Zootaxa 3313: 34–43 (2012) www.mapress.com / zootaxa/ Copyright © 2012 · Magnolia Press

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Description of a new Psychoda Latreille species from Fennoscandia (Diptera: Psychodidae) JUKKA SALMELA1,4, GUNNAR M. KVIFTE2,3 & ANNA MORE1 1

Zoological Museum, Department of Biology, FI-20014, University of Turku, Finland Department of Entomology, Bergen University Museum, P.O.Box 7800, University of Bergen, 5020 Bergen, Norway. E-mail: [email protected] 3 Ecological and Environmental Change Research Group, Institute of Biology, University of Bergen, Norway 4 Corresponding author. E-mail: [email protected] 2

Abstract Psychoda cultella sp. n. is described. The new species is characterized by 16-segmented antennae with the three apical flagellomeres globular, very short and partially fused; labellum with five terminal digital projections; aedeagus ca 1.7 times the length of gonostylus; distiphallus bipartite, composed of a ventral phallomere with a roundish blunt tip and a dorsal phallomere with pointed tip; basiphallus in lateral view sub-basally widened. The new species is known from Finland (29 localities) and Norway (6 localities), ranging from the southern boreal ecoregion to the subalpine zone. Psychoda cultella sp. n. is mainly found in Malaise trap samples collected from moist coniferous forests and spruce mires (riparian forests, springs) and rarely from fens and subalpine heaths. Two male specimens from Norway were reared from fruiting bodies of polypore fungi (either Fomes fomentarius or Piptoporus betulinus). Key words: Finland, Norway, Boreal zone, moth flies, taxonomy, saproxylic species, DNA barcode

Introduction The Psychodinae form the most species-rich subfamily of Psychodidae with more than 2000 species described from all over the world. The most widespread and abundant genus in the group is Psychoda Latreille, 1796; which comprises about 450 species worldwide and around 40 in Europe. Many of the most common Psychoda species have a global distribution, and populations of certain species may attain very high densities in artifical or natural habitats (e.g. Vaillant 1973; Ali & Koh-Yokomi 1991; Svensson 2009). Their larvae are mostly saprotrophic, living in vertebrate excrements or in other habitats rich in organic material. However, some species have been found in fungal fruiting bodies, carrion or phytotelmata (Withers 1988; Svensson 2009). Since Psychoda Latreille, 1796 is a very species-rich and diverse taxon, it will probably eventually need subdivision. Several alternative classifications have been proposed for the group, however in the absence of a good phylogenetic basis these are to be considered premature (see discussion in Cordeiro et al. 2011). Ježek (1977; 1983; 1984; 2007) and Ježek & van Harten (1996; 2005) subdivide the genus into a total of 13 genera; which are treated as subgenera by Withers (1986) and Bravo et al. (2006). However, Withers (1989), in his treatise on British Psychodidae, considered the splitting of Psychoda unnecessary and recently Cordeiro et al. (2011) chose to disregard this subgeneric classification altogether. A majority of species, especially in the tropics, are not readily placed in any described subgenus and no phylogenetic framework exists for a robust new classification (Cordeiro et al. 2011). In the present paper we describe a new European Psychoda species that cannot with confidence be placed in any of the (sub)genera diagnosed by Ježek (2007 and references therein). Its affinity to other members of Psychoda is briefly discussed.

34 Accepted by G. Curler: 26 Mar. 2012; published: 15 May 2012

Material and methods Measurements are given as means followed by the ranges and number of specimens on which the measurement is based. All measurements are given in μm, except wing length which is given in mm. Wing length is measured from the base/end of the costal node to the apex of R5. Morphological terminology is according to Curler and Courtney (2009). For the genitalia, the terms "dorsal" and "ventral" refer to the orientation of the animal after the inversion of the genitalia has taken place, rather than the morphologically dorsal and ventral sides. Most of the Finnish coordinates given below follow the national grid 27°E system, other coordinates are according to the WGS84 system Some specimens (see Material examined) were permanently slide mounted either in Euparal or Canada balsam. In most cases specimens were first macerated in KOH and then cleaned in acetic acid and ethanol. Only male specimens were studied. Finnish specimens are arranged in biogeographical provinces, from north to south. Li=Lapponia inariensis, Le=Lapponia enontekiensis, Lkoc=Lapponia kemensis pars occidentalis, Ks=Regio kuusamoensis, Oba=Ostrobottnia borealis pars australis, Om=Ostrobottnia media, Oa=Ostrobottnia australis, St=Satakunda, Sb=Savonia borealis, Kb=Karelia borealis, Tb=Tavastia borealis, Ab=Regio aboensis. DNA barcodes were obtained in cooperation with the Barcode of Life Datasystems (http://www.boldsystems.org). Legs of specimens were placed in 96–100% ethanol in a 96-well lysis microplate and shipped to the Biodiversity Institute of Ontario where DNA was extracted and sequenced using standard protocol and primers. See Ratnasingham & Hebert (2007) for details on the BOLD project. The following acronyms for museums and collections are used in the text: MZHF—Finnish Museum of Natural History (Zoological Museum), University of Helsinki, Finland; ZMLU—Zoological Museum, Lund University, Sweden; ZMUB—Zoological Collections, University Museum of Bergen, University of Bergen, Norway ZMUT— Zoological Museum, University of Turku, Finland.

