New Oriental tribe Iscini, new non-dilatognathan species of Notophlebia Peters & Edmunds 1970 and independent origin of Dilatognathus-type mouth apparatus in Atalophlebiinae (Ephemeroptera: Leptophlebiidae) Author KLUGE, NIKITA J. text Zootaxa 2014 2014-02-04 3760 4 522 538 http://dx.doi.org/10.11646/zootaxa.3760.4.2 journal article 5951 10.11646/zootaxa.3760.4.2 20e9fca4-d11b-4f5e-b0aa-40ffd6edd0a1 1175-5326 5040928 A180D4A6-78B7-4FFB-A291-728B2FAF3495 Notophlebia ganeshi Kluge sp. n. ( Figs 1–38 ) Material. Holotype : L-S-I ♂ {specimen [XV] (20)}: INDIA , state Karnataka , Shivamogga district , tributaries of rivers Agumbe-hole and Yele-hole near Agumbe , 1.II.2013 , coll. N. Kluge , L. Sheyko. Paratypes : the same locality, 11.I.–1.II.2013 : 2 L-S-I ♂ , 2 L-S-I ♀ , 54 larvae . Part of larvae were collected in river Yele-hole above Onake Abbe Falls , about 4 km WNW of Agumbe. Most part of larvae were collected in a small mountain forest stream—tributary of river Agumbe-hole between village Malandur and the road Agumbe—Sringeri , about 4 km ESE of Agumbe ; there is a confluence of two streams, at which Ganesha mini temple is situated (in honor of which this species is named). Imagos were reared in cages, and larvae from these two localities have been mixed . Larva. CUTICULAR COLORATION: Ocher-brownish, without contrasting markings; head, labrum, lateral parts of mandibles, pronotum, mesonotum, thoracic pleurites and abdominal terga darker; thoracic and abdominal sterna lighter. Legs ocher-brownish; fore femur with diffusive, large, roundish lighter blank in proximal half; middle and hind femora with diffusive, longitudinal, arched blank parallel to outer margin. HYPODERMAL COLORATION: Pronotum and mesonotum at most ocher, with lateral margins darker brownish. Femora either entirely ocher, or with darker brown macula at apex. Abdomen ocher. Both lamellae of each tergalius dark gray. SHAPE AND SETATION : Labrum ( Fig. 5 ) as wide as clypeus, widest in proximal half; fore margin convex; median incision small, with smooth shelf instead of denticles (unlike N. jobi , whose median incision and shelf are lost); distal transverse setal row dense and irregular, its setal bases are situated close one to another and form a stripe 5–6 setal bases width; instead of proximal transverse setal row, a wide field of irregularly situated setae [see Notophlebia (1)]. Mandibles ( Fig. 10 ) with outer margin moderately convex, with a few setae not forming arched field (unlike N. jobi ). Hypopharynx and superlinguae not widened (unlike N. jobi ), inner lobes of superlinguae projected distad of hypopharynx ( Fig. 4 ) [see Notophlebia (2)]. Maxilla ( Fig. 7 ) without tusk, with well-developed apical flange (see Kluge 2012 : Fig. 3 ), dentiseta and subapical row of pectinate setae (unlike N. jobi ); number of subapical pectinate setae reduced to 3, with marginal one smaller than two others ( Fig. 6 ) [see Iscini (1)]. 2 nd segment of maxillary palp with one long straight pointed stout seta near apex on ventro-lateral side, and with one placoid sensilla mediad of this seta. 3 rd (apical) segment of maxillary palp with numerous, moderately long, slender setae, situated densely and irregularly; their bases occupy apical 2/3 of ventral side ( Fig. 7 ) and apical 1/3 of dorsal side of the 3 rd segment (in contrast to N. jobi , they do not form regular rows); inner margin with longitudinal row of 3–6 shorter spine-like setae. Labium ( Fig. 8 ) with glossae small and not projected dorsad or ventrad of paraglossae; paraglossae moderately widened, roundish (unlike greatly widened in N. jobi ). Labial palp non-specialized; 2 nd segment not shortened; 3 rd segment less than twice longer than 2 nd segment; filtering setae on dorsal side of 3 rd segment moderately long and directed apically-inward ( Fig. 9 ) (unlike very long and directed apically-outward in N. jobi ); besides these setae, 1 st , 2 nd and 3 rd segments bear a longitudinal row of sparse setae on outer margins. Thorax ( Fig. 36 ) much narrower than in N. jobi ; fore protoptera brought together, in intact position their tornoapical margins contiguous up to tips. Hind legs much larger than fore and middle legs; fore and hind femora much thicker than middle femur ( Fig. 3 ). Femora wide, narrowed toward apex; fore femur wide, widest in middle part ( Fig. 14 ); middle femur less wide, widest in distal half; hind femur wide, widest near middle—in younger larva proximad of middle ( Fig. 3 ), in mature larva distad of middle. All femora with irregularly arranged stout long blunt setae mainly on outer and dorsal sides, and with long slender hair-like setae on outer side (not shown on Figs 3 and 14 ). Patella-tibial suture absent on fore and middle legs, present on hind leg ( Fig. 3 ) [see Iscini (3)]. Anterior (dorsal) side of fore, middle and hind tibia with a longitudinal