Inventory of prong-gilled mayflies ( Ephemeroptera : Leptophlebiidae ) of India with records of endemic taxa

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recognized species of mayflies.In understudied regions like Madagascar, taxonomic work on leptophlebiids is expected to yield upwards of 15 genera and 100 species new to science (Benstead et al. 2003).Leptophlebiid mayflies are considered to have undergone extensive adaptive radiation resulting in their present occupation of different aquatic microhabitats (Tsui & Peters 1975) and highly diverse gill morphologies.Previously, gill morphology has been linked to ecological factors (Peters et al. 1964;Riek 1973;Towns & Peters 1996).Leptophlebiid has maximum diversity in the Southern Hemisphere (Edmunds, 1972).It represents one of the major stem groups within the Ephemeroptera consisting of relatively ancestral and highly derived components (McCafferty & Edmunds 1979).Leptophlebiidae is a basal lineage and a sister group to a relatively derived clade that includes a pair of sister groups, Scapphodonta and Pannota (McCafferty & Wang 2000), in addition to a more basal lineage represented by the Behningiidae (McCafferty 2004).
Faunistic studies on Leptophlebiidae have progressed significantly in India.Sporadic taxonomic studies on Leptophlebiidae were conducted in Himalayan streams by Dubey in the early 1970s and two species viz., Atalophlebia chialhnia (Dubey, 1971) and Thraulodes marhienus (Dubey, 1970) were described from imagoes.The genus Atalophlebia Eaton, l881 is known only from Australia, and the genus Thraulodes Ulmer, 1920 is known only from the New World and hence the species are probably misplaced at the generic level (Hubbard & Peters 1978).Detailed studies using standardized generic delineations of Eastern Hemisphere Leptophlebiidae (Peters & Edmunds 1970) have resulted in the discovery of several new species belonging to ten genera, from India.Two genera viz., Choroterpes Eaton, l881 and Thraulus Eaton, 1881 are widely distributed, two genera viz., Gilliesia Peters andEdmunds, 1970 andIsca Gillies, 1951 have an Oriental distribution and six genera viz., Edmundsula Sivaramakrishnan, 1985, Indialis Peters & Edmunds, 1970, Klugephlebia Selvakumar, Subramanian & Sivaramakrishnan, 2016, Nathanella Demoulin, 1955, Notophlebia Peters & Edmunds, 1970and Petersula Sivaramakrishnan, 1984 are endemic to the Western Ghats and probably many of them are of a Gondwanan in origin.Presently, 26 species belonging to 12 genera under this family are reported in India (Sivaramakrishnan 2016;Selvakumar et al. 2016Selvakumar et al. , 2017a, b), b).The aim of the present study is to provide diagnostic characters, extension of distribution, endemic status and comprehensive knowledge of Leptophlebiidae species from India.

Material and Methods
Collections were made in streams and river basins of the all over India during 2009 to 2015.The present study investigated leptophlebiid mayfly in 48 sampling sites from 11 states and one union territory of India including earlier report (Table 1).Sampling area is mountainous with waterfalls and streams, and holds promise as harboring taxa.Collecting was conducted with an aquatic D-frame net.In streams, the substrate was kick-sampled, allowing the current to carry organic debris, including insects, into the net.Waterfalls were sampled by scouring the rock surfaces by hand, allowing the current to carry insects into the net.Along stream margins and in ponds, vegetation was swept with the aquatic D-frame net.All insects were preserved into 70% ethyl alcohol.Mayfly nymphs are particularly fragile because the gills and terminal filaments detach from the body very easily.Therefore, when possible, series of specimens were collected to maximize the likelihood of obtaining intact specimens and accurate determinations.To minimize damage to specimens, mayflies were collected in containers separate from other aquatic insects.Collected samples were brought to laboratory and were examined using a Leica M205A microscope and identified using published taxonomic literature and type specimens in the Zoological Survey of India (ZSI) and Southern Regional Centre (ZSI/SRC), Chennai.Identified specimens were deposited in ZSI, Kolkata, ZSI, SRC, Chennai and Department of Zoology (DZ), The Madura College (MC), Madurai.

Systematic account
Twenty six species belonging to 12 genera under two subfamilies of Leptophlebiidae from India are documented.All genera and species are presented alphabetically for convenience.This order should in no way be regarded indicating phylogeny.
Status: Endemic to the Himalaya.Remarks: Diagnostic characters are provided based on original description.Larva and imago are unknown.Further detailed study is required to assign this species to suitable genus.

