Sphaeronemoura siveci, Murányi, Dávid & Li, Weihai, 2013

Murányi, Dávid & Li, Weihai, 2013, Two new species of stoneflies (Plecoptera: Nemouridae) from Northeastern India, with a checklist of the family in the Indian Subcontinent, Zootaxa 3694 (2), pp. 167-177 : 169-176

Sphaeronemoura siveci sp. n.

( Figs. 5–8 )

Diagnosis. Male tergite VIII lacks a protruding lobe. Male tergite X bears paired tergal projections near the apex of epiproct. Male paraproct: inner lobe longer than the projection of median lobe, both similarly sclerotized. Male epiproct: flagellum double curved horizontally, the acute, claw-like apex bent downwards.

Type material. Holotype male: INDIA: Meghalaya State, West Garo Hills District, Nokrek National Park, 1100 ± 150 m a.s.l., N 25°29.6’ E 90°19.5’, 9- 17.05.1996, leg. E. Jendek, O. Šauša (WNHM; glued on paper sheet, abdomen cleared in KOH and pinned beneath the specimen in microvial with glycerine).

Description. Medium sized species, macropterous. Forewing length: holotype 8.0 mm. Head uniformly dark brown; palpi lighter, scape and pedicel brown, rest of antenna lacking. Pronotum dark brown, trapezoid, wider than long; rugosities not conspicuous. Legs uniformly dark brown; wings dark brown with some metallic hue, venation black. Abdomen pale but with transverse, sclerotized dark bands on terga, terminal segments dark brown. Pilosity generally short but with long hairs on terminal segments.

Male terminalia ( Figs. 5–8 ): Hypoproct trapezoid with square corners, basal part as long as wide but apically expanding, then apical projection tapering with upcurved, blunt tip; vesicle of the holotype broken but probably typical of the genus. Paraproct inner lobe finger-like, longer than projection of the median lobe, apex blunt. Median lobe with short, rounded projection of the same brown colour as the longer inner lobe; membranous part with conspicuous, short hairs. Outer lobe darker than the other two lobes, forming a slender strip curved from the cercal base to the median lobe, apex forked. Cerci simple, sausage-like, moderately sclerotized and slightly bent inwards. Tergite VIII bears no lobe but with distinct antecosta, slightly sclerotized posteriorly and with a posterior row of strong hairs. Tergite IX well sclerotized, antecosta medially thickened before a wide, posteriomedial membranous field that is delimited and partly covered with strong hairs. Tergite X with antecosta interrupted in the medial third of the segment width, medial third of the segment completely membranous but covered with very short hairs between the anterior edge and the tergal projections; well sclerotized lateral thirds bear long and strong hairs, longest ones delimiting the membranous area. Tergal projections situated next to the apex of epiproct, rather strong and paired, anterior one acute while posterior one blunt. Epiproct as usual for the genus, tongue-shaped in dorsal view, with double curved, acute flagellum. Dorsal sclerite forked after a strong, trapezoid base; branches short and terminating before the epiproct’s apex, running laterally and slightly bent downwards, delimiting a large, membranous dorsal area. Ventral sclerite trapezoid, apically slightly dilated, bears no teeth nor setae. Membranous apex tapering in dorsal view, covered base of the flagellum visible by transparency and attached to the ventral sclerite. Protruding part of the flagellum double curved horizontally, the acute, claw-like apex bent downwards the tergite X antecosta. Basal half of the flagellum dark brown but gradually lightened towards the pale brown apex; a conspicuous central filament terminates before the apex.

Female: Unknown.

Affinities: The new species differs from S. paraproctalis Aubert, 1967 , the only other species of the genus known from the Himalayan region, by bearing projections on tergite X and having median lobe of the paraproct shorter than the inner lobe. The paired projections on tergite X are unique in the genus distinguishing the new species from the eleven other species of this small, Southeastern Asian genus. Additional characters useful to distinguish S. siveci are the combination of features of the paraproct, enlarged flagellum of the epiproct and the lack of a lobe on tergite VIII.

Distribution and ecology. The species was collected in a national park on the western edge of the Meghalaya Plateau, close to the border of Bangladesh ( Fig. 9 ). This range is well separated from the Himalayas by the Brahmaputra valley and forms a western directed chain of the Patkai-Arakan ranges that run North to South on the bordering areas of India and Myanmar. The only other stonefly collected at the type locality was a female of a Phanoperla .

Etymology. The species is dedicated to Dr. Ignac Sivec, Ljubljana, Slovenia, in recognition of his work on the Oriental stoneflies, regarding especially the genus Sphaeronemoura . The name is used as the genitive of a noun of male gender.

