taxonID	type	description	language	source
1D7287CDFFD0D670FF090C9F0B19FA05.taxon	description	(Fig. 1)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD0D670FF090C9F0B19FA05.taxon	diagnosis	Diagnosis. This species can be distinguished from other Teloganopsis species by the following combination of characters: larva: (i) dorsal surface of body often with light, longitudinal, yellowish band formed by spots on head, pronotum, mesonotum and on abdominal terga V and VIII, but sometimes with only small spots on thorax and abdomen being presented, and therefore no obvious longitudinal band (Fig. 1 A, B; fig. 1 in Tong & Dudgeon 2000); (ii) labrum, mandibles and labium without long hair-like setae (figs 5, 6, 9 – 11 in Tong & Dudgeon 2000); (iii) mandible canine not elongated (figs 9, 10 in Tong & Dudgeon 2000); (iv) inner margin of maxilla with a continuous row of setae (Fig. 1 C; fig. 11 in Tong & Dudgeon 2000); (v) maxillary palp absent (Fig. 1 C; fig. 11 in Tong & Dudgeon 2000); (vi) thorax without tubercles (fig. 1 in Tong & Dudgeon 2000); (vii) inner margins of fore tibia and fore tarsus without long hair-like setae; (viii) paired tergal projections absent; (ix) posterolateral projections of abdominal segments IV – IX strongly marked (Fig. 1 A); (x) dorsal surface of forefemur with transversal row of long spatulate setae and with the same kind setae along outer margin of the femur (Fig. 1 D; fig. 8 in Tong & Dudgeon 2000); (xi) caudal filaments light yellowish-brown with dark brown medial and apical bands (Fig. 1 B; fig. 12 in Tong & Dudgeon 2000). Male imago: (xii) penis short, projection of penis small, not visible in ventral view (fig. 2 in Tong & Dudgeon 2000; fig. 8 D in Zhang et al. 2017).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD0D670FF090C9F0B19FA05.taxon	distribution	Distribution. China (Xu et al. 1984; Tong & Dudgeon 2000; Zhou et al. 2006), India and India-China border region (new data), and Thailand (Jacobus & McCafferty 2008, new data).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD0D670FF090C9F0B19FA05.taxon	discussion	Remarks. This species originally was described based on adults from China (Xu et al. 1984). Zhou et al. (2006) provided revisions to this species, including the recognition of new junior synonyms. The larva of this species (as Serratella albostriata Tong & Dudgeon, 2000, junior synonym) was described and illustrated sufficiently by Tong & Dudgeon (2000); the male imago was illustrated by Xu et al. (1984) and Zhang et al. (2017: figs 8 b, d). Our new data represent the first report of this species from India and the India-China border region, in particular. Ubero-Pascal & Sartori (2009) revised the genus, and additional contributions have been made by Jacobus (2009), Sartori (2014), Zhang et al. (2017) and Gorovaya (2019). Other Indomalayan species include Teloganopsis brocha (Kang & Yang, 1995), T. gracilis (Tshernova, 1952), T. media Ulmer, 1939, T. oriens Jacobus & McCafferty, 2006, T. puigae Ubero-Pascal & Sartori, 2009, and T. setosa Zhou (in Zhang et al.), 2017. Several potential new species remain undescribed.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD1D679FF090A520E01FDDC.taxon	description	(Figs 2 – 5; Jacobus & McCafferty 2008: fig. 14)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD1D679FF090A520E01FDDC.taxon	materials_examined	Type material. Holotype: larva (slide # 623, mounted with Canada balsam), THAILAND, Chiang Mai Province, Chom Thong District, stream — main source of the Klang Phat River, 18.577542 ° N, 98.527056 ° E, h ~ 1370 m a. s. l., 18. xi. 2009, Palatov D. M. & Chertoprud M. V. leg. — IN Thai 10 Sersp [NMNH NASU]. Other material: INDIA: 1 larva, Arunachal Pradesh, Lower Subansiri District, Tale Valley, 27.537201 ° N, 93.959883 ° E, h ~ 2370 m a. s. l., 14. iv. 2015, Coll. K. A. Subramanian — Reg. No. 5603 / H 13 [ZSI]. NEPAL: 1 larva, Nawakot & Sindu Districts, 1 / 2 mi north of Gulbhanjyang (on lower trail), 18. ix. 1968, Coll. C Wiens [PERC] (previously reported as Serratella uenoi (Allen & Edmunds, 1963) by Jacobus & McCafferty 2008).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD1D679FF090A520E01FDDC.taxon	description	Description. Larva: Body length 5.0 – 5.3 mm; caudal filaments 4.5 – 4.8 mm. Body pale brown (Fig. 2 A). Head: With pair of small suboccipital tubercles and pair of distinct, blunt, occipital protuberances (Fig. 2 B) bearing short, stout setae with divergent margins and feathered apices (Fig. 2 C, D). The same scattered stout setae cover head surface, they also presented on compound eyes, but they are smaller. Genae moderately developed. Mouthparts: Labrum (Fig. 3 C) densely covered with long, hair-like setae; anterior margin with numerous feathered and hair-like setae; anteromedian emargination shallow. Median part of mandibles with numerous, long, hair-like setae; basal part of lateral margin with smaller number of middle-sized and short, hair-like setae. Right mandible (Fig. 3 A): outer incisor trifurcate, inner incisor bifurcate; prostheca consisting of dorsal process, smaller than on left mandible, and bunch of long and short hair-like setae; row of 10 – 15 long, stout, hair-like setae under mola; bunch of short, hair-like setae above mola. Left mandible (Fig. 3 B): outer incisor trifurcate, inner denticle small; inner incisor with two central denticles and one small lateral denticle; prostheca consisting of process and bunch of relatively long and short hair-like setae; inner surface with distinct denticles near mola. Maxilla with two dentisetae (Fig. 3 F), their inner margins serrate. Apex of maxilla with group of long, thin and stout, hair-like setae; apical part of inner margin with row of long, stout, hair-like setae; base of galea-lacinia with group of 4 – 6 long, stout, pointed, not bifurcated or bifurcated, hair-like setae. Maxillary palp 3 - segmented (Fig. 3 E), short; segment III elongate, narrowed from middle, rounded apically; segment I somewhat broader than segments II and III; apex of segment III with several fine setae. Superlinguae of hypopharynx with long, stout, hair-like setae on apices, dorsal and ventral surfaces with fine setae, short and hair-like setae, and stout and hair-like setae; apex of lingua convex, with