Genus: Choroterpes Eaton, 1881
Type species: Choroterpes lusitanica Eaton, 1881 Diagnosis: The Choroterpes complex is recognised by the following characters: in the larvae (i) a pair of slender filaments of first abdominal gill different from gills 2-6; (ii) apex of glossae provided with broad spatulate setae and (iii) posterior row of setae on the labrum arises close to its middle (except in some Neochoroterpes).In the adults (i) in forewing, MP (Media Posterior) symmetrical fork while in MP 2 asymmetrical; (ii) cubital area broad with four (sometimes three) intercalaries; (iii) forceps in the male abruptly widened in its basal and (iv) penes as two simple lobes, very short to elongate and lacking spines or accessory lobes (Selvakumar et al. 2013).
Status: Wide distribution.

Remarks:
The genus encompasses the six subgenera viz., Choroterpes s.s.Diagnosis: This species can be differentiated from other species by the following combination of characters: In the larvae (i) abdominal gill 1 slender with dorsal and ventral portions;(ii) median projection of gills 2-7 plate-like and markedly larger and longer than laterals and (iii) labrum with three transverse rows of setae on dorsal surface, middle row without setae medially; anteromedian margin of labrum with a deep U-shaped ventral incision.In the adults (i) male genital penes each with a finger-like process on the top, acute costal projection of the hindwings and (ii) apex located approximately 2/3 distance from base (Tong & Dudgeon 2003).
Distribution: India (Karnataka, Kerala and Tamil Nadu) and China: Hong Kong.
Himachal Pradesh Status: Oriental distribution.Remarks: This species was originally described from Hong Kong based on reared larvae and adults in the laboratory (Tong & Dudgeon 2003).Larvae of this species from southern Western Ghats is extension of its distributional range down south to 8-11 0 north of equator by Selvakumar et al. (2015).

Subgenus: Dilatognathus Kluge, 2012
Diagnosis: This subgenus can be differentiated from other subgenera by the following combination of characters: In the larvae (i) abdominal gills 2-7; (ii) labrum widened with median incision; (iii) maxilla with inner-apical projection stretched or not stretched to a tusk-like process with ventro-apical flange, palp elongated and bears long filtering setae; (iv) labial palp elongated and bears long filtering setae which form regular longitudinal rows.Diagnosis: This species can be distinguished from other species by the following combination of characters.In the larvae: (i) labrum anteromedian emargination well broad and 'U′ shaped; (ii) mandibles with outer margin slightly angled a tuft of setae at angle; and (3) apical, median and basal part of femora with dark brown spots.In the imagoes: (i) hind wing dark brown marking on nodus; (ii) forceps and penes yellow and (iii) penis lobes short (Dinakaran et al. 2009) Diagnosis: This species can be identified by the following combination of characters: In the larvae (i) anteromedian emargination of labrum comparatively narrow and deeply cleft ('V′ shaped), lateral margin broadly acute; (ii) mandibles with outer margin comparatively deeply angled with a tuft of setae at angle; apical and median dark brown dark brown maculae at femora and (iii) dorsal and ventral lamellae platelike narrow and terminated in three slender subequal process, tracheae unbranched (Selvakumar et al. 2013).
Distribution: Karnataka, Kerala and Tamil Nadu.
Status: Endemic to the Western Ghats.
Remarks: Adult stage is unknown.

Choroterpes (Euthraulus) parvula (Gillies, 1951)
Material examined: 5480/H13, 1 male imago, 2 male subimagoes and 2 female subimagoes, 29.iii.(Gillies, 1951) can be distinguished from other species by the following combination of characters: In imago (i) foreceps base not divided, but extended to cover base of penes in a gently rounded curve; forceps stout, four segmented, jointed, basal broad and rounded, second long and curved, arising from the outer half of the basal, third segment incompletely divided from preceding, fourth segment thick and elongate; (ii) penes simple, without appendages, skittle-shaped, continuous at the base, separated apically; and (iii) ninth sternum of female with subanal plate well developed and with a very slight apical notch (Gillies 1951).
Distribution: Chhattisgarh and Madhya Pradesh.
Status: Endemic to India.Remarks: Larva is unknown.
Status: Endemic to the Western Ghats.Remarks: Adult is unknown.

