Kimminsula, Peters & Edmunds, 1970

Kluge, Nikita J., 2022, Review of the Kimminsula-complex (Ephemeroptera, Leptophlebiidae), Zootaxa 5212 (1), pp. 1-140 : 6-10

Kimminsula -complex

( Figs 3 , 7–710 )

The Kimminsula -complex belongs to subsequently subordinated taxa: Leptophlebiidae >Atalophleboadentata Kluge 2009 > Atalophlebopectinata Kluge 2009 > Atalophleboculata Kluge 2009 > Atalophlebomaxillata Kluge 2009 > Atalophlebolinguata Kluge 2009. General system of the taxon Atalophlebolinguata remains poorly elaborated.

The Kimminsula -complex is distributed only in the Indian Peninsula and the Island of Sri Lanka and includes Kimminsula Peters & Edmunds 1970 (with 3–5 species), Petersula Sivaramakrishnan 1984 (with 2–3 species), Ghatula gen. n. (with 2 species), Ceylonula gen. n. (with 1 species) and Hubbardula gen. n. (with 1 species). Larvae of all these species are similar in general appearance [see (1) below]; characters separating larvae of the Kimminsula -complex from other taxa with a similar larval appearance are the presence of fronto-clypeal incisions [see (2) below] and a characteristic shape and setation of femora [see (10) below]. Imagines of the Kimminsula -complex differ from other Oriental Leptoplebiidae by well developed hind wings of plesiomorphic shape and venation [see (16) below] and by the claw structure [see (17) below].

Common characters of the Kimminsula -complex

(1) General appearance. Larva has flattened body, widened head and enlarged legs with broad femora and claws bent perpendicular to the leg flatness ( Figs 202 , 440–443 ). Superficially, they resemble larvae of Heptageniidae , but in contrast to them have no frontal shield; instead, the wide dorsal head surface contains surfaces of closely pressed together clypeus, labrum and mandibles ( Fig. 235 ) (the same in other Atalophlebomaxillata). Initially such larval structure is an adaptation for inhabiting stone surfaces in rapidly running water ( Fig. 219 ), but larvae of Ceylonula femoralis inhabit soft substrates in stagnant water ( Fig. 220 ).

(2) Fronto-clypeal incisions. Lateral margins of the clypeus are separated from lateral margins of the frons by a pair of sharp incisions leading into the anterior tentorial pits; within these incisions, the margins of the frons overlap the margins of the clypeus dorsally ( Figs 3 , 14 , 68 , 105 , 235 , 335 , 389 , 444 , 485 , 552–553 , 616 ). This differs from many other Atalophlebolinguata, where the frons and the clypeus have an integral dorsal surface, being separated only by a pair of smooth concavities of the lateral margins ( Figs 5–6 ) (see below, discussion of systematic position of the Kimminsula -complex).

(3) Labrum. As in most other Atalophlebopectinata, the dorsal surface of the labrum bears the posterior transverse setal row and the anterior transverse setal row; in the Kimminsula -complex both rows stretch nearly across the labrum width; each row is dense, either regular, i.e. with one seta per section ( Figs 16 , 73 , 232 : anter.r, 488, 619), or has the form of a strip with a few setae per section ( Figs 102 , 232 : poster.r, 617). In all species of Kimminsula -complex, the anterior margin of labrum is hooded, i.e. the initial anterior margin is bent ventrally, so that in dorsal view the posterior transverse setal row appears to be located on the visible anterior margin or close to it, while the anterior transverse setal row is often hidden ventrally ( Fig. 101 ). Setae of the true ventral side of labrum are situated irregularly (i.e. not forming rows), and form a distinctly outlined transverse ventro-anterior strip close to the anterior margin ( Figs 101, 103 ). In all species except Ghatula rufa gen. sp. n., labrum is wider than clypeus. Shape of the median incision is variable: either shallow with more or less developed denticles on the initial anterior margin (in the Ceylonese taxa Ceylonula gen. n., Kimminsula and Hubbardula gen. n., Figs 233 , 487 , 619 ), or cleft-like and lacking denticles (in the Indian taxa Ghatula gen. n. and Petersula , Figs 17 , 103 ). Petersula differs from other taxa by peculiar labral depressors [see below, Petersula (2)]

(4) Mandibles. As in majority of Atalophlebolinguata, mandibles are articulated at ventral side of the head capsule some distance from its lateral margins ( Fig. 235 ); mandibles are flattened, with outer margins convex and projected laterally. The outer margin of mandible bears the distal tuft of setae, which occupies an area adjacent to the base of the incisor, and the middle tuft of setae which is more compact and located more laterally ( Figs 22–23 , 75–76 , 229 , 563 , 620 ). Among members of the Kimminsula -complex, the middle tuft is absent only in Petersula ( Figs 145–146 ). The same in some other leptophlebiid taxa ( Kluge 2020).

(5) Maxillae. Maxilla is wide. The subapical ventral row of comb-like setae (peculiar for Leptophlebiidae ) has more or less expressed curvature, which divides it into two portions: a lateral portion consisting of setae with smaller sockets, and a median portions consisting of setae with larger sockets (length of all setae is equal). In the Ceylonese taxa, i.e. Ceylonula gen. n., Kimminsula and Hubbardula gen. n., the lateral portion is very long and the median portion is short, consisting of 5–10 setae ( Figs 332–334 ). In the Indian taxa, i.e. Ghatula gen. n. and Petersula , the whole subapical ventral setal row is shorter, and its both portions are subequal in length ( Figs 112–113 ) (the same in many other leptophlebiid taxa).

(6) Maxillary palps. Maxillary palp bears long setae on outer sides of the 2nd and 3rd segments, dense setae on ventral side and on apex of the 3rd segment. The inner side of the 3rd segment bears a regular row of setae, whose size and number vary individually and do not provide taxonomic characters within the Kimminsulacomplex. The inner side of the 2nd segment bears longer setae which are either concentrated at its apex (in both species of Petersula , Fig. 111 ) or form a more or less long longitudinal row (in other species, Figs 19 , 236 , 334 , 493 , 566 , 623 ).