hair-like setae on dorsal and ventral surfaces (Fig. 3 D). Labial palp 3 - segmented (Fig. 3 G); segments I and II subequal in length; surfaces, inner and outer margins of segment I and II covered with long, thin, hair-like setae and less numerous long, stout, hair-like setae. Segment III distinctly elongated (length / width ratio in last larval instar = 2.19 – 2.45). Glossae rounded; apices of paraglossae and glossae covered with long, stout, hair-like setae. Thorax: Pronotum without anterolateral and posterolateral projections (Fig. 2 E); with one pair distinct submedian tubercles and three pairs small indistinct tubercles (Fig. 2 F). Mesothorax with two pairs of indistinct protuberances and ridges, and one distinct protuberance between wingpads. Protuberances and ridges of prothorax and mesothorax, veins of wingpads covered with short, stout setae with divergent margins and feathered apices. Femora of legs moderately flattened (length / width ratio in last larval instar: forefemur 2.04 – 2.05; middle femur 2.39 – 2.49; hind femur 2.38 – 2.57) (Fig. 4 A – C). Femora longer than tibiae, and tibiae longer than tarsi. Dorsal surfaces of all femora covered mainly with short, feathered, usually bifurcate, stout setae (Fig. 4 E – H) (most numerous on middle and hind femora), and with scattered short or middle-sized hair-like setae; also, irregular rows of middle-sized, hair-like setae situated along inner margins. Distal tips (distal margins and adjacent areas of dorsal surfaces) of all femora with groups of short, feathered, sometimes bifurcate, setae. Dorsal surface of forefemur with irregular rows of middle-sized and long, hair-like setae along outer and inner margins; with group of 4 – 5 mainly long, rounded or pointed, stout setae; most of these stout setae located near outer margin (Fig. 4 A, D). Outer margin of forefemur with few different-sized hair-like setae and few short, feathered, sometimes bifurcate, setae and two chalazae bearing long, pointed or rounded, stout setae. Inner margin of forefemur with short, hair-like setae only. Outer margins of fore tibia and tarsus with a few thin, long hair-like setae. Inner margin of fore tibia with several short stout setae along margin and group of several elongated setae near distal end, some with serration of one margin. Inner margin of fore tarsus with middle-sized and long, pointed stout setae; their number increases towards claw. Outer margins of mid- and hindfemora with long hair-like setae (most numerous and forming regular row in basal part), few short, feathered, sometimes bifurcate, setae and row of 6 – 9 long, pointed or rounded apically, stout setae; some chalazae forming serration of margins (Fig. 4 B, C). Inner margins of mid- and hindfemora with no stout setae or chalazae. Setation of middle and hind tarsi as those on fore leg. Dorsal surface of middle and hind tibiae of row of few stout setae continue on inner margin of tibiae; setae bluntly pointed of rounded apically; on hind tibia setae more numerous and longer. Inner margin of tibiae also with group of several elongated stout setae (some with one serrated margin) near distal end. Outer margin of middle and hind tibiae with few hair-like setae only; hind tibia additionally bears few long stout setae along margin. Tarsal claw with 5 – 7 denticles, distal one largest, and up to 5 subapical setae (Fig. 4 I, J). Abdomen. Pairs of projections present on terga III – IX, with those on terga IV – IX more developed; largest on tergum VIII (Fig. 5 A – C). All projections with spatulate, stout setae; most apical, stout setae grouped in bunches (Fig. 5 A, B). Dorsal surfaces of terga IV – IX with areas of short, stout setae above projections. Lateral surfaces of paired projections of tergum VIII and adjacent part of posterior margin with greatly elongated, apically rounded, stout setae (Fig. 5 A – C). Posterior margin of tergum IX (excluding area between projections) with several spatulate, stout setae with rounded apices. Distinct posterolateral projections on segments IV – IX; lateral margins covered with spatulate, stout setae. Sterna VIII – IX and lateral areas of sterna IV – VII covered with short, stout setae. Gills (Fig. 5 D – G). Gill III with elongate posterolateral angle; with well-defined, brown, trilobed pattern; somewhat truncate; and without medial transverse band of weakened membrane (Fig. 5 D). Ventral lamellae of gills III – VI bifurcate and multifoliate; medial cleft of gills VI ventral lamella deep. Caudal filaments subequal in length (Fig. 2 A). Segments with rows of elongated, rounded apically, stout setae on posterior margins alternate with segments bears rows of long, stout, hair-like setae on posterior margins; all these setae shorter than corresponding segment (Fig. 5 H). Adult: Unknown Egg. Chorion smooth, without reticulations (Jacobus & McCafferty 2008: fig. 14; Nepal specimen herein).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD1D679FF090A520E01FDDC.taxon	diagnosis	Diagnosis. The species can be distinguished from larvae of other Serratella Edmunds, 1959 species by the following combination of characters: (i) head with pair of small suboccipital tubercles and pair of distinct, blunt, suboccipital protuberances (Fig. 2 A, B); (ii) pronotum without anterolateral and posterolateral projections (Fig. 2 E), with one pair distinct submedian tubercles and three pairs small indistinct tubercles (Fig. 2 F); (iii) mesothorax with two pairs of indistinct protuberances and ridges, and one distinct protuberance between wingpads; (iv) two pairs of head protuberances, protuberances and ridges of prothorax and mesothorax, veins of wingpads covered with short, stout setae with divergent margins and feathered apices (as in Figs 2 C, D, 4 F, G); (v) maxilla with short 3 - segmented palp (Fig. 3 E); third palpal segment elongated; (vi) tarsal claw with 5 – 7 denticles, distal one largest, and up to 5 subapical setae (Fig. 4 I, J); (vii) pairs of projections present on terga III – IX, with those on terga IV – IX more developed; largest on tergum VIII (Fig. 5 A – C); (viii) all paired projections of terga with spatulate, stout setae; most apical, stout setae grouped in bunches (Fig. 5 A, B); (ix) lateral surfaces of the paired projections of tergum VIII and the adjacent part of the posterior margin (excluding area between projections) with greatly elongated, apically rounded, stout setae (Fig. 5 A – C). Despite this new species being assigned to Serratella, it should be noted that similar tergum VIII setation is present in Quatica paradinasi (Gonzalez del Tanago & Garcia de Jalon, 1981), but the setae are much shorter (fig. 2 in Studemann & Tomka 1987).