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Pulmonata, Helicoidea, Hygromiidae) Ruthenica, 2019, vol. 29, No. 2: 77-86. © Ruthenica, 2019 Published online March 5, 2019 http: www.ruthenica.com On the phylogenetic relationships of Elbasania Schileyko et Fehér, 2017 (Pulmonata, Helicoidea, Hygromiidae) Marco T. NEIBER Universität Hamburg, Centrum für Naturkunde (CeNak), Zoologisches Museum, Abteilung Biodiversität der Tiere, Martin-Luther-King-Platz 3, 20146 Hamburg, GERMANY. E-Mail [email protected]; [email protected] ABSTRACT. The genus-group taxon Elbasania Schi- mainly on the basis of similarities of the dart appara- leyko et Fehér, 2017 has recently been introduced as a tus. subgenus of Metafruticicola Ihering, 1892 for a spe- In a comprehensive molecular phylogenetic study cies occurring in north-western Greece and Albania. Using mitochondrial and nuclear markers, the phyloge- of western Palearctic Helicoidea Rafinesque, 1815, netic relationships of Elbasania within Metafruticico- Razkin et al. [2015] classified the clade to which lini (Hygromiidae) are reconstructed. The results of hygromiids and related groups belong into three these analyses suggest that Elbasania is more closely newly delimited families: Canariellidae Schileyko, related to Hiltrudia Nordsieck, 1993, which has a range 1991, Geomitridae Boettger, 1909 and Hygromii- adjacent to that of Elbasania from Croatia to northern dae. The Hygromiidae were classified into three Albania, than to Metafruticicola. Elbasania shares subfamilies, Hygromiinae (including Trochulinae with Hiltrudia and also Cyrnotheba Germain, 1929 a Lindholm, 1927 and Monachainae Wenz, 1930 very characteristic microsculpture of the shell and an (1904)), Ciliellinae Schileyko, 1970 and Leptaxinae overall similar genital system, which however differs Boettger, 1909. However, the sampling of Hygromi- among these three taxa with regard to its internal struc- idae was focused on West European taxa and repre- tures, especially those of the penis. Therefore, it is suggested regarding Elbasania as a distinct genus sentatives of several family-group taxa were not here. included in the analyses. Neiber et al. [2017], on the basis of a sampling of almost all genus-group taxa, analysed the phylogenetic relationships of the Hy- Introduction gromiidae in the sense of Razkin et al. [2015] and showed that homoplasies are frequent and wide- The systematics of the Hygromiidae Tryon, 1866 spread in the family, but also that several geograph- has traditionally been based mainly on characters of ically largely coherent, although morphologically the genital system, especially on characters of the diverse, radiations can be recognized. On the basis dart apparatus. The systematic importance of these of their results, Neiber et al. [2017] proposed to genital characters has been controversially discussed divide the Hygromiidae into three subfamilies: Hy- and different classification schemes have been pro- gromiinae (with the tribes Hygromiini and Perfora- posed for the family and related groups [Hesse, tellini Neiber, Razkin et Hausdorf, 2017), Trochuli- 1921, 1931; Schileyko, 1970, 1972a, b, 1978a, b, nae (with the tribes Trochulini, Archaicini Schi- 1991, 2004, 2005; Nordsieck, 1987, 1993, 2017] leyko, 1978, Ashfordiini Neiber, Razkin et Hausdorf, resulting in an intricate history of classifications 2017, Caucasigenini Neiber, Razkin et Hausdorf, [for an overview, see Neiber et al., 2017: Supple- 2017, Ciliellini, Ganulini Neiber, Razkin et Hausdorf, mentary Text 1 and Supplementary Table S1]. The 2017, Halolimnohelicini Nordsieck, 1986, Monach- molecular genetic studies of Steinke et al. [2004] aini and Urticicolini Neiber, Razkin et Hausdorf, (see also Groenenberg et al. [2011] for correc- 2017) and Leptaxinae (with the tribes Leptaxini, tions), Koene and Schulenburg [2005] and Man- Cryptosaccini Neiber, Razkin et Hausdorf, 2017 ganelli et al. [2005] hinted at the possibility that the and Metafruticicolini Schileyko, 1972). basic premise of the systems of Schileyko [1972b, Schileyko [1972b] originally proposed Metafru- 1978b, 1991, 2004, 2005] and Nordsieck [1987, ticicolinae for Metafruticicola Ihering, 1892, Creti- 1993] may be too simplistic, i.e. to group species gena Schileyko, 1972 and the Caucasian to north- 78 M.T. Neiber east Anatolian genus Caucasocressa Hesse, 1921 al. [2017] that included representatives of all four with a completely reduced dart apparatus, i.e. with- subgenera and was also questioned by Schileyko out dart sac and accessory sac and without glandu- and Fehér [2017] and Bitzilekis et al. [2017] who lae mucosae. Nordsieck [1993] included additional- observed incongruencies with regard to anatomical ly Fruticocampylaea Kobelt, 1871, Shileykoia Hu- and/or conchological characters. However, Bank et dec, 1969 (= Fruticocampylaea; see