species of Chopralla and cannot be confused with other species. In southern India C. (Ch.) ceylonensis can be confused with the closely related species C. (Ch.) ghatensis sp. n., whose larvae differ only by presence of denticles on abdominal terga V–VIII and winged stages are indistinguishable from C. (Ch.) ceylonensis .

Selvakumar et al. (2017) determined larvae of Chopralla from southern India as C. pusilla . Resolution of the total photo ( Selvakumar et al. 2017: fig. 6) allows to see a pair of large denticles on lateral angles of the posterior projection of tenth abdominal tergum, that differs C. ceylonensis ( Fig. 433 ) from C. pusilla ( Fig. 630 ); other characters of the larvae reported by Selvakumar et al. (2017) (i.e. coloration, mandibular structure, arrangement of denticles on abdominal terga) also agree with the characteristics of C. ceylonensis . The photo indicated as «Foreleg» ( Selvakumar et al. 2017: fig. 10) actually belongs to the hind leg.

3.2. Centroptella (Chopralla) ghatensis sp. n.

( Figs 489–533 )

Etymology. Distributional area of this species is located in the mountains of Western Ghats.

Material examined. Holotype: L-S-I ♂ {number [XXI](1)2013}: INDIA, State Tamilnadu, Tirunelveli District, Courtallam , Chittar river near Peraruvi (= Main Falls ), 5.II.2013, coll. N. Kluge & L. Sheyko. Paratypes: the same locality and collectors, 3–7.II.2013: 6 L-S-I ♂ , 3 L-S-I ♀, 45 larvae; Theni District , Suruli, 24–25.I.2016, coll. N. Kluge & L. Sheyko: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 498 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 499 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless ( Fig. 514 ). Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 502–504 ; as in Figs 554–559 ). Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 500 ); medio-anterior and medioposterior sigilla lighter than background (as in Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than on Fig. 500 Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Fig. 496 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Fig. 497 ).

SHAPE: Labrum either equally wide at base and middle ( Fig. 507 ), or slightly wider at base. Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Figs 505–506 ). Prostheca of left mandible with 3 blunt processes and 2 pointed processes ( Fig. 505 ). Prostheca of right mandible directed medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes ( Fig. 506 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 508 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them; in some individuals differently on left and right sides ( Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 502–504 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [as in Figs 560–563 ; see Centroptella s. l. (2)]. Claw with two rows of 3–6 denticles in each [(as in Figs 547–548 ; see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 518 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles ( Fig. 501 ), with or without 1 or 2 pairs of smaller denticles by sides of them (as in Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 519 ). Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially (as in Fig. 450 ), in male without denticles between protogonostyli ( Fig. 529 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 501 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 489–495 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment), located on every 4th or every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 496 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 513 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga elongate, oval, with longitudinal striation ( Fig. 518 ). Scales on abdominal sterna, metanotum, thoracic pleura and leg coxa shorter and partly colorless ( Fig. 519 ).

STOUT SETAE: Outer side of femur with regular row of 6–17 long, clavate setae and 2 subapical setae of same form ( Figs 502–504 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets ( Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia,; on hind leg longitudinal (as in Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 529 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 524 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 525 ). Legs mostly colorless or light brownish, with brown articulations. Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 520 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely dark orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 521–533 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 522–523 ; as in Fig. 392 ). Abdominal terga mostly reddish- brown with lighter ochre areas; abdominal sterna ochre; lateral tracheal trunks and some other abdominal tracheae bordered with blackish ( Fig. 526 ). Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Fig. 527 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Fig. 530 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 532–533 ). Oval; chorion with isoedric net-like relief, without pores, Sperm guide small, occupying one cell of net-like relief.

Dimension. Fore wing length 4.8 mm (male); 5.4 mm (female).

Distribution. Western Ghats ( India).

Comparison. Larva and imago of the south Indian species Centroptella (Chopralla) ghatensis sp. n. are very similar to C. (Chopralla) ceylonensis , which inhabit both southern India and Sri Lanka. Larva of C. ghatensis sp. n. differs from C. ceylonensis in having well developed denticles on abdominal terga V–VIII. In India, C. ghatensis sp. n. and C. ceylonensis were collected in different localities, never being found together.

3.3. Centroptella (Chopralla) rufostriata sp. n.

( Figs 1 , 534–590 )

Etymology. Allusion to hypodermal coloration of abdmen, which consists of reddish cross tripes stripes on posterior margins of terga ( Figs 539 , 566–567 ).

Material examined. Holotype: L-S-I ♂ {specimen [XXXVII](14)2009}: INDONESIA, Lombok , Senaru, 23.IX.2009, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 19–26.IX.2009: 11 L-S-I ♂ , 10 L-S-I ♀, 3 L-S ♂, 3 L-S-I ♀, 45 larvae; Java, Bogor, Mount Sulak, Ciapus , altitude 624 m, 24.II.2008, coll. V. Ivanov: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; frons either colorless, or be bordered with brown (i.e. with brown arc connecting paired ocelli) and/or with median brown spot just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 536 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 534 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless; legs with coxae partly colorless, partly brown; trochanteres colorless ( Fig. 537 ). Fore femur with long brown macula on posterior side and brown outer side; other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 551–559 . Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Figs 535, 537–538 ); medio-anterior and medioposterior sigilla lighter than background ( Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than in Figs 535–538 . Caudalii light brownish ( Fig. 541 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and on each abdominal tergum ( Fig. 539 ).

SHAPE: Labrum subrectangular, either equally wide at base and near apex, or slightly wider at base ( Fig. 544 ). Mandibles ( Figs 542–543 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 549 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with extremely small vestiges of hind protoptera (28 exuviae examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 551–553 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 560–563 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–5 denticles in each ( Figs 547–548 ) [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles; tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 564 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–4 moderately long denticles ( Fig. 550 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 565 ), in male without denticles between protogonostyli ( Fig. 571 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 584–590 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 541 ).

SCALES: Scales on pronotum, mesonotum and protoptera short (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 564 ). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 565 ).

STOUT SETAE: Outer side of femur with regular row of 6–11 long, clavate setae and 2 subapical setae of same form ( Figs 551–553 ). Inner-anterior side of each femur with small (10–15 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 560–563 ). Outer-apical seta of tibia small, blunt and elongate ( Figs 560–563 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (j) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows ( Figs 562–563 ) [see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 ( Fig. 571 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 668 ). Thoracic sterna and pleura with brown and colorless areas; legs light brownish, nearly colorless ( Figs 669–670 ). Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago, see below.

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 566 ). Head ochre with reddish- brown. Turbinate eyes relatively high and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Legs either entirely pale ochre ( Fig. 566 ), or tinged with reddish: femora with reddish apically, middle and hind tibiae with longitudinal reddish stripe (as in female— Figs 578–580 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in female— Figs 579–580 ). Abdominal terga I–VI whitish or light ochre, terga VII–VIII reddish, terga IX–X ochre; each tergum I–VIII with narrow, contrasting, reddish-brown stripe on posterior margin; sterna whitish or ochre ( Fig. 567 ). Cerci unicolor pale ochre.

Genitalia ( Fig. 572 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with inner margin of 1st segment forming blunt angle; 2nd segment thickest at middle; 3rd segment elongate, not thickened apically. Penial bridge medially with sclerotized, conic projection ( Fig. 574 ). Gonovectes shallowly bent S-like, i.e. with apices bent cranially; membrane adjacent to their apices forming unpaired sack ( Fig. 572 ).

Imago, female ( Figs 575–580 ). Coloration of head, thorax, legs, wings, cerci and abdominal sterna as in male. Abdominal terga contrastingly darker than sterna: each tergum II–VIII extensively colored with reddish and ocher, with narrow reddish-brown stripe on posterior margin. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 308–309 ).

Egg ( Figs 581–583 ). Oval; chorion with small, shallow cavities forming isoedric net-like relief; minute pores, if present, surround cells of this net. Sperm guide wide, occupying several cell of net-like relief.

Dimension. Fore wing length 4.0– 4.5 mm (male).

Distribution. Sunda Islands: Java and Lombok ( Indonesia).

Comparison. Larva of Cenroptella (Chopralla) rufostriata sp. n. resembles C. (Ch.) ceylonensis and C. (Ch.) pusilla in having a row of denticles on posterior margin of abdominal tergum IX only and in lacking denticles on median part of abdominal tergum X. It can be distinguished from these species by denticles on the trapezoid projection of tergum X ( Fig. 550 ), which are larger than in C. pusilla , but smaller and more numerous (usually 3 on each side) than in C. ceylonensis (which usually has one large denticle on each side).

Male imago of C. rufostriata sp. n. differs from C. ceylonensis by absence of extensive reddish coloration on abdominal terga I–VI and by triangular median projection of penial bridge ( Fig. 574 ). In contrast to other species, male imago of C. rufostriata sp. n. has unusual unpaired membranous sack connected with apices of gonovectes ( Fig. 572 ).

3.4. Centroptella (Chopralla) papuanica sp. n.

( Figs 591–625 )

Material examined. Holotype: L-S-I ♀ {specimen [IV](4)B}: INDONESIA, Papua, Jayapura, Waena , 13.VIII.2012, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 8–13.VIII.2012, coll. N. Kluge & L. Sheyko: 1 L-S-I ♀ , 2 L/S ♂, 1 larva.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; genae brown; dorsal side of head brown. Pronotum and mesonotum with diffusive darker and lighter brown areas ( Fig. 599 ). Thoracic pleura and metanotum brown with lighter areas; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. All femora on outer side mostly colorless, with diffusive longitudinal darkening at midlength nearer outer margin; fore and middle femora with outer side brown and with large brown macula on posterior side; hind femur with outer side colorless, with brown longitudinal macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 611–613 ). Abdominal terga with diffusive darker and lighter brown areas; each tergum II–III and V–VI with pair of transverse blanks located posteriad-laterad of medioposterior sigilla and with smaller, more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light, with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown; medio-anterior and medioposterior sigilla lighter than background ( Fig. 598 ). Caudalii light brownish ( Fig. 617 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and each abdominal tergum (as in winged stages— Fig. 620 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base ( Fig. 604 ). Mandibles: incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially-distally, pressed to kinetodontium, curved and dentate apically, without long branch; median margin of right mandible proximad of prostheca with or without short seta-like processes ( Figs 607–608 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 611–613 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 614–616 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–4 denticles in each [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with long triangular denticles; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 606 ). Posterior projection of tergum X with small, irregular denticles on median part, laterally with paired groups of 2–5 long denticles ( Fig. 610 ). Sterna I–IV without denticles; sterna V–VIII with regular, pointed, triangular denticles ( Fig. 602 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 603 ), in male without denticles between protogonostyli ( Fig. 609 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 610 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 591–597 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 617 ).

SCALES: Scales on occipit, median parts of pronotum and mesonotum very small (with sockets 3–4 mkm and length 4–5 mkm) ( Fig. 600 ). Lateral parts of pronotum, lateral parts of mesonotum and protoptera nearly lack scales. Thoracic pleura and median part of metanotum with scales in opercula-bearing sockets, with longitudinal striation ( Fig. 601 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales in operculae-bearing sockets (sockets 5–6 mkm). Scales on abdominal terga elongate and pointed, with longitudinal striation ( Fig. 606 ). Scales on abdominal sterna either similar ( Fig. 603 ), or partly colorless ( Fig. 602 ).

STOUT SETAE: Outer side of femur with regular row of 5–8 long, clavate setae and 2 subapical setae of same form ( Figs 611–613 ). Inner-anterior side of each femur with small (about 10 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 614–616 ). Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia ( Figs 614–615 ); on hind leg longitudinal ( Fig. 616 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 609 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture lighter ( Fig. 622 ). Thoracic sterna and pleura with brown and colorless areas ( Fig. 621 ). Legs light brownish, nearly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and sterna brown, cerci colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimago developing under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); abdomen light with posterior margins of some terga narrowly bordered with reddish-brown ( Fig. 623 ); median projection of penial bridge shallow and truncate; sterno-styligeral muscle present ( Fig. 609 ) [see Chopralla (7)].

Imago, female ( Figs 618–620 ). Head ochre. Thorax and abdomen contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; pronotum and first 4 intersegmental membranes of abdominal terga with pair of transverse bright red stripes; posterior margins of terga narrowly bordered with reddish-brown. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, femur of fore leg slightly darker apically; tibiae of all legs ochre, slightly darkened in proximal part ( Fig. 618 ). Patella-tibial suture present on middle and hind legs, absent on forelegs ( Fig. 618 ) [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 691–692). Cerci unicolor pale ochre.