Taxonomy Psychoda Latreille, 1796. Diagnosis (after Cordeiro et al. 2011): Labellum flattened, with 3–6 apical teeth; premental apodeme absent; antennae with 12–14 flagellomeres, flagellomeres distal to the 11th diminutive, often displaying fusions; ascoids with one posterior and two or three anterior branches; gonocoxites of male broadly separated; cercopod with a single retinaculum; female genital plate with two or (rarely) three lobes, usually carrying a median genital digit.

Psychoda cultella sp. n. Figs 1–3, 4a Psychoda sp (Salmela et al. 2007: 47, sampling locality) Chodopsycha sp (Salmela 2008: 54, sampling locality) Psychoda sp 1 (Ilmonen & Salmela 2010: 20; Salmela 2011: 25, sampling localities)

Material examined: Holotype, male. FINLAND, Li: Utsjoki, Kevo, Kutuniemi 7740:3500, 7.VIII. 2000 S. Neuvonen leg. (in alcohol, ZMUT, Fig. 1a). Left antenna broken, right antenna intact. Legs of the left side are missing, right mid and hind legs are present. Paratypes, 176 males: FINLAND, Li: Utsjoki, Kevo, Kutuniemi 7740:3500, 7.VIII. 2000 S. Neuvonen leg., 2 males (in alcohol, ZMUT); the same locality but 1.VI.–12.VII. 2000, 2 males (in alcohol, ZMUT); Li: Utsjoki, Skallovaara 7740:3500, 13.VII. 2000 S. Neuvonen leg., 1 male (slide mounted, ZMUT); Le: Enontekiö, Pahtavaara SE, 7625151:3321084, 11.VI.–19.VII. 2009 J. Salmela leg, 7 males (in alcohol, ZMUT); Lkoc: Kittilä, Narkivaara NE, 7584693:3409932, 1.–31.VIII. 2007, J. Salmela leg., 1 male (in alcohol, ZMUT); Ks: Taivalkoski, Ruokamaanoja, 7268984:3541491, 2.VII.–1.VIII. 2006 J. Salmela leg., 8 males (in alcohol, MZHF); Ks: Taivalkoski, Mäntymaanoja, 7269129:3542210, 3.VII.–1.VIII. 2006, J. Salmela leg., 12 males (in alcohol, ZMUT); Ks: Taivalkoski, Kurjenoja, 7282803:3553388, 2.VII.–1.VIII. 2006 J. Salmela leg., 11 males (in alcohol, ZMUT); Ks: Taivalkoski, Karhuoja 65.4161N° 27.9536E° 25.V.–23.VI.2007 J. Salmela leg., 1 male (slide mounted, ZMUT); Ks: Kuusamo, Saaripuro, 7357336:3611517, 1.VII.–3.VIII. 2005 J. Salmela leg., 1 male (in alcohol, ZMUT); Ks: Kuusamo, Kalliojoki, 7344909:3610594, 22.VI.–3.VIII. 2005 J. Salmela leg., 1

