© Entomologica Fennica. 28 October 2009 Identification and taxonomy of the West Palaearctic species of Tachina Meigen (Diptera: Tachinidae) based on male terminalia and molecular analyses Hana Novotna, Jaromir Vafihara*, Andrea Tothova, Natalia Murarikova, Petr Bejdak, Michal Tkoc & Rudolf Rozkosny J M. & Novotna, H., Vafihara, ., Tothova, A., Murarikova, N., Bejdak, P., Tkoé, Rozkosny, R. 2009: Identification and taxonomy ofthe West Palaearctic species of Tachina Meigen (Diptera: Tachinidae) based on male terminalia and molecu- lar analyses. — Entomol. Fennica 20: 139—169. The male postabdominal structures of the West Palaearctic species of the genus Tachina are described. A new identification key is given. Characters are illus- trated by original pen drawings and deep focus micrographs, some ofthem for the first time. The results are documented by molecular analyses (based on CO°I, Cyt°b, 12S, and 16S rDNA). This approach solves old taxonomical discrepan- cies, which resulted in these conclusions: 1) the taxonomic concept ofthe genus was evaluated; 2) the position ofthe present subgenus Tachina s.str. seems to be untenable: T. grossa (Linnaeus, 1758) could be categorized inside existing sub- genus Tachina s.str. and a new subgenus could be created for T. magna (Giglio- Tos, 1890); 3) an expected new species from subgenus Eudoromyia was con- firmed; 4) T. nigrolzirta (Stein, 1924) having been resurrected from synonymy was confirmed as a valid species; 5) some differences between central European and Japanese specimens of T. nupta (Rondani, 1859) were found. H. Novotna, J. Vannara, A. Tothova, N. Murarikova, P. Bejdak, M. T[(06 & R. Rozko§ny, Masaryk University, Faculty ofScience, Department ofBotany and Zoology, Kotlafska 2, 611°37 Brno, Czech Republic; >"Corresponding author ’5 e—mail.‘ [email protected] Received 1 7 October 2008, accepted 18 May 2009 1 . Introduction fauna appears to be richer than the West Palaearc- tic. The recorded number of species described Species of the genus Tachina occur in the Neo- from the former USSR and present China (Zimin tropical, Nearctic, Palaearctic, and Oriental Re- & Kolomietz 1984, Chao et al. 1998, O’Hara et gions but they are apparently missing in the al. 2009) comprises 44 and 51 taxa respectively, Afrotropical and Australasian Regions (O’Hara though both lists include a number ofvague syn- 2006). There is no known species with a Holarc- onyms. The West Palaearctic fauna is much more tic distribution (O’Hara & Wood 2004). The re- limited; the Fauna Europaea database includes 12 cent concept ofthe genus embraces 42 species in known species only (Tschorsnig et al. 2004). The the Palaearctic Region (Herting 1984, Herting & knowledge of Tachina taxonomy and phylogen- Dely-Draskovits 1993). The East Palaearctic etic relationships ofits species is still insufficient. 140 Novotnd et al. ° ENTOMOL. FENNICA Vol. 20 Table 1. Male postabdomen examined (for the DNA analyses see also Table 2). Abbreviations for countries: AT- Austria, CZ-Czech Rep., DE-Germany, ES-Spain, Fl-Finland, FR-France, GR-Greece, lT-ltaly, RU-European Russia, SE-Sweden, SK-Slovakia. For abbreviations of collectors see Material and methods. “*”: other speci- mens for DNA validation without postabdominal analysis. Taxon Provenance Males/DNA validation Collection Tachina (Eudoromyia) canariensis ES-Canary ls|s 2/1 TSCH, ZlE caste ES, IT, Serbia 4/3 CER, TSCH, VAN, ZlE corsicana GR, Tunisia 2/2 CER, ZlE fera AT, CZ, Fl, IT, SE, SK 20/2 BER, CER, VAN magnicornis CZ, Fl, IT, SE, SK 13/2 BAR, BER, TSCH, VAN nupta CZ, lran, lT, Japan 7/4 CER, lCH, VAN sp. FI, FR, SE, SK 12/5 BER, CEP, TSCH, VAN, ZlE (Tachina) magna ES, GR, IT 3/2 CER, TSCH, VAN (Servillia) lurida cz, Morocco 7/2 BAR, VAN nigrohirta DE, SK 4/2 CEP, TSCH, VAN ursina CZ, lT, RU, SK 6/1 BAR, CER, TSCH, VAN (Tachina) grossa CZ, SK 2/1 TSCH, VAN (Echinogaster) praeceps Kirgizia 2/2* VAN Outgroups Tachinidae Germaria ruflceps CZ, SK 2/1* CEP, VAN Gonia divisa SK 1/— CEP Gymnocheta viridis CZ 1/1* CEP Linnaemya picta SK 3/1 * CEP Nemoraea pellucida SK 1/— CEP Nowickia ferox CZ 1/1* BAR Peleteria rubescens SK 1/— CEP Phasia hemiptera CZ 1/— CEP Schineria tergestina SK 1/— CEP Winthemia variegata SK 1/— CEP Rhinophoridae Stevenia atramentaria CZ 1/— BAR The hitherto available keys are often based on eX- (Herting & Dely-Draskovits 1993, Tschorsnig et tensively variable structures and/or colour char- al. 2004) comprises besides 12 valid species also acters, while the male and female terminalia are 45 available synonymic names. The present con- only rarely considered. Moreover, some present cept of species taxonomy and higher taxa of the keys have demonstrated a distinct species over- genus Tachina was published by Herting (1984) lapping for some ofthe frequently used key char- in his Catalogue where he recognized four sub- acteristics, see e.g. morphometry of fore claws genera. and tarsi, female frons of T. magnicornis and T. The significance of the specific differences fem, etc. in the identification key in Tschorsnig & found in structures constituting the male termina- Herting (1994). The large