Egg ( Figs 624–625 ). Oval; chorion with net-like relief stretched longitudinally. Micropile without sperm guide.

Dimension. Fore wing length of female 4.3 mm.

Distribution. New Guinea.

Comparison. Centroptella (Chopralla) papuanica sp. n. differs from other species of Chopralla in structure of right prostheca, which is pressed to kinetodontium and lacks long branches ( Fig. 608 ). The same structure of right prostheca occurs in all Crassolus , from which Centroptella (Chopralla) papuanica sp. n. differs by the characters listed in Table 2.

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.1. Centroptella (Chopralla) ceylonensis Müller-Liebenau 1983

( Figs 429–488 )

Centroptella ceylonensis Müller-Liebenau 1983: 486 (larva); Centroptella similis Müller-Liebenau 1983: 487 (larva), syn. n.

Chopralla ceylonensis Waltz & McCafferty 1987a: 183 ;

Chopralla similis: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) ceylonensis: Salles, Gattolliat & Sartori 2016: 104 , figs 4E, 5G, 6G, 6M;

Bungona (Chopralla) similis: Salles, Gattolliat & Sartori 2016: 104 ;

Bungona (Chopralla) pusilla and Bungona (Centroptella) pusilla: Selvakumar, Kubendran, Chandra Kailash & Sidhu Avtar Kaur 2017: 274 (larva).

Material examined. SRI LANKA: border of Uva and Central Provinces, tributary of river Uma near Randenigala dam, 13–14.I.2011, coll. N. Kluge & L. Sheyko: 5 larvae ; Uva Province, Badulla District, Haputale , 15–23.I.2011, coll. N. Kluge & L. Sheyko: 1 L-S-I ♂ ; Uva Province, Badulla District, Lemastota Oya , 20–21.I.2011, coll. N. Kluge & L. Sheyko: 2 larvae ; Sabaragamuwa Province, river Seetha Sangula in Dalhausie near Sri Pada ( Adam’s Peak ), 24–26.I.2011, coll. N. Kluge & L. Sheyko: 2 S-I ♂, 22 larvae ; the same place and collectors, 1–10.II.2020: 5 L-S-I ♂, 11 L-S-I ♀, 1 S-I ♀, 31 larvae; river Battulu Oya, Fishing Hut (S of Sri Pada = Adam’s Peak ), 8–9.II.2020: L/S ♀, 1 larva ; Central Province, Nuwara Eliya District, river Mahaveli Ganga in Ginigathhena (17 km N Hatton), 31.I–3.II.2011, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 9 S-I ♂, 6 I ♂, 5 S-I ♀, 3 I ♀, 1 S ♀, 3 larvae; Belihuloya , 15– 26.I.2020, coll. N. Kluge & L. Sheylo: 1 L-S-I ♀ , 1 I ♀, 6 larvae; Wellawaya , 22.I.2020, coll. N. Kluge & L. Sheyko: 2 larvae . INDIA: State Karnataka, Udupi District, river Seethanadhi-hole near Someswar , 11.I–1.II.2013, coll. N. Kluge & L. Sheyko: 2 L-S-I ♂ , 3 L-S-I ♀, 1 L-S ♀, 9 larvae. State Kerala, Kottayam District, Erumeli , 22.I.2016, coll. N. Kluge & L. Sheyko: 3 L-S-I ♀ , 3 L; Ernakulam District, Malayatoor , 14–15.II.2016, coll. N. Kluge & L. Sheyko: 10 L-S-I ♂ , 1 L-S ♂, 1 L-S-I ♀, 6 larvae.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 430 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 431 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs. 434–436 ; as in Figs 554–559 ). Abdominal terga with contrasting brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 429 ; Müller-Liebenau 1983: figs 4, 6); medio-anterior and medioposterior sigilla lighter than background (as in Fig 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Figs 429, 437–438 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Figs 456–457, 460 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base (as in Fig. 507 ; Müller-Liebenau 1983: fig.1a). Mandibles ( Figs 439–448 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 432 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them (in some individuals differently on left and right sides, as in Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 434–436 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1983: figs 1i–l). Claw with two rows of 1–6 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1983: figs 1m –n).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles (in single female from Malayatoor terga VII–VIII with irregular small denticles); tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 449 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles (as in Fig. 501 ), with or without 1 pair (rarely 2 pairs) of smaller denticles by sides of them ( Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially ( Fig. 450 ), in male without denticles between protogonostyli ( Fig. 469 ). Paraprocts with thin and colorless antero-median apodeme, with large pointed denticles on posterior margin (as in Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 475–488 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment) located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Figs 437–438 ).

SCALES: Certain areas of pronotum and mesonotum with small, colorless scales. Femora, tibiae and tarsi with colorless, short, oval scales. Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 449 ; Müller-Liebenau 1983: figs 5, 7). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 450 ).

STOUT SETAE: Outer side of femur with regular row of 4–15 long, clavate setae and 2 subapical setae of same form ( Figs 434–436 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Müller-Liebenau 1983: figs 1i–l) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 469 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 453 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 452 ). Legs mostly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and cerci light brownish, sterna nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 451 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 454–455 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in Fig. 392 ). Abdominal terga dark ochre with extensive reddish-brown hypodermal markings ( Figs 456–463 ); abdominal sterna ochre, some sterna with pair of reddish-brown longitudinal stripes sublaterally ( Figs 456–458 ; lateral tracheal trunks and some other abdominal tracheae bordered with blackish. Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Figs 465–468 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment either short, or elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Figs 470–472 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 473–474 ). Oval; chorion with isoedric net-like relief, with very sparsely and irregularly arranged minute pores. Sperm guide small and deep.

Dimension. Fore wing length of male and female 3.2–5 mm.

Synonymy of C. ceylonensis and C. similis . Müller-Liebenau (1983) described Centroptella ceylonensis and C. similis bases on larvae only and stated that «The main difference between both species is the lack of hind wing pads in C. similis sp. n. (in both sexes)». In reality, presence and size of larval vestiges of hind protoptera varies individually in this species (while hind wings are always completely absent in winged stages); among examined larvae of C. ceylonensis from one and the same population (Delhausie near Sri Pada), some individuals have wellexpressed vestiges of hind protoptera, some have very small convexities instead of them, some have not vestiges of hind protoptera at all, and some have vestige of hind protopteron on one side and no such vestige on another side (as in Figs 510–511 ).

Besides this character, Müller-Liebenau (1983) noted: «Also the size ratio of turbinate eyes and the lateral eyes appears somewhat different in the male nymphs of both species». Actually, the drawings of male larval heads ( Müller-Liebenau 1983: figs 1p and 2a) do not allow to distinguish these species.

Müller-Liebenau (1983) also noted that scales on abdominal terga are «in general a little more elongate in C. ceylonensis sp. n. ». Actually, the scales significantly vary in size and proportions on terga of one and the same individual, that is visible from the photos in the original description ( Müller-Liebenau 1983: figs 5, 7).

According to the original description of C. similis , its cuticular coloration of larval abdomen somewhat differs from that of C. ceylonensis . Actually, the coloration of abdominal cuticle varies individually in greater degree than it is shown on the pictures in the original descriptions of C. ceylonensis and C. similis . Thus, the name C. similis should be regarded as a junior synonym of C. ceylonensis .

Distribution. Southern India (Western Ghats) and Sri Lanka.

Variability. Hypodermal coloration of abdomen significantly varies individually within each population ( Figs 456–463 , 470–472 ). The examined specimens from Sri Lanka are larger (fore wing length 4.5–5 mm) then the examined specimens from India, whose fore wing length is 3.5–4 mm. However, examination of many imagines of both sexes individually associated with their larval and subimaginal exuviae allowed to conclude that populations from Sri Lanka and India belong to one and the same species.

Species determination. In Sri Lanka Centroptella (Chopralla) ceylonensis is the single species of Chopralla and cannot be confused with other species. In southern India C. (Ch.) ceylonensis can be confused with the closely related species C. (Ch.) ghatensis sp. n., whose larvae differ only by presence of denticles on abdominal terga V–VIII and winged stages are indistinguishable from C. (Ch.) ceylonensis .

Selvakumar et al. (2017) determined larvae of Chopralla from southern India as C. pusilla . Resolution of the total photo ( Selvakumar et al. 2017: fig. 6) allows to see a pair of large denticles on lateral angles of the posterior projection of tenth abdominal tergum, that differs C. ceylonensis ( Fig. 433 ) from C. pusilla ( Fig. 630 ); other characters of the larvae reported by Selvakumar et al. (2017) (i.e. coloration, mandibular structure, arrangement of denticles on abdominal terga) also agree with the characteristics of C. ceylonensis . The photo indicated as «Foreleg» ( Selvakumar et al. 2017: fig. 10) actually belongs to the hind leg.

3.2. Centroptella (Chopralla) ghatensis sp. n.

( Figs 489–533 )

Etymology. Distributional area of this species is located in the mountains of Western Ghats.

Material examined. Holotype: L-S-I ♂ {number [XXI](1)2013}: INDIA, State Tamilnadu, Tirunelveli District, Courtallam , Chittar river near Peraruvi (= Main Falls ), 5.II.2013, coll. N. Kluge & L. Sheyko. Paratypes: the same locality and collectors, 3–7.II.2013: 6 L-S-I ♂ , 3 L-S-I ♀, 45 larvae; Theni District , Suruli, 24–25.I.2016, coll. N. Kluge & L. Sheyko: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 498 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 499 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless ( Fig. 514 ). Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 502–504 ; as in Figs 554–559 ). Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 500 ); medio-anterior and medioposterior sigilla lighter than background (as in Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than on Fig. 500 Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Fig. 496 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Fig. 497 ).

SHAPE: Labrum either equally wide at base and middle ( Fig. 507 ), or slightly wider at base. Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Figs 505–506 ). Prostheca of left mandible with 3 blunt processes and 2 pointed processes ( Fig. 505 ). Prostheca of right mandible directed medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes ( Fig. 506 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 508 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them; in some individuals differently on left and right sides ( Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 502–504 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [as in Figs 560–563 ; see Centroptella s. l. (2)]. Claw with two rows of 3–6 denticles in each [(as in Figs 547–548 ; see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 518 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles ( Fig. 501 ), with or without 1 or 2 pairs of smaller denticles by sides of them (as in Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 519 ). Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially (as in Fig. 450 ), in male without denticles between protogonostyli ( Fig. 529 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 501 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 489–495 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment), located on every 4th or every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 496 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 513 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga elongate, oval, with longitudinal striation ( Fig. 518 ). Scales on abdominal sterna, metanotum, thoracic pleura and leg coxa shorter and partly colorless ( Fig. 519 ).

STOUT SETAE: Outer side of femur with regular row of 6–17 long, clavate setae and 2 subapical setae of same form ( Figs 502–504 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets ( Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia,; on hind leg longitudinal (as in Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 529 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 524 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 525 ). Legs mostly colorless or light brownish, with brown articulations. Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 520 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely dark orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 521–533 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 522–523 ; as in Fig. 392 ). Abdominal terga mostly reddish- brown with lighter ochre areas; abdominal sterna ochre; lateral tracheal trunks and some other abdominal tracheae bordered with blackish ( Fig. 526 ). Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Fig. 527 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Fig. 530 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 532–533 ). Oval; chorion with isoedric net-like relief, without pores, Sperm guide small, occupying one cell of net-like relief.

Dimension. Fore wing length 4.8 mm (male); 5.4 mm (female).

Distribution. Western Ghats ( India).

Comparison. Larva and imago of the south Indian species Centroptella (Chopralla) ghatensis sp. n. are very similar to C. (Chopralla) ceylonensis , which inhabit both southern India and Sri Lanka. Larva of C. ghatensis sp. n. differs from C. ceylonensis in having well developed denticles on abdominal terga V–VIII. In India, C. ghatensis sp. n. and C. ceylonensis were collected in different localities, never being found together.

3.3. Centroptella (Chopralla) rufostriata sp. n.

( Figs 1 , 534–590 )

Etymology. Allusion to hypodermal coloration of abdmen, which consists of reddish cross tripes stripes on posterior margins of terga ( Figs 539 , 566–567 ).