Checklist of the Nemouridae species known from the Indian Subcontinent

We provide a list of species from Pakistan, Nepal, Bhutan and the states of India. Species of Nemouridae have not yet been recorded from Bangladesh or from Myanmar (Sivec & Yang 2001). The distribution of the Nemouridae in bordering countries of Russia has been noted by Zhiltzova (2003) and in China by Yang et al. (2013). If otherwise not indicated in the checklist, distribution outside the Indian Subcontinent refers to these two works.

The presence of the genus Protonemura Kempny, 1898 requires confirmation for the region, as the single species, P. scutigera Kimmins, 1950 b, is known only from females and possibly belongs to Indonemoura Baumann, 1975 or Mesonemoura Baumann, 1975 . On the basis of its original description, Protonemura mira Harper, 1974 is transferred to Indonemoura herein due to its paraproctal and epiproctal features, and the lack (or reduction) of gills: I. mira (Harper, 1974) comb. n. Most species of Illiesonemoura Baumann, 1975 were reclassified as Nemoura Latreille, 1796 in Zwick & Sivec, 1980. We list these as Illiesonemoura herein, because still treated as such in the Plecoptera Species File (DeWalt et al 2013). Nemoura bituberculata Kimmins, 1950 and N. rahlae Jewett, 1958 were described on the basis of females, both their status and generic assignment are questionable. The numerous females described but not named in the papers of Aubert (1959, 1967), Harper (1974), Zwick (1977) and Zwick & Sivec (1980) are not included in the checklist.

Species endemic to the Himalayas are marked with an asterisk (*) while those endemic to the Meghalaya Plateau and Patkai-Arakan ranges are marked with a ring (°). At present, there are 12 species known from Pakistan, 29 from Nepal, 16 from Bhutan and 69 from India (regarding states: 4 from Jammu and Kashmir, 10 from Himachal Pradesh, 1 from Tamil Nadu, 12 from West Bengal, 17 from Meghalaya, 7 from Assam, 29 from Arunachal Pradesh, 12 from Manipur and 2 from Mizoram).

Amphinemurinae

* Amphinemura albifasciata Sivec, 1981 Nepal (Sivec 1981a)

A. amatulai Aubert, 1967 India: Meghalaya (Aubert 1967), Arunachal Pradesh (Aubert

1967)

* A. baumanni sp. n. India: West Bengal (present paper)

* A. bella Zwick, 1977 Bhutan (Zwick 1977)

* A. bilolai Aubert, 1967 India: Arunachal Pradesh (Aubert 1967) * A. bomdilai Aubert, 1967 India: Assam (Aubert 1967), Arunachal Pradesh (Aubert

1967)

° A. cherrapunjii (Aubert, 1967) India: Meghalaya (Aubert 1967, Zwick & Sivec 1980) * A. dentata Zwick, 1977 Bhutan (Zwick 1977)

* A. elegans Zwick, 1980 (in Zwick & Sivec 1980) India: West Bengal (Zwick & Sivec 1980) * A. exigua Zwick, 1977 Bhutan (Zwick 1977)

* A. fusca Zwick, 1977 Bhutan (Zwick 1977)

* A. fuscipes Zwick, 1977 Bhutan (Zwick 1977)

* A. laguncula Harper, 1975 Nepal (Harper 1975)

* A. lebezi Sivec, 1981 Nepal (Sivec 1981a)

° A. lithami Aubert, 1967 India: Manipur (Aubert 1967)

* A. lurida Zwick, 1977 Bhutan (Zwick 1977)

A. luteipes Kimmins, 1947 India: West Bengal (Kimmins 1947, Zwick & Sivec 1980), Assam (Aubert 1967), Meghalaya (Aubert 1967), Arunachal Pradesh (Aubert 1967), Manipur (Aubert 1967); Nepal (Harper 1975, sub. nom. A. paraluteipes Aubert, 1967 ; Sivec 1980, 1981a)

° A. manipurensis Aubert, 1967 India: Meghalaya (Aubert 1967), Manipur (Aubert 1967) * A. martensi Zwick, 1980 (in Zwick & Sivec 1980) Nepal (Zwick & Sivec 1980, Sivec 1980, 1981a) * A. minor Sivec, 1982 Nepal (Sivec 1982)

A. mirabilis mirabilis (Martynov, 1928) Kaukasus, Transcaucasia , Anatolia, Iran; Afghanistan (Aubert

1966); Pakistan (Aubert 1959); China: Xinjiang * A. moshingi Aubert, 1967 India: Arunachal Pradesh (Aubert 1967) * A. nepalensis Harper, 1975 Nepal (Harper 1975)