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD1D679FF090A520E01FDDC.taxon	etymology	Etymology. This species is named in honor of Dr. Dmitry M. Palatov, friend of the first author and specialist in aquatic invertebrates of the Palearctic and Indomalayan realms, who collected this species in Thailand.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD1D679FF090A520E01FDDC.taxon	distribution	Distribution. Thailand, India-China border region, and Nepal. Habitats. In Thailand, the new species was collected from a stream that is a main source of the Klang Phat River. The stream is situated in forest, has a high current velocity and rapids, and has sandy and stony bottom (Fig. 20 B). In India, the species was collected from a first order stream in Rhododendron and Bamboo forest. The stream has a sandy bottom and rapids in some sections (Fig. 20 C). The Nepal specimen is covered with sandy silt, suggesting a similar habitat.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD1D679FF090A520E01FDDC.taxon	discussion	Remarks. Initially Serratella uenoi (Allen & Edmunds, 1963) was described as a representative of the subgenus Drunella Needham, 1905 (then part of the genus Ephemerella Walsh 1862) by Allen & Edmunds 1963, based only on description and illustrations of “ Ephemerella sp. ” by Ueno (1955), as the whereabouts of Ueno’s specimen was then — and remains — unknown. Later, a second species, Ephemerella (Acerella) undatella Allen, 1971, was described based on the same specimen (or rather description and illustrations). Subsequently, the name was recognized as an objective junior synonym (Allen 1973). Ephemerella (Drunella) uenoi (Allen & Edmunds, 1963) was transferred to other genera (Allen 1986; Paclt 1994) before being placed most recently in Serratella Edmunds, 1959 based on phylogenetic analysis of morphological data (Jacobus & McCafferty 2008: fig. 98). The characters used to analyze the species’ relationships were scored from a single specimen from Nepal and from historical literature (Jacobus & McCafferty 2008). During the course of this study, we discovered that the specimen from Nepal differed from the species described by Ueno (1955), especially in regards to the morphology of the maxilla (Ueno 1955: figs 6, 6 b); the two also differ considerably in size, with Ueno’s species being larger, even though the Nepal specimen is a mature female with black wingpads. Intraspecific variation in body size and maxillary palp development has been assumed for many ephemerellid species (e. g., Jacobus et al. 2004); however, in this case, the discovery of additional material reveals that the characters in question show little variation between individuals. Thus, we specifically reject the intraspecific variation hypotheses previously implied for S. uenoi and therefore no longer consider the Nepal specimen to be conspecific with Ueno’s species. Therefore, the operational taxonomic unit (OTU) labeled “ uenoi ” by Jacobus & McCafferty (2008) should be considered an erroneous amalgamation, and the species hypothesis that it represents is rejected. In light of this, we restrict the name Serratella uenoi (Allen & Edmunds, 1963) to the specimen described by Ueno (1955). Very clear illustrations of the species show some differences from other species of the genus Serratella, especially: the apex of the maxilla, the length ratio and shape of segments of the labial palp; the number of head tubercles; the shape of projections on terga (posterolateral and paired); setation of all femora; and presence of anterolateral projections (medially notched) on mesonotum. Modalities of some characters of S. uenoi are unusual for Hyrtanellini, and more typical of some Ephemerellini, viz. representatives of the genera Notacanthella Jacobus & McCafferty, 2008, Spinorea Jacobus & McCafferty, 2008, Ephacerella Paclt, 1994, Adoranexa Jacobus & McCafferty, 2008, and Cincticostella Allen, 1971. As part of the tribe Ephemerellini, each of the latter five genera have a ventral lamella of gill VI that lacks a deep medial cleft. The original illustration of S. uenoi shows a distinct cleft on the ventral lamella of gill VI (fig. 18 in Ueno 1955), which excludes it from Ephemerellini. Although we find it reasonable to question the generic placement of S. uenoi, we leave it in Serratella until fresh material from the type locality, as precisely indicated by Ueno (1955), can be examined in detail. Serratella fusongensis (Su & You, 1988) (north-east of China) and Serratella longipennis (Zhou, Gui & Su, 1997) (China, east-central mainland) are the only species of Serratella from East and Southeast Asia unknown in the larval stage, and based on biogeography, we consider them unlikely to be conspecific with our new species, which is unknown as alates. Serratella palatovi sp. nov. is the third representative of the genus known from the Indomalayan realm; the two others are S. uenoi, which has a questionable generic position, and S. brevicauda Jacobus, Zhou & McCafferty, 2009, a species whose generic placement was provisional (Jacobus et al. 2009). Thus, it is clear that more data, especially for the male adults, are needed for these species and the genus in the region.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD8D67EFF090EBA0F7CFED8.taxon	description	(Figs 6 – 9)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD8D67EFF090EBA0F7CFED8.taxon	materials_examined	Material examined: INDIA: 4 larvae (two in slides ## 688, 689, with Euparal) Uttarakhand State, Chamoli district, Alaknanda River (700 m above mouth of the Pindar River), 30.267728 N, 79.220931 E, h ~ 800 m a. s. l., 5. ii. 2011, Palatov D. M. and Chertoprud M. V. leg. — IN Indi 6 Hyrsp 1 / 1 - 4 [NMNH NASU].	