Material examined. Holotype: L-S-I ♂ {specimen [XXXVII](14)2009}: INDONESIA, Lombok , Senaru, 23.IX.2009, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 19–26.IX.2009: 11 L-S-I ♂ , 10 L-S-I ♀, 3 L-S ♂, 3 L-S-I ♀, 45 larvae; Java, Bogor, Mount Sulak, Ciapus , altitude 624 m, 24.II.2008, coll. V. Ivanov: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; frons either colorless, or be bordered with brown (i.e. with brown arc connecting paired ocelli) and/or with median brown spot just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 536 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 534 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless; legs with coxae partly colorless, partly brown; trochanteres colorless ( Fig. 537 ). Fore femur with long brown macula on posterior side and brown outer side; other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 551–559 . Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Figs 535, 537–538 ); medio-anterior and medioposterior sigilla lighter than background ( Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than in Figs 535–538 . Caudalii light brownish ( Fig. 541 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and on each abdominal tergum ( Fig. 539 ).

SHAPE: Labrum subrectangular, either equally wide at base and near apex, or slightly wider at base ( Fig. 544 ). Mandibles ( Figs 542–543 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 549 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with extremely small vestiges of hind protoptera (28 exuviae examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 551–553 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 560–563 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–5 denticles in each ( Figs 547–548 ) [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles; tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 564 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–4 moderately long denticles ( Fig. 550 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 565 ), in male without denticles between protogonostyli ( Fig. 571 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 584–590 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 541 ).

SCALES: Scales on pronotum, mesonotum and protoptera short (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 564 ). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 565 ).

STOUT SETAE: Outer side of femur with regular row of 6–11 long, clavate setae and 2 subapical setae of same form ( Figs 551–553 ). Inner-anterior side of each femur with small (10–15 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 560–563 ). Outer-apical seta of tibia small, blunt and elongate ( Figs 560–563 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (j) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows ( Figs 562–563 ) [see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 ( Fig. 571 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 668 ). Thoracic sterna and pleura with brown and colorless areas; legs light brownish, nearly colorless ( Figs 669–670 ). Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago, see below.

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 566 ). Head ochre with reddish- brown. Turbinate eyes relatively high and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Legs either entirely pale ochre ( Fig. 566 ), or tinged with reddish: femora with reddish apically, middle and hind tibiae with longitudinal reddish stripe (as in female— Figs 578–580 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in female— Figs 579–580 ). Abdominal terga I–VI whitish or light ochre, terga VII–VIII reddish, terga IX–X ochre; each tergum I–VIII with narrow, contrasting, reddish-brown stripe on posterior margin; sterna whitish or ochre ( Fig. 567 ). Cerci unicolor pale ochre.

Genitalia ( Fig. 572 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with inner margin of 1st segment forming blunt angle; 2nd segment thickest at middle; 3rd segment elongate, not thickened apically. Penial bridge medially with sclerotized, conic projection ( Fig. 574 ). Gonovectes shallowly bent S-like, i.e. with apices bent cranially; membrane adjacent to their apices forming unpaired sack ( Fig. 572 ).

Imago, female ( Figs 575–580 ). Coloration of head, thorax, legs, wings, cerci and abdominal sterna as in male. Abdominal terga contrastingly darker than sterna: each tergum II–VIII extensively colored with reddish and ocher, with narrow reddish-brown stripe on posterior margin. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 308–309 ).

Egg ( Figs 581–583 ). Oval; chorion with small, shallow cavities forming isoedric net-like relief; minute pores, if present, surround cells of this net. Sperm guide wide, occupying several cell of net-like relief.

Dimension. Fore wing length 4.0– 4.5 mm (male).

Distribution. Sunda Islands: Java and Lombok ( Indonesia).

Comparison. Larva of Cenroptella (Chopralla) rufostriata sp. n. resembles C. (Ch.) ceylonensis and C. (Ch.) pusilla in having a row of denticles on posterior margin of abdominal tergum IX only and in lacking denticles on median part of abdominal tergum X. It can be distinguished from these species by denticles on the trapezoid projection of tergum X ( Fig. 550 ), which are larger than in C. pusilla , but smaller and more numerous (usually 3 on each side) than in C. ceylonensis (which usually has one large denticle on each side).

Male imago of C. rufostriata sp. n. differs from C. ceylonensis by absence of extensive reddish coloration on abdominal terga I–VI and by triangular median projection of penial bridge ( Fig. 574 ). In contrast to other species, male imago of C. rufostriata sp. n. has unusual unpaired membranous sack connected with apices of gonovectes ( Fig. 572 ).

3.4. Centroptella (Chopralla) papuanica sp. n.

( Figs 591–625 )

Material examined. Holotype: L-S-I ♀ {specimen [IV](4)B}: INDONESIA, Papua, Jayapura, Waena , 13.VIII.2012, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 8–13.VIII.2012, coll. N. Kluge & L. Sheyko: 1 L-S-I ♀ , 2 L/S ♂, 1 larva.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; genae brown; dorsal side of head brown. Pronotum and mesonotum with diffusive darker and lighter brown areas ( Fig. 599 ). Thoracic pleura and metanotum brown with lighter areas; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. All femora on outer side mostly colorless, with diffusive longitudinal darkening at midlength nearer outer margin; fore and middle femora with outer side brown and with large brown macula on posterior side; hind femur with outer side colorless, with brown longitudinal macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 611–613 ). Abdominal terga with diffusive darker and lighter brown areas; each tergum II–III and V–VI with pair of transverse blanks located posteriad-laterad of medioposterior sigilla and with smaller, more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light, with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown; medio-anterior and medioposterior sigilla lighter than background ( Fig. 598 ). Caudalii light brownish ( Fig. 617 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and each abdominal tergum (as in winged stages— Fig. 620 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base ( Fig. 604 ). Mandibles: incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially-distally, pressed to kinetodontium, curved and dentate apically, without long branch; median margin of right mandible proximad of prostheca with or without short seta-like processes ( Figs 607–608 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 611–613 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 614–616 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–4 denticles in each [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with long triangular denticles; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 606 ). Posterior projection of tergum X with small, irregular denticles on median part, laterally with paired groups of 2–5 long denticles ( Fig. 610 ). Sterna I–IV without denticles; sterna V–VIII with regular, pointed, triangular denticles ( Fig. 602 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 603 ), in male without denticles between protogonostyli ( Fig. 609 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 610 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 591–597 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 617 ).

SCALES: Scales on occipit, median parts of pronotum and mesonotum very small (with sockets 3–4 mkm and length 4–5 mkm) ( Fig. 600 ). Lateral parts of pronotum, lateral parts of mesonotum and protoptera nearly lack scales. Thoracic pleura and median part of metanotum with scales in opercula-bearing sockets, with longitudinal striation ( Fig. 601 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales in operculae-bearing sockets (sockets 5–6 mkm). Scales on abdominal terga elongate and pointed, with longitudinal striation ( Fig. 606 ). Scales on abdominal sterna either similar ( Fig. 603 ), or partly colorless ( Fig. 602 ).

STOUT SETAE: Outer side of femur with regular row of 5–8 long, clavate setae and 2 subapical setae of same form ( Figs 611–613 ). Inner-anterior side of each femur with small (about 10 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 614–616 ). Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia ( Figs 614–615 ); on hind leg longitudinal ( Fig. 616 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 609 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture lighter ( Fig. 622 ). Thoracic sterna and pleura with brown and colorless areas ( Fig. 621 ). Legs light brownish, nearly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and sterna brown, cerci colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimago developing under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); abdomen light with posterior margins of some terga narrowly bordered with reddish-brown ( Fig. 623 ); median projection of penial bridge shallow and truncate; sterno-styligeral muscle present ( Fig. 609 ) [see Chopralla (7)].

Imago, female ( Figs 618–620 ). Head ochre. Thorax and abdomen contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; pronotum and first 4 intersegmental membranes of abdominal terga with pair of transverse bright red stripes; posterior margins of terga narrowly bordered with reddish-brown. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, femur of fore leg slightly darker apically; tibiae of all legs ochre, slightly darkened in proximal part ( Fig. 618 ). Patella-tibial suture present on middle and hind legs, absent on forelegs ( Fig. 618 ) [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 691–692). Cerci unicolor pale ochre.

Egg ( Figs 624–625 ). Oval; chorion with net-like relief stretched longitudinally. Micropile without sperm guide.

Dimension. Fore wing length of female 4.3 mm.

Distribution. New Guinea.

Comparison. Centroptella (Chopralla) papuanica sp. n. differs from other species of Chopralla in structure of right prostheca, which is pressed to kinetodontium and lacks long branches ( Fig. 608 ). The same structure of right prostheca occurs in all Crassolus , from which Centroptella (Chopralla) papuanica sp. n. differs by the characters listed in Table 2.

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.2. Centroptella (Chopralla) ghatensis sp. n.

( Figs 489–533 )

Etymology. Distributional area of this species is located in the mountains of Western Ghats.

Material examined. Holotype: L-S-I ♂ {number [XXI](1)2013}: INDIA, State Tamilnadu, Tirunelveli District, Courtallam , Chittar river near Peraruvi (= Main Falls ), 5.II.2013, coll. N. Kluge & L. Sheyko. Paratypes: the same locality and collectors, 3–7.II.2013: 6 L-S-I ♂ , 3 L-S-I ♀, 45 larvae; Theni District , Suruli, 24–25.I.2016, coll. N. Kluge & L. Sheyko: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless, frons can be bordered with brown (i.e. with brown arc connecting paired ocelli), median brown spot can be located just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 498 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 499 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless ( Fig. 514 ). Legs with coxae partly colorless, partly brown; trochanteres colorless. Fore femur with long brown macula on posterior side and brown outer side, other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 502–504 ; as in Figs 554–559 ). Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Fig. 500 ); medio-anterior and medioposterior sigilla lighter than background (as in Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than on Fig. 500 Caudalii light at base (from colorless to light brown), diffusely darkened distally ( Fig. 496 ).

HYPODERMAL COLORATION: Abdomen of larvae of both sexes with composite and extensive reddishbrown hypodermal color pattern as in winged stages ( Fig. 497 ).

SHAPE: Labrum either equally wide at base and middle ( Fig. 507 ), or slightly wider at base. Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Figs 505–506 ). Prostheca of left mandible with 3 blunt processes and 2 pointed processes ( Fig. 505 ). Prostheca of right mandible directed medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes ( Fig. 506 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 508 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum either with short vestiges of hind protoptera, or without them; in some individuals differently on left and right sides ( Figs 510–511 ). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 502–504 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [as in Figs 560–563 ; see Centroptella s. l. (2)]. Claw with two rows of 3–6 denticles in each [(as in Figs 547–548 ; see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 518 ). Posterior projection of tergum X without denticles on median part, laterally with pair of long denticles ( Fig. 501 ), with or without 1 or 2 pairs of smaller denticles by sides of them (as in Fig. 433 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 519 ). Posterior margin of sternum IX in female convex, with row of triangular denticles interrupted medially (as in Fig. 450 ), in male without denticles between protogonostyli ( Fig. 529 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 501 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 489–495 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment), located on every 4th or every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 496 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 513 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga elongate, oval, with longitudinal striation ( Fig. 518 ). Scales on abdominal sterna, metanotum, thoracic pleura and leg coxa shorter and partly colorless ( Fig. 519 ).

STOUT SETAE: Outer side of femur with regular row of 6–17 long, clavate setae and 2 subapical setae of same form ( Figs 502–504 ). Inner-anterior side of each femur with few very short (5–8 mkm length), stout, blunt setae. On fore leg inner-anterior side of trochanter and/or proximal part of femur with or without one or several longer (15–20 mkm length), stout, pointed, setae with small denticles by sides (as in Figs 41 , 362–363 ). Tibia and tarsus with small stout setae on inner side. Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets ( Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia,; on hind leg longitudinal (as in Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 529 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head nearly colorless, antennae light brown. Pronotum light brown. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 524 ). Thoracic sterna and pleura mostly colorless, with certain sclerites brown ( Fig. 525 ). Legs mostly colorless or light brownish, with brown articulations. Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 520 ). Head ochre with reddish-brown. Turbinate eyes relatively low and wide, entirely dark orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, apically more or less intensively tinged with reddish; fore tibia light ochre, apically tinged with reddish; middle and hind tibiae ochre, with more or less expressed longitudinal reddish stripe; tarsi light ochre ( Figs 521–533 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 522–523 ; as in Fig. 392 ). Abdominal terga mostly reddish- brown with lighter ochre areas; abdominal sterna ochre; lateral tracheal trunks and some other abdominal tracheae bordered with blackish ( Fig. 526 ). Cerci ochre, in proximal part with joinings reddish-brown.