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male (in alcohol, ZMUT); Oba: Pudasjärvi, Murto–oja, 7267183:3525642, 1.VI.–3.VII. 2006 J. Salmela leg., 1 male (slide mounted, ZMUT); Oba: Pudasjärvi, Säynäjäoja, 7248094:3539234, 25.V.–23.VI. 2007 J. Salmela leg., 1 male (in alcohol, MZHF); Om: Kälviä, Kaakkurinneva, 7093674:3322690, 27.VII.–23.IX. 2008 J. Salmela leg., 1 male (in alcohol, ZMUT); Oa: Kauhajoki, Kärkikeidas SW, 6905266:3247527, 1.VIII.–14.IX. 2008 J. Salmela leg., 4 males (in alcohol, ZMUT); Oa: Kauhajoki, Kauhalammi SE, 6907401:3263801, 1.VII. 2008 J. Salmela leg., 1 male (slide mounted, ZMUT); the same locality and date, 11 males (in alcohol, ZMUT); St: Honkajoki, Hautakorpi, 6886031:3244032, 1.–30.VI. 2008 J. Salmela leg., 19 males (in alcohol, ZMUT); the same locality but 2.V.– 1.VI. 2008, 6 males (in alcohol, ZMUT); St: Kankaanpää, Kiviahde, 6877278:3260596, 1.–31.V. 2008 J. Salmela leg., 3 males (in alcohol, ZMUT); St: Kankaanpää, Lähdetneva 6864869:3261866, 1.VII.–10.VIII. 2008 J. Salmela leg., 2 males (in alcohol, ZMUT); St: Kankaanpää, Hevoshaankeidas, 6876609:3260476, 1.VII.–9.VIII. 2008 J. Salmela leg., 3 males (in alcohol, ZMUT); Sb: Vieremä, Kurkipuro, 7093441:3483640, 18.VI.–13.VII. 2008, J. Salmela leg., 3 males (in alcohol, ZMUT); Kb: Nurmes, Niinimäki, 7053724:3615945, 21.VI.–16.VII. 2008 J. Salmela leg., 17 males (in alcohol, ZMUT); Kb: Nurmes, Rajapuro, 7055314:3624158, 20.VI.–15.VII. 2008 J. Salmela leg., 26 males (in alcohol, ZMUT); Kb: Rautavaara, Risupuro 7033949:3565089, 15.VIII.–21.IX. 2008 J. Salmela leg., 1 male (in alcohol, ZMUT); Kb: Rautavaara, Nurmespuro, 7034411:3560105, 19.VI.–14.VII. 2008 J. Salmela leg., 14 males (in alcohol, ZMUT); Kb: Rautavaara, Pankapuro, 7060512:3571797, 15.VIII.–21.IX. 2008 J. Salmela leg., 1 male (in alcohol, ZMUT); the same locality but 18.V.–21.VI. 2008, 1 male (slide mounted, ZMUT); Kb: Rautavaara, Ukonpuro, 7055557:3572303, 19.VI.–16.VII. 2008 J. Salmela leg., 3 males (slide mounted, ZMUT); the same locality but 15.VIII.–21.IX. 2008, 1 male (in alcohol, ZMUT); Tb: Jyväskylä, Sarvivuori 62.1820N° 25.7473E°, 8.VI.2002 J. Salmela leg., 1 male (slide mounted, ZMUT); Tb: Petäjävesi, Purokoski 62.3768 N° 25.0683E°, 1.VIII.–9.X.2007 J. Salmela & L. Mikonranta leg., 1 male (slide mounted, ZMLU); Ab: Kiikala, Varesjoki, 670746:331380, 1.VIII.–1.IX. 2006 J. Ilmonen leg., 1 male (in alcohol, ZMUT). NORWAY, Finnmark: Sør–Varanger, Pasvik, 96–høyden, Russevann, 69.44497°N, 29.89904 °E, 60 m.a.s.l., 19.–24.VI.2010, Finnmarksprosjektet leg., 1 male (slide mounted, ZMUB, GenBank accession number JQ349634), same locality but 24.VI–20.VII.2010, Finnmarksprosjektet leg., 1 male (slide mounted, ZMUB, GenBank accession number JQ349633). Finnmark: Sør–Varanger, Pasvik, Sametijohka, 69.40106°N, 29.71923°E, 43 m.a.s.l., 24.VI– 20.VII.2010, Finnmarksprosjektet leg., 1 male (slide mounted, ZMUB), same locality but 21.VIII–6.IX.2010, 1 male (slide mounted, ZMUB), Finnmark: Porsanger, Baukop Vuolit Gealbbotjavri, 70.20469°N, 24.90605°E, 26 m.a.sl., 26.VII–25.VIII.2010, Finnmarksprosjektet leg., 2 males (slide mounted, ZMUB), Finnmark: Kautokeino, Nahpoljohka, 69.21029°N, 23.76200° E, 320 m.a.s.l., 6–20.VIII.2010, Finnmarksprosjektet leg., 1 male (slide mounted, ZMUB). Hordaland: Lindås, Vollom 60.6382 °N 5.2142 °E), 23.7–3.8.1991, K.H. Thunes leg. 1 male, ZMUB #8419. Hordaland: Os, Raudli (60.274 °N, 5.484 °E), 25.VII–1.VIII.1991. G.A. Halvorsen leg. 1 male, ZMUB #B8416. Diagnosis. Small species, wing length 1.4–1.9 mm. Antennae 16-segmented, last four flagellomeres globular. Flagellomere 11 with very short neck, flagellomeres 12–14 without necks; last three flagellomeres very small, fused or almost so. Labellum with five terminal digital projections. Gonostylus elongated, almost straight. Aedeagus ca. 1.7 times longer than gonostylus. Distiphalli bipartite: ventral phallomere apically roundish or truncated, resembling blunt knife in lateral view; dorsal phallomere narrow and apically pointed, basally curved. Basiphallus (in lateral view) widened sub-basally. Cercopod ca. 1.9 times longer than gonostylus. Description. Male. Head (Fig. 2a). Posterior interocular distance 27 (21–36, n=8), anterior distance 42 (34– 47, n=8). Width of clypeus 75 (63–89, n=8), length 67 (59–76, n=8). Labrum length 76 (56–119, n=8). Labellum with five terminal digital projections (Fig. 2b). Antennae 16-segmented. Scape prolonged (length 69, 57–81, n=8), pedicel globular (length 52, 45–60, n=8). Flagellomeres 1–10 pitcher-shaped, flagellomere 11 with very short neck. Flagellomeres 12–14 globular, wholly or partially fused (Fig. 2c). Ascoids 3-branched, present in flagellomeres 1– 11. Lengths of flagellomeres: f1 83 (70–96, n=8), f2 86 (75–99, n=8), f3 87 (80–101, n=8), f4 87 (80–98, n=8), f5 85 (77–100, n=7), f6 85 (75–100, n=7), f7 84 (73–85, n=6), f8 78 (73–85, n=6), f9 72 (59–79, n=6), f10 66 (50–81, n=5), f11 34 (30–39, n=4), f12 14 (11–16, n=4), f13 11 (9–13, n=4), f14 10 (9–12, n=4). Palpi 4-segmented, elongated, palpomeres 1–3 almost equal in length, p4 longest. Lengths of palpomeres: p1 68 (54–77, n=8), p2 64 (57–72, n=8), p3 63 (50–82, n=8), p4 78 (63–93, n=8). Thorax brownish, without distinctive features. Wing length 1.6 mm (1.4–1.9 mm, n=8). Wing venation, see Fig. 1b. Halteres whitish, hyaline. Legs brownish. Tarsal claws curved and sharply pointed. Abdomen brownish. Hypandrium ribbon-like, nearly straight but slightly expanded medially. Gonocoxite slightly bulbous in basal part. Gonostylus elongated, almost straight (Fig. 3a), length 118

