The family Lecithoceridae in Bhutan is reviewed, with descriptions of one new species in Torodora Meyrick, 1894, T. namgaydemae Park & Gielis, sp. n., and a new species in Thubana Walker, 1864, Th. bhutanica Park & Gielis, sp. n. An unnamed species of Torodora is presented and discussed. In addition eight species are reported for the first time from Bhutan: Torodora multispinata Park, 2020; T. macrosigna Gozmány, 1973; T. trigona Gozmány, 1973; T. quadrangulata Wadhawan & Walia, 2007; Lecithocera cornutima Park, 2009; L. nepalica Gozmány, 1973; Homaloxestis cicatrix Gozmány, 1973 and Eccedoxa selena Wu, 1994. Images of the adults and genitalia are provided for all species.
Bhutan is a country in the Himalaya mountains, wedged between Tibet and India. Most of the northern part of the country is mountainous, with summits up to 7000 m, and with a fauna showing Palaearctic characteristics. At high altitudes alpine meadows and scrubs are the main vegetation, and at lower altitudes coniferous forests are the dominant habitat. In the south of the country, there is a narrow stretch of land along the border with India, with an altitude of 125–300 m. The fauna in the lowland and the lower parts of the valleys is Oriental. The vegetation in these lower parts consists of broad-leaved forests. The demarcation between these faunal areas is a zone along the mountains between 1500 and 2500 m. Due to the high precipitation which locally can exceed 3000 mm per year, especially in the monsoon period from May to August, the mountains show strong erosion. Slopes are covered with a thick layer of loose soil and sediments accumulate at the bottom of the valleys. An additional factor is the temperature. In the low regions, especially in the spring and early summer, monsoon winds bring heated air from the Indian subcontinent, often with temperatures over 40°C and high humidity. The temperatures drop with increasing altitude, but the winds still bring thick clouds up to high altitudes. Agriculture is very small-scaled, with rice fields along the slopes up to 2500 m. Cattle herding takes place in temperate camps in the mountains during the summer period. In autumn these herds migrate to lower altitudes to overwinter.
A cooperation between the Naturalis Biodiversity Center in Leiden, The Netherlands, and the National Biodiversity Center in Serbithang, Bhutan, started in 2016, and field work was enabled, in combination with an educational programme for Bhutanese students and field workers. In the following years, visits to Bhutan were made during April, mid-May to mid-June, and in August.
The Bhutanese moth fauna, including the family Lecithoceridae, has been poorly explored. Prior to this report, only a single lecithocerid species, Torodora arcifera (Meyrick, 1907), was known from Bhutan: a female specimen belonging to its syntype series (S. India, Bhutan, and Birma) was designated as its lectotype (Clarke 1955).
Material and methods
Specimens were collected at night from sheets and walls illuminated with mercury vapour lamps, or actinic tube lights, and in the daytime by sweeping the vegetation with a net. The localities where collecting took place varied in altitude from 125 to 3995 m. After collecting and killing, moths were stored in glassine envelopes, and dried for shipping. For further research the material was relaxed, pinned and the wings spread. Genitalia were dissected and mounted on glass slides in Euparal. Photographs of the moths were made with a Nikon D300 camera with bellows extension on which a Zeiss Luminar 100-mm lens was mounted, and illuminated in a “light-box” with a full-spectrum tube light; genitalia were photographed with an Olympus BH2 microscope and a Nikon D800 digital camera. Images were processed with Adobe Photoshop CC (Adobe Systems, San Jose, CA, USA).
Cornelis (Cees) Gielis, Lexmond, The Netherlands
Hokkaido University, Sapporo, Japan
Institute of Zoology, Chinese Academy of Science, Beijing, China
National Biodiversity Center, Serbithang, Bhutan
The Natural History Museum, London, UK
National Institute of Biological Resources, Incheon, Korea
Naturalis Biodiversity Center, Leiden, The Netherlands
Ugyen Wangchuck Institute for Conservation, Environment and Research, Bumthang, Bhutan
Zoologische Sammlung des Bayerischen Staates, München, Germany
Genus Lecithocera Herrich-Schäffer, 1853
Lecithocera cornutima Park, 2009
Lecithocera cornutima Park, 2009: 63. – Thailand, Chiang Mai. Holotype in NIBR.