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD8D67EFF090EBA0F7CFED8.taxon	diagnosis	Diagnosis. Hyrtanella grandipennis can be distinguished from other Indomalayan Hyrtanellini by the following combination of characters in the larval stage: (i) entire body covered with numerous rounded excrescences (Fig. 6 B – D, 8 B, C, E, 9 B); (ii) head with pair of small, rounded occipital tubercles; (iii) maxilla with or without (even in one specimen) very short, single-segmented palp bears hair-like seta (Fig. 7 E – G); (iv) pro- and mesothorax with variable developed blunt dorsal protuberances; (v) mesothorax with notched anterolateral projections bears few rounded apically stout setae (Fig. 6 B, C); (vi) claw with no stout, preapical denticle that is larger than the rest, (vii) claw usually with distal palisade of denticles (if absent, probably worn down); (viii) pairs of flattened, rounded apically, widely spaced submedian projections present on terga III (VI, V) – VIII, with those on terga VI – VIII more developed; largest on tergum VIII (Fig. 9 A, B, E, F); tergum IX without submedian projections or with wide, short and rounded apically ones; (ix) abdomen with distinct and deep chamber for gills, its inner margin presented with posterolateral projections of terga (Fig. 9 A, E, F); (x) segment IX elongated, with large and pointed posterolateral projections reached posterior margin of tergum X (Fig. 9 A); (xi) posterior margin of sternum VIII with pair of triangular projections that visible even dorsally (Fig. 9 A, C – F); (xii) gills III – VI distinctly elongated, especially gill III (Fig. 9 G). Male imago has (xiii) broad forceps and penis; (xiv) genital forceps segment III elongate, with its length being more than 2 x its width; (xv) genital forceps segment II not swollen or bowed, but with a distal bend; (xvi) penes lobes with dorsolateral projections. Imagoes and subimagoes of H. grandipennis also differ from H. pascalae Jacobus & Sartori, 2004 (figs 15 – 17 in Jacobus & Sartori 2004) from Borneo based on the venation and shape of hind wings. In contrast to H. pascalae, in fore wing of H. grandipennis one of the iCuA divided and attached to CuA; stigmatic area subdivided by a secondary vein into upper and lower rows of several cellules between C and Sc; CuP and AA of fore wing subparallel, not converge toward margin; hind wing with small costal process in middle of margin.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD8D67EFF090EBA0F7CFED8.taxon	distribution	Distribution. Southern mainland China (Zhou et al. 2000; Jacobus et al. 2004), Vietnam, Thailand (Jacobus et al. 2004) and India (new data).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFD8D67EFF090EBA0F7CFED8.taxon	discussion	Remarks. Hyrtanella grandipennis was described originally in Torleya Lestage, 1917 based on the male imago stage (Zhou et al. 2000). Later, the larval stage of the species was described based on specimens from China, Vietnam and Thailand (Jacobus et al. 2004), and considerable morphological variation within and between these populations was noted. Jacobus & McCafferty (2008) placed this species in Hyrtanella Allen & Edmunds, 1976 based on detailed study and analysis of imaginal and larval characters. Xu et al. (2020) reported this species in Torleya. In their study of mitogenomes, H. grandipennis and T. tumiforceps Zhou & Su, 1997 (junior syn. of Torleya nepalica (Allen & Edmunds, 1963 )) were placed in the same clade, but other species of Hyrtanella and Torleya were not included in the investigation, so no new conclusions can be made about relationships and generic affinities of the various species in Torleya and Hyrtanella beyond those hypothesized by Jacobus & McCafferty (2008: fig. 99) and Ogden et al. (2009: figs 1,2). The male imagoes of H. pascalae and H. christineae Allen & Edmunds, 1976 are not described yet, so adequate comparisons and diagnoses are not yet practical. Among our specimens of H. grandipennis, we note that a small maxillary palp is variably present or absent (even in one specimen) among the single known population in India. Jacobus et al. (2004) previously hypothesized this feature was variable, even though Vietnam specimens consistently had the palp, and China and Thailand specimens lacked such a palp. In our opinion, taxonomic status of three close forms of H. grandipennis and their generic position need proper investigation with integrative approach, with molecular, morphological and possibly ecological study. Unfortunately, the Indian specimens at hand were collected in 4 % Formaldehyde, and later placed in Ethanol, ruling out the chance of obtaining good DNA sequence data; moreover, only the Chinese form has the male imago described.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFDDD663FF090CD70ED4FBD4.taxon	description	(Fig. 10)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFDDD663FF090CD70ED4FBD4.taxon	materials_examined	Material examined. INDIA: 14 larvae, Arunachal Pradesh, Lower Subansiri District, Tale Valley, 27.537201 ° N, 93.959883 ° E, h ~ 2370 m a. s. l., 14. iv. 2015, Coll. K. A. Subramanian — Reg. No. 5608 / H 13 [ZSI]; 1 larva, Arunachal Pradesh, Lower Subansiri District, Ranga River, 27.396404 ° N, 93.757378 ° E, h ~ 625 m a. s. l., 21. iv. 2015, Colls. K. A. Subramanian & Bikramjit Sinha — Reg. No. 5347 / H 13 [ZSI]; 13 larvae, Arunachal Pradesh, Papum Pare District, Dibru River, 27.147655 ° N, 93.74908 ° E, h ~ 128 m a. s. l., 22. iv. 2015, Colls. K. A. Subramanian & Bikramjit Sinha — Reg. No. 5609 / H 13 [ZSI]; 1 larva, West Bengal, Darjeeling (Sikkim border), Rishikhola, Rishi River, 27.169677 ° N, 88.635109 ° E, h ~ 554 m a. s. l., 23. iii. 2013, Coll. Srimoyee Basu — Reg. No. 5349 / H 13 [ZSI]. NEPAL: 3 larvae, Bagmati Zone, East Rapti River, 27.571631, 84.668756, h — 230 m a. s. l., 27. i. 2014, Chertoprud M. V. leg. — IN Nepa 16 Torsp 1 [NMNH NASU].	