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FIGURE 1. Psychoda cultella sp. n. a) Holotype male, habitus, lateral view. b) Paratype male, wing. Scale bars: a 0.5 mm, b 0.2 mm.

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FIGURE 2. Psychoda cultella sp. n., paratype male. a) head, dorsal view. b) Labellum. c) Apical flagellomeres (f 11–f 16) of antenna. Scale bars: 0.1 mm.

(100–136, n=8). Length of aedeagus 202 (168–250, n=8). Distiphalli bipartite: ventral phallomere apically roundish or truncated, resembling blunt knife in lateral view; dorsal phallomere narrow and apically pointed, basally curved (Fig. 3a,b, 4a). Basiphallus widened sub-basally in lateral view (Fig. 3b), narrow in dorsal view (Fig. 3a). Subaedeagal plate short, distally setose (Fig. 3a, 4a). Cercopod elongated (Fig. 3c), length 229 (210–262, n=8), retinaculum length 65 (57–72, n=8). Epandrium as in Fig. 3d, internal structures not illustrated. Female unknown. Etymology. From latin cultella, knife, referring to the shape of the ventral phallomere. DNA barcode. Two Norwegian Psychoda cultella sp. n. specimens (GenBank accession numbers JQ349633 and JQ349634), have been sequenced for their COI–5P barcoding region (658 bp). Kimura parameter 2 (K2P) distance between these specimens was small, only 0.46 %, i.e. three nucleotides between the sequences were different in the following sites: 118 (C/T), 256 (A/C), 616 (C/T). These differences have no effect in the coding of amino acids. Compared to some Fennoscandian Psychoda species, P. cultella sp. n. seems to have rather isolated position: its pair-wise K2P distances between P. satchelli Quate, P. grisescens Tonnoir, P. phalaenoides Linnaeus and P. lobata Tonnoir are each about 10 % (G.M. Kvifte, unpublished data).

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FIGURE 3. Psychoda cultella sp. n., paratype male. a) Gonocoxites, gonostyli and aedeagus, dorsal view. b) Aedeagus, lateral view. c) Cercopod and retinaculum, lateral view. d) Epandrium, ventral view. Scale bars: 0.1 mm.