Genitalia ( Fig. 527 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with 1st segment narrowed apically; 2nd segment thickened toward apex; 3rd segment elongate, thickened toward apex. Penial bridge medially with large, sclerotized, hemispheric or conic projection. Gonovectes shallowly bent S-like, i.e. with apices bent cranially.

Imago, female ( Fig. 530 ). Coloration of head, thorax, abdomen, wings and cerci as in male. Fore tibia ochre, tinged with reddish on inner side; other leg coloration as in male. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 391–392 ).

Egg ( Figs 532–533 ). Oval; chorion with isoedric net-like relief, without pores, Sperm guide small, occupying one cell of net-like relief.

Dimension. Fore wing length 4.8 mm (male); 5.4 mm (female).

Distribution. Western Ghats ( India).

Comparison. Larva and imago of the south Indian species Centroptella (Chopralla) ghatensis sp. n. are very similar to C. (Chopralla) ceylonensis , which inhabit both southern India and Sri Lanka. Larva of C. ghatensis sp. n. differs from C. ceylonensis in having well developed denticles on abdominal terga V–VIII. In India, C. ghatensis sp. n. and C. ceylonensis were collected in different localities, never being found together.

3.3. Centroptella (Chopralla) rufostriata sp. n.

( Figs 1 , 534–590 )

Etymology. Allusion to hypodermal coloration of abdmen, which consists of reddish cross tripes stripes on posterior margins of terga ( Figs 539 , 566–567 ).

Material examined. Holotype: L-S-I ♂ {specimen [XXXVII](14)2009}: INDONESIA, Lombok , Senaru, 23.IX.2009, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 19–26.IX.2009: 11 L-S-I ♂ , 10 L-S-I ♀, 3 L-S ♂, 3 L-S-I ♀, 45 larvae; Java, Bogor, Mount Sulak, Ciapus , altitude 624 m, 24.II.2008, coll. V. Ivanov: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; frons either colorless, or be bordered with brown (i.e. with brown arc connecting paired ocelli) and/or with median brown spot just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 536 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 534 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless; legs with coxae partly colorless, partly brown; trochanteres colorless ( Fig. 537 ). Fore femur with long brown macula on posterior side and brown outer side; other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 551–559 . Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Figs 535, 537–538 ); medio-anterior and medioposterior sigilla lighter than background ( Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than in Figs 535–538 . Caudalii light brownish ( Fig. 541 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and on each abdominal tergum ( Fig. 539 ).

SHAPE: Labrum subrectangular, either equally wide at base and near apex, or slightly wider at base ( Fig. 544 ). Mandibles ( Figs 542–543 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 549 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with extremely small vestiges of hind protoptera (28 exuviae examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 551–553 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 560–563 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–5 denticles in each ( Figs 547–548 ) [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles; tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 564 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–4 moderately long denticles ( Fig. 550 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 565 ), in male without denticles between protogonostyli ( Fig. 571 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 584–590 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 541 ).

SCALES: Scales on pronotum, mesonotum and protoptera short (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 564 ). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 565 ).

STOUT SETAE: Outer side of femur with regular row of 6–11 long, clavate setae and 2 subapical setae of same form ( Figs 551–553 ). Inner-anterior side of each femur with small (10–15 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 560–563 ). Outer-apical seta of tibia small, blunt and elongate ( Figs 560–563 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (j) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows ( Figs 562–563 ) [see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 ( Fig. 571 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 668 ). Thoracic sterna and pleura with brown and colorless areas; legs light brownish, nearly colorless ( Figs 669–670 ). Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago, see below.

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 566 ). Head ochre with reddish- brown. Turbinate eyes relatively high and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Legs either entirely pale ochre ( Fig. 566 ), or tinged with reddish: femora with reddish apically, middle and hind tibiae with longitudinal reddish stripe (as in female— Figs 578–580 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in female— Figs 579–580 ). Abdominal terga I–VI whitish or light ochre, terga VII–VIII reddish, terga IX–X ochre; each tergum I–VIII with narrow, contrasting, reddish-brown stripe on posterior margin; sterna whitish or ochre ( Fig. 567 ). Cerci unicolor pale ochre.

Genitalia ( Fig. 572 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with inner margin of 1st segment forming blunt angle; 2nd segment thickest at middle; 3rd segment elongate, not thickened apically. Penial bridge medially with sclerotized, conic projection ( Fig. 574 ). Gonovectes shallowly bent S-like, i.e. with apices bent cranially; membrane adjacent to their apices forming unpaired sack ( Fig. 572 ).

Imago, female ( Figs 575–580 ). Coloration of head, thorax, legs, wings, cerci and abdominal sterna as in male. Abdominal terga contrastingly darker than sterna: each tergum II–VIII extensively colored with reddish and ocher, with narrow reddish-brown stripe on posterior margin. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 308–309 ).

Egg ( Figs 581–583 ). Oval; chorion with small, shallow cavities forming isoedric net-like relief; minute pores, if present, surround cells of this net. Sperm guide wide, occupying several cell of net-like relief.

Dimension. Fore wing length 4.0– 4.5 mm (male).

Distribution. Sunda Islands: Java and Lombok ( Indonesia).

Comparison. Larva of Cenroptella (Chopralla) rufostriata sp. n. resembles C. (Ch.) ceylonensis and C. (Ch.) pusilla in having a row of denticles on posterior margin of abdominal tergum IX only and in lacking denticles on median part of abdominal tergum X. It can be distinguished from these species by denticles on the trapezoid projection of tergum X ( Fig. 550 ), which are larger than in C. pusilla , but smaller and more numerous (usually 3 on each side) than in C. ceylonensis (which usually has one large denticle on each side).

Male imago of C. rufostriata sp. n. differs from C. ceylonensis by absence of extensive reddish coloration on abdominal terga I–VI and by triangular median projection of penial bridge ( Fig. 574 ). In contrast to other species, male imago of C. rufostriata sp. n. has unusual unpaired membranous sack connected with apices of gonovectes ( Fig. 572 ).

3.4. Centroptella (Chopralla) papuanica sp. n.

( Figs 591–625 )

Material examined. Holotype: L-S-I ♀ {specimen [IV](4)B}: INDONESIA, Papua, Jayapura, Waena , 13.VIII.2012, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 8–13.VIII.2012, coll. N. Kluge & L. Sheyko: 1 L-S-I ♀ , 2 L/S ♂, 1 larva.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; genae brown; dorsal side of head brown. Pronotum and mesonotum with diffusive darker and lighter brown areas ( Fig. 599 ). Thoracic pleura and metanotum brown with lighter areas; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. All femora on outer side mostly colorless, with diffusive longitudinal darkening at midlength nearer outer margin; fore and middle femora with outer side brown and with large brown macula on posterior side; hind femur with outer side colorless, with brown longitudinal macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 611–613 ). Abdominal terga with diffusive darker and lighter brown areas; each tergum II–III and V–VI with pair of transverse blanks located posteriad-laterad of medioposterior sigilla and with smaller, more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light, with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown; medio-anterior and medioposterior sigilla lighter than background ( Fig. 598 ). Caudalii light brownish ( Fig. 617 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and each abdominal tergum (as in winged stages— Fig. 620 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base ( Fig. 604 ). Mandibles: incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially-distally, pressed to kinetodontium, curved and dentate apically, without long branch; median margin of right mandible proximad of prostheca with or without short seta-like processes ( Figs 607–608 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 611–613 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 614–616 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–4 denticles in each [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with long triangular denticles; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 606 ). Posterior projection of tergum X with small, irregular denticles on median part, laterally with paired groups of 2–5 long denticles ( Fig. 610 ). Sterna I–IV without denticles; sterna V–VIII with regular, pointed, triangular denticles ( Fig. 602 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 603 ), in male without denticles between protogonostyli ( Fig. 609 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 610 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 591–597 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 617 ).

SCALES: Scales on occipit, median parts of pronotum and mesonotum very small (with sockets 3–4 mkm and length 4–5 mkm) ( Fig. 600 ). Lateral parts of pronotum, lateral parts of mesonotum and protoptera nearly lack scales. Thoracic pleura and median part of metanotum with scales in opercula-bearing sockets, with longitudinal striation ( Fig. 601 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales in operculae-bearing sockets (sockets 5–6 mkm). Scales on abdominal terga elongate and pointed, with longitudinal striation ( Fig. 606 ). Scales on abdominal sterna either similar ( Fig. 603 ), or partly colorless ( Fig. 602 ).

STOUT SETAE: Outer side of femur with regular row of 5–8 long, clavate setae and 2 subapical setae of same form ( Figs 611–613 ). Inner-anterior side of each femur with small (about 10 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 614–616 ). Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia ( Figs 614–615 ); on hind leg longitudinal ( Fig. 616 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 609 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture lighter ( Fig. 622 ). Thoracic sterna and pleura with brown and colorless areas ( Fig. 621 ). Legs light brownish, nearly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and sterna brown, cerci colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimago developing under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); abdomen light with posterior margins of some terga narrowly bordered with reddish-brown ( Fig. 623 ); median projection of penial bridge shallow and truncate; sterno-styligeral muscle present ( Fig. 609 ) [see Chopralla (7)].

Imago, female ( Figs 618–620 ). Head ochre. Thorax and abdomen contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; pronotum and first 4 intersegmental membranes of abdominal terga with pair of transverse bright red stripes; posterior margins of terga narrowly bordered with reddish-brown. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, femur of fore leg slightly darker apically; tibiae of all legs ochre, slightly darkened in proximal part ( Fig. 618 ). Patella-tibial suture present on middle and hind legs, absent on forelegs ( Fig. 618 ) [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 691–692). Cerci unicolor pale ochre.

Egg ( Figs 624–625 ). Oval; chorion with net-like relief stretched longitudinally. Micropile without sperm guide.

Dimension. Fore wing length of female 4.3 mm.

Distribution. New Guinea.

Comparison. Centroptella (Chopralla) papuanica sp. n. differs from other species of Chopralla in structure of right prostheca, which is pressed to kinetodontium and lacks long branches ( Fig. 608 ). The same structure of right prostheca occurs in all Crassolus , from which Centroptella (Chopralla) papuanica sp. n. differs by the characters listed in Table 2.

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.3. Centroptella (Chopralla) rufostriata sp. n.

( Figs 1 , 534–590 )

Etymology. Allusion to hypodermal coloration of abdmen, which consists of reddish cross tripes stripes on posterior margins of terga ( Figs 539 , 566–567 ).

Material examined. Holotype: L-S-I ♂ {specimen [XXXVII](14)2009}: INDONESIA, Lombok , Senaru, 23.IX.2009, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 19–26.IX.2009: 11 L-S-I ♂ , 10 L-S-I ♀, 3 L-S ♂, 3 L-S-I ♀, 45 larvae; Java, Bogor, Mount Sulak, Ciapus , altitude 624 m, 24.II.2008, coll. V. Ivanov: 2 larvae .

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; frons either colorless, or be bordered with brown (i.e. with brown arc connecting paired ocelli) and/or with median brown spot just posteriad median ocellus; genae brown or colorless; dorsal side of head brown ( Fig. 536 ). Pronotum and mesonotum with contrasting brown, ochre and/or colorless areas, forming characteristic pattern ( Fig. 534 ). Thoracic pleura and metanotum partly brown, partly colorless; sterna colorless; legs with coxae partly colorless, partly brown; trochanteres colorless ( Fig. 537 ). Fore femur with long brown macula on posterior side and brown outer side; other sides mostly colorless; middle femur with smaller brown macula on posterior side and brown outer side; hind femur mostly colorless, with brown macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 551–559 . Abdominal terga with brown, ochre and/or colorless areas forming characteristic pattern; each tergum II–VI with unpaired, median blank; among them, each tergum II–III and V–VI with pair of large, transverse blanks located posteriad-laterad of medioposterior sigilla; among them, terga III and V with more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light (from colorless to light brown), with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown ( Figs 535, 537–538 ); medio-anterior and medioposterior sigilla lighter than background ( Fig. 540 ). In some individuals coloration of abdominal terga mostly dark, and blanks not expressed. In some individuals brown areas either more extensive, or less contrasting than in Figs 535–538 . Caudalii light brownish ( Fig. 541 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and on each abdominal tergum ( Fig. 539 ).