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFDDD663FF090CD70ED4FBD4.taxon	diagnosis	Diagnosis. This species can be distinguished from other Torleya species by the following combination of characters: Larva: (i) dorsal surface of body covered by long hair-like setae (Fig. 10 A; fig. 1 in Allen & Edmunds 1963); (ii) maxilla with rudimentary palp (fig. 13 in Allen & Edmunds 1963); (iii) anteromedian emargination very shallow, almost absent (fig. 14 in Allen & Edmunds 1963); (iv) labium highly reduced (fig. 17 in Allen & Edmunds 1963); (v) abdominal segment VIII with a unique shape and setation (Fig. 10 A, C; fig. 1 in Allen & Edmunds 1963; fig. 1 in Jacobus et al. 2007); (vi) posterolateral projections of segment IX extremely elongated (Fig. 10 A, C; fig. 1 in Allen & Edmunds 1963); (vii) tarsal claw with distinct distal palisade of denticles (Fig. 10 B; fig. 15 in Allen & Edmunds 1963); (viii) lateral margins of abdominal segments IV – VIII with short stout setae; stout setae on segment VIII pass over acute posterior projection (fig. 1 in Allen & Edmunds 1963); (ix) paired submedial projections on abdominal terga are absent (Fig. 10 C; fig. 1 in Allen & Edmunds 1963); (x) gills III – VI with outer posterolateral protuberances, gill III not covering following gills entirely (figs 1, 16 in Allen & Edmunds 1963). Male imago: (xi) penis short, lobes apically rounded (fig. 2 in Jacobus et al. 2007); (xii) dorsal projection of penis small, rounded, with broad apical cleft (figs 2, 4 in Jacobus et al. 2007) (xiii) genital forceps segment II robust, with relatively straight lateral profile, not expanded apically and segment III subovoid (fig. 2 in Jacobus et al. 2007); (xiv) abdomen with light general coloration and purple shading, lacking a prominent, middorsal, longitudinal stripe (figs 3, 4 in Jacobus et al. 2007).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFDDD663FF090CD70ED4FBD4.taxon	distribution	Distribution. India (Allen 1980, Jacobus & McCafferty 2003, Jacobus et al. 2007, new data), India-China border region (new data), Malaysia (Jacobus et al. 2007), Nepal (Allen & Edmunds 1963, new data), Thailand (Wang & Sites 1999; Jacobus et al. 2007) and Vietnam (Jacobus et al. 2007).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFDDD663FF090CD70ED4FBD4.taxon	discussion	Remarks. The larva of this species was adequately described and illustrated from Nepal by Allen & Edmunds (1963) and from Thailand by Wang & Sites (1999) (as Crinitella permkami Wang & Sites, 1999, junior synonym). Larvae of this species already were reported from Kashmir, and the adult description and details of species variability were provided by Jacobus et al. (2007). Following Jacobus & McCafferty (2008), we consider Crinitella Allen & Edmunds, 1963 as a junior synonym of Torleya Lestage, 1917. Ogden et al. (2009: figs 1, 2) could not confirm monophyly of Crinitella + Torleya, however, based on their partial sampling of these genus groups. The other Indomalayan species of Torleya include T. dibruensis sp. nov., T. lacuna (Jacobus, McCafferty & Sites, 2007), T. longforceps (Gui, Zhou & Su, 1999), T. lutosa Kang & Yang, 1995, T. naga Jacobus & McCafferty (in Jacobus et al.), 2004, T. nepalica (Allen & Edmunds, 1963) and T. simbalbarensis sp. nov.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC2D660FF09086A0852F8AE.taxon	description	(Figs 11 – 13)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC2D660FF09086A0852F8AE.taxon	materials_examined	Type material. Holotype: larva, INDIA, Arunachal Pradesh, Papum Pare District, Dibru River, 27.147655 ° N, 93.74908 ° E, h ~ 128 m a. s. l., 22. iv. 2015, Colls. K. A. Subramanian & Bikramjit Sinha — Reg. No. 5606 / H 13 [ZSI]. Paratypes: 1 larva, same data as holotype — Reg. No. 5607 / H 13 [ZSI]; 2 larvae, INDIA, Arunachal Pradesh, Lower Subansiri District, Paniya stream, 27.81791 ° N, 94.09502 ° E, h ~ 993 m a. s. l., 14. vi. 2017, Colls. K. A. Subramanian & M. Vasanth — Reg. No. SRC-I / E 510 [ZSI].	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC2D660FF09086A0852F8AE.taxon	description	Description. Late larval instar: Length of body 3.5 – 3.8 mm; cercus and median caudal filament subequal, each 1.5 – 1.8 mm. General color light brown, with variable brown shading and markings; front of head, outer and inner margins of legs and lateral margins of abdominal terga covered with long, hair-like setae (Figs 11 A, 12 A, B). Head: Head with one pair of distinct, blunt occipital tubercles bear small short stout setae. The same scattered stout setae cover head. Color light brown with variable brown markings; frons pale; two dark brown, rounded spots situated between occipital protuberances and compound eyes. Areas around paired ocelli yellowish. Antennal scape and pedicel light brown; flagellar segments yellowish-brown, with fine, hair-like setae at apex of each segment; hair-like setae nearly one-third length of respective segment. Clypeus and genae densely covered with long hair-like setae (Fig. 12 A). Labrum (Fig. 11 B) brown; anterior notch shallow and wide; anterolateral angles weakly expressed, rounded; dorsal surface with dense, transverse row of cilia-like setae. Superlinguae of hypopharynx with row of setae on anterior margin; lingua with very sparse and tiny setae on dorsal and ventral surfaces, apex convex (Fig. 11 C). Mandible external margin with scattered hair-like setae basally and pair of long cilia-like setae medially (Fig. 11 D, E). Maxilla (Fig. 11 F) with few distal hair-like setae; palp 3 - segmented, with visible articulation; spinous processes short. Labium with cilia-like setae on ventral surface; segment III of maxillary palp rounded, about one-third length of segment II (Fig. 11 G). Thorax: Pronotum without distinct protuberances. Mesonotum with several small, protuberances. Mesonotum with transverse brown shading medially (Fig. 11 A); tip of fore wingpads pale to white. Legs pale, distinctly flattened, especially fore femur (Fig. 13 B – D). Dorsal surface of forefemur with irregular transverse row of long, pointed, stout setae proximally; outer and inner margins with rows of long hair-like setae and long, pointed, stout setae (Fig. 13 A, B); dorsal surface of middle and hind femora smoky brown, with a few short stout setae; outer and inner margins with long, pointed, stout setae and long, hair-like setae (Fig. 13 C, D). Tibiae of all legs distinctly wider than tarsi. Each claw with 2 – 3 medial denticles, palisade of four to five long subdistal denticles, and subapical seta (Fig. 13 E). Abdomen: Terga with dark longitudinal medial line and lateral shading (Figs 11 A, 12 B). Dorsal lamella of gill III rounded, extending to middle of tergum VIII (Fig. 