Genus: Edmundsula Sivaramakrishnan, 1985
Type species: Edmundsula lotica Sivaramakrishnan, 1985 Diagnosis: This genus can be distinguished from other genera of Leptophlebiidae by the following combination characters: In adults: (i) fork of MP and fork of Icu 1 from CuA in the forewings occur about 1/3 of the Diagnosis: Edmundsula lotica can be distinguished by the following combination of characters: In adults: (i) fork of MP and fork of Icu 1 from CuA in the forewings occur about 1/3 of the distance from the base of wings to margin; both forks symmetrical; (ii) costal margin of hind wings possesses a blunt costal projection; apex of costal projection located less than ½ distance from base of wings; (iii) claws of a pair dissimilar, one apically hooked, the other obtuse, padlike; (iv) penes divided, tubular, broader at base and tapering towards apex; apex of each penis lobe curved ventrally and (v) ninth sternum of female cleft apically.In larvae: (i) gills present on abdominal segments 1-7; (ii) dorsal and ventral portions of lamellae of gill 1 slender and lanceolate with few tracheae; (iii) dorsal and ventral portion of lamellae of gills 2-7 lanceolate, long, and smoothly tapered near apex; (iv) segment 3 of labial palpi with a row of short heavy spines on inner dorsal margin; (v) denticles on claws progressively larger apically, apical denticle much larger and (vi) posterolateral spines occur on abdominal segments 4-9, spines progressively larger posteriorly, apices of spines on segments 8-9 sharp (Sivaramakrishnan 1985).
Distribution: Tamil Nadu and Karnataka.
Status: Endemic to the Western Ghats.
Remarks: Occurs in the some part of the Western Ghats.

Genus: Indialis Peters & Edmunds, 1970
Type species: Indialis badia Peters and Edmunds, 1970 Diagnosis: This genus can be differentiated from all other Leptophlebiid genera by the following combination of larval characters: In the larvae (i) abdominal gills 1-7 alike and slender with tracheae branched; (ii) a large tooth-like projection present on inner anterior margin of the maxillae; (iii) tarsal claws hooked, with a row of denticles that progressively larger apically and (iv) five denticles present on anteromedian emargination of labrum.In the adults: (i) more than two intercalaries in cubital area of forewings; (ii) vein of MP forked less than 1/2 of distance from base to margin and (iii) apex of each penis lobe bulbous and reduced tip, outer margin of apical half each penis lobe without a row of spinules (Peters & Edmunds 1970).
Status: Endemic to southern India.Remarks: Peters and Edmunds (1970) established Indialis for the species I. badia based on one male subimago and forty nine larvae collected by W.L. Peters and J.G. Peters in Kerala and Andhra Pradesh states of India.Sivaramakrishnan (1985a) described the female imago and egg structure of I. badia from the Tamiraparani River, southern Western Ghats.Peters (1975)
Status: Oriental distribution.Remarks: Gillies (1951) established the genus Isca from male and female imagoes of I. purpurea that collected in Hong Kong and India.Larva of Isca was described by Peters & Edmunds (1970) and two new species of Isca also described.Larvae of these two species were congeneric with those of I. purpurea.However, the adults are so morphologically distinct from I. purpurea and each other that two new subgenera viz., Minyphlebia Peters &Edmunds, 1970 andTanycola Peters &Edmunds, 1970 were established for these species by Peters & Edmunds (1970).
Subgenus Isca s.s.Gillies, 1951 Diagnosis: This subgenus can be differentiated from all other leptophlebiid genera by the following combination of characters: (i) vein MA forked a little more than 1/2 of distance from base to margin, fork asymmetrical; cilia present along posterior margin of wings; (ii) abdominal terga extend around onto venter of abdomen, this most marked on segment 7 but scarcely at all on segments 1 and 2; (iii) penes divided, tubular, broad, apex of each penis lobe curved inwardly and ventrally; (iv) ninth sternum of female apically cleft.Diagnosis: This species can be differentiated from all other leptophlebiid genera by the following combination of characters.In the imago (i) hind wings absent; (ii) cross veins absent in basal 1/2 of cell C in forewings; (iii) tarsal claws dissimilar; and (iv) segments 2 and 3 of male genital forceps short.In the larvae (i) abdominal segments extend around to venter of abdomen; (ii) dorsal and ventral portion of abdominal gills 2-6 slender and tracheae unbranched; gill 7 consists of 1 slender lamella and tracheae unbranched; (iii) claws apically hooked, and with a row of denticles; apical denticle larger; and (iv) small posterolateral spinas present on abdominal segments 7-9, and spines progressively larger posteriorly (Gillies 1951).