SHAPE: Labrum subrectangular, either equally wide at base and near apex, or slightly wider at base ( Fig. 544 ). Mandibles ( Figs 542–543 ): incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment ( Fig. 549 ).

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with extremely small vestiges of hind protoptera (28 exuviae examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 551–553 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 560–563 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–5 denticles in each ( Figs 547–548 ) [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII without denticles; tergum IX with long triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 564 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–4 moderately long denticles ( Fig. 550 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 565 ), in male without denticles between protogonostyli ( Fig. 571 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 550 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 584–590 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 541 ).

SCALES: Scales on pronotum, mesonotum and protoptera short (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, with longitudinal striation ( Fig. 564 ). Scales on abdominal sterna, thoracic pleura and metanotum shorter and partly colorless ( Fig. 565 ).

STOUT SETAE: Outer side of femur with regular row of 6–11 long, clavate setae and 2 subapical setae of same form ( Figs 551–553 ). Inner-anterior side of each femur with small (10–15 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 560–563 ). Outer-apical seta of tibia small, blunt and elongate ( Figs 560–563 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (j) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal ( Figs 560–563 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows ( Figs 562–563 ) [see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 ( Fig. 571 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture colorless ( Fig. 668 ). Thoracic sterna and pleura with brown and colorless areas; legs light brownish, nearly colorless ( Figs 669–670 ). Wings light, without brown circles at bases of microtrichia. Abdominal terga light brownish, sterna and cerci nearly colorless.

HYPODERMAL COLORATION: As in imago, see below.

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male ( Fig. 566 ). Head ochre with reddish- brown. Turbinate eyes relatively high and wide, entirely orange. Thorax contrastingly darker (ochre with reddish-brown) in dorsal half and lighter (pale ochre) in ventral half. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal and subcostal veins proximad of costal brace brownish. Legs either entirely pale ochre ( Fig. 566 ), or tinged with reddish: femora with reddish apically, middle and hind tibiae with longitudinal reddish stripe (as in female— Figs 578–580 ). Tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) (as in female— Figs 579–580 ). Abdominal terga I–VI whitish or light ochre, terga VII–VIII reddish, terga IX–X ochre; each tergum I–VIII with narrow, contrasting, reddish-brown stripe on posterior margin; sterna whitish or ochre ( Fig. 567 ). Cerci unicolor pale ochre.

Genitalia ( Fig. 572 ). Sterno-styligeral muscle slender and paired [see Chopralla (7)]. Gonostylus with inner margin of 1st segment forming blunt angle; 2nd segment thickest at middle; 3rd segment elongate, not thickened apically. Penial bridge medially with sclerotized, conic projection ( Fig. 574 ). Gonovectes shallowly bent S-like, i.e. with apices bent cranially; membrane adjacent to their apices forming unpaired sack ( Fig. 572 ).

Imago, female ( Figs 575–580 ). Coloration of head, thorax, legs, wings, cerci and abdominal sterna as in male. Abdominal terga contrastingly darker than sterna: each tergum II–VIII extensively colored with reddish and ocher, with narrow reddish-brown stripe on posterior margin. Patella-tibial suture present on middle and hind legs, absent on forelegs [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 308–309 ).

Egg ( Figs 581–583 ). Oval; chorion with small, shallow cavities forming isoedric net-like relief; minute pores, if present, surround cells of this net. Sperm guide wide, occupying several cell of net-like relief.

Dimension. Fore wing length 4.0– 4.5 mm (male).

Distribution. Sunda Islands: Java and Lombok ( Indonesia).

Comparison. Larva of Cenroptella (Chopralla) rufostriata sp. n. resembles C. (Ch.) ceylonensis and C. (Ch.) pusilla in having a row of denticles on posterior margin of abdominal tergum IX only and in lacking denticles on median part of abdominal tergum X. It can be distinguished from these species by denticles on the trapezoid projection of tergum X ( Fig. 550 ), which are larger than in C. pusilla , but smaller and more numerous (usually 3 on each side) than in C. ceylonensis (which usually has one large denticle on each side).

Male imago of C. rufostriata sp. n. differs from C. ceylonensis by absence of extensive reddish coloration on abdominal terga I–VI and by triangular median projection of penial bridge ( Fig. 574 ). In contrast to other species, male imago of C. rufostriata sp. n. has unusual unpaired membranous sack connected with apices of gonovectes ( Fig. 572 ).

3.4. Centroptella (Chopralla) papuanica sp. n.

( Figs 591–625 )

Material examined. Holotype: L-S-I ♀ {specimen [IV](4)B}: INDONESIA, Papua, Jayapura, Waena , 13.VIII.2012, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 8–13.VIII.2012, coll. N. Kluge & L. Sheyko: 1 L-S-I ♀ , 2 L/S ♂, 1 larva.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; genae brown; dorsal side of head brown. Pronotum and mesonotum with diffusive darker and lighter brown areas ( Fig. 599 ). Thoracic pleura and metanotum brown with lighter areas; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. All femora on outer side mostly colorless, with diffusive longitudinal darkening at midlength nearer outer margin; fore and middle femora with outer side brown and with large brown macula on posterior side; hind femur with outer side colorless, with brown longitudinal macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 611–613 ). Abdominal terga with diffusive darker and lighter brown areas; each tergum II–III and V–VI with pair of transverse blanks located posteriad-laterad of medioposterior sigilla and with smaller, more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light, with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown; medio-anterior and medioposterior sigilla lighter than background ( Fig. 598 ). Caudalii light brownish ( Fig. 617 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and each abdominal tergum (as in winged stages— Fig. 620 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base ( Fig. 604 ). Mandibles: incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially-distally, pressed to kinetodontium, curved and dentate apically, without long branch; median margin of right mandible proximad of prostheca with or without short seta-like processes ( Figs 607–608 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 611–613 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 614–616 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–4 denticles in each [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with long triangular denticles; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 606 ). Posterior projection of tergum X with small, irregular denticles on median part, laterally with paired groups of 2–5 long denticles ( Fig. 610 ). Sterna I–IV without denticles; sterna V–VIII with regular, pointed, triangular denticles ( Fig. 602 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 603 ), in male without denticles between protogonostyli ( Fig. 609 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 610 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 591–597 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 617 ).

SCALES: Scales on occipit, median parts of pronotum and mesonotum very small (with sockets 3–4 mkm and length 4–5 mkm) ( Fig. 600 ). Lateral parts of pronotum, lateral parts of mesonotum and protoptera nearly lack scales. Thoracic pleura and median part of metanotum with scales in opercula-bearing sockets, with longitudinal striation ( Fig. 601 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales in operculae-bearing sockets (sockets 5–6 mkm). Scales on abdominal terga elongate and pointed, with longitudinal striation ( Fig. 606 ). Scales on abdominal sterna either similar ( Fig. 603 ), or partly colorless ( Fig. 602 ).

STOUT SETAE: Outer side of femur with regular row of 5–8 long, clavate setae and 2 subapical setae of same form ( Figs 611–613 ). Inner-anterior side of each femur with small (about 10 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 614–616 ). Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia ( Figs 614–615 ); on hind leg longitudinal ( Fig. 616 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 609 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture lighter ( Fig. 622 ). Thoracic sterna and pleura with brown and colorless areas ( Fig. 621 ). Legs light brownish, nearly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and sterna brown, cerci colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimago developing under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); abdomen light with posterior margins of some terga narrowly bordered with reddish-brown ( Fig. 623 ); median projection of penial bridge shallow and truncate; sterno-styligeral muscle present ( Fig. 609 ) [see Chopralla (7)].

Imago, female ( Figs 618–620 ). Head ochre. Thorax and abdomen contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; pronotum and first 4 intersegmental membranes of abdominal terga with pair of transverse bright red stripes; posterior margins of terga narrowly bordered with reddish-brown. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, femur of fore leg slightly darker apically; tibiae of all legs ochre, slightly darkened in proximal part ( Fig. 618 ). Patella-tibial suture present on middle and hind legs, absent on forelegs ( Fig. 618 ) [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 691–692). Cerci unicolor pale ochre.

Egg ( Figs 624–625 ). Oval; chorion with net-like relief stretched longitudinally. Micropile without sperm guide.

Dimension. Fore wing length of female 4.3 mm.

Distribution. New Guinea.

Comparison. Centroptella (Chopralla) papuanica sp. n. differs from other species of Chopralla in structure of right prostheca, which is pressed to kinetodontium and lacks long branches ( Fig. 608 ). The same structure of right prostheca occurs in all Crassolus , from which Centroptella (Chopralla) papuanica sp. n. differs by the characters listed in Table 2.

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.4. Centroptella (Chopralla) papuanica sp. n.

( Figs 591–625 )

Material examined. Holotype: L-S-I ♀ {specimen [IV](4)B}: INDONESIA, Papua, Jayapura, Waena , 13.VIII.2012, coll. N. Kluge & L. Sheyko. Paratypes: the same locality, 8–13.VIII.2012, coll. N. Kluge & L. Sheyko: 1 L-S-I ♀ , 2 L/S ♂, 1 larva.

Descriptions.

Larva. CUTICULAR COLORATION: Head with anterior side mostly colorless; genae brown; dorsal side of head brown. Pronotum and mesonotum with diffusive darker and lighter brown areas ( Fig. 599 ). Thoracic pleura and metanotum brown with lighter areas; sterna colorless. Legs with coxae partly colorless, partly brown; trochanteres colorless. All femora on outer side mostly colorless, with diffusive longitudinal darkening at midlength nearer outer margin; fore and middle femora with outer side brown and with large brown macula on posterior side; hind femur with outer side colorless, with brown longitudinal macula on posterior side near inner margin; tibia of each leg diffusively darkened with brown along rows of long, fine setae; tarsus of each leg brown proximally and along rows of long, fine setae; claws brown ( Figs 611–613 ). Abdominal terga with diffusive darker and lighter brown areas; each tergum II–III and V–VI with pair of transverse blanks located posteriad-laterad of medioposterior sigilla and with smaller, more anterior pair of transverse blanks located laterad of medioanterior sigilla; on terga II and IV blanks occupy most part of tergum; terga VII–VIII mostly light, with posterior and/or anterior margins bordered by darker brown; terga IX–X mostly brown; medio-anterior and medioposterior sigilla lighter than background ( Fig. 598 ). Caudalii light brownish ( Fig. 617 ).

HYPODERMAL COLORATION: Reddish-brown stripe on posterior margin of metanotum and each abdominal tergum (as in winged stages— Fig. 620 ).

SHAPE: Labrum either equally wide at base and middle, or slightly wider at base ( Fig. 604 ). Mandibles: incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle; prostheca of left mandible with 3 blunt processes and 3 pointed processes; prostheca of right mandible directed medially-distally, pressed to kinetodontium, curved and dentate apically, without long branch; median margin of right mandible proximad of prostheca with or without short seta-like processes ( Figs 607–608 ). Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore leg tarsus (measured on outer side) as long as tibia; on middle and hind legs tarsus slightly shorter than tibia ( Figs 611–613 ). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 614–616 ) [see Centroptella s. l. (2)]. Claw with two rows of 1–4 denticles in each [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with long triangular denticles; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 606 ). Posterior projection of tergum X with small, irregular denticles on median part, laterally with paired groups of 2–5 long denticles ( Fig. 610 ). Sterna I–IV without denticles; sterna V–VIII with regular, pointed, triangular denticles ( Fig. 602 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially ( Fig. 603 ), in male without denticles between protogonostyli ( Fig. 609 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 610 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII ( Figs 591–597 ). Each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching or not reaching midlength of next segment), located on every 2nd segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 617 ).

SCALES: Scales on occipit, median parts of pronotum and mesonotum very small (with sockets 3–4 mkm and length 4–5 mkm) ( Fig. 600 ). Lateral parts of pronotum, lateral parts of mesonotum and protoptera nearly lack scales. Thoracic pleura and median part of metanotum with scales in opercula-bearing sockets, with longitudinal striation ( Fig. 601 ). Femora, tibiae and tarsi densely covered with colorless, short, oval scales in operculae-bearing sockets (sockets 5–6 mkm). Scales on abdominal terga elongate and pointed, with longitudinal striation ( Fig. 606 ). Scales on abdominal sterna either similar ( Fig. 603 ), or partly colorless ( Fig. 602 ).