Material. 1♂, Bhutan, Chhukha, 8 km S Gedu, Darla, 26°51’30″ N 89°33’ 30″ E, 1750 m, 17–18.v.2017 (leg. C. & F.K. Gielis & K. Wangdi), prep CG7473, wing vein prep CG7571, photo Bhutan 1193, sta 40 (NBC). New for Bhutan.
Wingspan 13 mm. The species can be distinguished from congeners by the presence of five small, yellowish-orange costal spots from basal 1/3 of costa of the forewing, and hindwing with well visible veins covered by dark brown scales. The characteristic male genitalia are described below.
Male genitalia (Fig. 4; see also Park 2009, Fig. 12a, b). Basal lobes of uncus uniquely developed with long, narrow stalk in basal half and widely broadened in distal half, deeply concave on caudal margin. Gnathos small and short. Valva broad basally, with crescent concavity ventrally; costa concave; cucullus elongate with round apex, with dense long setae on surface and short, conic spines along ventral margin. Juxta heavily sclerotized, horn-shaped latero-caudal processes not exceeding apex of vinculum. Aedeagus as long as valva, bifurcate apically, with large, strong, horn-shaped cornutus.
Bhutan, Thailand (Chiang Mai).
Lecithocera nepalica Gozmány, 1973
Lecithocera nepalica Gozmány, 1973: 421. – Nepal, nr East Jubing. Holotype in ZSM.
Gozmány 1978: 100.
Material. 1♀, Bhutan, Trashi Yangtse, Trashi Yangtse, 27°36’30″ N 91°29’30″ E, 1765 m, 26–27.v.2017 (C. & F.K. Gielis & K. Wangdi), prep CG7482, photo Bhutan 1208, sta 47 (NBC); 1♀, same locality and date, prep CG7484, prep forewing CG7573, photo Bhutan 1211 (RMNH). New for Bhutan.
Wingspan 15–17 mm. The species is characterized by the pale brown forewing, with well-developed discal spots: a smaller at middle, and a larger which often faintly spreads towards dorsum at end of cell. The forewing has R3 free from R4+5 at base, and CuA1 and CuA2 short-stalked.
Female genitalia (see also Gozmány 1978, Pl. 64, Fig. 36). Antrum large, cup-shaped, weakly sclerotized. Ductus bursae narrowed in distal 1/5, broadened anteriorly; ductus seminalis broad, arising from about distal 1/3. Corpus bursae ovate, rather small; signum somewhat elongate and rounded, with numerous conic spines on surface.
Genus Homaloxestis Meyrick, 1910
Homaloxestis cicatrix Gozmány, 1973
Homaloxestis cicatrix Gozmány, 1973: 415. – Nepal, Rati-Tal, Monahari Khola. Holotype in ZSM.
Material. 1♂, Bhutan, Sarpang, Gelephu, 26°53′ N 90°28′ E, 275 m, 10.iv.2019 (C. & F.K. Gielis & K. Wangdi), prep CG7499, photo Bhutan 1226, sta 100 (NBC). New for Bhutan.
Wingspan 13–14 mm. The forewing is pale yellow, scattered with yellowish brown scales and without distinct markings. The hindwing and abdomen of the holotype are destroyed.
Male genitalia (see also Gozmány 1978, Pl. 19, Fig. 12); Wu 1997, Pl. 15, Fig. 5; Park et al. 2008, Figs. 47, 47a). Gnathos rather slender, strongly bent downward beyond 2/3. Valva broad basally; cucullus elongated, with truncate apex; ventral margin abruptly oblique from middle with minute conic spines along margin beyond. Juxta emarginate medially on caudal margin. Aedeagus stout, with small apical spines ventrally; cornuti consisting of two large sacs bearing spines.