* A. nigrifrons Zwick, 1977 India: West Bengal (Zwick & Sivec 1980); Bhutan (Zwick

1977)

° A. nongrimi Aubert, 1967 India: Assam (Aubert 1967), Meghalaya (Aubert 1967),

Manipur (Aubert 1967)

A. nubila Kimmins, 1950 India: Tamil Nadu (Kimmins 1950a)

A. paraluteipes Aubert, 1967 India: Assam (Aubert 1967), Meghalaya (Aubert 1967), Arunachal Pradesh (Aubert 1967), Manipur (Aubert 1967), Mizoram (1967)

* A. pseudoluteipes Aubert, 1967 India: Arunachal Pradesh (Aubert 1967) * A. pulchra Zwick, 1977 India: West Bengal (Zwick & Sivec 1980); Bhutan (Zwick

1977)

A. rahungi Aubert, 1967 India: West Bengal (Zwick & Sivec 1980), Meghalaya (Aubert 1967, Zwick & Sivec 1980), Arunachal Pradesh (Aubert 1967), Manipur (Aubert 1967); Nepal (Zwick & Sivec 1980); Bhutan (Zwick 1977)

* A. renata Kimmins, 1950 India: Assam (Aubert 1967), Arunachal Pradesh (Kimmins

1950 b, Aubert 1967)

A. schmidi (Aubert, 1959) Pakistan (Aubert 1959)

* A. simplex Zwick, 1977 Bhutan (Zwick 1977)

* A. stangeli Sivec, 1980 (in Zwick & Sivec 1980) Nepal (Zwick & Sivec, 1980, Sivec 1980)

* A. talungdzongi Aubert, 1967 India: Arunachal Pradesh (Aubert 1967)

* A. tricantha (Jewett, 1958) Pakistan (Zwick & Sivec 1980); India: Jammu and Kashmir (Zwick & Sivec 1980), Himachal Pradesh (Jewett 1958, 1960, 1970, 1975)

* A. wittmeri Zwick, 1980 (in Zwick & Sivec 1980) Nepal (Zwick & Sivec 1980)

* Indonemoura adunca (Harper, 1974) India: West Bengal (Zwick & Sivec 1980); Nepal (Harper 1974, Zwick & Sivec 1980, Sivec 1980, 1981a, 1981b, 1982) I. assami (Aubert, 1967) India: West Bengal (Zwick & Sivec 1980), Meghalaya (Aubert 1967), Arunachal Pradesh (Aubert 1967); Nepal (Harper 1974, sub. nom. Protonemura godavariensis Harper, 1974 ; Zwick & Sivec 1980, Sivec 1980, 1981a); Bhutan (Zwick 1977); China: Tibet (Li & Yang 2008)

* I. dirangdzongi (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

* I. geminus Zwick, 1977 Bhutan (Zwick 1977)

* I. gigaoni (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

* I. indica (Kimmins, 1947) India: West Bengal (Kimmins 1947), Arunachal Pradesh

(Aubert 1967); Nepal (Sivec 1980, 1981a, 1981b) * I. kamengi (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

* I. langtangi Sivec, 1980 (in Zwick & Sivec 1980) Nepal (Zwick & Sivec 1980, Sivec 1980)

° I. loebli Zwick, 1980 (in Zwick & Sivec 1980) India: Meghalaya (Zwick & Sivec 1980)

° I. mclachlani (Kimmins, 1950) India: Meghalaya (Kimmins 1950b, Aubert 1967) ° I. manipuri (Aubert, 1967) India: Manipur (Aubert 1967)

* I. mira (Harper, 1974) comb. n. Nepal (Harper 1974)

* I. nahkui (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

* I. nyukmadongi (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

° I. quadridentata (Kimmins, 1950) India: Meghalaya (Kimmins 1950b), Manipur (Aubert 1967) * I. rubrifasciata Zwick, 1977 Bhutan (Zwick 1977)

* I. sangtii (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

* I. shergaoni (Aubert, 1967) India: West Bengal (Zwick & Sivec 1980), Arunachal Pradesh (Aubert 1967); Bhutan (Zwick 1977); China: Tibet (Li & Yang 2008)

* Mesonemoura brachyfiligera (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

* M. falcata (Kimmins, 1950) India: Arunachal Pradesh (Kimmins 1950b)

* M. filigera (Kimmins, 1947) India: West Bengal (Kimmins 1947, Zwick & Sivec 1980);

Nepal (Zwick & Sivec 1980, Sivec 1980, 1981a)

* M. funicula (Harper, 1974) Nepal (Harper 1974, Zwick & Sivec 1980, Sivec 1980, 1981a,