STOUT SETAE: Outer side of femur with regular row of 5–8 long, clavate setae and 2 subapical setae of same form ( Figs 611–613 ). Inner-anterior side of each femur with small (about 10 mkm length), stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side ( Figs 614–616 ). Outer-apical seta of tibia small, blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia ( Figs 614–615 ); on hind leg longitudinal ( Fig. 616 ) [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: 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 and inserted into 2nd segment ( Fig. 609 ) [see Chopralla (5)].

RESPIRATORY MOVEMENTS: Tergalii able to make rhythmic respiratory movements.

Subimago. CUTICULAR COLORATION: Head and antennae colorless, with microtrichia dark brown. Pronotum light brownish. Mesonotum light brown with certain sutures darker and medioparapsidal suture lighter ( Fig. 622 ). Thoracic sterna and pleura with brown and colorless areas ( Fig. 621 ). Legs light brownish, nearly colorless. Wings light, without brown circles at bases of microtrichia. Abdominal terga and sterna brown, cerci colorless.

HYPODERMAL COLORATION: As in imago (see below).

TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimago developing under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); abdomen light with posterior margins of some terga narrowly bordered with reddish-brown ( Fig. 623 ); median projection of penial bridge shallow and truncate; sterno-styligeral muscle present ( Fig. 609 ) [see Chopralla (7)].

Imago, female ( Figs 618–620 ). Head ochre. Thorax and abdomen contrastingly darker (ochre with reddishbrown) in dorsal half and lighter (pale ochre) in ventral half; pronotum and first 4 intersegmental membranes of abdominal terga with pair of transverse bright red stripes; posterior margins of terga narrowly bordered with reddish-brown. Wing with membrane colorless, veins pale ochre or colorless, extreme base of costal vein proximad of costal brace brownish. Femora of all legs ochre, femur of fore leg slightly darker apically; tibiae of all legs ochre, slightly darkened in proximal part ( Fig. 618 ). Patella-tibial suture present on middle and hind legs, absent on forelegs ( Fig. 618 ) [as in male; see Centroptella s. l. (2)]. Tarsus of each leg with 1 apical spine on initial 3rd tarsomere (on foreleg—on tarsomere next after 2nd tarsomere, on middle and hind leg—on tarsomere next after 1st+2nd tarsomere) (as in Figs 691–692). Cerci unicolor pale ochre.

Egg ( Figs 624–625 ). Oval; chorion with net-like relief stretched longitudinally. Micropile without sperm guide.

Dimension. Fore wing length of female 4.3 mm.

Distribution. New Guinea.

Comparison. Centroptella (Chopralla) papuanica sp. n. differs from other species of Chopralla in structure of right prostheca, which is pressed to kinetodontium and lacks long branches ( Fig. 608 ). The same structure of right prostheca occurs in all Crassolus , from which Centroptella (Chopralla) papuanica sp. n. differs by the characters listed in Table 2.

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.5. Centroptella (Chopralla) pusilla Müller-Liebenau 1984

( Figs 626–631 )

Centroptella pusilla Müller-Liebenau 1984a: 94 (larva);

Chopralla pusilla: Waltz & McCafferty 1987a: 183 ;

Bungona (Chopralla) pusilla Salles, Gattolliat & Sartori 2016: 104 ;

« Cloeodes Sabah-1» (unpublished label).

Material examined. MALAYSIA, Negeri Sabah, Karagasan, Ranau , 6.IX.1994, coll. S.C. Kang: 3 L/S ♂ (other specimens in Entomological Institute in Česke Budejovice ) .

Descriptions.

Larva. CUTICULAR COLORATION: Pronotum and abdominal terga I–VI as figured by Müller-Liebenau (1984a: fig. 11).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum short, widest at base and narrowed distally ( Fig. 625 ). Incisor of left mandible terminated with 4 denticles decreasing from most distal to most proximal; incisor of right mandible terminated with 3 denticles of subequal width, without 4th denticle; left and right incisors with outer-ventral denticle ( Müller-Liebenau 1984a: fig. 5e). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes. Prostheca of right mandible directed medially or medially-proximally, bifurcate, with longest branch directed medially-proximally or proximally; median margin of right mandible proximad of prostheca with short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented, with 2nd much longer than 1st segment.

Pronotum without protuberances (in contrast to C. ingridae ). Metanotum with vestiges of hind protoptera (in all 6 individuals examined). Forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia ( Müller-Liebenau 1984a: fig. 5h). All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia [see Centroptella s. l. (2)] ( Müller-Liebenau 1984a: fig. 5h). Claw with two rows of 2–5 denticles in each [see Chopralla (2)] ( Müller-Liebenau 1984a: fig. 5i).

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–VIII, without denticles; tergum IX with triangular denticles interrupted medially, behind pair of submedian setae ( Fig. 626 ). Posterior projection of abdominal tergum X trapezoid, without denticles or with small denticles on angles only ( Fig. 631 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles. Posterior margin of sternum IX in male without denticles between protogonostyli ( Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 630 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea (see Centroptella s. l.).

Enlarged denticles on outer side of cercus not large (reaching midlength of next segment), located on every 4th segment and arranged by 1–2 on each side of secondary swimming setal row [see Centroptella s. l. (3)].

SCALES: Scales on pronotum, mesonotum and protoptera very small (with sockets 3–4 mkm and length 4–5 mkm). Femora, tibiae and tarsi densely covered with colorless, short, oval scales (sockets 5–6 mkm). Scales on abdominal terga numerous, elongate, oval, wider than their sockets ( Fig. 626 ). Scales on abdominal sterna, thoracic pleura and metanotum smaller.

STOUT SETAE: Outer side of femur with regular row of 7–13 long, clavate setae and 2 subapical setae of same form ( Müller-Liebenau 1984a: fig. 5h). Inner-anterior side of each femur with small, stout, blunt setae; no longer setae on inner-anterior side of fore femur. Tibia and tarsus with stout setae on inner side. Outer-apical seta of tibia blunt and elongate (as in Fig. 364 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). Posterior arm of U-shaped row of long setae on fore- and middle leg situated across tibia, on hind leg longitudinal [see Chopralla (1)] ( Müller-Liebenau 1984a: fig. 5h). Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, V-shaped setae sublaterally (g) (as in Fig. 604 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated [see Chopralla (4)].

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 627 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere); sterno-styligeral muscle present [see Chopralla (7)].

Imago, female. Unknown.

Egg. Unknown.

Dimension. Body length about 3.5 mm.

Comments. In the original description, labrum is figured as relatively long, with unusual shape of lateral sides ( Müller-Liebenau 1984a: fig. 5a). Type specimens were kept in Purdue University ( USA), but now are probably missing (L. Jacobus, personal communication). My determination of the specimens examined as C. pusilla is based on the assumption that Müller-Liebenau took the internal arms of labrum for lateral-proximal margins of the labrum ( Figs 628–629 ).

Distribution. Borneo. Selvakumar et al. (2017) erroneously reported C. pusilla from southern India; actually the larvae from India reported by them belong to C. ceylonensis (see above).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.6. Centroptella (Chopralla) kangi sp. n.

( Figs 632–672 )

« Cloeodes Sabah-4» (unpublished label).

Etymology. The species in named in honor to S.C. Kang, whose material is used for the description.

Material examined. Holotype: L/S ♂: MALAYSIA, Negeri Sabah, Karagasan, Ranau , 4.IX.1994, coll. S.C. Kang. Paratypes: the same locality, date and collector: 1 L/S ♂, 1 L/S ♀, 1 L ♀. Other material (not examined): the same locality, date and collector: larvae in Entomological Institute, Česke Budejovice, Czech Republic).

Descriptions.

Larva. CUTICULAR CLORATION: Pronotum and mesonotum with brown, ochre and/or colorless areas ( Fig. 633 ). Abdominal terga II–VI with medioanterior and medioposterior sigilla lighter than background, without contrasting transverse blanks; terga VII–VIII lighter than others ( Fig. 632 ).

HYPODERMAL COLORATION: Unknown.

SHAPE: Labrum widest at base and narrowed distally ( Fig. 659 ). Mandibles ( Figs 655–658 ): incisor of left mandible terminated with 3 denticles, among which distal denticle much wider than two proximal ones; incisor of right mandible terminated with 2 denticles, among which distal denticle much wider than proximal one; left and right incisors with outer-ventral denticle ( Fig. 656 ). Prostheca of left mandible with 3 blunt processes and 2–3 pointed processes; prostheca of right mandible directed medially or medially-proximally, arched proximally, without bifurcation ( Fig. 657 ); median margin of right mandible proximad of prostheca with or without short seta-like processes. Maxillary palp nearly as long as lacinia, 2-segmented ( Fig. 661 ).

Pronotum without protuberances (in contrast to C. ingridae ). Vestiges of hind protoptera very small or absent. Legs longer than in other species ( Figs 651–653 ), length of outer side of patella equal to tibia width ( Figs 662–666 ); forelegs longest, hind legs shortest; on fore and middle legs tarsus (measured on outer side) as long as tibia; on hind legs tarsus slightly shorter than tibia. All femora parallel-sided, outer margin straight or slightly concave, inner margin slightly convex. Foreleg without patella-tibial suture, middle and hind legs with patella-tibial suture greatly shifted to apex of tibia ( Figs 662–666 ) [see Centroptella s. l. (2)]. Claw with two rows of 0–3 denticles in each; selected claws with denticle(s) on one side or without denticles [see Chopralla (2)].

Denticles on posterior margins of abdominal terga and sterna flat (not conic). Terga I–IV without denticles; terga V–IX with triangular denticles, small and irregular on tergum V and long on posterior terga; on tergum IX row of denticles interrupted medially, behind pair of submedian setae ( Fig. 667 ). Posterior projection of tergum X without denticles on median part, laterally with paired groups of 3–6 long or short denticles ( Fig. 650 ). Sterna I–IV without denticles; sterna V–VIII with regular, small, pointed, triangular denticles ( Fig. 668 ). Posterior margin of sternum IX in female convex, with row of triangular denticles diminished medially, in male without denticles between protogonostyli (as in Fig. 627 ). Paraprocts with thin and colorless antero- median apodeme, with large pointed denticles on posterior margin ( Fig. 649 ).

Tergalius I narrow, elongate-ellipsoid; other tergalii wider, gradually changing in shape from tergalius II to tergalius VII; each tergalius II–VII, besides costal and anal ribs, with straight and narrow middle rib, located on dorsal surface on background of main trachea ( Figs 635–648 ) (see Centroptella s. l.).

Enlarged denticles on outer side of cercus relatively large (reaching or exceeding midlength of next segment arranged), located on every 2nd or every 4th segment and arranged by 1–3 on each side of secondary swimming setal row [see Centroptella s. l. (3)] ( Fig. 634 ).

SCALES: Scales on thoracic terga varying from short to band-like: certain areas of pronotum and mesonotum with colorless scales 2 mkm width and 5–15 mkm length ( Fig. 669 ). Scales on abdominal terga numerous, elongateoval, with longitudinal striation, significantly wider than their sockets, widely overlapping one another ( Fig. 667 ). Scales on most anterior abdominal terga, on abdominal sterna, metanotum and thoracic pleura smaller ( Fig. 668 ).

STOUT SETAE: Row of long, clavate, setae on outer side of femur dense and irregular at proximal part, sparse and regular at distal part, consists of 9–13 setae and 2 subapical setae of same form ( Figs 651–653 ). Tibia and tarsus with few very small stout setae on inner side ( Fig. 662, 665 ). Outer-apical seta of tibia blunt and elongate ( Figs 662–666 ).

LONG, FINE SETAE: Bifurcate setae of metasternum (i) (see Centroptella s. l.) form pair of dense, regular rows partly with contiguous sockets (as in Fig. 517 ). U-shaped row of long setae distanced from tibia-femur articulation; its posterior arm on fore- and middle legs situated across tibia ( Figs 662–666 ); on hind leg longitudinal [see Chopralla (1)]. Dorsal side of each tarsus with long, fine setae situated irregularly and partly forming two longitudinal rows (as in Figs 562–563 ; see Centroptella s. l.). Tergum VII with pair of fine, bifurcate (Y-shaped) setae sublaterally (g) (as in Fig. 406 ). Each sternum IV–V with pair of regular, transverse rows of long, fine, bifurcate setae with spaced sockets (j); other sterna either without such setae, or with few setae irregularly situated.