Bhutan, Nepal, Myanmar, China (Jiangxi, Hainan), Vietnam (North).
Genus Torodora Meyrick, 1894
Torodora namgaydemae Park & Gielis, sp. n.
Type material. Holotype, ♀, Bhutan, Trasigang, Wamrong, 27°07′ N 91°34′ E, 2050 m, 17.iv.2019 (C. & F.K. Gielis & K. Wangdi), prep CG7474, photo Bhutan 1197, sta 107 (NBC).
Paratypes: 1♀, locality and date as holotype, prep CG7478, photo Bhutan 1202 (RMNH); 1♀, Bhutan, Trashi Yangtse, Trashi Yangtse, 27°36’49″ N 91°29’34″ E, 1760 m, 18.viii. 2016 (C. Gielis & K. Wangdi), prep CG7506, wing vein prep CG7570, photo Bhutan 1236, sta 28 (NIBR).
The new species can be distinguished from the known species of the genus with more or less similar wing pattern and markings, including T. cordisigna Park, 2020 and T. multispinata from Laos, T. crassidigitata Li, 2010 (see Zhang & Li 2010) from China, and T. parasciadosa Park, 2015 (see Park et al. 2015) from Taiwan, by the female genitalia. It differs by the specifically modified ostium bursae of the new species which broadens like a bowl having 3–5 unique grooves on both sides.
Female. Wingspan 17 mm. Head with dark brown appressed scales, with ochreous scales laterally above eyes. Antenna about 4/5 the length of forewing, ochreous with brown scales at regular intervals. Labial palp arched, strongly upturned; 2nd segment slightly thickened, long, dark brown on outer surface; 3rd segment slender, as long as 2nd segment, ochreous-grey. Thorax and tegulae dark brown. Hind tibia dark brown, speckled with ochreous scales; apex pale yellow; tarsi pale yellow. Forewing dark brown, with a pair of blackish discal spots at middle of wing and at end of cell; postmedian line faintly developed, strongly zigzagged; costa slightly concave beyond middle, with small, triangular, yellowish-white costal patch at 5/6; apex produced; termen oblique, concave medially, sinuate; venation with R3, R4 and R5 on a common stalk: R3 and R4 stalked for basal 1/4, R4 and R5 stalked for about 2/3; R5 to termen; M1 remote from R3+4 at base; M2 close to M3; M3 approximate to CuA1+2; CuA1 and CuA2 long-stalked; cell closed; fringes dark brown, with narrow yellowish-white basal line. Hindwing pale brown-grey; fringes pale grey, with narrow, yellowish basal line. Abdomen dark brown dorsally; segments I–III much paler.
Female genitalia. Apophyses anteriores about half the length of apophyses posteriores. Ostium broadened, bowl-shaped, with several irregular furrows both sides. Antrum elongate, about 1/6 length of ductus bursae, weakly sclerotized. Ductus bursae long, slightly coiled; ductus seminalis incepted in the middle of length of ductus bursae. Corpus bursae elongate; signum semi-ovate, width about twice its length, with strong spines along caudal margin and numerous, smaller conic spines on surface.
The moth flies in April and August, at an altitude of 1760–2050 m. Feeding pattern unknown.
Bhutan: Trashigang, Trashi Yangtse.
The species is dedicated to, and to honour, Mrs. Namgay Dem.
Material. ♀, Bhutan, Tsirang, Damphu, 27°00′ N 90°07′ E, 1530 m, 9–10.vi.2017 (C. & F.K. Gielis & K. Wangdi), prep CG7487, wing vein prep CG7569, sta 58 (NBC).
