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Systematics of Pipizini and Taxonomy of European Pipiza Fallã©N See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/264505907 Systematics of Pipizini and taxonomy of European Pipiza Fallén: Molecular and morphological evidence (Diptera, Syrphidae) Article in Zoologica Scripta · May 2013 DOI: 10.1111/zsc.12005 CITATIONS READS 13 300 6 authors, including: Gunilla Ståhls Jelena Ačanski University of Helsinki BioSense Institute, University of Novi Sad 138 PUBLICATIONS 867 CITATIONS 37 PUBLICATIONS 58 CITATIONS SEE PROFILE SEE PROFILE Ana Stefanović Grković University of Novi Sad 7 PUBLICATIONS 43 CITATIONS SEE PROFILE All content following this page was uploaded by Ana Stefanović Grković on 15 September 2014. The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. Zoologica Scripta Systematics of Pipizini and taxonomy of European Pipiza Falle´ n: molecular and morphological evidence (Diptera, Syrphidae) † ANTE VUJIC´ ,GUNILLA STA˚ HLS,JELENA ACˇ ANSKI,HANS BARTSCH ,RUNE BYGEBJERG &ANA STEFANOVIC´ Submitted: 4 June 2012 Vujic´, A., Sta˚hls, G., Acˇanski, J., Bartsch, H., Bygebjerg, R. & Stefanovic´, A. (2013). Sys- Accepted: 21 November 2012 tematics of Pipizini and taxonomy of European Pipiza Falle´n: molecular and morphological doi:10.1111/zsc.12005 evidence (Diptera, Syrphidae). —Zoologica Scripta, 42, 288–305. In the present work the monophyly and molecular phylogenetic relationships of the genera of tribe Pipizini (Syrphidae) were investigated based on mitochondrial cytochrome c oxi- dase subunit I (COI) and nuclear 28S rDNA sequences, and the relationships among spe- cies of genus Pipiza Falle´n, 1810 based on mtDNA COI sequences. Molecular phylogenetic analyses of Pipizini supported Pipiza as monophyletic and as sister group to all other Pipizini, and resolved other Pipizini genera as monophyletic lineages except for genus Heringia Rondani, 1856. To recognize the distinctness and maintain the monophyly the genus Heringia was redefined, generic rank was assigned to Neocnemodon Goffe, 1944 stat. n., and the genus Claussenia Vujic´ & Sta˚hls gen. n., type-species Claussenia hispanica (Strobl, 1909), was described. A revision of the European Pipiza species, including a discus- sion of taxonomic characters and a morphological redefinition of all included species, is presented. One new species, Pipiza laurusi Vujic´ & Sta˚hls sp. n. was described. The taxa Pipiza carbonaria Meigen, 1822; Pipiza fasciata, Meigen 1822; Pipiza lugubris (Fabricius, 1775), Pipiza noctiluca (Linneaues, 1758), Pipiza notata Meigen, 1822 were redefined. Lecto- types are designated for 17 taxa, and neotypes were designated for seven taxa. Fourteen new synonymies were proposed. Male genitalia were illustrated for all the species, and a key of the 12 European species for males and females was provided. Geometric morpho- metrics of wing landmarks and extended sampling of mtDNA COI sequences was employed to delimitate taxa of the P. noctiluca and P. lugubris complexes. Despite subtle morphological differences, wing geometric morphometrics variables of wing size and shape showed highly significant differences among species within P. noctiluca and P. lugubris com- plexes, which were supported by the molecular data. Corresponding author: Ante Vujic´, Department of Biology and Ecology, Faculty of Science, University of Novi Sad, Trg Dositeja Obradovic´a 2, 21000 Novi Sad, Serbia. E-mail: ante. [email protected] Gunilla Sta˚hls, Entomology Department, Finnish Museum of Natural History, PO Box 17, FI-00014 University of Helsinki, Finland. E-mail: gunilla.stahls@helsinki.fi Jelena Acˇanski, Department of Biology and Ecology, Faculty of Science, University of Novi Sad, Trg Dositeja Obradovic´a 2, 21000 Novi Sad, Serbia. E-mail: [email protected] Rune Bygebjerg, Museum of Zoology, Lund University, Helgonava¨gen 3, SE-223 62 Lund, Sweden. E-mail: [email protected] Ana Stefanovic´, Department of Biology and Ecology, Faculty of Science, University of Novi Sad, Trg Dositeja Obradovic´a 2, 21000 Novi Sad, Serbia. E-mail: [email protected] †Ja¨rfa¨lla, Sweden. Deceased 4 April, 2011. Introduction some species have spots on the abdomen. A pilose postpr- The monophyletic tribe Pipizini (Diptera, Syrphidae) is a onotum, a flat face lacking a facial knob and entirely cov- sister group to subfamily Syrphinae (Sta˚hls et al. 2003). All ered in long pilosity, and wing vein R4 + 5 straight, not Pipizini are small to medium sized, dark coloured, but sinuate, and crossvein R-M perpendicular ending before 288 ª 2013 The Authors d Zoologica Scripta ª 2013 The Norwegian Academy of Science and Letters, 42, 3, May 2013, pp 288–305 A. Vujic´ et al. d Systematics of Pipizini and