Isca
Distribution: India (Andhra Pradesh, Karnataka, Tamil Nadu and West Bengal) and Hong Kong.
Remarks: Larva and adult stages are known.

Genus: Klugephlebia Selvakumar, Subramanian & Sivaramakrishnan, 2016
Type species: Klugephlebia kodai Selvakumar, Subramanian & Sivaramakrishnan, 2016 Diagnosis: This genus can be differentiated from all other genera of Atalophlebiinae by the following combination of characters: In the imago: (i) vein MP forked slightly less than half of distance from base to margin, MP2 attached at base to vein MP1 by a crossvein; (ii) costal margin of hindwings with bluntly convex projection; apex of costal projection located less than half distance from base; (iii) claws of a pair dissimilar, one apically hooked, the other obtuse, padlike and (iv) segments 2 and 3 of forceps short, apex of segment 3 rounded, base of forceps broad, inner margin forming a smooth bend near middle of forceps; penis divided, tubular, broader at base and tapering towards apex.In the larvae: (i) gills present on abdominal segments 1-7; dorsal and ventral portions of lamellae of gill 1 slender and lanceolate with branched tracheae, dorsal and ventral portions of lamellae of gills 2-7 wider and lanceolate, long and suddenly tapering at apex; (ii) fore and mid femora with a regular row of long, thin setae on outer margin; denticles on claws progressively larger apically; (iii) length of the labrum more than half of the width, lateral lobes rounded, anteriomedian emargination deeply cleft, apparently with two denticles; proximal transverse setal row laterally curved distally; (iv) maxillary palp short, with long setae on third segment and third segment of labial palp with 5-6 thick, spine-like setae on dorsal surface, inner and outer margins with short, thin setae (Selvakumar et al. 2016).
Status: Endemic to the Western Ghats.

Remarks:
The genus was established for the single species, Klugephlebia kodai Selvakumar, Subramanian & Sivaramakrishnan, 2016.Diagnosis: This species can be differentiated from all other genera of Atalophlebiinae by the following combination of characters: In the imago: (i) vein MP forked slightly less than half of distance from base to margin, MP2 attached at base to vein MP1 by a crossvein; (ii) costal margin of hindwings with bluntly convex projection; apex of costal projection located less than half distance from base; (iii) claws of a pair dissimilar, one apically hooked, the other obtuse, padlike and (iv) segments 2 and 3 of forceps short, apex of segment 3 rounded, base of forceps broad, inner margin forming a smooth bend near middle of forceps; penis divided, tubular, broader at base and tapering towards apex.In the larvae: (i) gills present on abdominal segments 1-7; dorsal and ventral portions of lamellae of gill 1 slender and lanceolate with branched tracheae, dorsal and ventral portions of lamellae of gills 2-7 wider and lanceolate, long and suddenly tapering at apex; (ii) fore and mid femora with a regular row of long, thin setae on outer margin; denticles on claws progressively larger apically; (iii) length of the labrum more than half of the width, lateral lobes rounded, anteriomedian emargination deeply cleft, apparently with 2 denticles; proximal transverse setal row laterally curved distally; (iv) maxillary palp short, with long setae on third segment and third segment of labial palp with 5-6 thick, spine-like setae on dorsal surface, inner and outer margins with short, thin setae (Selvakumar et al., 2016).
Status: Endemic to the Western Ghats.Remarks: The species was described both larva and adult.