12 B), apically rounded, with fine distal setae, brown, trilobed pattern indistinct (Fig. 13 F); gill III operculate; ventral lamellae of gills III – VI bifurcate and multifoliate (Fig. 13 G). Terga I – V and VIII – X mostly pale; terga VI – VII usually brown (Fig. 12 B). Terga V – IX with spatulate, stout setae on posterior margins (setae progressively shorter on segments VI – VIII); terga VII and VIII with hair-like and spatulate, stout setae; terga IV – VII and IX with paired posterior protuberances, bears small stout setae with rounded apices; terga VIII and X without paired posterior protuberances; posterior protuberances very small on tergum IV and IX; largest on terga V – VII, usually bearing 4 – 6 short spatulate stout setae; Lateral parts of posterior margin of tergum VIII with row of spatulate, stout setae. Segments IV – IX with posterolateral projections. Segment IX distinctly narrower than segment VIII (Fig. 12 B). Sterna pale, with few short, hair-like setae. Caudal filaments pale to white, with broad, dark brown band medially; tips dark brown; apex of each segment with spatulate, stout setae and fine hair-like setae. Adults. Unknown.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC2D660FF09086A0852F8AE.taxon	diagnosis	Diagnosis. Larvae of Torleya dibruensis sp. nov. can be distinguished from other representatives of the genus by the following combination of characters: (i) head with one pair of blunt occipital tubercles; (ii) head, legs and abdomen with areas densely covered with long, hair-like setae (Figs 11 A, 12 A, B, 13 A – D); (iii) claw with 2 – 3 medial denticles, palisade of four to five long subdistal denticles on the inner margin, and subapical seta (Fig. 13 E); (iv) terga V – VII with paired blunt protuberances, usually bearing four to six short spatulate, stout setae (Fig. 11 A); (v) gills rounded, gill III entirely cover following gills (Figs 11 A, 12 B, 13 F, G); (vi) foreleg with irregular transverse row of long pointed, stout setae at proximal part of dorsal surface; same setae presented on outer and inner margins (Fig. 13 A, B); (vii) maxillary palp present, short (Fig. 11 F); (viii) labial palp segment III about one-third length of segment II (Fig. 11 G).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC2D660FF09086A0852F8AE.taxon	etymology	Etymology. This species is named after the type locality, Dibru River, Arunachal Pradesh.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC2D660FF09086A0852F8AE.taxon	distribution	Distribution. India-China border region. Habitat. Larvae of the species were collected in rivers and large streams with cobble and sandy bottom and numerous boulders (Fig. 20 D).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC2D660FF09086A0852F8AE.taxon	discussion	Remarks. Torleya longforceps from Fujian, in far eastern China (Gui et al. 1999), is the only species of Torleya unknown in the larval stage, and we consider it unlikely (based on biogeography) to be conspecific with this new species, which is unknown as alates.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC6D666FF090C9F0E07FEA0.taxon	description	(Fig. 14)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC6D666FF090C9F0E07FEA0.taxon	materials_examined	Material examined. Holotype: larva, THAILAND, Kanchanaburi, stream, Amphur Thong Pha Phum, Heuy Ka Yaeng at Ban Padsadoo Klang, 14 ° 33 ’ N, 98 ° 34 ’ E, 296 m altitude, 9 / IV / 2003, L- 457, Sites, AV, Prommi, Setaphan [UMRM].	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC6D666FF090C9F0E07FEA0.taxon	diagnosis	Diagnosis. The larva of this species can be distinguished from congeners by the following combination of characters: (i) one pair of blunt occipital protuberances present (Fig. 14 B; fig. 5 in Jacobus et al. 2007); (ii) head, thorax and legs densely covered with long hair-like setae (Fig. 14 A, B; fig. 5 in Jacobus et al. 2007); (iii) pronotum with four dorsal protuberances (fig. 5 in Jacobus et al. 2007); (iv) mesonotum with transverse brown shading medially; tip of fore wingpads pale to white, terga with dark longitudinal medial line and lateral shading; (v) terga III – IX, and sometimes tergum II, with paired posterior protuberances; on terga IV – VIII protuberances largest, usually with up to two spatulate, stout setae (fig. 5 in Jacobus et al. 2007); (vi) maxillary palp absent; (vii) posterolateral projections of terga IV – VIII produced dorsolaterally, forming the abdominal gill chamber; (viii) gills distinctly elongated, gill III entirely cover following gills (fig. 5 in Jacobus et al. 2007); (ix) tarsal claws with one or two basal denticles, two or three medial denticles, two to four long subdistal denticles on the inner margin, and subdistal setae; (x) foreleg with submedial row of long spatulate setae.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC6D666FF090C9F0E07FEA0.taxon	distribution	Distribution. Thailand and Vietnam (Jacobus et al. 2007). The previous record from India (Jacobus et al. 2007) is now questionable (see Remarks, immediately below).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC6D666FF090C9F0E07FEA0.taxon	discussion	Remarks. This species was described by Jacobus et al. (2007) based on larvae and subimagoes from Thailand and Vietnam. These authors also reported male and female subimagoes from Tamil Nadu (India). The imago stages of T. lacuna are unknown. Previous reports of T. lacuna from India should be reviewed after subimagoes and imagoes of T. dibruensis sp. nov. and T. simbalbarensis sp. nov. have been discovered.