Female. Wingspan 20 mm. Head pale orange appressed scales. Antenna as long as forewing, pale orange throughout. Labial palp arched, strongly recurved; 2nd segment thickened, pale orange on outer surface and grey ventrally. Thorax and tegulae violet-brown; hind tibia dark brown; tarsi and spurs pale orange. Forewing violet-brown, without distinct markings, but with yellowish-orange scales along inner margin between basal 3/5 and 4/5; costa slightly oblique beyond distal 1/6; apex smoothly rounded; termen slightly oblique, with faint yellowish-white marginal line; venation with R3, R4 and R5 on a common stalk; R3 and R4 stalked for basal 1/4, R4 and R5 stalked for about 2/3; R5 to termen; M1 remote from R3+4 at base; M2 close to M3, nearly parallel to M1; M3 free; CuA1 and CuA2 short-stalked; cell closed; fringes concolorous with ground colour. Hindwing dark brown; costa with yellowish-white streak in basal 2/3; fringes concolorous with ground colour. Abdomen dark brown dorsally, with pale orange distal end.
Female genitalia. Apophyses anteriores very short, nearly atrophied. Ostium uniquely formed, with well-developed lateral lobes directed inward; caudal margin nearly straight. Ductus bursae wide tubular, as long as corpus bursae, with row of six isolated spines. Bursa copulatrix vesicular, signum more or less pepper-shaped, slightly broadened distally, with dense conic spines on surface.
This species is superficially very similar to Thubana onyx Gozmány, 1978, which was described from a single male from Nepal, by having a similar orange head dorsally and deep violet-brown forewing ground colour. It differs from the latter by the forewing venation, with M3 free and CuA1 and CuA2 short-stalked, a diagnostic character of Torodora; the apex is sharply produced and the termen less oblique, whereas in Thubana onyx, the forewing with costa is very oblique beyond its distal 1/6, the apex obtuse, and the termen very oblique. As Gozmány (1978) did not describe its venation there is no certainty that the species was correctly placed in Thubana. To avoid creating a synonym we await describing it as a new species till further material becomes available.
Torodora multispinata Park & Bae, 2020
Torodora multispinata Park & Bae, 2020: 313. – Cambodia. Holotype in NIBR.
Material. 1♀, Bhutan, Samdrup Jongkhar, Phuentshog thang, 26°53′ N 91°41′ E, 345 m, 16.iv.2019 (C. & F.K. Gielis & K. Wangdi), prep CG7471, photo Bhutan 1198, sta 106 (RMNH). New for Bhutan.
Female. Wingspan 14–16 mm (Bhutan specimen: 16 mm). This species is superficially similar to T. occidentalis Park, 2013 which was described from Cambodia, but the female genitalia of this species are unique with numerous strong spines in the corpus bursae.
Female genitalia (see also Park & Bae 2020, Fig. 6D). Antrum membranous, funnel-shaped; ductus bursae narrow, shorter than length of corpus bursae; ductus seminalis arising from near middle. Corpus bursae extremely large, ovate, with two series of unique spine-like scales distally; signum heart-shaped, with numerous conic spines on surface, deeply incised on anterior margin.
Bhutan, Laos (Bolikhamsai Prov.), Cambodia (Kep Prov.).
Torodora macrosigna Gozmány, 1973
Torodora macrosigna Gozmány, 1973: 440. – Nepal. Holotype in ZSM.
Gozmány 1978: 219.
Material. 1♀, Bhutan, Zhemgang, 3 km N Tingtibi, 27°09′ N 90°40′ E, 610 m, 12.iv.2019 (C. & F.K. Gielis & K. Wangdi), prep CG7477, photo Bhutan 1201, sta 102 (NBC). New for Bhutan.
Female genitalia. Ductus bursae narrowed in its distal 1/6, beyond antrum, with short spines; corpus bursae with elongated signum, shaped like a bell pepper, with dense conic spines. Differences compared to the illustration by Gozmány (1973), are considered to be infraspecific variation (see also Gozmány 1978, Pl. 83, Fig. 144; Park & Bae 2020a, Fig. 6H).
Bhutan, Nepal (Bhimpedi), Cambodia (Pursat).