taxonomy of Pipiza middle of discal cell readily separates this taxon from all Pipiza noctiluca (Linnaeus, 1758) and Pipiza quadrimaculata other hoverflies. The family-group name Pipizini was first (Panzer, 1802). In The Biosystematic Database of World introduced by Williston (1885) (as Pipizinae). Pipiza is the Diptera (Pape & Thompson 2012), available names were type genus of the tribe. The tribe comprises six genera, listed for 15 species occurring in Europe. Cryptopipiza Mutin, 1998, Heringia Rondani, 1856, Pipiza Molecular phylogenetic studies of Syrphidae taxa have Falle´n, 1810, Pipizella Rondani, 1856, Trichopsomyia Willis- frequently employed parsimony analyses of the mitochon- ton, 1888 and Triglyphus Loew, 1840, with altogether drial cytochrome c oxidase subunit I (COI) and nuclear approx. There were 63 species in Europe. Cryptopipiza is 28S ribosomal RNA (28S) gene regions, and these gene an Old World monotypic genus, and with Pipiza, Pipizella regions have shown levels of variability informative for and Heringia sensu lato are confined to the Holarctic both generic level and species level phylogenies (e.g. Sta˚hls region. Triglyphus and Trichopsomyia comprise only few et al. 2004; Mengual et al. 2008a; Vujic´ et al. 2012). The species in the Old World, but are speciose in the New mtDNA COI gene was informative in several studies of World and ⁄ or Australian regions. Neocnemodon Goffe, hoverfly species level molecular phylogenetic relationships, 1944 was originally described as the genus Cnemodon Eg- but has showed low and ⁄ or overlapping uncorrected pair- ger, 1865 preoccupied by Cnemodon Schoenherr, 1823 wise sequence divergences between closely related taxa (synonym of Phytoscaphus Schoenherr), but subsequently (e.g. Claussen & Sta˚hls 2007; Mengual et al. 2008b; Milan- treated as subgenus of Heringia (Rotheray & Gilbert kov et al. 2008; Sta˚hls et al. 2008). 1989). Recent taxonomic revisions or reviews including Geometric morphometrics is a useful tool to detect European Pipizini taxa are those of Claussen et al. (1994) minimal morphological variations, which often are unde- on the genus Heringia s.l., Vujic´ (1999) on the Heringia tectable by classical morphometrical studies (Zelditch subgenus Neocnemodon, Vujic´ et al. (2008) on the Pipiza et al. 2004; Vilemant et al. 2007). Geometric morpho- luteitarsis group and van Steenis & Lucas (2011) on the metrics has mostly been used to discriminate between genus Pipizella. The generic relationships of Pipizini groups defined a priori using morphological or biologi- remain, however, unstudied. cal information (Baylac et al. 2003). Wing shape in Species of genus Pipiza are medium-sized, blackish hov- insects (e.g. Diptera, Hymenoptera and Odonata) has erflies, although some species have one or two pairs of been shown to be highly heritable and less sensitive to pale spots on the abdomen. Adults prefer habitats on the environmental changes than the wing size (Birdsall et al. forest edge and larvae are predators of gall-forming aphids 2000; Moraes et al. 2004; Mezey & Houle 2005; Dwor- (Speight 2010). Diagnostic characters for this genus are kin & Gibson 2006; Yeaman et al. 2010). Wings of flies the following: anterior anepisternum and katepisternum usually have strong and clear veins and vein junctions, bare or with very short pilosity, katepimeron with short with very little positional variation. Geometric morpho- hairs, male with distinct conically produced frontal promi- metrics applied to wing vein junctions was successfully nence, hind trochanter of male simple, without ventral used to resolve taxonomic uncertainties in species-pairs spur; male genitalia with two clear apomorphies: surstylus of the family Drosophilidae (Moraes et al. 2004), of clo- with well-developed dorsal semicircular lobe, lower gonoc- sely related the family Syrphidae (Ludosˇki et al. 2008; ercus sickle-cell like shape (adapted based on Thompson Francuski et al. 2011) and for other insects, for example, & Rotheray 1998). Hymenoptera (Baracchi et al. 2011; Kandemir et al. The catalogue of Palaearctic Diptera (Peck 1988) listed 2011). In our study, wing morphometry was used to dis- 17 species for the region, of which 12 were distributed in cern between and delimit species of the Pipiza lugubris Europe. Violovitsh (1988) revised the Palaearctic Pipiza and P. noctiluca complexes. species including 15 species, but the identity of most The aims of the present study are fourfold, to (i) eluci- European Pipiza still remained convoluted. Hence, the date the molecular phylogenetic relationships of the Pipiz- genus was still in need of a revision, as pointed out by ini genera; (ii) to assess the monophyly
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