Psychomyiinae, Walker, 1852

Tachet, Henri, Coppa, Gennaro & Forcellini, Maxence, 2018, A comparative description of the larvae of Psychomyia pusilla (Fabricius 1781), Metalype fragilis (Pictet 1834), and Paduniella vandeli Décamps 1965 (Trichoptera: Psychomyiidae) and comments on the larvae of other species belonging to these three genera, Zootaxa 4402 (1), pp. 91-112 : 101-103

Distribution of the Psychomyiinae

Metalype ( Fig. 34 )—The genus Metalype was described by Klapálek (1898). Almost a century later, Malicky (1995), based on some characters of adults, considered Metalype and Psychomyia synonyms. Schmid (1997), after examination of Psychomyia collected in India between 1958 and 1962, described 55 new species that he divided into 6 Groups. His Psy. mahayinna Group included Psy. mahayinna Schmid 1961 , Psy. fragilis Pictet 1834 , Psy. klapaleki Malicky 1995 , Psy. uncatissima Botosaneanu 1970 , Psy. anaktujuh Malicky 1995, Psy. holzenthali Schmid 1997, and Psy. kumara Schmid 1997. Schmid (1997) considered that the Psy. mahayinna Group is “le plus primitif du genre ( Psychomyia ) par la séparation complète des appendices préanaux et des pointes du IX° tergite et la forme simple des appendices inférieurs.” Independently, Li & Morse (1997a), from a phylogenetic analysis, divided the Psychomyiidae into two subfamilies: the Psychomyiinae and the Tinodinae . They considered that the genus Metalype is valid. Schmid (1997) considered that the Psy. mahayinna Group belongs to the genus Psychomyia , but, de facto, the M. mahayinna Group corresponds to the genus Metalype . In the discussions by these specialists of adults, none of them refers to the publication of Edington & Alderson (1973), who argued that the larvae of Psychomyia pusilla and Metalype fragilis have characters that are too different for these two species to belong to the same genus. Waringer & Graf (2011) distinguished the larvae of these two species, but included them both in genus Psychomyia . Based on the works of Edington & Alderson (1973), Schmid (1997), and Li & Morse (1997a), we treat Psychomyia and Metalype as distinct genera. Metalype is a genus with eight species ( Schmid 1997; Morse 2016; Fig. 34 ). There are two species in the West Palearctic Region: M. fragilis and M. klapaleki , considered to be sister species (Malicky 1995; Urbanic et al. 2003). There are six other species in Oriental Asia: one in Pakistan, another in Nepal, two others in India (Uttarakhand and Assam States), and two East Palearctic species: M. nithaiah Malicky 2014 in Taiwan and M. uncatissima (Botosaneanu 1970) in North Korea (Botosaneanu 1970), South Korea (Jung et al. 2011), and Japan ( Torii 2011a, 2013). Recently, Qiu et al. (2017) described and redescribed four species: three new species for China: M. hubeiensis Qiu & Morse 2017 (from Hubei Province), M. shexianensis Qiu & Morse 2017 (An-hui Province), and M. truncata Qiu & Morse 2017 (Sichuan Province), and M. uncatissima (Botosaneanu 1970) in the Hei-long-jiang Province.

Psychomyia ( Fig. 35 )—With about 150 species ( Morse 2016), this is the genus of Psychomyiinae with the greatest number of species. The genus is missing from the Afrotropical Region. There are three species in the Nearctic Region, two of which are rather confined: Psy. nomada in the east, from South Carolina to Maine and Psy. lumina in the northwest from California to Washington. The third species Psy. flavida occurs in most of North America, from western British Colombia to Quebec in the north, southward to Arizona and North Carolina and probably all provinces and states between them ( Rasmussen & Morse 2014). Psychomyia flavida also occurs in the East Palearctic Region from Japan to Mongolia ( Torii 2011b; Chuluunbat & Morse 2007), including parts of Asian Russia (Amur Oblast, Altay, southern Khabarovsk, South Primorye, and Magadan Oblast; Ivanov 2011). In the West Palearctic Region ( Wiberg-Larsen 2004; Malicky 2004, 2005; Uherkovich & Nogradi 1991; Ujvarosi et al. 2008), there are five species, one of which ( P. pusilla ) has a broad distribution throughout Europe from Ireland ( O’Connor 2015) to Finland in the north ( Tobias & Tobias 1981), southward to the western Maghreb (Dakki 1982; Malicky & Lounaci 1987) and to the Caucasus (Ivanov 2011) and Iran (Chvojka 2006) and the Levant (Flint 1967a, Sipahiler 2005), and probably all areas between them. The other four species are more confined. In the East Palearctic Region, there are, apart from P. flavida , nine species with six species endemic in Japan ( Torii 2011b). The Oriental Region is a complex set: China, Southeast Asia, and the Indian subcontinent (including Sri Lanka). Five species occur in China ( Yang et al. 2005), but the limit between the Oriental and East Palearctic Regions remains blurred in that country. Southeast Asia can be divided into two parts: Mainland Southeast Asia (MLSEA) including Myanmar ( Burma), Thailand, Laos, Cambodia, and Vietnam, and Maritime Southeast Asia (MASEA) corresponding to Malaysia, Indonesia (except for West Papua which is administratively a part of Indonesia, but tectonically belongs to the Australian plate) and the Philippines. There are 30 Psychomyia species on the Mainland part ( Malicky 2010) and 45 on the Maritime part ( Morse 2016).