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC7D665FF090DBE0B1EF831.taxon	description	(Figs 15 – 17)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC7D665FF090DBE0B1EF831.taxon	materials_examined	Material examined: Holotype, larva, NEPAL, Pelung, 5850 ’ elev., 17 - iv- 1957, EI Coher — IN PERC 0064041 [PERC]. Other material: NEPAL: 2 larvae, Bagmati zone, Shivapuri Nagarjun National Park, small stream (3 km Northwards of the village Mulcharka), 27.806097 ° N, 85.435522 ° E, h ~ 1800 m a. s. l., 28. ii. 2007, Chertoprud M. V. leg. — IN Nepa 3 Tornep [NMNH NASU]; 2 larvae, Bagmati Zone, East Rapti River, 27.571631, 84.668756, h — 230 m a. s. l., 27. i. 2014, Chertoprud M. V. leg. — IN Nepa 16 Torsp 2 [NMNH NASU]; 2 larvae, Bagmati Zone, Dhading District, stream in valley of Malekhu River, 27.742472, 84.807750, h — 564 m a. s. l., 23. i. 2014, Chertoprud M. V. leg. — IN Indi 17 Torsp 1 [NMNH NASU]. INDIA: one larva, Karnataka, Someshwara Wildlife Sanctuary, Authi, 13.57431 ° N, 75.12207 ° E, h ~ 639 m a. s. l., 16. xi. 2015, Coll. S. Ramya Roopa — Reg. No. 5351 / H 13 [ZSI]; 1 larva, Meghalaya, East Garo Hills, Upper Rongbu Village, 25.91615 ° N, 90.83157 ° E, h ~ 101 m a. s. l., 26. vi. 2016, Coll. E. Eyarin Jehamalar — Reg. No. 5340 / H 13 [ZSI]; 7 larvae, INDIA, Arunachal Pradesh, Lower Subansiri District, Talley Valley, 27.537201 ° N, 93.959883 ° E, h ~ 2370 m a. s. l., 14. iv. 2015, Coll. K. A. Subramanian — Reg. No. 5610 / H 13 [ZSI]; one larva, Tamil Nadu, Theni, Kurangani stream, 10.083611 ° N, 77.248611 ° E, h ~ 1744 m a. s. l., 21. i. 2010, Coll. C. Selvakumar — Reg. No. 5611 / H 13 [ZSI]; one larva, Tamil Nadu, Kodaikanal, Gundar river, 10.226692 ° N, 77.451117 ° E, h ~ 2323 m a. s. l.; 31. iii. 2012, Coll. C. Selvakumar — Reg. No. 5351 / H 13 [ZSI]; 3 larvae, Tamil Nadu, Tirunelveli District, Papanasam, 8.710278 ° N, 77.367500 ° E, h ~ 108 m a. s. l., 18. vii. 2009, Coll. C. Selvakumar — Reg. No. MCDZ / E- 52 [MCDZ]; 1 larva, Karnataka, Nandini hole, 13.389722 ° N, 77.179722 ° E, h ~ 640 m a. s. l., 3. v. 2013, Coll. C. Selvakumar — Reg. No. MCDZ / E- 53 [MCDZ]; one larva, Meghalaya, East Jaintia Hills, Umpung village stream, 25.30767 ° N, 92.63658 ° E, h ~ 1010 m a. s. l., 12. iii. 2016, Coll. E. Eyarin Jehamalar — Reg. No. 5339 / H 13 [ZSI]; one larva, Nagaland, Intanki National Park, Intanki River, 25.39883 ° N, 93.30686 ° E, h ~ 181 m a. s. l., 23. iii. 2016, Coll. C. Selvakumar — Reg. No. 5535 / H 13 [ZSI]; one larva, Nagaland, Intanki National Park, Intanki River, 25.39048 ° N, 93.31913 ° E, h ~ 206 m a. s. l., 24. iii. 2016, Coll. C. Selvakumar — Reg. No. 5542 / H 13 [ZSI]; one larva (in slide number 633) Uttarakhand, Almora district, 2 - nd left tributary of the river Ramganga-left (in Dwarahat forest, 10.1 km North-Eastwards of the Chaukhutia town), 29.925608 ° N, 79.445983 ° E, h ~ 1200 m a. s. l., 2. ii. 2011, Palatov D. M. leg. — IN Indi 1 Tornep [NMNH NASU].	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC7D665FF090DBE0B1EF831.taxon	diagnosis	Diagnosis. This species can be distinguished from other Torleya species by the following combination of characters. Larva: (i) maxilla without palp (Fig. 16 G; fig. 26 in Allen & Edmunds 1963); (ii) head with one pair of distinct protuberances each bearing a few short, stout setae (Fig. 16 B, C; fig. 21 in Allen & Edmunds 1963); (iii) both prothorax and mesothorax with two pairs of distinct protuberances, all of them and dorsal surface of thorax bearing short and elongated, stout setae (Fig. 16 D); (iv) dorsal surface of forefemur with transverse row of long, pointed, stout setae; outer margin of forefemur with one chalaza bearing a long, pointed, stout seta (Fig. 17 C; fig. 20 in Allen & Edmunds 1963); (v) outer margin of middle and hind femora with two or three chalazae each bearing a long, pointed, stout seta, and long, hair-like setae; dorsal surface of middle and hind femora with scattered spatulate, stout setae and long, hair-like setae (Fig. 17 D – F); (vi) claw with 6 – 8 denticles, two denticles arranged somewhat separately (subdistal), distinct distal palisade of denticles absent (Fig. 17 G, H; fig. 22 in Allen & Edmunds 1963); (vii) posterior margins of terga III – IX with submedian protuberances bearing spatulate, stout setae; terga V – VII with the largest protuberances (Figs 15 A, B, 16 A, 17 A, B); (viii) gills distinctly elongated, but not covering following gills entirely (Figs 15 B, C, 16 I; fig. 23 in Allen & Edmunds 1963); (ix) distinct abdominal gill chamber absent (Figs 15 B, 17 A; fig. 23 in Allen & Edmunds 1963). Male imago: (x) penis elongated, lobes apically rounded (fig. 5 in Zhou & Su 1997); (xi) dorsal projection of penis protrude beyond the lateral margins of penis shaft (fig. 5 in Zhou & Su 1997) (xii) genital forceps segment II elongated, expanded apically; segment III subovoid (fig. 5 in Zhou & Su 1997).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC7D665FF090DBE0B1EF831.taxon	distribution	Distribution. Indomalayan [southern mainland China (Zhou & Su 1997) including Hong Kong (Tong & Dudgeon 2000), Taiwan (Kang & Yang 1995), India (Jacobus et al. 2004, new data), India-China border region (new data), Japan [Okinawa] (Ishiwata 2018), Malaysia (Jacobus et al. 2004), Nepal (Allen & Edmunds 1963; Jacobus et al. 2004, new data), Pakistan (Jacobus et al. 2004), Thailand (Jacobus et al. 2004) and Vietnam (Jacobus et al. 2004)].	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFC7D665FF090DBE0B1EF831.taxon	discussion	Remarks. The larvae of this species were described from Nepal by Allen & Edmunds (1963). Adults were described as Torleya tumiforceps from China by Zhou & Su (1997). Discussions of larval and adult variability were provided by Jacobus et al. (2004). The complete mitogenome of this species (as Torleya tumiforceps) was recently reported by Xu et al. (2020). This species was reported earlier from Karnataka (India) by Jacobus et al. (2004). Jacobus et al. (2004) reported a wide range of morphological variation for this species, some of which may prove to correlate with geography; it is possible this species represents a cryptic species complex, requiring evaluation of its junior synonyms.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFCAD66AFF090AB10BB3F818.taxon	description	(Figs 18, 19)	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFCAD66AFF090AB10BB3F818.taxon	materials_examined	Type material. Holotype: larva, INDIA, Himachal Pradesh, Sirmour District, Simbalbara Wildlife Sanctuary, stream, 30.44 ° N, 77.53 ° E, 16. xi. 2012, Coll. K. A. Subramanian — Reg. No. 5602 / H 13 [ZSI].	