Torodora trigona Gozmány, 1978
Torodora trigona Gozmány, 1978: 199. – Nepal, Godavari. Holotype in HUJ.
Material. 1 ♂, Bhutan, Punakha, 2 km N Rimchu, 1440 m, 27°41’40″ N 89°46’06″ E, 24.iv.2019 (C. & F.K. Gielis & K. Wangdi), prep CG7472, photo Bhutan 1192, sta 114 (NBC). New for Bhutan.
Male. Wingspan 20 mm. This species is superficially similar to the following species, T. quadrangulata, which was described from Himachal Pradesh, North India, based on two male specimens, but it can be distinguished by the male genitalia
Male genitalia. Uncus broad at base, gradually narrowed toward apex. Gnathos with nearly squarish basal plate and the median process nearly atrophied. Valva with rounded cucullus. Juxta with digitate lateral lobes. This kind of gnathos is exceptional in Torodora, only shared with T. quadrangulata.
The photo (Fig. 23) of the holotype of this species in Hokkaido University, Japan, is given for comparison.
Torodora quadrangulata Wadhawan & Walia, 2007
Torodora quadrangulata Wadhawan & Walia, 2007: 2079. – India, Himachal, Chandigarh. Holotype in Panjab University, Chandigarh, India.
Material. Without abdomen, Bhutan, Trashigang, Kanglung, 1850 m, 27°17’11″ N 91°31’17″ E, 19.viii.2016 (C. Gielis & K. Wangdi), photo Bhutan 0114, sta 29 (NBC). New for Bhutan.
Male. Wingspan 15 mm. This species is superficially similar to T. trigona with similar markings on the forewing, but can be easily distinguished by the male genitalia with more elongate cucullus and the juxta in V-shape on caudal margin. The abdomen of the Bhutanese specimen is missing.
Bhutan, India (North),
The specimen collected in Bhutan is much smaller than the holotype, with a wingspan of 15 mm (vs 20 mm in the type). Male genitalia (Fig. 25) are reproduced after Wadhawan & Walia, 2007, with kind permission of Prof. Dr. Virinder Kumar Walia.
Torodora arcifera(Meyrick, 1907)
Brachmia arcifera Meyrick, 1907: 738. – Bhutan. Lectotype in NHMUK.
Material. 1♀, Bhutan, Tsirang, Damphu, 27°00′ N 90°07′ E, 1530 m, 9–10.vi.2017 (C. & F.K. Gielis & K. Wangdi), prep CG7493, photo Bhutan 1221, sta 58 (NBC); 1♀, Bhutan, Zhemgang, Panbang, 26°50′ N 90°56′ E, 135 m, 13.iv.2019 (C. & F.K. Gielis & K. Wangdi), photo Bhutan 1222, sta 103 (RMNH). Female genitalia see Clarke (19565, Pl. 55, Fig. 1a-c); Gozmány (1978, Pl. 81, Fig. 136).
Bhutan, India (South), Nepal.
Genus Thubana Walker, 1864
Thubana bhutanica Park & Gielis, sp. n.
Type material. Holotype, ♀, Bhutan, Samdrup Jongkhar, 1 km W Phuentshogthang, 28 km E Deothang, 26°52′ N 91°41′ E, 340 m, 22–23.v.2017 (C. & F.K. Gielis & K. Wangdi), prep CG7492, wing vein prep CG7567, photo Bhutan 1220, sta 44 (NBC).
This species is superficially similar to Torodora serpentina (Diakonoff, 1952), which was described from NE Myanmar, based on a male. The species is characterized by its forewing with a unique pattern of a dark purplish-brown area on pale orange colour, divided by a large triangular pale orange patch beyond middle of costa. The new species can be distinguished from T. serpentina by the broad dark violet-brown area reaching costa and divided by an orange triangularly concave spot medially, whereas in T. serpentina, it does not reach the costa and is smoothly concave medially.