row of stout elongate blunt setae ( Fig. 2a ; 14 ) [see Notophlebia (3)]; fore tibia with dense pointed bipectinate stout setae on inner side and very stout setae in distal part of posterior (ventral) side ( Fig. 2p ) [see Iscini (2)]; middle tibia with spine-like setae on inner side; hind tibia with spine-like setae on inner side and stout elongate blunt setae on outer side ( Fig. 3 ); middle and hind tibiae with numerous long slender hair-like setae on outer side (as in Fig. 54 ), fore tibia with a few such setae (not shown in Fig. 3 ). Tarsus of each leg with row of spine-like setae on inner side ( Fig. 20 ). Claw with 2–4 denticles on proximal portion, 5–8 denticles in anterior row and 1 larger denticle apically-posteriorly ( Fig. 35 ). Abdomen has characteristic form [see Iscini (5)], lacks posterolateral projections on terga I– VI , have short blunt projections on terga VII and VIII and long blunt projections on tergum IX [see Notophlebia (4)] ( Fig. 3 ). Posterior margins of abdominal terga with a regular row of pointed denticles alternated with setae ( Fig. 38 ); denticles become longer from tergum I to tergum IX. Posterior margins of abdominal sterna without denticles. All tergalii I– VI bilamellate, with both lamellae blunt; tergalius I a little shorter than tergalius II; tergalii from II to V gradually shorter; tergalius VI much smaller [see Notophlebia (5)] ( Figs 31–34 ); all tergalii narrower than in N. jobi . Protopenis of mature male larva much shorter than subimaginal penis, with gonopore located dorsally and lined with cuticle ( Fig. 37 ); before molt to subimago, most part of subimaginal penis projects proximad of cuticular larval protopenis. Female larva with small shallow incision on posterior margin of sternum IX (Fig. 21). Cerci longer than body, paracercus 1.5 times longer than cerci. FIGURES 1–3. Notophlebia ganeshi sp. n. 1, fore wing (holotype); 2, distal part of larval fore tibia, posterior view (stout setae on anterior side shown by interrupted lines); 3, male larva of penultimate instar, lateral view (long hair-like setae on legs not shown). Abbreviations: a , row of stout setae on anterior side; p , row of stout setae on posterior side. FIGURES 4–10. Notophlebia ganeshi sp. n. Larval mouthparts (at the same magnification, except 6). 4, hypopharynx and superlinguae; 5, labrum (setae of distal transverse row not shown, their bases shown as dots and levels of their apices shown by dotted line); 6, apex of maxilla, ventral view; 7, maxilla, ventral view (apical and median setae not shown, their apices shown by dotted lines); 8, labium (dorsal view at left, ventral view at right); 9, apex of labial palp, dorsal view; 10, left mandible (5, 10, holotype). FIGURES 11–20. Notophlebia ganeshi sp. n. 11, male imaginal mesonotum; 12, male subimaginal exuviae of right half of mesonotum. 13–19, legs at the same magnification (arrows show ontogenesis): 13, fore leg of male imago; 14, fore leg of male larva (long hair-like setae not shown); 15–17, fore, middle and hind legs of male subimago; 18, hind leg of male imago; 19, fore leg of female imago. 20, tarsus of larval hind leg before molt to subimago (larval cuticle shown as optic section by black; subimaginal cuticle shown by integral lines; imaginal claw developing under subimaginal cuticle shown by interrupted lines). 21, posterior margin of abdominal sternum IX of mature female larva; 22, the same, of female imago (12–18, holotype). Abbreviations: LPs , lateroparapsidal suture; MNS , mesonotal suture; MPs , medioparapsidal suture. FIGURES 23–25. Notophlebia ganeshi sp. n. (holotype). 23, genitals of male imago, ventral view (at left, brown pigmentation of sternum, styliger and gonostylus shown by dots; at right, brown pigmentation of penis shown by dots, muscles shown by interrupted lines); 24, penis lobe, dorsal view; 25, exuviae of subimaginal gonostylus. FIGURES 26–30. Notophlebia ganeshi sp. n. , genitals of male imago. 26, genitals at rest, ventral view; 27 penis apex, ventral side; 28, the same, dorsal side; 29, genitals in excited condition, with gonostylus turned ventrally and penes projected; 30, penis apex, apical-dorsal side (26–28, holotype). Subimago. CUTICULAR COLORATION: Cuticle nearly entirely colorless, thoracic sutures and legs partly diffusively washed with light brownish. Mesonotum without pigmented areas present in most Leptophlebiidae ; lateral portion of mesonotal suture does not serve as boundary of pigmented area, but has a form of longitudinal suture running between medioparapsidal and lateroparapsidal sutures ( Fig. 12 ). SHAPE AND TEXTURE : Fore femur widened proximally, middle and hind femora narrower ( Figs 15–17 ). On fore leg of male tibia and tarsus as short as on other legs, tarsus with swollen 1 st –4 th segments ( Fig. 15 ). Tarsi of all legs of male and female covered with microtrichiae, like other parts of legs and body. Caudalii 5–6 times shorter than body, cerci shorter than paracercus. FIGURES 31–45. 