(7) Labium. Shape and setation of labium is uniform in all representatives ( Figs 26 , 77 , 116–118 , 241 , 492 , 567 , 624–625 ). Submentum is bare without prominent setae: either without visible setae at all ( Figs 116 , 567 , 625 ), or with very small occasional setae. The glossae are not large, not projected from paraglossae neither ventrally, nor dorsally; long setae of their ventral side are situated densely and irregularly ( Fig. 625 ); long setae on the dorsal side of glossa form irregular row or strip of a horseshoe shape ( Fig. 242 , 624 ) [as in Thaulodes (Kuge 2020: fig. 20)].

(8) Labial palps. The last (3rd) segment of labial palp is shorter and narrower than others and apically blunt. A small, regular row of stout claw-like setae crosses apex of this segment. In Petersula , these claw-like setae are relatively long ( Fig. 117 ); in other taxa they are very short, with length about twice exceeding width at base ( Figs 25 , 240 , 492 , 567 , 626 ). The same condition occurs in most other Atalophleboculata (see discussion about systematic position below).

(9) Larval thorax. The pronotum is nearly rectangular, with rounded antero-lateral angles and a free anterior margin (i.e. transverse margin between antero-lateral angles and the neck membrane); a transverse ridge runs parallel to the free anterior margin; humeral setae are located between the free anterior margin and the ridge, being absent in other places ( Figs 9 , 91 , 95 , 228 , 324 , 391 , 481 , 556 , 610 ) (the same in many other Atalophlebolinguata).

(10) Larval femora. Femora are widened, so the groove on inner side (into which the tibia can be partly inserted) is bordered with prominent flanges (called here the inner-anterior flange and the inner-posterior flange); at least on the middle femur, the inner-anterior flange is expanded in its distal part, so the femur appears widest in distal part ( Fig. 28 ); the forefemur has similar shape in all species except Ghatula rufa gen sp. n., in which the inner-anterior flange of forefemur is reduced, being narrower than the inner-posterior flange ( Fig. 27 ); the hind femur retains the usual elongate-ellipsoid shape, being widest in the middle ( Fig. 29 ).

The forefemur is not widened proximally (in contrast to many other Leptophlebiidae ), but the cuticle of its anterior surface bears the proximal blank peculiar for mayfly larvae whose forefemur is proximally wider ( Kluge 2020: p. 15). The proximal blank of foreleg differs from blanks occurring on the middle and hind legs ( Fig 27 , 119 , 243 , 336 , 392 , 449 , 496 , 571 , 626 ).

The anterior surface of each femur bears more or less numerous, irregularly situated, stout setae, which can be either blunt ( Fig. 494 ), or pointed ( Fig. 495 ), or variable. On the inner side of the femur the stout setae form a more or less regular row running by margin of the inner-anterior flange. On middle and hind femora (but never on forefemur) the stout setae form a regular, arched, transverse row situated near the femur base ( Figs 28–29 , 79–80 , 120–121 , 245 , 337–338 , 497–498 , 572–573 , 627 ). On outer side of each femur the stout setae are enlarged and form two more or less regular rows (which can be called the outer-anterior row and the outer-posterior row), between which the outer strip of thin hairs is located; hairs of this strip are situated densely and irregularly ( Fig. 629 ). Hairs of the outer strip are generally longer and/or denser on the fore- and the middle femora, than on the hind femur ( Figs 27–29 , 78–80 , 119–121 , 336–338 , 392–394 , 449–451 , 496– 498 , 571–573 , 612–614 ); in Ceylonula gen. n. they are completely absent on the hind femur ( Fig. 245 ). The posterior side of the hind femur bears curved setae —these are small, pointed, usually pectinate setae, curved toward the inner margin of the femur and forming an irregular strip along inner margin of the femur ( Figs 36 , 123 , 247 ).

The curved setae are probably initial for Leptophlebiidae ; other features of femur setation, especially the transverse proximal rows on middle and hind femora, distinguish the Kimminsula -complex from many other Leptophlebiidae (see below, discussion about systematic position of the Kimminsula -complex).

(11) Larval tibiae. Patella-tibial suture, which initially for mayflies is absent on the forelegs and present on the middle and the hind legs ( Kluge 2004), retains this condition in Ghatula gen. n., Ceylonula gen. n. and Hubbardula gen.n.; it is differently modified in Petersula and in Kimminsula [see below, Petersula (7) and Kimminsula (7)].

As in most other Leptophlebiidae , tibiae of three leg pairs are differentiated: the pointed pectinate setae of inner side are preferably developed on the foreleg, and the stout setae of outer side are preferably developed on the hind leg.

The foretibia bears stout pointed setae, which are often bipectinate; these setae occupy a longer or shorter part of the inner side of the foretibia, being always present at least near the apex of the tibia ( Figs 124–126 , 253–254 , 339–340 , 574–575 , 632–633 ); on the middle tibia such setae are present only in Ceylonula gen. n. ( Figs 255–256 ); on the hind tibia such setae are absent in all representatives of the Kimminsula -complex.

The hind tibia differs from other tibiae by the presence of stout setae located on all sides, forming two longitudinal rows on outer side of the tibia—the outer-posterior and the outer-anterior ones ( Figs 258 , 504–505 , 639 ) and forming a transverse row on posterior side at the tibia apex ( Figs 35 , 259 ).

As in many other Leptophlebiidae , the outer side of each tibia bears a longitudinal row of hairs (long and slender setae). In Ghatula gen. n. and Ceylonula gen. n. this row of hairs is single, but Petersula , Kimminsula and Hubbardula gen. n. have a second row located more anteriorly. Each of these rows is either regular, i.e. with one seta per section ( Figs 124–125 , 342 , 501 ), or has a form of strip with a few setae per section ( Figs 576 , 639 ). On the hind tibia, which has two rows of stout setae on the outer side, the initial outer row of hairs is located between them, and the additional anterior row of hairs is located anteriad of the outer-anterior row of stout setae ( Fig. 639 ).

(12) Larval tarsus. The tarsus of each leg pair bears hairs on the outer side similar to hairs on the tibia; these hairs are longer and denser on the hind leg. Stout setae on inner sides of tarsi are differentiated as follows: on the fore- and middle legs they are small and few, can be absent; on the hind leg stout setae always form a longitudinal row at least on the distal part of the tarsus; in proximal part of the row setae are sparse and small, but they become denser, larger and hooked toward apex of the tarsus ( Fig. 35 ). Sometimes inner side of the foretarsus bears several stout, pointed, pectinate setae of the same structure as the setae on inner side of the tibia of the same leg; such setae occur in Ceylonula femoralis ( Figs 251, 253 ) and some others.