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FIGURE 4. a) Psychoda cultella sp. n. and b) Psychoda grisescens Tonnoir, male aedeagal complex, dorsal view.

Discussion Distribution and ecology. Psychoda cultella sp. n. is quite common and locally abundant in the middle and northern boreal ecoregions of Finland, especially in the provinces Oa, St, Kb and Ks. All Finnish records are based on Malaise trap samples. In Norway the species seems to be less abundant; however it appears to be rather widespread in the Northernmost county Finnmark (see Fig. 5). The southernmost known locality for the species is in the southern boreal zone (Finland: Kiikala) and the northernmost localities in the subalpine zone (Norway: Baukop Vuolit Gealbbotjavri). Two of the Norwegian records are from the oceanic southwestern part of the country (Hordaland). Raudli is a heterogenous mixed forest site dominated by Pinus sylvestris, Betula pubescens, Picea sp., Quercus robur and Sorbus aucuparia whereas Vollom is the northernmost beech forest (Fagus sylvatica) in Europe (Thunes 1994).

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Psychoda cultella sp. n. has been collected from mesic riparian coniferous forests of headwater streams, spruce mires and springs and rarely from fens and subalpine mountain birch forests (Betula pubescens ssp. czerepanovii). The specimens from Norway: Hordaland were reared from one of the wood-decaying polyporous fungi Fomes fomentarius or Piptoporus betulinus (see Thunes 1994 for details); it is thus likely that Psychoda cultella sp.n. is a saproxylic species with larval development in fungal fruiting bodies. Immature stages of most other Psychodini are saprobionts, dwelling in animal excrements or in habitats rich of organic material (e.g. Satchell 1947; Withers 1988; Svensson 2009). European Psychoda and other Psychodidae species breeding in fungal fruiting bodies were reviewed in Bernotiené & Rimšaité (2009).

FIGURE 5. Known localities for Psychoda cultella sp. n.

Systematics. We refrain from placing P. cultella sp. n. in any described subgenus. In Ježek and van Harten's (2005) key to world Psychoda s.lat. genera, Psychoda cultella sp. n. keys to couplet 7, coming closest to Psychodula Ježek, 1984 (type species Psychoda minuta Banks, 1894). P. cultella sp. n. does not, however, have the conspicuous sclerotizations flanking the aedeagus in Psychodula s.str. Apart from the highly variable antennal apex, no obvious characters link P. cultella sp. n. to Psychodula.

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Salmela (2008) supposed P. cultella sp. n. to be a species of Chodopsycha Ježek, 1984 (type species Psychoda lobata Tonnoir, 1940), which is not unlikely given its likely mycobiont larvae. However, the new species does not fit completely with Ježek's (1984) diagnosis of Chodopsycha: in P. cultella sp. n. flagellomeres 13 and 14 are occasionally fused and flagellomere 11 has a clear distal border. Psychoda cultella sp.n. keys to Psychoda grisescens Tonnoir, 1922 in Withers (1989) as the hypandrium is nearly square-shaped. But the two species are clearly distinct and can be separated by comparison of the following characters: Psychoda grisescens (Fig. 4b) has the hypandrium comparatively broad, the subaedeagal plate elongate and hairless and the distiphallus short, reaching the tip of gonocoxite only; whereas Psychoda cultella sp. n. has the hypandrium comparatively narrow (Fig. 4a), the subaedeagal plate short and setose (Fig. 3a, 4a) and the distiphallus elongate, reaching longer than tip of the gonocoxite (Fig. 3a, 4a). Due to the immense diversity of the genus, it will probably eventually be necessary to subdivide Psychoda. This new classification should be based on detailed examination of a wider range of characters (Ježek & van Harten 2005, p. 207). We recommend that future authors seek to build a firm phylogenetic framework using both morphological and molecular characters in order to establish a stable and systematically sound classification.

Acknowledgements We are grateful to Torbjørn Ekrem (Vitenskapsmuseet, Trondheim, Norway) and Valerie Levesque-Beaudin (Biodiversity Institute of Ontario, University of Guelph, Canada) for help in obtaining the sequence data. We thank Phil Withers and two anonymous referees for constructive criticism and comments on the manuscript. Gunnar Mikalsen Kvifte was partially funded by Artsdatabanken (the Norwegian Taxonomy Initiative) through their project “Kartlegging av insekter knyttet til vann og fuktige habitater i Finnmark” (“Finnmarksprosjektet”). Jukka Salmela was supported by a grant from Finnish Cultural Foundation (Suomen Kultuurirahasto).

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