1981b)

* M. mastigophora (Harper, 1974) Nepal (Harper 1974, Zwick & Sivec 1980, Sivec 1980, 1981a) * M. metafiligera (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967); Bhutan (Zwick

1977)

* M. mishmica (Kimmins, 1950) India: Arunachal Pradesh (Kimmins 1950b)

* M. parafiligera (Aubert, 1967) India: Arunachal Pradesh (Aubert 1967)

° M. pseudofiligera (Aubert, 1967) India: Manipur (Aubert 1967)

M. skardui (Aubert, 1959) Afghanistan (Aubert 1966); Pakistan (Aubert 1959, Zwick & Sivec 1980); India: Jammu and Kashmir (Zwick & Sivec 1980); Himachal Pradesh (Jewett 1970, 1975)

M. vaillanti (Navás, 1922) Afghanistan, Tajikistan; Pakistan (Aubert 1959); India: Himachal Pradesh (Jewett 1970, 1975); China: Gansu (Navás 1922, Li & Yang 2009)

* Protonemura ? scutigera Kimmins, 1950 India: Arunachal Pradesh (Kimmins 1950b, Aubert 1967) Nemourinae

* Illiesonemoura ampula (Jewett, 1958) India: Himachal Pradesh (Jewett 1958, 1970) * I. atripes (Aubert, 1959) Pakistan (Aubert 1959)

* I. battakundi (Aubert, 1959) Pakistan (Aubert 1959)

* I. besali (Aubert, 1959) Pakistan (Aubert 1959, Zwick & Sivec 1980); Nepal

(Zwick & Sivec 1980)

* I. cordata (Jewett, 1958) India: Himachal Pradesh (Jewett 1958, 1960, 1970); Nepal

(Zwick & Sivec 1980)

* I. falcifera (Harper, 1975) Nepal (Harper 1975)

* I. gosainkundensis (Harper, 1975) Nepal (Harper 1975, Zwick & Sivec 1980, Sivec 1980,

1981 a, 1981 b)

* I. lilami (Aubert, 1959) Pakistan (Aubert 1959); India: Jammu and Kashmir (Zwick

& Sivec 1980)

* I. maluksari (Aubert, 1959) Pakistan (Aubert 1959)

* I. pakistani (Aubert, 1959) Pakistan (Aubert 1959)

I. polystigma (Aubert, 1959) Pakistan (Aubert 1959); India: Jammu and Kashmir (Zwick

& Sivec 1980)

* I. punctata (Jewett, 1958) India: Himachal Pradesh (Jewett 1958, 1960, 1970, 1975) * I. punjabensis (Jewett, 1958) India: Himachal Pradesh (Jewett 1958, 1960, 1970, 1975,

Zwick & Sivec 1980)

* I. verrucosa (Harper, 1975) Nepal (Harper 1975)

* Nemoura abscissa Zwick, 1977 Bhutan (Zwick 1977)

° N. auberti Zwick, 1977 India: Manipur (Aubert 1967, sub. nom. N. schmidi Aubert,

1967; Zwick 1977)

*N.? bituberculata Kimmins, 1950 India: Arunachal Pradesh (Kimmins 1950b) * N. bokhari Aubert, 1967 India: Arunachal Pradesh (Aubert 1967) N. brevicauda Zwick, 1980 (in Zwick & Sivec 1980) India: Meghalaya (Zwick & Sivec 1980); Nepal (Zwick &

Sivec 1980)

° N. chattriki Aubert, 1967 India: Manipur (Aubert 1967)

* N. chugi Aubert, 1967 India: Arunachal Pradesh (Aubert 1967) * N. indica (Needham, 1909) India: Assam? (Needham 1909, Zwick 1977; locality is

‘Upper Assam’)

° N. khasii Aubert, 1967 India: Meghalaya (Aubert 1967)

° N. kuhleni Aubert, 1967 India: Manipur (Aubert 1967)

° N. lahkipuri Aubert, 1967 India: Assam (Aubert 1967)

* N. magnicauda Zwick, 1980 (in Zwick & Sivec 1980) India: West Bengal (Zwick & Sivec 1980) ° N. mawlangi Aubert, 1967 India: Meghalaya (Aubert 1967)

* N. nepalensis Zwick, 1980 (in Zwick & Sivec 1980) Nepal (Zwick & Sivec 1980)

*N.? rahlae Jewett, 1958 India: Himachal Pradesh (Jewett 1958) ° N. serrarimi Aubert, 1967 India: Meghalaya (Aubert 1967)

N. spinosa Wu, 1939 India: Mizoram (Aubert 1967); China: Guizhou, Shaanxi,

Yunnan.

