Hymenoptera, Tenthredinidae)
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Zootaxa 4614 (3): 511–528 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4614.3.5 http://zoobank.org/urn:lsid:zoobank.org:pub:0D818E35-181B-4F2C-845C-3E456D45804B The West Palaearctic Pseudodineura and Endophytus species (Hymenoptera, Tenthredinidae) ANDREW LISTON1,4, MARKO PROUS1,2 & HEGE VÅRDAL3 1Senckenberg Deutsches Entomologisches Institut, Eberswalder Straße 90, 15374 Müncheberg, Germany. E-mail: [email protected] 2Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia. E-mail: [email protected] 3Swedish Museum of Natural History, Box 50007, SE-10405 Stockholm, Sweden. E-mail: [email protected] 4Corresponding author Abstract Six valid species of Pseudodineura are now recognised as occurring in the West Palaearctic, and the only described species of the related genus Endophytus. Larvae of all species are leaf-miners in Ranunculaceae. An identification key to adults is provided, followed by species commentaries which include summarised data on taxonomy, larval host plants, and distribution, with particular reference to Sweden. Whereas identification of some specimens using morphological characters may not be possible, each species apparently has a distinct COI barcode sequence. Pseudodineura heringi (Enslin, 1921) is a new junior synonym of P. parvula (Klug, 1816). Pseudodineura mocsaryi Zombori, 1976 and P. scaligera Zombori, 1979 are new junior synonyms of P. clematidisrectae Hering, 1935. Lectotypes are designated for: Dolerus minutus Hartig, 1837, Pelmatopus clematidis Hering, 1932, P. enslini Hering, 1923, P. heringi Enslin, 1921, and P. mentiens var. konowi Enslin, 1921. Key words: Symphyta, Nematinae, sawflies, Sweden, key, taxonomy, distribution Introduction Pseudodineura Konow, 1885 is a small, Holarctic genus of nematine sawflies, containing only ten valid species (Smith 1976, Smith et al. 2010, and this work). Endophytus Hering, 1934 is monotypic, and only known from the Palaearctic (Taeger et al. 2010). These genera form together with the Nearctic Kerita (3 species) a monophyletic lineage which is sometimes called the tribe Pseudodineurini Benson, 1938 (Prous et al. 2014). The larvae of Pseu- dodineura and Endophytus mine the leaves of Ranunculaceae, whereas the only known host of a Kerita species is Mertensia virginica, Boraginaceae (Smith 2009). With the exception of fuscula, adults of the European Pseudodin- eura species are rarely collected, probably because of their small size, short flight period, and generally narrow host plant range (Altenhofer & Pschorn-Walcher 2006). Furthermore, some species are mono- or oligophagous on plant taxa such as Pulsatilla species, Anemone sylvestris and Clematis recta, which have suffered massive declines in large parts of their European ranges. On the other hand, where they do occur, Pseudodineura species can sometimes reach high population levels, made evident by abundance of their conspicuous leaf-mines. Two species, enslini and mentiens, are regularly found in parks and gardens, and not infrequently regarded as a nuisance, because at high population densities they render the leaves of their hosts unsightly, and can reduce the production of inflorescences (Buhr 1941, Gerlach 2005). Material and methods The names of collections referred to in the text are abbreviated as follows: Accepted by A. Taeger: 23 Apr. 2019; published: 11 Jun. 2019 511 FMNH Finnish Museum of Natural History, Helsinki, Finland HNHM Hungarian Natural History Museum, Budapest, Hungary MNHN Muséum national d’Histoire naturelle, Paris, France MSNG Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy MZLU Lunds universitet, Entomology Collection, Lund, Sweden NHRS Naturhistoriska riksmuseet, Stockholm, Sweden SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany VVCT Private collection of Veli Vikberg, Turenki, Finland ZMHB Naturkundemuseum, Berlin, Germany ZSM Zoologische Staatssammlung, Munich, Germany Registration numbers of specimens allocated by the SDEI (five digits, omitting the prefix „DEI-GISHym“) are cited in the figure captions, thus enabling the corresponding collection data to be located in the list of specimens examined. An asterisk (*) preceding a country name in the list of specimens examined indicates that we know of no previous published records from that country. Morphological terminology follows Viitasaari (2002), except for the use of sawtooth instead of serrula (see Malagón-Aldana et al. 2017). Genitalia were macerated in room-temperature 10–15% KOH for four to five hours before being mounted temporarily in glycerine on glass slides for photography. After photography, the dissected parts were gummed with