MALE GENITALIA IN LARVA: In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis - type » pose, with 2nd and 3rd segments directed caudally and inserted into previous ones ( Fig. 654 ) [see Chopralla (5)].

Subimago. TEXTURE: On all legs of both sexes all tarsal segments entirely covered with pointed microlepides (as in Fig. 391 ).

Imago, male. Unknown. Judging by subimaginal leg developed under larval cuticle, tarsus of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Figs 652–653 ); sterno-styligeral muscle present ( Fig. 654 ) [see Chopralla (7)].

Imago, female. Unknown. Judging by subimaginal leg developing under larval cuticle, tarsus of fore leg without apical spines (as in males of this and other species); tarsi of middle and hind legs with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere).

Egg ( Figs 670–672 ). Oval; chorion with sharply projected net-like relief. Micropile not found, probably located under ridge of net-like relief.

Dimension. Body length about 5 mm.

Distribution. Borneo.

Comparison. Larva of Centroptella (Chopralla) kangi sp. n. differs from other known species of Centroptella s. l. by longer tibia with the U-shaped row of long setae located fare from the femur-tibia articulation ( Figs 561–563 , 662–666 ). Winged females of C. kangi sp. n. differ from other examined species of Centroptella s. l. by absence of apical spines on fore tarsi.

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.7. Centroptella (Chopralla) colorata Soldán, Braasch & Luu-Tham-Muu 1987

( Figs 673–686 )

Centroptella colorata Soldán, Braasch & Luu-Tham-Muu 1987: 346 (larva, ♀ subimago); Kluge, Godunko & Svitok 2020: 121 (larva).

Chopralla colorata: Tong & Dudgeon 2003: 17 ;

Bungona (Chopralla) colorata: Salles, Gattolliat & Sartori 2015: 104 .

Centroptella (Chopralla) fusina (Tong & Dudgeon 2003) syn. n.:

Chopralla fusina Tong & Dudgeon 2003: 17 (larva, ♂ & ♀ imagines);

Bungona (Chopralla) fusina: Salles, Gattolliat & Sartori 2016: 104 ; Shi & Tong 2019: 581 (larva);

Centroptella fusina: Kluge, Godunko & Svitok 2020: 92 .

Material examined. Holotype of C. colorata : immature male larva of last instar (see Kluge et al. 2020). Holotype and paratypes of C. fusina : photos provided by Xiaoli Tong.

Distribution. Vietnam and Hong Kong.

Comments: type specimens of C. colorata . Originally, C. colorata was described as larvae and female subimago extracted from mature larva ( Soldán et al. 1987). At present, only the holotype, male larva of last instar, is retained, wile the paratypes are lost.

According to the original description, «posterior margin of terga ... IV–X with stout pointed triangular teeth arranged into a simple row» ( Soldán et al. 1987: 346); however, in the holotype tergum IV has no denticles (teeth), and the denticles are present beginning from the tergum V ( Fig. 682 ). Structure of tergum X was not reported in the original description; the holotype has one pair of large denticles on angles of its posterior projection ( Fig. 683 ).

The combination of larval characters of C. colorata (right prostheca V-shaped, denticles on posterior margins of abdominal terga V–IX, tenth abdominal tergum with one pair of large denticles only) agrees with that of the new Indian species C. ghatensis sp. n. Both larvae and winged stages of C. colorata well differ from C. ghatensis sp. n. by absence of hypodermal coloration on abdominal terga. The holotype of C. colorata examined by me has very contrasting brown markings on cuticle of thoracic terga and pleura and on abdominal terga ( Fig. 681 ), but its hypoderm under cuticle is entirely colorless. According to the original description, the female subimago extracted from larva had «Body pale whitish .... Abdominal terga without markings» ( Soldán et al. 1987: 346). This means that the absence of hypodermal coloration observed in the larva kept in alcohol during 36 years was not caused by some selective depigmentation of the hypoderm (with cuticular pigmentation well preserved), but was initially inherent to these specimens. In contrast to C. colorata , male and female larvae and winged stages of C. ghatensis sp. n. have very intensive and contrasting hypodermal coloration of abdominal terga, which have not become paler after seven years preservation in alcohol.

Comments: type specimens of C. fusina . C. fusina was described from Hong Kong based on 4 larvae, one male and one female imagines and one male subimago, with female larva designated as the holotype. Both imagines and the subimago reported in the original description, were individually reared from larvae, and their subimaginal exuviae were preserved and examined, but this fact was not reported in the original description (personal communication by Xiaoli Tong). On the drawing of male imaginal genitalia given in the original description (Tong & Dudgeon 2003: fig. 16), terminal segments of gonostyli are somewhat longer than on the photo of the slide from which this drawing was done ( Fig. 684 ).

Larva of C. fusina was said to differ from C. colorata in the color pattern of pronotum and mesonotum and in three morphological characters as follows ( Shi & Tong 2019):

(1) «The prostheca of right mandible bifurcated and plumose ... (vs. the right prostheca plumose but not bifurcated in B. colorata )». In the original description of C. colorata the prostheca is really characterized as «plumose but not branched», but figured as bifurcate with branches brought together ( Soldán et al. 1987: fig. 17).

(2) «Tergites V–X with acute triangular spines on posterior margins ... (vs. tergites IV–X with such spines in B. colorata )». This statement was based on the original description of C. colorata , which disagrees with the holotype structure (see above). Actually both the holotype of C. colorata and the type specimens of C. fusina have rows of denticles (i.e. spines, or teeth) on posterior margins of abdominal terga V–IX and one pair of large denticles on posterior projection of tergum X ( Figs 682–683 , 685–686 ).

(3) «Gill I less than 4 times as long as wide, and other gills relatively broad, much less than 2.5 times as long as wide) ... (vs. in B. colorata , gill I more than 5 times as long as wide and other gills narrow-elongate and at least 2.5 times as long as wide)». Actually, tergalii (so called «gills») of C. colorata never had been figured, and this comparison was based on the wordily description only, which stated that «gill 1 ... more than 5 times longer than broad; gills 2–5 apically rounded or bluntly pointed ... about 2.5–3.5 times as long as broad» ( Soldán et al. 1987: 346). Actually, tergalii of the holotype of C. colorata are wider than reported in the original description ( Figs 673–679 ).

Color pattern of pronotum and mesonotum somewhat differs in the original description of C. colorata ( Soldán et al. 1987: fig. 26), on the holotype ( Fig. 681 ) and in the redescription of C. fusina ( Shi & Tong 2019: figs 52, 59), that can be explained as individual variability.

Based on this, I propose the following subjective synonymy: C. colorata = C. fusina syn. n.

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

3.8. Centroptella (Chopralla) bintang ( Marle et al. 2016)

Bungona (Chopralla) sp. 1: Salles, Gattolliat & Sartori 2016: 104 (Salles, personal communication);

Bungona (Chopralla) bintang Marle, Salles & Gattolliat 2016: 227 (larva);

Centroptella bintang: Kluge, Godunko & Svitok 2020: 92 .

Distribution. Borneo (known from Indonesian part).

Comments. Centroptella bintang is known as larvae only. Its larva well differs from other Centroptella s. l. in having denticles on posterior margins of abdominal terga VIII–IX (rather than on tergum IX, terga V–IX or II–IX in other species of Centroptella s. l.).

Figures

FIGURE 421. Centroptella (Crassolus?) bifida, mature male larva (photo by Xiaoli Tong).

FIGURES 395–398. Centroptella (Crassolus) ingridae, larva. 395, labrum (SEM picture by R. Godunko); 396–397, left and right mandibles; 398, male abdominal terga with reddish and orange hypodermal coloration visible through cuticle.

FIGURES 422–428. Centroptella (Crassolus?) sp. «Nepal», larva. 422–423, left and right mandibles with developing mandibles of next instar inside (arrow shows tuft of setiform processes on proximal angle of mola); 424–425, maxilla; 426, abdominal tergum X; 427, abdominal tergum IX; 428, labrum.

FIGURES 1–3. Heads of female larvae (frontal view). 1, Centroptella rufostriata sp. n.; 2, Potamocloeon edentatum; 3, Cloeodes vibratorius.

FIGURES 429–438. Centroptella (Chopralla) ceylonensis, larva. 429, abdomen; 430, head; 431, pronotum and mesonotum; 432, maxilla; 433, abdominal tergum X; 434–436, fore, middle and hind legs; 437–438, cerci of different specimens.

FIGURES 439–448. Centroptella (Chopralla) ceylonensis, mandibles. 439–440, left mandible, dorsal and median view; 441– 442, left and right mandibles of penultimolarva before molt to ultimolarva (with mandibles of next instar inside); 443–444, the same, another specimen; 445–448, right prostheca of different species. Abbreviations: in1–in4, denticles of incisor; kd1–kd3, denticles of kinetodontium; o-v.d, outer-ventral denticle of incisor; prs, prostheca.

FIGURES 449–450. Centroptella (Chopralla) ceylonensis, exuviae of larval abdomen. 449, tergum IX; 450, sternum IX of female.

FIGURES 451–463. Centroptella (Chopralla) ceylonensis. 451, male imago; 452, subimaginal exuviae of mesopleuron; 453, subimaginal exuviae of mesonotum; 454–455, fore and middle legs of male imago; 456–457, abdomina of male imagines extracted from larvae; 458–463, abdomina of male imagines (451–459, specimens from Sri Lanka; 460–463, specimens from India).

FIGURES 464–469. Centroptella (Chopralla) ceylonensis. 464, subimaginal exuviae of gonostylus; 465–468, genitalia of male imagines; 469, subimaginal gonostyli crumpled under larval cuticle. Abbreviations as in Figs 303–307.

FIGURES 470–474. Centroptella (Chopralla) ceylonensis. 470–472, female imagines; 473–474, egg.

FIGURES 475–495. Tergalii I–VII. 475–481 and 482–488, Centroptella (Chopralla) ceylonensis; 489–495, Centroptella (Chopralla) ghatensis sp. n.

FIGURES 496–504. Centroptella (Chopralla) ghatensis sp. n., larva. 496, cercus (holotype); 497, subimaginal abdomen extracted from larva, to show hypodermal coloration; 498–504, larval exuviae: 498, head; 499, pronotum and mesonotum; 500, abdomen; 501, abdominal segment X; 502–504, fore, middle and hind legs.

FIGURES 505–511. Centroptella (Chopralla) ghatensis sp. n., larva. 505–506, left and right mandibles of penultimolarva before molt to ultimolarva (with mandibles of next instar inside); 507, labrum; 508, maxilla; 509, labium; 510–511, left and right halves of exuviae of larval metanotum (holotype). Abbreviation: ppt, vestige of hind protopteron; letter g corresponds to bases of bifurcate setae in Fig. 14.

FIGURES 512–517. Centroptella (Chopralla) ghatensis sp. n., exuviae of larval thorax. 512–513, scales and bifurcate setae on anterior part of mesonotum (corresponding to e on Fig. 14); 514–517, bifurcate setae on thoracic sternum (corresponding to h and i on Fig. 15).

FIGURES 518–519. Centroptella (Chopralla) ghatensis sp. n., exuviae of larval abdomen. 518, tergum IX; 519, sternum VII.

FIGURES 520–526. Centroptella (Chopralla) ghatensis sp. n. 520, male imago; 521–523, fore, middle and hind legs of male imago; 524, exuviae of subimaginal mesonotum; 525, exuviae of subimaginal mesopleuron; 526, abdomen of male imago (521–524, 526, holotype).

FIGURES 527–529. Centroptella (Chopralla) ghatensis sp. n. 527, genitalia of male imago (left gonostylar and gonovectal muscles not shown); 528, their subimaginal exuviae; 529, (left gonostylar and gonovectal muscles not shown). Abbreviations: gd, gonoduct; gs1–gs3, segments of gonostyli; gv, gonovectis; m.IX-X, base of intersegmental muscle; m.gs, gonostylar muscle; m.gv.d, dorsal muscle of gonovectis; m.gv.v, ventral muscle of gonovectis; m.sg, styligeral muscle; pb, median projection of penial bridge; pgs, larval protogonostylus; usg, unistyliger.