(13) Larval claw. Claw is bent perpendicular to the leg flatness and distinctly divided into the small articulatory portion and the large rigid portion ( Figs 129 , 506 ) ( Kluge 2020).

(14) Larval abdomen. The surface of terga and sterna is always smooth, without prominent relief or significant setae. Posterior margin of each abdominal tergum bears a row of sharply pointed denticles greatly varying in size; medium size of denticles is either subequal on all terga I–X ( Figs 38–41 ), or significantly differs on different terga. Posterior margin of the 10th tergum always has shallow median concavity ( Figs 41 , 398 , 586 , 631 ). Posterior margins of all sterna I–IX are always smooth, without denticles ( Fig. 94 ).

Caudalii are about 2–3 times longer than the body; all 3 caudalii (the cerci and the paracercus) are equally developed both in Ceylonula femoralis (whose paracercus is vestigial in the winged stages, Figs 269–271 ) and in other species (where the paracercus is equal to the cerci in all stages); posterior margin of each segment of cercus and paracercus bears whorls of pointed denticles and setae in all species ( Figs 41–42 ).

(15) Tergalii. Tergalii always retain 2 lamellae (initial for Leptophlebiidae ); tergalii of all pairs have similar structure, with both lamellae similar. Tergalii make rhythmic undulate movements with the wave going from front to back; either all pairs of tergalii participate equally in respiration ( Figs 440–442 ) or tergalii I do not participate in respiratory movement ( Fig. 202 ). In other respects, structure of tergalii significantly differs in different taxa [see below, Ghatula (9), Petersula (9), Ceylonula (9), Kimminsula (9) and Hubbardula (9)]. All 7 pairs of tergalii are retained in Petersula , Ceylonula gen. n., Kimminsula and Hubbardula gen. n., while Ghatula gen. n. has no tergalii of 7th pair, thus retains only 6 pairs of tergalii.

(16) Forewings. Furcation of MA is symmetrical or slightly asymmetrical. Furcation of MP is more proximal than furcation of RS and is sharply asymmetrical: vein MP 2 is attached at base to veins MP 1 and CuA either by equally oblique crossveins ( Fig. 411 ), or the crossvein connecting MP 2 with MP 1 is more oblique and may be interpreted as asymmetric arising of MP 2 from MP 1 ( Figs 51 , 68 , 151 , 156 , 279 , 453 , 539 , 670 ); this difference may be individual ( Figs 216, 218 ). Cubital field has 2 intercalaries between Cu 1 and Cu 2; in Kimminsula latifolia sp. n. a third, additional intercalary is present between them ( Fig. 581 ). Crossveins are numerous; some crossveins reach posterior margin of the wing between ends of the longitudinal and the intercalary veins. Crossveins in the area of pterostigma are numerous, complete and non-anastomosed; in Ceylonula femoralis the costal field is wider than the subcostal field, and the pterostigmatic crossveins are nearly perpendicular to Sc ( Fig. 279 ); in other taxa the costal field is not wider than the subcostal field, and the pterostigmatic crossveins vary from sharply oblique ( Fig. 414 ) to perpendicular to Sc ( Figs 670 ).

(17) Hind wings. Forewing has well expressed tornus, and the hind wing is relatively large, about as long as the basi-tornal margin of the forewing. Costal projection of the hind wing is blunt and slightly projected; Sc is continued far distad of the costal projection ( Figs 52 , 153 , 157–158 , 217 , 278 , 365 , 412 , 454 , 540 , 671 ). The vein MA is non-bifurcate as in all other Furcatergaliae (see Kluge 2004); other veins are either fully developed ( Figs 157–158 ), or MP is non-bifurcate (in Ghatula gen. n., Fig. 52 ).

(18) Claws of winged stages. On each leg of imago and subimago, both claws are pointed and hooked ( Figs 289 , 676 ) (see below, discussion about systematic position of the Kimminsula -complex).

(19) Penis. The pair of penis lobes are movably connected at base, so that are able to move apart and be brought together under action of the sterno-penial muscles. In Petersula the penial lobes are separated nearly to the base ( Figs 179 , 183 ), while in Ceylonula gen. n., Kimminsula and Hubbardula gen. n., the median sides of penis lobes are connected one with another at significant distance ( Figs 297 , 301 , 375 , 377 , 418 , 421 , 469–470 , 534 , 545 , 690 , 697 ); in the last case their mobility is ensured by the fact that the cuticle of these median areas is delicate, soft and elastic.

In Petersula , Kimminsula and Ceylonula gen. n., apex of each paired penis lobe bears a pointed process directed medio-ventrally; such processes are not present in in Hubbardula gen. n. ( Figs 690 , 697 ); genitalia of Ghatula gen. n. are unknown. In taxa with the medio-ventral processes, these processes arise from the penis apices and look similarly, but their structure is significantly different: in Kimminsula they are conic, i.e. round in cross section ( Figs 374–375 , 377, 380 , 417–418 , 421 , 469–470 , 545 ), in Ceylonula gen. n. they are grooved with concavities facing medio-dorsally ( Figs 300–301 ), and in Petersula they are grooved with concavities faced latero-ventrally ( Figs 172–173, 179 ).

In Petersula , the pointed projections are already present on larval protopenis, where they have the same position as in imago ( Figs 175–177 , 209 ). In Ceylonula gen. n. and Kimminsula , larval protopenis has no any precursors of these projections ( Figs 304–305 , 379 , 425–426 , 549 , 602 ).

In all cases (in Petersula , Kimminsula , and Ceylonula gen. n.) the medio-ventral pointed processes arise from the ventral sides of penis lobes, significantly distad to the gonopores, while the gonopores are opened on the dorsal sides of the penis lobes. The medio-ventral pointed processes have no connection with the gonoducts: in Petersula and Ceylonula gen. n. their grooves do not touch the gonopores ( Figs 172 , 300 ), and in Kimminsula they have no any grooves or canals at all. The pair of gonoducts passing inside the penis lobes, have no musculated sperm pumps, in contrast to Thraulodes ( Kluge 2020: figs 101, 111) and some others.