Berlese fluid to a card fixed to the specimen’s pin. Results Character assessment Antennae. We counted the number of flagellomeres of 79 females and 25 males of the six West Palaearctic Pseudo- dineura species in the SDEI collection. A few additional specimens lacked antennae and could not be scored. The majority have seven flagellomeres (both antennae: 7/7), except: three females of 48 fuscula, two with 7/8 flagel- lomeres, and one with 8/8; two females and one male of 24 parvula, the females with 8/8 and 7/8 flagellomeres, the male with 8/8. The “additional” tenth antennomere is variable in proportions, even between each antenna of the same specimen. It seems that the presence of an additional flagellomere may be rather infrequent in European populations of Pseudodineura. Until the taxonomic significance, if any, of the number of flagellomeres in this genus has been better clarified, this character should not be given undue weight. Smith et al. (2010) compared the single Alaskan specimen described in that paper as P. kasatochi D. R. Smith with the otherwise apparently similar P. heringi, and attributed high diagnostic significance to the holotype of kasatochi having both antennae with eight flagellomeres. More significant are the proportions of the flagellomeres, which in some cases show clear differences between species in the length of a basal flagellomere compared to a more apical one, or the ratio of width to length in a single flagellomere. Colour pattern. The species are similarly coloured. The head and body are black, with more or less pale mouth- parts and legs, and in some species with pale markings (yellow-brown) on the thorax and abdomen. Although the range of variability in body coloration within most species is rather narrow, P. mentiens is an exception: the abdo- men may be almost completely black, to entirely pale. The colour of the metafemur is constant within most species, except clematidis: entirely pale in mentiens and clematidisrectae, and clearly partly black in all the others. Males of Pseudodineura clematidis can have entirely pale, or slightly dark-marked femora. We have used femur colour as the entry character in our identification key, but one should examine the specimen from more than one angle, or look at more than one leg, because an entirely pale femur, which is both translucent and reflective, may still look dark if it is near to black areas of the body. Genetic data. During the past few years, genetic data (mainly COI barcodes) for all of the European Pseudodin- eura species, and Endophytus anemones, have become available (Boevé et al. 2009, Schmidt et al. 2017). In BOLD Systems (boldsystems.org), all specimens of each species group in single BINs (Barcode Index Number Uniform Resource Identifier), except that there is no BIN assigned yet to P. clematidis, because of the lack of sufficiently long barcode sequences. Based on the partial COI barcode of P. clematidis, the closest is P. clematidisrectae differ- ing at least by 4.6%. Barcoding can therefore be recommended for species identification in these genera. 512 · Zootaxa 4614 (3) © 2019 Magnolia Press LISTON ET AL We retain Endophytus as a separate genus from Pseudodineura, even though most of the morphological char- acters which distinguish them could be regarded as representing species-level variability within a single genus. Although the differences in wing venation have the quality traditionally associated with genus-level classification, these character states are very unstable in Endophytus. The main reason for treating the genera as distinct is their large genetic distance from each other (see Prous et al. 2014, fig. 2). At present, the genetic distances between En- dophytus, Pseudodineura and Kerita appear to be rather similar, but their relationships could still change when more data becomes available. FIGURES 1–8. Pseudodineura species. 1, enslini larva (17050). 2, fuscula leaf-mine in Ranunculus sp. (31922). 3, clematidis- rectae ♀ (mocsaryi holotype, 31821). 4, parvula ♀ (31886). 5, clematidisrectae ♀ (11441) antenna. 6, clematidis ♀ (31812) antenna. 7, mentiens ♀ (19816) leg colour. 8, clematidis ♀ (88876) leg colour. Scale bars 1mm. WEST PALAEARCTIC PSEUDODINEURA Zootaxa 4614 (3) © 2019 Magnolia Press · 513 Genitalia. For the sawsheath, see under that term, below. The sawteeth of the lancet of Endophytus anemones are flat (Fig. 33), contrasting with the prominently pro- jecting lobes of all Palaearctic Pseudodineura (Figs 27–32). Within Pseudodineura, examination of the lancet is of limited use for identification: only that of fuscula is readily recognisable, by its relatively much larger sawteeth. The lancets of the other species are very similar,