FIGURES 530–533. Centroptella (Chopralla) ghatensis sp. n. 530, female imago; 531, abdomen of female imago; 532–533, egg.

FIGURES 534–540. Centroptella (Chopralla) rufostriata sp. n. 534–536, larval exuviae (534, pronotum and mesonotum; 535, abdomen; 536, head); 537–538, female larva of last instar; 539, younger larva with lighter cuticular coloration (grayish) and better visible hypodermal coloration (reddish brown); 540, exuviae of abdominal tergum III (MA, medioanterior sigillum; MP, medioposterior sigillum).

FIGURES 541–550. Centroptella (Chopralla) rufostriata sp. n., larva. 541, cercus; 542–543, left and right mandibles; 544, labrum; 545–546, labium; 547–548, claw; 549, maxilla; 550, abdominal segment X without caudalii.

FIGURES 551–559. Centroptella (Chopralla) rufostriata sp. n., larval legs. 551–553, translucent exuviae of fore, middle and hind legs with cuticular coloration visible on both sides; 554–556, outer sides of fore, middle and hind legs; 557–559, their inner sides.

FIGURES 560–565. Centroptella (Chopralla) rufostriata sp. n., larva. 560–561, fore and middle tibiae, anterior view (setae and sutures on opposite side shown by interrupted lines); 562–563, hind tibia and tarsus, anterior and outer view (bases of long, fine setae shown as rings). 564, abdominal tergum IX; 565, abdominal sternum VII.

FIGURES 566–570. Centroptella (Chopralla) rufostriata sp. n. 566, male imago; 567, abdomen of male imago; 568–570, subimaginal exuviae: 568, mesonotum, 569, other parts of thorax and head; 570, mesopleuron.

FIGURES 571–574. Centroptella (Chopralla) rufostriata sp. n. 571, subimaginal gonostyli crumpled under larval cuticle; 572, genitalia (holotype; left gonostylar and gonovectal muscles not shown); 573, their subimaginal exuviae; 574, penial bridge. Abbreviations: gd, gonoduct; gs1–gs3, segments of gonostyli; gv, gonovectis; m.IX-X, base of intersegmental muscle; m.gs, gonostylar muscle; m.gv.d, dorsal muscle of gonovectis; m.gv.v, ventral muscle of gonovectis; m.sg, styligeral muscle; pb, median projection of penial bridge; pgs, larval protogonostylus; usg, unistyliger.

FIGURES 575–580. Centroptella (Chopralla) rufostriata sp. n., female imago. 575, dorsal view; 576, head; 577, abdomen; 578–580, fore, middle and hind legs.

FIGURES 581–583. Centroptella (Chopralla) rufostriata sp. n., eggs.

FIGURES 584–597. Tergalii I–VII. 584–590, Centroptella (Chopralla) rufostriata sp. n.; 591–597, Centroptella (Chopralla) papuanica sp. n.

FIGURES 598–606. Centroptella (Chopralla) papuanica sp. n., larval exuviae (holotype). 598, abdomen; 599, pronotum and mesonotum; 600, area of pronotum close to median suture with bifurcate seta corresponding to a on Fig. 14; 601, scale on metanotum with same magnification; 602, abdominal sternum VI and enlarged scales; 603, abdominal sternum IX and enlarged scales; 604, labrum; 605, labium; 606, abdominal terga VIII–IX and enlarged scales.

FIGURES 607–610. Centroptella (Chopralla) papuanica sp. n. 607–608, left and right mandibles; 609, subimaginal gonostyli crumpled under larval cuticle; 610, larval abdominal segment X without caudalii (holotype). Abbreviations: gs1–gs3, segments of gonostyli; m.gs, gonostylar muscle; m.sg, styligeral muscle; pgs, larval protogonostylus; usg, unistyliger.

FIGURES 611–617. Centroptella (Chopralla) papuanica sp. n., larval exuviae. 611–613, fore, middle and hind legs, with focus on anterior side; 614–616, their tibiae, with focus on posterior side (arrows show patella-tibial suture); 617, cercus.

FIGURES 618–623. Centroptella (Chopralla) papuanica sp. n. 618, fore leg of female imago; 619, abdomen of female imago; 620, female imago; 621, subimaginal exuviae of mesopleuron; 622, subimaginal exuviae of mesonotum; 623, abdomen of male subimago extracted from larva (618, 619, 621, holotype).

FIGURES 624–625. Centroptella (Chopralla) papuanica sp. n., egg (extracted from holotype).

FIGURES 626–631. Centroptella (Chopralla) pusilla, larva. 626, abdominal tergum IX; 627, subimaginal gonostyli crumpled under larval cuticle; 628, labrum; 629, labrum according to original description (Müller-Liebenau 1984a: fig. 5a) with presumable basal margin added with red line; 630, paraprocts; 631, posterior margin of abdominal tergum X. Abbreviations: gs1–gs3, segments of gonostyli; pgs, larval protogonostylus; usg, unistyliger.

FIGURES 632–650. Centroptella (Chopralla) kangi sp. n., larvae. 632, cuticle of abdomen (holotype); 633, cuticle of pronotum and mesonotum; 634, cercus; 635–641 and 642–648, tergalii I–VII of two specimens; 649, paraprocts; 650, abdominal tergum X.

FIGURES 651–654. Centroptella (Chopralla) kangi sp. n., larva ready to molt to subimago (holotype). 651–653, fore, middle and hind legs; 654, subimaginal gonostyli crumpled under larval cuticle. Abbreviations as in Figs 571–572, 609).

FIGURES 655–661. Centroptella (Chopralla) kangi sp. n. 655–657, right mandible; 658, left mandible; 659, labrum (holotype); 660, labium; 661, maxilla.

FIGURES 662–666. Centroptella (Chopralla) kangi sp. n. 662–664, fore tibia from anterior, outer and posterior views; 665– 666, middle tibia from anterior and outer views. Abbreviation: pt-ext, outer side of patella.

FIGURES 667–669. Centroptella (Chopralla) kangi sp. n. 667, abdominal tergum IX and its enlarged fragment; 668, abdominal sternum VIII and its enlarged fragment; 669, latero-posterior area of pronotum with row of bifurcate setae corresponding to d on Fig. 14.

FIGURES 670–672. Centroptella (Chopralla) kangi sp. n., eggs.

FIGURES 673–683. Centroptella (Chopralla) colorata, larva (holotype). 573–679, tergalii I–VII; 680, claw; 681, male larva of last instar; 682, abdominal terga IV–V; 683, abdominal tergum X. Abbreviation: MP, medioposterior sigilla.

FIGURES 684–686. Centroptella (Chopralla) colorata: paratypes of C. fusina (photos by Xiaoli Tong). 684, genitalia; 685– 686, exuviae of larval abdomen.

FIGURES 23–32. Centroptella (s. str.) longisetosa cinerea subsp. n., larval exuviae. 23, frons and labrum; 24–25, pronotum and mesonotum; 26–28, abdomen; 29–31, fore, middle and hind legs; 32, labial palp (23, 24, 27, 32, holotype).

FIGURES 75–86. Centroptella (s. str.) femorata sp. n. (holotype), larval exuviae. 75, abdomen; 76, head; 77, pronotum and mesonotum; 78, remainder thorax and legs; 79, glossae and paraglossae, focus on ventral side; 80, the same, focus on dorsal side; 81, labial palp; 82–83, caudalii; 84, maxilla; 85, abdominal tergum X; 86, paraprocts.

FIGURES 100–107. Centroptella (s. str.) soldani. 100–106, larval exuviae: 100, caudalii, 101–102, abdominal tergum X; 103, head; 104, pronotum and mesonotum; 105, remainder thorax and legs; 106, abdomen. 107, larval middle leg with reddish hypodermal coloration visible through cuticle.

FIGURES 125–131. Centroptella (s. str.) ornatipes sp. n. larval exuviae. 125, cercus (dorsal view); 126, cercus (lateral view); 127, head; 128, pronotum and mesonotum; 129, abdomen; 130, maxilla and labium; 131, labrum.

FIGURES 185–191. Centroptella (s. str.) breviseta sp. n. 185–189, larval exuviae (holotype): 185, head; 186, pronotum and mesonotum; 187, part of thorax with fore, middle and hind legs; 188, abdomen; 189, fragment of cercus. 190, young larva. 191, male larva ready to molt to subimago (photo by J.-L. Gattolliat).

FIGURES 273–280. Centroptella (Crassolus) saxophila, larval exuviae. 273, abdomen; 274, pronotum and mesonotum; 275, head; 276, abdominal tergum V; 277, abdominal tergum V of another specimen; 278, abdominal tergum X; 279, paraprocts; 280, apex of abdomen of female.

FIGURES 337–344. Centroptella (Crassolus) ludmilae sp. n. 337–342, larval exuviae: 337, head; 338–340, pronotum and mesonotum; 341–342, abdomen; 343, immature larva with cuticular and hypodermal coloration; 334, abdomen of larva shortly after molt, to show reddish-brown hypodermal coloration (338, holotype).

FIGURES 49–52. Centroptella (s. str.) longisetosa cinerea subsp. n. 49, male imago; 50, subimaginal abdomen extracted from mature larva; 51, subimaginal mesonotum; 52, subimaginal pleuron (holotype).

FIGURES 90–97. Centroptella (s. str.) femorata sp. n. (holotype). 90, larval abdominal sternum IV (letters j and k correspond to Fig. 14); 91, tergum II; 92, tergum VIII; 93, male imago; 94, its abdomen; 95, genitalia; 96, subimaginal pleuron; 97, subimaginal mesonotum.

FIGURES 143–148. Centroptella (s. str.) ornatipes sp. n. 143, male imago; 144–145, fore and middle leg of male imago; 146, subimaginal mesopleuron; 147, subimaginal mesonotum; 148, abdomen of male subimago extracted from mature larva.

FIGURES 158–163. Centroptella (s. str.) ornatipes sp. n., female imago. 158, fore leg; 159–160. dorsal and ventral view; 161– 162, egg and its micropile with sperm guide; 163, margin of sperm guide of another egg, with external layer partly detached.

FIGURES 298–302. Centroptella (Crassolus) saxophila. 298–300, male imagines; 301, abdomen of male imago; 302, abdomen of male subimago extracted from mature larva.

FIGURES 373–379. Centroptella (Crassolus) ludmilae sp. n. 373–374, fore and middle legs of male imago; 375, subimaginal exuviae of abdomen; 376, subimaginal exuviae of wing; 377–379, male imagines.

FIGURES 38–42. Centroptella (s. str.) longisetosa cinerea subsp. n., larva. 38, claw; 39, caudalii; 40, cercus (holotype); 41, fore femur; 42, abdominal terga IX–X.

FIGURES 352–363. Centroptella (Crassolus) ludmilae sp. n., exuviae of larval legs. 352, fore femur; 353, seta on outer side of femur; 354–355, claws; 356–358, fore, middle and hind legs; 359–361, fore, middle and hind tibiae (focus on posterior side); 362, stout setae on proximal part of inner-anterior side of fore femur; 363, the same on hind femur.

FIGURES 364–368. Centroptella (Crassolus) ludmilae sp. n., larval exuviae. 364, apex of fore tibia (arrow shows outer-apical seta); 365, abdominal tergum V; 366, abdominal tergum III of holotype (MA, medioanterior sigilla; MP, medioposterior sigilla); 367, abdominal tergum X; 368, female abdominal sternum IX and paraprocts.

FIGURES 404–406. Centroptella (Crassolus) ingridae, larval cuticle of abdomen. 404, tergum III of holotype (MA, medioanterior sigillum; MP, medioposterior sigillum); 405, terga and sterna; 406, tergum VII (letter g corresponds to Fig. 14).

FIGURES 387–392. Centroptella (Crassolus) ludmilae sp. n. 387–388, female imago; 389, exuviae of subimaginal pleuron; 390, exuviae of subimaginal mesonotum; 391, exuviae of female subimaginal fore and hind legs. Abbreviations: T1–T5, tarsomeres.

FIGURES 308–315. Centroptella (Crassolus) saxophila. 308–309, tibia of fore and hind legs of female subimago; 310, abdomen of female imago; 311, subimaginal exuviae of mesonotum; 312, subimaginal exuviae of mesopleuron; 313–315, eggs. Abbreviations: T1–T5, tarsomeres.