(20) Styliger and gonostyli. The posterior-dorsal margin of styliger is distinct, and its postero-ventral margin is not expressed, so the gonostyli are attached on ellipsoid membranous cavities faced ventro-caudally and well visible from ventral view ( Figs 181 , 184 ). In other respects styligers and gonostyli are significantly diverse: in Petersula and Kimminsula the styliger is short and simple, and the gonostyli retain division into 4 segments ( Figs 184 , 376 , 420 , 472 ); in Ceylonula femoralis the styliger is projected medially, and the gonostyli lack boundary between the initial 1st and 2nd segments ( Fig. 296 ); in Hubbardula heterolepida sp. n. the styliger is elongated and modified, the membranous cavities of gonostyli attachments are connected medially, and the gonostyli lack boundary between the initial 2nd and 3rd segments ( Figs 689, 691 , 698 ).

Composition of the Kimminsula -complex

This complex includes two south Indian genera, Petersula Sivaramakrishnan 1984 and Ghatula gen. n., and three Ceylonese genera, Ceylonula gen. n., Kimminsula Peters & Edmunds 1970 and Hubbardula gen. n. Key characters of these genera are given in Table 1 .

Figures

FIGURES 1–6. Junction of frons and clypeus in larvae of Leptophlebiidae (left margin, dorsal view). 1, Leptophlebia vesperina (Linnaeus 1758); 2. Terpides echinovaris Kluge 2015; 3, Ceylonula femoralis (Hagen 1858); 4, Thraulus bellus Eaton 1881; 5, Choroterpes picteti (Eaton 1871); 6, Meridialaris chiloeensis (Demoulin 1955). Abbreviations: cly, clypeus; c.md, anterior mandibular condyle; fr, frons; lbr, labrum.

FIGURES 7–15. Ghatula rufa gen. sp. n., larva. 7–13, exuviae (with the same magnifivation; holotype): 7, head with labrum and mandibles; 8, abdominal terga; 9, right half of pronotum and mesonotum, humeral setae enlarged; 10, abdominal sterna VII–IX; 11–13, fore-, middle and hind legs; 14, right-anterior part of head margin; 15, young female larva. Abbreviations: cly, clypeus; c.md.a and c.md.p, anterior and posterior mandibular condyles visible on ventral side of head through cuticle; in, incision laterad of posterior mandibular condyle; in-a and in-p, convexities anteriad and posteriad of incision.

FIGURES 16–26. Ghatula rufa gen. sp. n., larval mouthparts. 16–18, labrum (16, with focus on dorsal side; 17, with focus on ventral side); 19–21, maxilla; 22–23, left and right mandibles; 24, right half of hypopharynx with superlingua; 25–26, labium (16, 18, 19, holotype).

FIGURES 27–29. Ghatula rufa gen. sp. n., exuviae of larval femora of fore-, middle and hind legs, anterior view (holotype). Abbreviations: i-a.f, inner-anterior flange; i-p.f, inner-posterior flange; p.b, proximal blank; t.r., transverse row of stout setae.

FIGURES 30–35. Ghatula rufa gen. sp. n., exuviae of larval tibiae, tarsi and claws (holotype). 30, foreleg (anterior view); 31, its claw; 32, middle leg (anterior view); 33, hind leg (anterior view); 34–35, its enlarged fragments in posterior view. Arrows show point where patella-tibial suture crosses inner side of tibia.

FIGURES 36–42. Ghatula rufa gen. sp. n., details of larval exuviae (holotype). 36, curved setae on posterior side of hind femur; 37, stout setae on anterior side of forefemur; 38–41, denticles on posterior sides of abdominal terga I, VIII, IX and X; 42, middle part of cercus.

FIGURES 43–50. Ghatula rufa gen. sp. n., larva. 43–48, tergalii I–VI (holotype; costal margin facing up, ventral lamella turned at left, dorsal lamella at right); 49–50, fore- and hind protoptera of young larva. Abbreviations: c-pr, costal-proximal convexity of ventral lamella; a-pr, anal-proximal convexity of ventral lamella.

FIGURES 51–58. Ghatula rufa gen. sp. n., female (holotype). 51–52, fore- and hind wings of imago; 53–55, fore-, middle and hind legs of imago with enlarged fragment of hind tibia; 56, subimaginal exuviae of mesonotum; 57–58, subimago.Arrows show point where patella-tibial suture crosses inner side of tibia.

FIGURES 59–62. Ghatula rufa gen. sp. n., eggs (extracted from holotype).

FIGURES 63–67. Ghatula gen. n., posterior margin of sternum IX. 63–65, Ghatula rufa sp. n.: 63, female subimago (holotype); 64, last instar female larva (holotype); 65, penultimate instar male larva; 66–67, Ghatula quadrimaculata sp. n.: 66, last instar female larva; 67, penultimate instar male larva.Abbreviations: pgs, protogonostylus of male larva; pp, protopenis of male larva.

FIGURES 68–71. Ghatula quadrimaculata sp. n., larvae. 68–70, female larva of last instar (holotype); 71, denticles on posterior margin of abdominal segment VI.

FIGURES 72–77. Ghatula quadrimaculata sp. n., larvae. 72–73, labrum; 74, maxilla; 75–76, left and right mandibles; 77, labium. Abbreviations: d.t, distal tuft of setae; m.t, middle tuft of setae.

FIGURES 78–80. Ghatula quadrimaculata sp. n., larva. 78, forefemur, 79, middle femur, 80, hind femur.Abbreviations: i-a.f, inner-anterior flange; i-p.f, inner-posterior flange.

FIGURES 81–84. Ghatula quadrimaculata sp. n., larva. 81–83, fore-, middle and hind tibiae and tarsi; 84, claw. Arrow shows point where patella-tibial suture crosses inner side of tibia.

FIGURES 85–93. Ghatula quadrimaculata sp. n., larva. 85–90, tergalii of I–VI pairs (costal margin facing up, ventral lamella turned to left, dorsal lamella to right); 91, half of pronotum and mesonotum, humeral setae enlarged, foreprotopteron with cuticular and hypodermal coloration; 92, enlarged hind protopteron, the same; 93, hind protopteron with visible venation.

FIGURES 94–100. Petersula heptagenoides sp. n., larval exuviae (with the same magnification). 94, thoracic and abdominal sterna with thoracic pleuron and half of postnotum; 95, left half of pronotum and mesonotum, humeral setae enlarged; 96, head with labrum and mandibles; 97–99, fore-, middle and hind legs, 100, abdominal terga.