Genus Nathanella Demoulin, 1955
Type species: Nathanella indica Demoulin, 1955 Diagnosis: This genus can be distinguished from all other genera by the following combination of characters: In the larvae (i) abdominal gills present on segments 1-7, and dorsal and ventral portions of lamellae leaf-like and apicaly terminated three projections, median longer than laterals; (ii) outer margin of mandibles smoothly curved basally and straight apically with a row of hair in the apical half; (iii) anteromedian margin of labrum straight with 5 broad-based denticles and (iv) lateral margins of the head capsule broadly expanded.In the adults (i) hind wings absent; (ii) vein MP 2 of fore wings attached at base to vein MP 1 and CuA by a cross vein, and attachment of vein MP 2 to MP 1 greater than1/4 to 1/3 distance from base to margin; (iii) penes divided, straight with apex expanded dorsally and (iv) claw similar (Peters & Edmunds 1970).
Distribution: Karnataka, Kerala and Tamil Nadu.
Status: Endemic to the Western Ghats.Remarks: Demoulin (1955) established Nathanella for a distinctive species, N. indica known only from male imagoes collected in southern India.Sivaramakrishnan et al. (1996) described female imago and larvae of N. indica and male and female imagoes and larvae of N. saraswathiae from Kerala border, near Bodi Mettu.
Diagnosis: This species can be distinguished from N. saraswathiae by the following combination characters: In the larvae: (i) median projection of abdominal gills broad and approximately twice length of laterals; (ii) tracheation in gills uniformly distributed; and (iii) distal, irregular light brown maculae on femora of legs.In the adults: (i) membrane of fore wing golden brown, cross veins in cells C and Sc narrowly clouded with brown; (ii) abdominal terga 3-7 of male brown except irregularly pale apically; and (iii) dorsal margin of styliger plate of male broadly convex (Demoulin, 1955).
Status: Endemic to the Western Ghats.Remarks: Male imago was described by Demoulin (1955).Female imago and larva were described by Sivaramakrishnan et al. (1996).Diagnosis: This species be identified by the following combination of characters: In the larvae (i) median projection of abdominal gills narrow and approximately 1-1/2 length of laterals; (ii) main trunk of tracheae of gills forked near distal half of lamellae and (iii) medial 120m, coll.F. Schmid.

Nathanella saraswathiae
Diagnosis: This species can be distinguished by the following combination of characters: In the adult (i) cilia occur on posterior margin of fore wings; (ii) membrane of anal area of fore wings enlarged posteriorly; and (iii) penes of male genitalia tubular, straight, and pointed (Peters & Edmunds 1970).
Distribution: Known only from the type locality Kunjankhuzi, Tamil Nadu.
Status: Endemic to the Western Ghats.
Remarks: Diagnostic characters are provided based on the original description.Larva is unknown.Diagnosis: This species can be identified by following combination of characters: In the larvae (i) gills not narrower; (ii) third segment of maxillary palp with very long stout pointed setae directed apically and forming nearly regular rows; (iii) third segment of labial palp with long pointed setae on outer side, dorsal side also with long setae.In the adults (i) apically each penis lobe bears a slender pointed serrate projection curved (Sivaramakrishnan & Peters 1984).
Status: Endemic to the Western Ghats.

Remarks:
The larva of this species has a highly specialized mouth apparatus of the Dilatognathus-type.This type of mouth apparatus has evolved independently in several non-related leptophlebiid taxa.

Genus: Petersula Sivaramakrishnan, 1984
Type species: Petersula courtallensis Sivaramakrishnan, 1984 Diagnosis: This genus can be distinguished from other genera by the following combination of characters: In the larvae: (i) labrum expanded and angled laterally; (ii) anterior margin of lingua of hypopharynx deeply cleft; apex of submedian lobes of lingua possesses a rack like process; (iii) outer margin of basal ½ of mandibles smoothly curved, while apical ½ almost straight; a row of hairs extends from mid outer margin almost to base of incisors; (iv) abdominal gills occur on segments 1-7 and are plate-like with margins unevenly fringed with broad filamentous processes and (v) posterolateral spines occur on abdominal segments 3-9 and progressively larger posteriorly.In the adults: (i) vein MP 2 of fore wings attached at base to vein MP 1 more than 1/3 of the distance from base to margin; (ii) costal margin of hind wings convex or with a rounded costal projection; (iii) each penis lobe with ventromedially directed spine-like projection near apex; (iv) claws of a pair alike, apically hooked with an opposing hook and (v) 9 th sternum of female shallowly cleft apically (Sivaramakrishnan 1984).Distribution: Karnataka, Kerala and Tamil Nadu.
Status: Endemic to the Western Ghats.