Female. Wingspan 16 mm. Head with pale orange appressed scales dorsally, with erect scales along margin of eyes. Antenna slightly longer than forewing, pale orange. Labial palp arched, upturned, about six times eye diameter, pale orange on outer surface. Thorax and tegulae pale orange. Legs pale orange; hind tibia pale orange with fuscous scales at apex; first tarsus pale orange, with dark brown apex. Forewing pale orange, broadly occupied by the dark greyish-brown area with purplish gloss from base to tornus along dorsum and divided by large triangular pale orange patch beyond middle on costa, pale orange beyond; apex produced; termen oblique, slightly concave; venation with R3 short-stalked with R4+R5; R4 and R5 stalked for basal 1/3; M1 remote from R3+4 at base; M3, CuA1 and CuA2 on a common stalk; fringes pale orange. Hindwing dark grey-brown; fringes pale orange around apex, pale grey on termen and dorsum, pale orange near base.
Female genitalia. Ostium bursae broadened. Antrum short, weakly sclerotized, funnel-shaped. Ductus bursae broad, nearly parallel-sided, slightly longer than corpus bursae, bearing dense conic spines in posterior 1/4; ductus seminalis arising from near posterior 1/3. Corpus bursae large, ovate; signum plate transversally elongated, about three times as wide as long, with dense conic spines, bigger ones along posterior margin.
Bhutan (Samdrup Jongkhar).
The moth flies in May, at an altitude of 340 m. Feeding pattern unknown.
This specific name is derived from the type locality, the country of Bhutan.
Genus Eccedoxa Gozmány, 1973
Eccedoxa selena Wu, 1994
Eccedoxa selena Wu, 1994: 128. – China, Sichuan. Holotype in IZCAS.
Wu 1997: 100.
Material. 1♀, Bhutan, Trashigang, Wamrong, 27°07′ N 91°34′ E, 2040 m, 24.v.2017 (C. & F.K. Gielis & K. Wangdi), prep CG7475, photo Bhutan 1199, sta 45 (NBC); 1♀, same locality and date, prep 7476, photo Bhutan 1200, sta 45 (RMNH); 1♀, Bhutan, Mongar, 8 km E Kori-La Pass, 27°18′ N 91°20′ E, 1975 m, 28–29.v.2017 (C. & F.K. Gielis & K. Wangdi), prep CG7479, wing vein prep CG7575, photo Bhutan 1203, sta 49 (NIBR). New for Bhutan.
Female. Wingspan 17–19 mm. The forewing of the Bhutanese specimens is more brownish orange than in the holotype, but the female genitalia accord well with those of E. selena.
Female genitalia (see also Wu 1997, Pl. 27, Fig. 4). Characterized by the ductus bursae with dense conic spines medially and with broad ductus seminalis, and the corpus bursae with a pair of crescent signa.
Bhutan, China (Hainan, Sichuan).
The genus Eccedoxa is an Oriental genus, originally established for Cophomantis lysimopa Meyrick, 1933. The genus can be distinguished from related genera by the venation with the forewing R5 and the hindwing M2 being absent. Six species are known: four of them from India and Sri Lanka; one each from China and Australia.
We would like to thank Kees Klein and Namgay Dem (Thimphu, Bhutan) for their hospitality and support given, enabling the visits to Bhutan. They also helped to make us known with the capital Thimphu and the people of Bhutan. They introduced us to numerous people, making the Fauna Survey of Bhutan a success. Mrs. Sangay Dema, Choki Geyltshen, and Pema Leda (NBC, Serbithang) are thanked for their help and support in preparing the collecting trips and the loan of specimens. We thank Vincent Kalkman and Wim Klein (RMNH, Leiden) for their help in the preparations of the field work and Prof. Dr. Virinder Kumar Walia for his permission to use the illustration of the male genitalia of Torodora quadrangulata. The field work has been made possible by grants of the Bhutan Trust Fund for Environmental Conservation (2016), and the Uyttenboogaart-Eliasen Foundation, The Netherlands (2016, 2017 and 2019).
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