FIGURES 101–107. Petersula heptagenoides sp. n., larva. 101, labrum in longitudinal section; 102, anterior margin of labrum, focus on dorsal side; 103, the same, focus on ventral side; 104, proximal-lateral margin of labrum; 105, cuticle of head of male larva; 106, labrum with hypoderm and muscles; 107, its muscle-depressor enlarged. Abbreviations: a.m, initial anterior margin; anter.r, anterior transverse setal row; cly, clypeus; c.md.a and c.md.p, anterior and posterior mandibular condyles visible on ventral side of head through cuticle; depr, muscle-depressor; in, incision laterad of posterior mandibular condyle; in-a and in-p, convexities anteriad and posteriad of incision; poster.r, posterior transverse setal row; v.s, ventral setae; v-a.s, ventro-anterior strip of setae.

FIGURES 108–113. Petersula heptagenoides sp. n., larva. 108–109, left mandible (108, compressed; 109, in natural position); 110, right mandible (compressed); 111, maxillary palp; 112, maxilla; 113, subapical ventral row of comb-like setae. Abbreviations: inc, incisor; kd, kinetodontium; prs, prostheca.

FIGURES 114–118. Petersula heptagenoides sp. n., larva. 114–115, just molted female larva of last instar with non-pigmented cuticle, to show hypodermal coloration; 116–118, labium.

FIGURES 119–123. Petersula heptagenoides sp. n., larval femora. 119–121, femora of fore-, middle and hind legs; 122, stout setae on anterior side of femur; 123, curved setae on posterior side of hind femur.

FIGURES 124–129. Petersula heptagenoides sp. n., larval tibiae, tarsi and claws (anterior view). 124–126, foreleg; 127, middle leg; 128–129, hind leg. Abbreviations: a.p, denticles of articulatory portion; a.r, anterior row of hair-like setae; o.r, outer row of hair-like setae; p-t.s, apex of patella-tibial suture; r.p, denticles of rigid portion.

FIGURES 130–144. Petersula heptagenoides sp. n., larvae. 130–136, tergalii I–VII (costal margin facing up, ventral lamella turned at left, dorsal lamella at right; holotype); 137–138 and 139–141, fore- and hind protoptera; 140 and 143, hind protoptera enlarged; 144, posterior margin of abdominal tergum IX (black triangle shows median line).

FIGURES 145–153. Petersula heptagenoides sp. n. 145–146, right and left mandibles, ventral view; 147, hypopharynx and superlinguae; 148–150, tergalii I, IV and VII; 151–152, fore- and hind wings; 153, hind wing enlarged.

FIGURES 154–165. Petersula heptagenoides sp. n. 154, subimaginal exuviae of mesonotum; 155–156, fore- and hind wings; 157–158, hind wings enlarged; 159–161 and 162–164, fore-, middle and hind legs of two female imagines; 165, male subimago.

FIGURES 166–171. Petersula heptagenoides sp. n. 166, subimaginal exuviae at boundary of abdominal sterna VII and VIII with genital opening; 167–168, male imago; 169–171, female imago.

FIGURES 172–181. Petersula heptagenoides sp. n. 172, apex of left penis lobe of male imago, median view; 173, the same, ventral view; 174, posterior margin of abdominal sternum IX of female larva; 175–177, protogonostyli and protopenes of male larvae of antepenultimate, penultimate and last instars; 178, posterior margin of abdominal sternum IX of female subimago; 179–180, penis of male imago with focus on ventral and dorsal side (gonoduct shown by dotted lines); 181, left penis and gonostylus, median view cutting line shown with dotted line). Abbreviations: gp, gonopore; pgs, protogonostylus; pp, protopenis.

FIGURES 182–187. Petersula spp. 182–186, Petersula heptagenoides sp. n.: 182–184, genitalia of male imago; 185, subimaginal exuviae of male genitalia; 186, subimaginal exuviae of wing; 187, the same of P. courtallensis. Abbreviations: gd, gonoduct; gs1–gs4, 1st–4th segments of gonostylus; sg, styliger.

FIGURES 188–195. Petersula spp., eggs. 188–191, Petersula heptagenoides sp. n.; 192–195, P. courtallensis.

FIGURES 196–204. Petersula courtallensis, larvae. 196, exuviae of metathoracic and abdominal sterna; 197–199, fore- and hind protoptera; 200, foreprotoptera of another individual; 201, larva molting to last instar; 202, alive larva with moving tergalii; 203–204, just molted female larva of last instar with non-pigmented cuticle, to show hypodermal coloration.

FIGURES 205–208. Petersula courtallensis. 205–207, female imagines; 208, subimaginal abdomen extracted from male larva.

FIGURES 209–218. Petersula courtallensis. 209, protogonostyli and protopenes of last instar male larva; 210, posterior margin of abdominal sternum IX of female larva; 211, the same of female subimago; 212–214, fore-, middle and hind legs of female imago; 215, subimaginal exuviae of mesonotum; 216–217, fore- and hind wings of imago; 218, fragment of forewing of another individual.

FIGURES 219–220. Biotops in Sri Lanka. 219, collecting sites of Hubbardula heterolepida sp. n. and Kimminsula fasciata in river Kirikatu oya; 220, collecting site of Ceylonula femoralis in river Seethagangula.

FIGURES 221–228. Ceylonula femoralis, larva. 221–226, exuviae (same magnification): 221, head with labrum and mandibles; 222, abdominal terga and sterna; 223, right half of pronotum and mesonotum; 224–226, fore-, middle and hind legs; 227, just molted larva with non-pigmented cuticle, to show hypodermal coloration; 228, free anterior margin of pronotum and humeral setae.

FIGURES 229–235. Ceylonula femoralis, larva. 229, left mandible; 230–235, apices of left and right mandibles, compressed; 232, anterior margin of labrum; 233, margin of labrum, anterior view; 234, proximal-lateral margin of labrum; 235, cuticle of head of female larva. Abbreviations: anter.r, anterior transverse setal row; c.md.a and c.md.p, anterior and posterior mandibular condyles visible on ventral side of head through cuticle; cd, kinetodontium; dent, denticles; d.t, distal tuft of setae; in, incision laterad of posterior mandibular condyle; in-a and in-p, convexities anteriad and posteriad of incision; inc, incisor; m.t, middle tuft of setae; poster.r, posterior transverse setal row; prs, prostheca; v-a.s, ventro-anterior strip of setae.