Remarks:
The genus Petersula was established for P. courtrallensis from the southern Western Ghats by Sivaramakrishnan 1984.A second species, P. nathani described based on adult from the Anamalai hills of southern Western Ghats (Sivaramakrishnan & Hubbard 1984).The genus is widespread in the Western Ghats.Diagnosis: This species can be identified by following combination characters: In the larvae (i) labrum expanded and angled laterally; (ii) anterior margin of lingua of hypopharynx deeply cleft; apex of submedian lobes of lingua possesses a racklike process; (iii) outer margin of basal ½ of mandibles smoothly curved, apical ½ almost straight; a row of hairs extended from mid outer margin almost to base of incisors; (iv) abdominal gills occur on segments 1-7 and plate-like with margins unevenly fringed with broad filamentous processes and (v) posterolateral spines occur on abdominal segments 3-9 and progressively larger posteriorly.In adults (i) vein MP 2 of fore wings attached at base to vein MP 1 more than 1/3 of the distance from base to margin; (ii) costal margin of hind wings convex or with a rounded costal projection; (iii) each penis lobe ventromedially directed spine-like projection near apex; (iv) claws of a pair alike, apically hooked with an opposing hook and (v) 9 th sternum of female shallowly cleft apically (Sivaramakrishnan 1984).
Status: Endemic to the Western Ghats.
Remarks: This species is wide distribution and abundant in the Western Ghats.
Diagnosis: This species can be distinguished from the only other species in the genus, P. courtallensis by the following characters: In imago: (i) terga 1-7 translucent yellowish brown and washed with brown; terga 8-10 opaque brown; (ii) paired longitudinal, submedian lines present on terga 3-5; (iii) vein MP 2 of fore wings attached at base only to vein MP 1 with a crossvein and (iv) length of spinelike projection arising from near apex of each penis lobe nearly 1/3 length of penis (Sivaramakrishnan & Hubbard 1984).
Status: Endemic to the Western Ghats.
Remarks: Diagnostic characters are provided based on the original description.Larva is unknown.

Genus: Thraulodes Ulmer, 1920
Remarks: The genus Thraulodes Ulmer, 1920 is known only from the New World and hence the species, Thraulodes marhieus Dubey, 1970 probably misplaced at the generic level (Hubbard & Peters 1978).
Status: Endemic to the Himalaya.Remarks: Diagnostic characters are provided based on the original description.Larva is unknown.

Genus: Thraulus Eaton, 1881
Type species: Thraulus bellus Eaton, 1881 Diagnosis: The genus can be differentiated from all other genera of the Leptophlebiidae by the following combination of characters.In the imago, (i) fork of vein MP of fore wings is closer to base of wings than fork of vein Rs; (ii) 2 intercalaries occur in cubital area of fore wings; (iii) penes tubular, divided and simple; and (iv) costal projection of hind wings acute and well developed, except for the costal projection of T. bellus which is more rounded.In the nymph, (i) dorsal and ventral portions of abdominal gills 2-7 ovate with fringed margins; (ii) dorsal and ventral portions of abdominal gills 1 slender, lanceolate, or ovate with fringed margins, or dorsal portion slender, lanceolate and ventral portion ovate with fringed margins; (iii) lateral tips of superlingua of hypopharynx emarginated; and (iv) tarsal claws hooked and narrow and with a row of denticles that are progressively larger apically (Peters & Edmunds 1970).
Distribution: Oriental, Afrotropical and Palearctic.Status: Wide distribution.Remarks: Presently, this genus encompasses 15 valid species, of which three are from Palearctic, three from Afrotropical and nine from Oriental realms (Barber-James et al. 2013).In India, the genus Thraulus Eaton, 1881 has 3 species viz., T. gopalani Grant & Sivaramakrishnan, 1985 described from both imago and larvae, T. mudumalaiensis Soman, 1991 described only from larvae and T. semicastaneus (Gillies, 1951) described only from imago.Diagnosis: This species can be distinguished from all other described species of Thraulus by the following combination of characters: In the larva (i) labrum with a rectangular mesal emargination anteriorly; (ii) inner row of the dorsal setae located just anterior to middle of the labrum; (iii) outer margin of the mandibles lacks a tuft of setae at the base of the incisors; (iv) denticles on the claws decrease in size apically; and (v) abdominal gills 1 with a dorsal lanceolate portion and a ventral fimbriate lamellar portion and abdominal gills on segments 2-7 with dorsal and ventral fimbriate lamellar portions.In the adult (i) upper portion of male eyes separated; (ii) forewings with a narrow dark brown band between costal brace and vein A 2 ; (iii) bullae of veins Sc and R 2 have a small dark brown macula; (iv) basal ½ of hind wings brown and apex blundly rounded; and (v) each penis with single longitudinal row of spine-like setae on dorsal surface (Grant & Sivaramakrishnan 1985).
Status: Endemic to the Western Ghats.
Remarks: Larva and adult are known.
Diagnosis: Thraulus mudumalaiensis can be distinguished from all other known species by the following combination of characters: In larvae: (i) claws with five minute denticles in apical set and 10 larger denticles in basal row in which the size increases medially, then decreases apically; (ii) labrum without denticles in the emargination, two rows of setae on dorsal side and an irregular intermittent setae ventrally in between two dorsal rows, a cluster of setae of either of the anterolateral side of its venter; (iii) coastal area of forewing pads hyaline, without longitudinal brown streak; (iv) mandibles with lateral sides smoothly rounded with some setae on mid region; (5) segment 2 of maxillary palp almost equal to the length of segment 1, segment 3, 0.74 the length of segment 2 and (6) segment 2 of labial palp 0.7 the length of segment 1, segment 3 a little longer than segment 2 (Soman, 1991).
Distribution: Known only from Nilgri, Tamil Nadu.Status: Endemic to the Western Ghats.