* N. tamangi Sivec, 1980 (in Zwick & Sivec 1980) Nepal (Zwick & Sivec 1980, Sivec 1980) * N. triangulifera Zwick, 1980 (in Zwick & Sivec 1980) India: Himachal Pradesh (Zwick & Sivec 1980); Nepal

(Zwick & Sivec 1980, Sivec 1980)

* N. unicornis Jewett, 1975 India: Himachal Pradesh (Jewett 1975); Nepal (Zwick &

Sivec 1980)

Zoogeographical aspects

Although the distributional data are rather scarce and unevenly dispersed, some zoogeographical conclusions can be drawn on the Nemouridae species of the Indian Subcontinent. With the exception of a single species described from Tamil Nadu ( Amphinemura nubila Kimmins, 1950 a, known only from the female holotype), the family is unknown south of the Indo-Gangetic Plain. The Nemouridae fauna of the Himalayas show distinct differences between the western and eastern ranges. Among the 19 species known from the Western Himalayas ( Pakistan, Jammu and Kashmir, Himachal Pradesh), only 4 dispersed to Nepal and none to the Eastern Himalayas. Five of the 19 species are not Himalayan endemics. Four species are known also from the Karakorum and 2 from the Hindukush: Mesonemoura vaillanti (Navás, 1922) reaches also Gansu Province of China, while Amphinemura mirabilis mirabilis (Martynov, 1928) is distributed westwards to the Kaukasus and eastwards to Xinjiang Province of China (Yang et al. 2013).

Forty-seven species are known from the more studied Eastern Himalayas (West Bengal, Bhutan, Arunachal Pradesh and parts of Assam; all data from West Bengal refer to the Himalayan Darjeeling District), but only 7 of these species have dispersed westwards into Nepal which has 17 recorded endemic species in addition to the eastern and western Himalayan species. Despite their limited distribution in the more or less continuous Himalayan ranges, seven of the 47 Eastern Himalayan species have crossed the Brahmaputra Valley (considered a strong dispersal barrier) and dispersed into the Meghalaya Plateau. Three of these species are known also from the Patkai- Arakan ranges ( Amphinemura luteipes Kimmins, 1947 , A. paraluteipes Aubert, 1967 and A. rahungi Aubert, 1967 ).

In addition to the seven species shared with the Eastern Himalayas, the Meghalaya Plateau has seven species hitherto know only from this range, and three species that are known from the Meghalaya Plateau and the Patkai- Arakan ranges. Additional seven species are known only from the Indian part of the Patkai-Arakan ranges (Manipur, Mizoram and parts of Assam), while a single species, Nemoura spinosa Wu, 1939 is known from the Arakan Mts of Mizoram and from Guizhou, Yunnan and Shaanxi provinces of China (Yang et al. 2013).

The Holarctic and Oriental genus Amphinemura is widespread throughout the northern mountainous ranges of the Subcontinent, with a single species known also from the southern low mountains. The Oriental genera Indonemoura and Sphaeronemoura seem to be lacking from the Western Himalayas, their westernmost occurrences are in Nepal. The East Palaearctic and Oriental genus Illiesonemoura (sensu Baumann 1975) is not known from the mountain ranges east of Nepal. Another East Palaearctic and Oriental genus Mesonemoura is represented in the whole Himalayan range but with only a single species from Manipur in the Patkai-Arakan ranges. The Holarctic and Oriental Nemoura is also missing from the Western Himalayas and most of the Indian species belong to the Oriental and Transberinghian cercispinosa complex.

Acknowledgements

Visit of the senior author to the WNHM received support from the SYNTHESYS Project http:// www.synthesys.info/ which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program, application number AT-TAF-2660. The junior author acknowledges partial support by the National Natural Science Foundation of China (No. 31000977).

Figures

FIGURE 9. Type localities of Amphinemura baumanni sp. n. (dot) and Sphaeronemoura siveci sp. n. (ring) — grey areas are above 1000 m, Indian states of published Nemouridae records marked with numbers as follows: 1 — Jammu and Kashmir; 2 — Himachal Pradesh; 3 — Tamil Nadu; 4 — West Bengal; 5 — Meghalaya; 6 — Assam; 7 — Arunachal Pradesh; 8 — Manipur; 9 — Mizoram.

FIGURES 5 – 8. Male terminalia of Sphaeronemoura siveci sp. n.; holotype, India, Meghalaya, Nokrek NP — 5: dorsal view; 6: laterodorsal view; 7: ventral view; 8: lateral view — scale 0.5 mm.