FIGURES 236–242. Ceylonula femoralis, larva. 236–238, maxilla and its details; 239, hypopharynx with superlingua; 240, apex of labial palp; 241, labium, focus on ventral side; 242, glossae, focus on dorsal side.

FIGURES 243–250. Ceylonula femoralis, larval exuviae. 243–245, femora of fore-, middle and hind legs; 246, stout setae on anterior side of hind femur; 247, curved setae on posterior side of hind femur; 248–250, posterior margins of abdominal terga I, IX and X.

FIGURES 251–260. Ceylonula femoralis, larval exuviae: tibiae, tarsi and claws. 251–254, foreleg (251, anterior view; 253, view from inner side); 255–257, middle leg (255–256, anterior view; 257, posterior view); 258–260, hind legs (258–259, posterior view; 260, anterior view).

FIGURES 261–271. Ceylonula femoralis. 261, right side of larval abdominal segments I–V with tergalii in natural position; 262–268, tergalii I–VII (costal margin facing up, ventral lamella turned at left, dorsal lamella at right); 269–271, bases of caudalii of larva, subimago and imago of same individual.

FIGURES 272–277. Ceylonula femoralis, subimago. 272–276, subimaginal exuviae: 272, abdomen of male; 273, right half of mesonotum; 274–275, fore- and hind legs of female; 276, enlarged fragment of hind leg; 277, male subimago. Arrow shows point where patella-tibial suture crosses inner side of tibia.

FIGURES 278–289. Ceylonula femoralis, imago: 278–284, male; 285–289, female (286–288, fore-, middle and hind legs with enlarged fragment of hind leg). Arrows show point where patella-tibial suture crosses inner side of tibia.

FIGURES 290–293. Ceylonula femoralis, male imago, syntype (pictures from website of the Museum of Comparative Zoology, Cambridge).Arrow shows where left hind tibia is twisted along patella-tibial suture.

FIGURES 294–299. Ceylonula femoralis, male genitalia. 284, imago; 295, subimaginal exuviae; 296, styliger with gonostyli; 297, penis and left gonad (right gonad detached); 298, its fragment enlarged; 299, genitalia of subimago. Abbreviations: gd, gonoduct; gs1–gs4, 1st–4th segments of gonostylus, m.gs, muscle of gonostylus; m.p, muscle of penis; m.s, sterno-styligeral muscle; sg, styliger.

FIGURES 300–305. Ceylonula femoralis. 300, apex of right penis lobe, median view (gp, gonopore); 301, penis, ventral view (gonoducts shown by dotted lines); 302, posterior margin of abdominal sternum IX of female subimago; 303, the same of female larva; 304, protogonostyli and protopenis of male larval exuviae; 305, enlarged protopenis inverted as result of molt.

FIGURES 306–310. Ceylonula femoralis, female imago: 306–307, ventral view; 308–310, lateral view. Abbreviation: g.o, genital opening.

FIGURES 311–316. Ceylonula femoralis, development of wing coloration. 311, fore- and hind protoptera of young larva; 312, hind protopteron enlarged; 313–314, fore- and hind wings of subimago; 315–316, fore- and hind wings of imago.

FIGURES 317–320. Ceylonula femoralis, eggs.

FIGURES 321–331. Kimminsula fasciata, larval exuviae (with the same magnification). 321–323, parts of head; 324–325, pronotum and mesonotum of two individuals; 326–328, fore-, middle and hind legs; 329–331, abdominal terga of three individuals.

FIGURES 332–335. Kimminsula fasciata, larva. 332–334, maxilla and its subapical ventral row of comb-like setae; 235, cuticle of head of male larva. Abbreviations: c.md.a and c.md.p, anterior and posterior mandibular condyles visible on ventral side of head through cuticle; in, incision laterad of posterior mandibular condyle; in-a and in-p, convexities anteriad and posteriad of incision.

FIGURES 336–338. Kimminsula fasciata, exuviae of larval femora of fore-, middle and hind legs.

FIGURES 339–347. Kimminsula fasciata, exuviae of larval tibiae, tarsi and claws. 339–340, foreleg (339, anterior view, 340, posterior view); 341–343, middle leg (341–342, anterior view; 343, posterior view); 344–345, hind leg (anterior view); 346–347, claws. Abbreviations: a.r, anterior row of hair-like setae; o.r, outer row of hair-like setae; p-t.s, point where patellatibial suture crosses inner side of tibia.

FIGURES 348–368. Kimminsula fasciata. 348–354 and 355–361, tergalii I–VII of two different larvae of last instar; 362–363, fore- and hind protoptera; 364, hind protopteron enlarged; 365, hind wing; 366, posterior margin of larval abdominal tergum VI; 367, just molted last instar female larva with cuticle colorless, to show hypodermal coloration; 368, female imago.

FIGURES 369–373. Kimminsula fasciata. 369, male imago; 370–371, male imago and subimago of the same individual; 372, subimaginal exuviae of mesonotum; 373, abdominal sterna and terga of male imago.

FIGURES 374–376. Kimminsula fasciata, male imagines. 374, penis with styliger; 375–376, separated penis and styliger with gonostyli.

FIGURES 377–382. Kimminsula fasciata. 377–378, penis of male imago with focus on ventral and dorsal sides (gonoducts shown by dotted lines); 379, protogonostyli and protopenis of last instar male larva; 380, apices of penis of male imago; 381, posterior margin of abdominal sternum IX of female subimago; 382, the same of female larva.

FIGURES 383–386. Kimminsula fasciata, eggs.

FIGURES 387–394. Kimminsula taprobanes, larval exuviae (with the same magnification). 387–388, left and right mandibles; 389, head; 390, abdomen; 391, pronotum and mesonotum; 292–394, fore-, middle and hind legs.

FIGURES 395–405. Kimminsula taprobanes, larva. 395, outer margin of exuviae of forefemur; 396–398, posterior margins of abdominal terga VI, IX and X.; 399–405, tergalii I–VII (costal margin facing up, ventral lamella turned at left, dorsal lamella at right).

FIGURES 406–412. Kimminsula taprobanes. 406–407, last instar female larva just after molt, with cuticle non-pigmented, to show hypodermal coloration; 408–410, fore- and hind protoptera of last instar larva; 411–412, fore- and hind wings of imago.

FIGURES 413–416. Kimminsula taprobanes. 413, male imago, syntype of annulatus [Potamanthus] (picture from website of the Museum of Comparative Zoology, Cambridge); 414–415, male imago and subimago of the same individual; 416, abdominal sterna and terga of male imago.