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFCAD66AFF090AB10BB3F818.taxon	description	Description. Larva: body length 5.2 mm; caudal filaments 2.4 mm. Body color light brown (Figs 18 A, 19 A). Head: Vertex with pair of well-developed and elongated protuberances. No ocellar protuberances. Inner margin of antennal base with short projections. Frontal shelf not expanded. Genae slightly produced. Long hair-like setae present along anterior margin of frontal shelf and sparsely present below compound eye. Clypeus with anterior row of long hair-like setae. Labrum width nearly 2 × length (Fig. 18 B), with shallow, very broad notch anteriorly; dorsal face with scattered long, hair-like setae on lateral margin and with several rows of brush-like setae anteriorly; ventral face with simple or fimbriate setae on lateral margins and dense, transverse row of 7 – 9 long, stout setae. Hypopharynx (Fig. 18 C) with superlinguae size subequal to linguae size; lingua with row of short setae on anterior margin; superlingua with row of long, thin setae on margin, with setae decreasing in length anteriorly. Mandibles with lateral setae relatively long (Fig. 18 D, E). Maxilla without palp (Fig. 18 G); apex of galea-lacinia relatively sharp with two incisors, and with few long, stout setae on apex; with two dentisetae and double row of four stout setae. Labium with glossae longer than wide (Fig. 18 F), longer than paraglossae; palp segment III elongate, with length / width ratio 2.13 – 2.15. Thorax: Prothorax width subequal to head width, with no anterolateral projections and three pairs of distinct dorsal tubercles; ventral spines absent. Mesonotum with two pairs of distinct, elongated, tubercles between fore wing pads; fore wing pads base darkly pigmented. Dorsal surface of thorax without any bifid or starlike setae. Coxae with slight dorsolateral projections; projections with hair-like setae. All femora slightly flattened (Fig. 19 B – C). Dorsal surface of forefemur with incomplete, narrow, transverse band of long, pointed or bluntly pointed, apically stout setae (Fig. 19 E); outer margin with middle-sized and long, hair-like setae and single chalaza bearing long, stout seta medially; inner margin with middle-sized and long, hair-like setae only; anterior margin with no dorsal projections (Fig. 19 B). Dorsal surface of foretibia with two long, pointed, stout setae; outer margins of foretibia and tarsus with different-sized hair-like setae, most long; inner margins of fore tibia and fore tarsus with short, hair-like setae and row of short and middle-sized spine-like setae; distal end of inner margin of foretibia with group of stout setae. Foretibial projection short. Outer margins of middle and hind femora with several long, pointed, stout setae and different-sized hair-like setae, most long (Fig. 19 C, D). Dorsal surfaces of middle and hind tibiae with several long, pointed, stout setae. Outer margins of middle and hind tibiae and tarsi with hair-like setae, most long. Inner margins of middle and hind tibiae and tarsi with different-sized spine-like setae. Tarsal claws of all legs each with about 8 denticles and several subapical setae, without distal palisade of denticles (Fig. 19 F). Abdomen: Gill III brown, semi-operculate, without medial transverse band of weakened membrane (Fig. 19 G); gills IV – VII translucent. Dorsal lamellae of gills IV – VI with small projection on posterior margin. Gills III – VI ventral lamellae bifid, with dorsal and ventral lobules; lobules tips acute. Gills VII narrow, with medial point of attachment to tergum. Terga I and II with long hair-like setae on posterior margins. Terga III – IX with pairs of welldeveloped, bluntly pointed submedian projections (Fig. 19 A) bear few short and blunt stout setae; those of terga V – VIII most developed; on tergum VIII largest and directed most laterally; tergum IX with pair of short, blunt, median spines and weak, oblique ridges. Tergum X without paired submedian projections. Segments VI – VIII with lateral margins strongly upturned, forming outer portion of prominent gill chamber. Pairs of projections progressively larger posteriorly up to segment VIII and covered with short, blunt stout setae. Abdominal sterna flattened, with no maculation and no long setae. Caudal filaments pale, with dense whorls of long, stout, hair-like setae. Adults: Unknown.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFCAD66AFF090AB10BB3F818.taxon	diagnosis	Diagnosis. Torleya simbalbarensis sp. nov. is easily distinguished from other Torleya species by the following combination of characters: (i) head with two pairs of suboccipital and occipital protuberances, second pair of protuberances well-developed and elongated; (ii) maxilla without palp (Fig. 18 G); (iii) tarsal claws of all legs each with about 8 denticles and several subapical setae, without distal palisade of denticles (Fig. 19 F); (iv) terga III – IX with pairs of well-developed, bluntly pointed submedian projections (Fig. 19 A); those of terga V – VIII most developed; on tergum VIII largest and directed most laterally; tergum IX with pair of short, blunt, median spines and weak; (v) setation of femora; (vi) shape of gill III.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFCAD66AFF090AB10BB3F818.taxon	etymology	Etymology. This species is named after the type locality, Simbalbara Wildlife Sanctuary, Himachal Pradesh.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFCAD66AFF090AB10BB3F818.taxon	distribution	Distribution. India. Habitat. Fast flowing stream, with sand and cobble bottom, in Sal (Shorea robusta Roth) (Malvales: Dipterocarpaceae) forest (Fig. 20 A).	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
1D7287CDFFCAD66AFF090AB10BB3F818.taxon	discussion	Remarks. Notably, Torleya simbalbarensis sp. nov. has no distinct distal palisade of denticles on tarsal claw, which usually is present on other Asian species of the genus; however, it is possible the claws might be worn (Jacobus et al. 2004). Torleya longforceps from Fujian, in far eastern China (Gui et al. 1999), is the only species of Torleya unknown in the larval stage, and we consider it unlikely (based on biogeography) to be conspecific with this new species, which is unknown as alates.	en	Martynov, Alexander V., Selvakumar, C., Subramanian, K. A., Sivaramakrishnan, K. G., Vasanth, M., Sinha, Bikramjit, Jacobus, Luke M. (2021): Overview of Indian Hyrtanellini (Ephemeroptera: Ephemerellidae), with new species and records from related regions. Zootaxa 4975 (3): 451-482, DOI: https://doi.org/10.11646/zootaxa.4975.3.2