Thraulus semicastaneus
Diagnosis: This species can be identified by the following combination of characters: in the imago (i) penes simple, narrow and divided but closely appressed; (ii) forewing translucent colourless, main veins amber, cross veins fine and numerous, two cubital intercalaries only, stigma containing 9-11 simple, sinuous vein-lets; and (iii) hindwing short, somewhat triangular, with tall costal spur and sharply upturned subcosta (Gillies 1951).Distribution: Known only from the Mutha river, Pune (Maharashtra).
Status: Endemic to the Western Ghats.
Remarks: Diagnostic characters are provided based on the original description.Larva is unknown.

Subfamily: Leptophlebiinae
Diagnosis: Leptophlebiinae can be differentiated from Atalophlebiinae by a suite of mouthpart characters and an elongate and deeply cleft ninth sternum in adult females (Peters & Edmunds 1970;Peters 1980;Kluge 1994).

Genus Gilliesia Peters & Edmunds, 1970
Type species: Gilliesia hindustanica (Gillies) Diagnosis: This genus can be differentiated from other genera of the Leptophlebiidae by the following combination of characters: In imago (i) hind wings present and well developed; (ii) vein MP 2 of the fore wings with independent of vein MP 1 ; (iii) female without ovipositor or egg guide; and (iv) 9th sternum of the female deeply cleft apically.In larva (i) posterolateral expansions of on abdominal segments 9 only well developed; (ii) gills long, slender and slightly forked at 2/5 basally; (iii) glossae narrow taped, with dense thickened-long hairs on ventral surface; and (iv) length of maxilla palpi segment three more than 1.6 times length of segment 2; apical-blunted, with numerous setae (Peters & Edmunds 1970).
Distribution: China, India and Thailand.

: Choroterpes s. s. Eaton, 1881 Diagnosis:
Gillies, 1951cosmopolitan in  distribution, Euthraulus Barnard, 1932restricted to the Old World, Neochoroterpes Allen, 1974 restricted to the New World and CryptopenellaGillies, 1951,  Dilatognathus Kluge, 2012 and Monochoroterpes Kluge,  2012restricted to the Oriental region.This subgenus Choroterpes can be differentiated from other subgenera by the following combination of characters: In the larvae (i) a broad, terminal lobe on the lamina of gills 2-6 and indistinguishable characters in the adults between subgenera.

Prong-gilled mayflies of India Selvakumar et al. 12395 distance
from the base of wings to margin; both forks symmetrical; (ii) costal margin of hind wings possesses a blunt costal projection; apex of costal projection located less than ½ distance from base of wings; (iii) claws of a pair dissimilar, one apically hooked, the other obtuse, pad-like; (iv) penes divided, tubular, broader at base and tapering towards apex; apex of each penis lobe curved ventrally and (v) ninth sternum of female cleft apically.
Sivaramakrishnan, 1985.u and Karnataka.Status: Endemic to the Western Ghats.Remarks: The genus was established for the single species, Edmundsula loticaSivaramakrishnan, 1985.
described I. rossi from a male imago from Kerala state.