number of synonymic lia in Diptera is generally accepted (cf. McAlpine names found in each regional fauna could there- 1981, 1989). The first extensive modern and sig- fore be regarded, at least partly, as a consequence nificant study concerning the male postabdomen, ofthis situation. Thus, the West Palaearctic fauna with emphasis on the phallus and gonites of 240 ENTOMOL. FENNICA Vol. 20 ° West Palaearctic Tachina 141 species, was published by Verbeke (1962, 1963). was supported, but some genera from Exoristinae Four selected European species of the genus appear divergent. The monophyly of subfamilies Tachina were studied in detail. Verbeke’s inter- Tachininae and Phasinae was doubted. Besides pretation of specific structures corresponds with phylogeny in Tachinidae, DNA-markers to the the generally accepted epandrial hypothesis (Mc- species identification (barcoding) were also used. alpine 1981, Sinclair 2000). A subsequent impor- Augusti et al. (2005) offered species specific tant and extensive study was published by primers for Lydella thompsom' and Pseudo— Tschorsnig (1985). He discussed evolutionary perichaeta nigrolineata, which helped them to trends in characters currently used in identifica- find parasitoid larvae in caterpillars of Ostrinia tion keys to Tachinidae and stressed a necessary nubilalis (Lepidoptera, Crambidae), a pest of revision ofall the generally used characters. Alto- corn. Smith et al. (2006) studied an ability of spe- gether 423 species were studied and selected cific sequences ofDNA barcodes (CO I, ITS 1) to characters were compared for 32 species groups. differentiate cryptic species and their context The author’s attention was chiefly focused on the with their host specializations (Smith et al. 2007). groups not treated by Verbeke (1962). A key to Gariepy et al. (2007) summarized the PCR meth- tribes was also proposed as a result of Tschors- ods used in parasites and predators, and intro- nig’s comparative studies. Ofthe genus Tachina, duced Tachinidae as a suitable model group. T. fem, T. grossa and T. ursina were examined. Wood (1987) and Tschorsnig & Richter (1998) presented some structures of the male terminalia 2. Material and methods at the family level as a part of the Manual of Nearctic Diptera and that of the Palaearctic Dip- The nomenclature and systematic position of the tera, respectively. Pape (1992) examined several West Palaearctic Tachina species follow Herting characters ofthe male terminalia in his phylogen- & Dely-Draskovits (1993) and in several cases etic study concerning the Tachinidae family also Herting (1984). Terminology of dipteran Also & in their group. Tschorsnig Herting (1994) male terminalia was adopted from Sinclair (2000) to Central used some char- key European species but some of the terms used for specific structures acters of the male to a postabdomen although by Tschorsnig (1985) are also applied. Two terms lesser extent. are introduced as new: callus of syncercus and DNA were used rather analyses sporadically spine of syncercus. for the Tachinidae the last decade. family during The material was identified by C. Bergstrom, Vossbrinck & Friedman (1989) used primarily P. Cerretti, J. Cepelak, B. Herting, L. P. Mesnil, DNA sequences for tachinid phylogeny and did H. Novotna, R. Rozkosny, H.-P. Tschorsnig, J. not support a monophyletic status of Tachinidae Vanhara, and J. Ziegler. within Cyclorrhapha. Concerning rapid succes- sive it was also evolutionary separations, sug- The following collections were studied: gested that those relationships cannot be differen- tiated using gene sequences of 28S rDNA. BAR — Miroslav Bartak, Czech University ofAgricul— Nirmala et al. (2001) usedNemoraeapellucida as ture, Faculty of Agrobiology, Food and Natural Re— an example for Tachinidae for analysis of Calyp- sources, Department of Zoology and Fishery, 165 21 tratae. On the basis of 16S and 18S rDNA they Praha 6 — Suchdol, Czech Republic; [email protected] did not find differentiated relationships for fami- BER — Christer Bergstrom, saves vag 10, Uppsala, lies close to the Tachinidae. The largest phylo- SE—75263, Sweden; [email protected]— genetic studies of the family Tachinidae were nordia.se published by Stireman (2002, 2005). He used 55 CEP— Juraj Cepelak (late), coll. deposited partly with species of the subfamily Exoristinae and 2 nu- Jaromir Vafihara (corresponding author) clear rDNA and Different — genes (28S EF-1oc). CER Pierfilippo Cerretti, Universita degli Studi di types of analyses brought extensive but in many Roma “La Sapienza”, Dipartimento di Biologia cases contradictory results. Monophyly of the Animale e dell’Uomo, Viale dell’Universita 32, 00185 family Tachinidae and subfamily Exoristinae Roma, Italy; [email protected] 142 Novotnd et al. ° ENTOMOL. FENNICA Vol. 20 Table 2. Material under DNA analysis and GenBank accession numbers (for abbreviations used see Table 1 and Material and methods). Taxon Sex ldent. GenBank accession numbers /|oca|ity CO | Cyt B 12S 16S Tachina (Eudoromyia) canariensis 1 f CEP —— FJ222662 FJ222691 /ES: Canary Islands, Tenerife, N, Cruz de Luis; Apr.