Walther et al. al. [2013] expressly acknowledged that their revi- [2016]), Kalitinaia Hudec et Lezhawa, 1967, Cir- sion should be seen as a framework for testing cassina Hesse, 1921 and Hiltrudia Nordsieck, 1993 molecular versus morphological data and that their in the Metafruticicolini, although Fruticocampylaea, extensive distributional data may be used as the Kalitinaia and some Circassina have a single dart basis for biogeographical research programmes. sac with accessory sac and glandulae mucosae. Recently, Schileyko and Fehér [2017] described According to Neiber et al. [2017] Fruticocampy- Elbasania as another subgenus of Metafruticicola, laea and Circassina belong to the Caucasigenini with the disjunctly distributed nominal taxon Meta- (see also Neiber and Hausdorf [2013, 2015], Walther fruticicola occidentalis Subai, 1999 from Albania et al. [2016, 2018], Neiber et al. [2018]) and and north-western Greece [Subai, 1999; Bank et Kalitinaia to the Geomitridae. The Metafruticico- al., 2013] as type species, which differs from all lini, as delimited by Neiber et al. [2017], include other anatomically known Metafruticicola species aside from Metafruticicola (incl. Cretigena, Wester- by a very long vagina that has a very thick, well- lundia Kobelt, 1904 and Rothifruticicola Bank, Git- muscularized wall, a very short free oviduct and a tenberger et Neubert, 2013) also Hiltrudia and Cyr- rather long, tubular penial papilla. In the present notheba Germain, 1929. A relationship of Hiltrudia contribution, previously published data from Neiber with Metafruticicola has previously been supposed et al. [2017] and newly generated mitochondrial by Maassen [1979], Pintér and Szigethy [1979], and nuclear sequences are used to test the proposed Nordsieck [1993] and recently also by Schileyko classification of Elbasania as a subgenus of Meta- and Fehér [2017], whereas in other studies Hiltru- fruticicola. dia species were included in Ashfordia Taylor, 1917, which also has a completely reduced dart apparatus Material and methods [Hesse, 1934; Maassen, 1978] but was proposed to represent a tribe, Ashfordiini, in the Trochulinae by As representatives of the Metafruticicolini in the Neiber et al. [2017]. Cyrnotheba and Metafrutici- sense of Neiber et al. [2017], published nuclear and cola have been shown to form a mostly well- mitochondrial sequences belonging to the nominal supported clade by Neiber et al. [2017] and a close taxa Metafruticicola (Metafruticicola) pellita (Fé- relationship with Metafruticicola has also been sup- russac, 1832), Metafruticicola (Rothifruticicola) posed independently by Schileyko and Fehér [2017] nicosiana freytagi (Maltzan, 1883), Metafruticicola on the basis of similarities of the genital system (Westerlundia) noverca (Pfeiffer, 1853), Metafru- (penial papilla and flagellum) and the shell, whereas ticicola (Westerlundia) naxiana (Férussac, 1832), previous authors either suggested a possible rela- Metafruticicola (Cretigena) sublecta (Maltzan, tionship of Cyrnotheba with Monachoides or Hy- 1884), Cyrnotheba corsica (Shuttleworth, 1843), gromia [Giusti, Manganelli, 1987], the Hygromiini Hiltrudia mathildae (Westerlund, 1881) and Hiltru- [Nordsieck, 1993] or Monachainae [Schileyko, dia kusmici (Clessin, 1887) from the study of Neib- 2005]. er et al. [2017] were included in the analyses. Several questions with regard to the position of Additionally, nuclear and mitochondrial sequences the Metafruticicolini within the Hygromiidae, its were newly generated from a specimen of the type generic composition and the relationships of taxa species, Metafruticicola (Elbasania) occidentalis are at present not well-resolved or need further Subai, 1999, of the recently described subgenus investigations. Preliminary results by Caro et al. Elbasania from Albania and from another specimen [2017] based on an increased molecular sampling from Albania, here referred to as Metafruticicola suggest however that Hiltrudia, Cyrnotheba and (Elbasania) cf. occidentalis because of anatomical Metafruticicola indeed form a well-supported clade, differences. A specimen belonging to Hygromia (Hy- which may represent the sister group of all other gromia) cinctella (Draparnaud, 1801) was used as Hygromiidae. outgroup for rooting phylogenetic trees. Data on Bank et al. [2013] revised the genus Metafru- sampling sites, voucher numbers, GenBank acces- ticicola and recognized four subgenera, i.e. Meta- sion numbers and the classification of the speci- fruticicola s.str., Westerlundia, Cretigena and Rothi- mens used are compiled in Tables 1–2. fruticicola, which were characterized by differenc- Total genomic DNA was extracted from the es in the sculpture of the shell. The system of Bank tissue samples following a slightly modified version
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