FIGURES 417–420. Kimminsula taprobanes, male genitalia. 417, genitalia of imago; 418, imaginal penis with gonads; 419, subimaginal exuviae; 420, imaginal styliger with gonostyli. Abbreviations: gd, gonoduct; gs1–gs4, 1st–4th segments of gonostylus; sg, styliger.

FIGURES 421–427. Kimminsula taprobanes. 421–422, penis of male imago with focus on ventral and dorsal sides (gonoducts shown by dotted lines; arrow); 423, posterior margin of abdominal sternum IX of female subimago; 424, the same of female larva; 425, protogonostyli and protopenis of last instar male larva (dorsal view) with developing subimaginal genitalia inside (shown by interrupted lines); 426, protopenis in ventral view; 427, penis of holotype of taprobanes [Baetis] (from Kimmins 1960). Arrows show points of «breaking» on lateral margins.

FIGURES 428–431. Kimminsula taprobanes. 428–430, female imago; 431, right half of subimaginal exuviae of mesonotum.

FIGURES 432–435. Kimminsula taprobanes, female. 434, abdominal segments VII–X; 433–435, subimaginal cuticle of abdominal sterna under three different magnifications.

FIGURES 436–439. Kimminsula taprobanes, eggs.

FIGURES 440–443. Form intermediate between Kimminsula taprobanes and K. fasciata, larvae.

FIGURES 444–451. Form intermediate between Kimminsula taprobanes and K. fasciata, larval exuviae (with the same magnification). 444, head; 445, pronotum and mesonotum; 446–447, left and right mandibles; 448, abdomen; 449–451, fore-, middle and hind legs.

FIGURES 452–457. Form intermediate between Kimminsula taprobanes and K. fasciata. 452, last instar female larva just after molt, with cuticle non-pigmented, to show hypodermal coloration; 453–454, fore- and hind wings of imago; 455–457, fore- and hind protoptera of last instar larva.

FIGURES 458–463. Form intermediate between Kimminsula taprobanes and K. fasciata. 458, male imago; 459, its subimaginal exuviae of mesonotum; 560–462, its fore-, middle and hind legs; 463, its abdomen.

FIGURES 464–467. Form intermediate between Kimminsula taprobanes and K. fasciata, tarsus of foreleg of male subimago. 464, its 1st tarsomere; 465, boundary between 1st and 2nd tarsomeres; 466 apex of 3rd tarsomere; 467, 4th tarsomere.

FIGURES 468–473. Form intermediate between Kimminsula taprobanes and K. fasciata, male genitalia. 468–471, penis with gonads (471, ventral view; 472, dorsal view; gonoducts shown by dotted line); 472, styliger with gonostyli; 473, their subimaginal exuviae. Abbreviations: gd, gonoduct; gs1–gs4, 1st–4th segments of gonostylus.

FIGURES 474–475. Form intermediate between Kimminsula taprobanes and K. fasciata, egg.

FIGURES 476–486. Kimminsula podi sp. n., larvae. 476–484, larval exuviae (with the same magnification): 476–477, left and right mandibles; 478, head; 479, metanotum with hind protoptera; 480, abdominal sterna and terga; 481, pronotum and mesonotum, humeral setae enlarged; 482–484, fore-, middle and hind legs; 485, fragment of head; 486, young larva with cuticular and hypodermal coloration combined. Abbreviations: cly, clypeus; in, incision laterad of posterior mandibular condyle; in-a and in-p, convexities anteriad and posteriad of incision.

FIGURES 487–495. Kimminsula podi sp. n., larvae. 487–488, anterior margin of labrum with focus on ventral and dorsal sides; 489, proximal-lateral margin of labrum; 490, posterior margin of abdominal tergum IX; 491, labium; 492, apex of labial palp; 493, maxilla; 494, stout setae on anterior side of forefemur; 495, the same on hind femur (487–488, 492, holotype). Abbreviations: anter.r, anterior transverse setal row; dent, denticles; poster.r, posterior transverse setal row; v-a.s, ventro-anterior strip of setae.

FIGURES 496–498. Kimminsula podi sp. n., larva: exuviae of fore-, middle and hind femora.

FIGURES 499–506. Kimminsula podi sp. n., larvae: exuviae of tibiae, tarsi and claws. 499, foreleg; 500–502, middle leg (500–501, focus on anterior side; 502, focus on posterior side); 503–505, hind leg (503–504, focus on anterior side; 505, focus on posterior side); 505, claw (501, 502, 506, holotype). Double-sided arrows show point where patella-tibial suture crosses inner side of tibia; abbreviations: a.p, denticles of articulatory portion; a.r, anterior row of hair-like setae; o.r, outer row of hair-like setae; r.p, denticles of rigid portion.

FIGURES 507–527. Kimminsula podi sp. n., larvae: tergalii (costal margin facing up, ventral lamella turned at left, dorsal lamella at right; 507–513, holotype). 507–513 and 514–520, tergalii I–VII in dry condition, to show outlines; 521–527, tergalii I–VII in Canadian balsam, to show coloration.

FIGURES 528–538. Kimminsula podi sp. n. 528–536, male imago extracted from subimaginal cuticle (holotype): 528, subimaginal exuviae of mesonotum; 529–531, imaginal head and thorax; 532, imaginal abdomen; 533, subimaginal exuviae of abdomen; 534, imaginal genitalia; 535–536, fore- and hind legs of pharate imago. 537–538, fore- and hind legs of mature larva with developed hypodermal coloration.

FIGURES 539–544. Kimminsula podi sp. n. 539–540, fore- and hind wings of subimago; 541, subimaginal exuviae of wing; 542, fragment of imaginal wing extracted from subimago; 543–544, fore- and hind protoptera of immature last instar larva (541–542, holotype).

FIGURES 545–549. Kimminsula podi sp. n. 545–546, penis of male imago with focus on ventral and dorsal sides; 547, distal margin of styliger with bases of gonostyli; 548, posterior margin of abdominal sternum IX of female larva; 549, protogonostyli and protopenis of last instar male larva, dorsal view (545–547, holotype).

FIGURES 550–551. Kimminsula podi sp. n., egg.