31–38, Notophlebia ganeshi sp. n. ; 39–45, Notophlebia jobi . 31–34 and 40–43, tergalii of I, II, V and VI pairs (ventral lamella turned back); 35, claw; 36 and 44, pronotum and mesonotum of last instar larva; 37 and 45, posterior margin of abdominal sternum IX of mature male larva and protopenis, dorsal view (gonoducts lined by cuticle shown by interrupted lines); 38 and 39, posterior margin of larval abdominal tergum VI (37, 38, holotype). Imago, male. Head ocher with brown. Upper eyes not elevated, brownish-orange. Thorax with light brown and ocher maculation; mesonotum with brown pigmented areas and ocher stripes between them [see Iscini (7)] ( Fig. 11 ). On fore wing ( Fig. 1 ) proximal part of subcostal field and partly proximal part of costal fields brown, remainder membrane colorless; longitudinal veins ocher; cross veins in anterior-proximal part of wing bordered with brown, in posterior and distal parts of wing colorless. Oblique cross veins of pterostigma either simple, or branched, or incomplete; either bordered by brown, or colorless, or invisible. Fore leg: femur brown with ocher; tibia white with brownish line on inner side, 2.5 times longer than femur; tarsus white, nearly 2 times longer than femur ( Fig. 13 ). Middle and hind legs: femur white with brown apex; tibia white with brown base; tarsus white ( Fig. 18 ). Middle and hind legs with patella-tibial suture well-developed; tarsus secondarily 5-segmented [see Iscini (10)] ( Fig. 20 ). Abdominal terga I–VII white, with blackish dots on lateral tracheal trunks; pale gray median maculae can be present on terga III–VII or on most posterior of them. Terga VIII–X dark brown with anterior margin light. Sterna I–VIII white. Sternum IX brown, with darker posterior-lateral areas and light posterior-median area ( Figs 23 , 26, 29 ). Styliger brown, gonostyli white with brown bases. Penial arms dark brown, external penis lobes ocher. Styliger short. Each penis lobe with straight pointed serrate apical projection [see Notophlebia (7)]; laterad of it, penis lobe forms convexity; mediad of it, penis lobe has a small sharp incision; gonopore locates on dorsal side of penis lobe, just dorsad of the serrate apical projection and the incision ( Figs 32 , 24 , 26–29 ). Caudalii white, without markings; cerci less than ½ of body length. Imago, female. Thorax as in male; mesonotum with the same brown pigmented areas separated by ocher stripes (as in Fig. 11 ). Wings as in male. Fore femur widened proximally, middle and hind femora narrower (as in subimago). Fore femur light brownish, with darker brown maculae near base and at apex ( Fig. 19 ). Middle and hind femora with larger longitudinal brown maculae. Tibiae of all legs with light inner side and brown outer side ( Fig. 19 ). Tarsi of all legs brownish. Abdominal terga I–X uniformly brown; sides lighter, with blackish dots on lateral tracheae. Abdominal sterna I–VIII lighter brownish, sternum IX brown. Posterior margin of sternum IX with small shallow incision (Fig. 22). Egg (extracted from mature female larva). Oval, 0.13 mm length; surface with irregular rugosity. Molts and transformations. Molt from subimago to imago occurs in both sexes (unlike Isca , whose female subimago does not molt). All 5 individuals reared in cages ( 3 males and 2 females ) molted from larva to subimago late at night (later than other mayflies, which do it at beginning of darkness) and molted from subimago to imago at the same night (earlier than most other mayflies in the same conditions), so that in morning I found only larval and subimaginal exuviae and dead imagos; female imagos had no eggs. I was unable to see subimagos alive and don't know duration of subimaginal stage, but can state that it is much shorter than in most mayflies. Unlike shed subimaginal skin of other mayflies (whose cuticle of wings is crumpled and can't be spread), on shed subimaginal skin of Notophlebia ganeshi cuticle of each wing has a form of soft sack and can be easily spread. Subimaginal legs retain size and proportions of larval legs, and male fore legs undergo great changes in course of molt from subimago to imago: tarsus of subimaginal fore leg has peculiar swollen shape ( Fig. 15 ); after molt to imago, tibia and tarsus become several times longer ( Fig. 13 ) [see Iscini (9)]. Subimaginal caudalii are 3–4 times shorter than larval caudalii; when larva molts to subimago, each subimaginal caudalius develops from proximal 1/7–1/5 of larval caudalius, while in most part of larval caudalius tissues degenerate; when subimago molts to imago, caudalius elongates 2–3 times thanks to its crumpling condition under subimaginal cuticle [see Iscini (13)]. Dimensions. Fore wing length (and approximated body length) 6–7 mm .