The Indian subcontinent ( Fig. 35 ), like Madagascar, was situated alongside the African continent 250 mya. Today, there are no species of Psychomyia known in the Afrotropical Region. During its long journey to the northeast, the Indian subcontinent remained isolated. The arrival of Psychomyia in India was possible only after the collision between India and Eurasia 30 million years ago, probably via Myanmar, because the Deccan Traps (covering 500 0 0 0 km2) and the Thar Desert in northwest India form a natural barrier to Trichoptera . The occurrence of Psy. benyagai Malicky & Chantaramongkol 1993 in Thailand, Vietnam, Bhutan, and India (Uttar Pradesh and Himachal Pradesh) ( Malicky & Chantaramongkol 1993; Thapanya et al. 2004) supports that hypothesis, that the origin of the Indian species of Psychomyia probably was Southeast Asia.

With regard to the countries situated in the Himalayas, North Pakistan has three species of Psychomyia , Nepal has two species) ( Malicky 2006), and Bhutan has two species ( Malicky et al. 2008) ( Fig. 35 ).

Until now, no Psychomyia have been found in Sri Lanka although only 31 kilometres separate southeastern India and Sri Lanka. During the glacial periods, a large, wide tombolo existed between southeastern India and Sri Lanka, as a result of the lowering of the level of the oceans, but this tombolo, consisting of sand and gravel, did not have adequate habitat for Trichoptera to complete their life cycle.

The Andaman and Nicobar Islands belong to India ( Malicky 1979) administratively, but geologically and tectonically they belong to the Sunda plate.

Psychomyia is clearly an “into India ” genus sensu Datta-Roy et al. (2014).

Paduniella ( Fig. 36 )—In the Nearctic Region, Paduniella occurs in Arkansas and Missouri (Pad. nearctica ) (Flint 1967b; Rasmussen & Morse 2014); in the West Palearctic region, one species (Pad. vandeli ) occurs in France (Décamps 1965; Coppa 2004; Coppa et al. 2008, 2010), the Iberian Peninsula (Gonzalez & Mendez 2011), and the western Maghreb (Dakki 1982; Malicky & Lounaci 1987). In European Russia, Paduniella uralensis occurs in both Western and Eastern Palearctic Russia (Central Volga, Ural Mountains, Far East; Ivanov 2011), East Palearctic China (Heilongjiang; Yang et al. 2005), and Japan (Nishimoto 2011). Also in the East Palearctic Region, Pad. amurensis occurs in Russia (Amur area, Khabarovsk Province, and South Primorye; Ivanov 2011) and Oriental China (Sichuan; Yang et al. 2005). Paduniella communis occurs in Japan (Nishimoto 2011) and Oriental China. In Southeast Asia, there are 14 species on the Mainland and 11 in Maritime Southeast Asia. For the Indian subcontinent ( Fig. 37 ), the distribution of 20 known Paduniella species is different from those of Psychomyia . There are three species in Pakistan ( Schmid 1961) and three in Nepal ( Malicky 1997, 2004). For India, one species occurs in Bihar State (Pad. fissa Martynov 1935) and one species in the Gujarat State (Pad. sabarmata Johanson & Olah 2010). Eleven species have been described from Sri Lanka ( Schmid 1958; Chantaramongkol & Malicky 1986).

Five species of Paduniella are known from southern China ( Li & Morse 1997b), one from Taiwan ( Kobayashi 1987), three from the Philippines (Banks 1939; Mey 1998), and four from Vietnam (Johanson & Oláh 2010). In the Afrotropical Region ( Fig. 36 ), four species of Paduniella have been described (de Moor & Scott 2003; Tobias & Tobias 2008). In Madagascar, five species have been described recently (Johanson & Oláh 2010).

Figures

FIGURE 34. Metalype spp. world distribution.

FIGURE 35. Psychomyia spp. world distribution. EP = East Palearctic Region, MASEA = Maritime Southeast Asian Region, MLSEA = Mainland Southeast Asian Region, NE = Nearctic Region, WP = West Palearctic Region. In each circles, number of species.

FIGURE 36. Paduniella spp. world distribution. Loci typici of eight species indicated by black dots. Number of Paduniella species indicatedfor Japan, southern China,MASEA (Maritime SoutheastAsia), MLSEA(MainlandSoutheast Asia),India, Sri Lanka, Madagascar.

FIGURES 37–41. Psychomyiidae spp., submental sclerites, ventral apotomes and anterior margins of the heads, ventral. (37) Psychomyia lumina (adapted from a drawing by Wiggins 1996); (38) P. flavida (adapted from a photograph); (39) Psy. ctenophora(adaptedfrom a photographby Vieira-Lanero2000); (40) Psy. undetermined South Korean species (adapted from a photograph); (41) Metalype fragilis (re-used Fig. 11 for comparisons).