FIGURES 552–561. Kimminsula latifolia sp. n., larvae. 552–553, heads of female and male; 554–555, left and right mandibles; 556, pronotum and mesonotum; 567, abdominal segments I–IX and X; 558, fragment of left anterior margin of pronotum and humeral setae; 559–561, fore-, middle and hind legs (553–556, holotype).

FIGURES 562–568. Kimminsula latifolia sp. n., larvae. 562–563, left and right mandibles; 564, proximal-lateral margin of labrum; 565, hypopharynx and superlingua; 566, maxilla; 567, labium; 568, apex of labial palp (562–563 and 565, 567–568, holotype). Abbreviations: d.t, distal tuft of setae; m.t, middle tuft of setae.

FIGURES 569–573. Kimminsula latifolia sp. n., larval femora. 569–570, stout setae on anterior side of forefemur; 571–573, fore-, middle and hind femora.

FIGURES 574–580. Kimminsula latifolia sp. n., larvae. 574–575, foreleg; 576–577, middle leg; 578–579, hind leg; 580, young female larva with well-expressed hypodermal coloration. Abbreviations: a.r, anterior row of hair-like setae; o.r, outer row of hair-like setae.

FIGURES 581–586. Kimminsula latifolia sp. n., larvae. 581–583, fore- and hind protoptera; 584–586, posterior margins of abdominal terga V, IX and X.

FIGURES 587–600. Kimminsula latifolia sp. n. 687–593, tergalii I–VII (in Canadian balsam); 594–600, tergalii I–VII (dry, to show outlines).

FIGURES 601–606. Kimminsula latifolia sp. n. 601, posterior margin of abdominal sternum IX of female larva; 602, protogonostyli and protopenis of last instar male larva (dorsal view; holotype); 603–604, posterolateral spines of abdominal segments VIII and IX; 605–606, the same of another specimen.

FIGURES 607–615. Hubbardula heterolepida gen. sp. n., larva. 607–614, larval exuviae (with the same magnification): 607– 609, parts of head with labrum and mandibles; 610, pronotum and mesonotum, humeral setae enlarged; 611, abdominal sterna and terga; 612–614, fore-, middle and hind legs; 615, femur of foreleg, to show hypodermal coloration (610–614, holotype).

FIGURES 616–619. Hubbardula heterolepida gen. sp. n., larva. 616, cuticle of head of female larva; 617, anterior margin of labrum, dorsal view; 618, proximal-lateral margin of labrum; 619, anterior view of labrum (617, 619, holotype). Abbreviations: anter.r, anterior transverse setal row; c.md.a and c.md.p, anterior and posterior mandibular condyles visible on ventral side of head through cuticle; dent, denticles; in, incision laterad of posterior mandibular condyle; in-a and in-p, convexities anteriad and posteriad of incision; poster.r, posterior transverse setal row; v-a.s, ventro-anterior strip of setae.

FIGURES 620–625. Hubbardula heterolepida gen. sp. n., larva. 620–621, left and right mandibles; 622, hypopharynx and superlingua; 623, maxilla; 624–625, labium with focus on dorsal and ventral sides; 626, apex of labial palp. Abbreviations: d.t, distal tuft of setae; m.t, middle tuft of setae.

FIGURES 626–631. Hubbardula heterolepida gen. sp. n., larval exuviae. 626–627, femora of fore- and hind legs; 628, stout setae on anterior side of middle femur; 629, setae on outer side of femur; 630–631, posterior margins of terga VII and X (630–631, holotype).

FIGURES 632–640. Hubbardula heterolepida gen. sp. n., larval exuviae of tibiae, tarsi and claws. 632–633, foreleg (632, anterior view, 633, posterior view); 634–636, middle leg (634, posterior view, 635–636, anterior view); 637–638, hind leg (637, posterior view; 638, anterior view; 639, outer view); 640, claw (634–635, 637, holotype). Double-sided arrows show point where patella-tibial suture crosses inner side of tibia; abbreviations: a.r, anterior row of hair-like setae; o.r, outer row of hairlike setae.

FIGURES 641–663. Hubbardula heterolepida gen. sp. n., right tergalii with ventral lamella bent left. 641, tergalius III of last instar larva (in Canadian balsam, to show natural coloration); 642–648, tergalii I–VII of the same larva; 649, tergalius III of holotype (dry, to show outline); 650–656, tergalii I–VII of young larva with minimally developed protoptera (in falling light, to show natural coloration); 657–663, the same in transmission light, to show outlines.

FIGURES 664–672. Hubbardula heterolepida gen. sp. n. 664–666 and 667–669, fore- and hind protoptera of two individuals; 670–672, fore- and hind wings (holotype).

FIGURES 673–682. Hubbardula heterolepida gen. sp. n. 673–674, male imago; 675, male subimago; 676–677, hind tibia, tarsus and claws of subimago; 678, abdomen of male imago; 679–682, fore-, middle and hind legs of male imago; 682, subimaginal exuviae of mesonotum (678–681, holotype).

FIGURES 683–687. Hubbardula heterolepida gen. sp. n., tarsus of foreleg of male subimago. 683, total view; 684, 1st tarsomere; 685, fragment of 5th tarsomere; 686–687, fragments of 4th tarsomere.

FIGURES 688–693. Hubbardula heterolepida gen. sp. n. 688–692, genitalia of male imago (688, lateral view; 689–692, ventral view): 689, uromere IX with styliger and gonostyli; 690, penis with gonades; 691, styliger and bases of gonostyli; 692, general view; 693, subimaginal exuviae. Abbreviations: gs1, gs2+3, gs4, 1st–4th segments of gonostylus; sg, styliger.

FIGURES 694–699. Hubbardula heterolepida gen. sp. n. 694, posterior margin of abdominal sternum IX of female subimago; 695, the same of female larva; 696, protogonostyli and protopenis of last instar male larva (dorsal view); 697, penis of male imago, ventral view; 698–699, genitalia of male imago, ventral view.Abbreviations: gs1, gs2+3, gs4, 1st–4th segments of gonostylus; m.gs, muscle of gonostylus; m.p, muscle of penis; m.s, sterno-styligeral muscle; sg, styliger.

FIGURES 700–706. Hubbardula heterolepida gen. sp. n. 700–704, female imagines and genital opening; 705–706, subimaginal exuviae of female abdominal sterna V–IX and enlarged field posteriad of genital opening.

FIGURES 707–710. Hubbardula heterolepida gen. sp. n., eggs.