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Notes on Enidae, 8. Systematics of Some Turkish Enini Previously

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Notes on Enidae, 8. Systematics of Some Turkish Enini Previously B2016-21-Gargely:Basteria-2015 11/19/2016 8:40 PM Page 177 Notes on Enidae, 8. Systematics of some Turkish Enini previously assigned to Imparietula and Pseudochondrula , with the description of two new genera (Gastropoda, Pulmonata) Barna P Áll-Gergely Department of Biology, Shinshu University, Matsumoto 390-8621, Japan; [email protected] Ruud A. Bank Chopinlaan 21, 9603 AM Hoogezand, the Netherlands; [email protected] [corresponding author] Introduction 177 We have investigated the genital system of several Enidae taxa from the northeastern part of Turkey that The land snail family Enidae B.B. Woodward, 1903 have previously been assigned to the genera Impari - (1880) can be subdivided into two subfamilies, viz. etula and Pseudochondrula . Four distinct groups can be Buliminusinae Kobelt, 1880 , and Eninae (e.g. Bank & recognized, which are mainly characterized by the Neubert, 1998; Hausdorf, 2001). The Buliminusinae inner structure of the penis and to a lesser extent on are characterized by the presence of a penial caecum the shell. These are Imparietula (the flattened lobe is at - (missing in the Eninae) and the long epiphallar flagel - tached to the penis wall; penial papilla absent ; the lum (short or absent in the Eninae). Furthermore, an shell without a parietalis and a columellaris but with a epiphallar caecum is missing, whereas it is mostly palatalis superior ), Pseudochondrula (the elongated present in the Eninae. The following genera belong to lobe is free from the penis wall; penial papilla absent ; the Buliminusinae: Buliminus Beck, 1837 (synonyms: the shell with a well-developed parietalis, columellaris Bulimina Ehrenberg, 1831; Petraeus Albers, 1850; Seste - and palatalis superior ), Anatolya gen. nov. (presence of ria Bourguignat, 1884 ); Pene Pallary, 192 9; Paramastus a penial papilla that is covered with irregular folds P. Hesse, 1933 (with the subgenus Cyrenaeus Heller, and narrow grooves; the shell without a parietalis and 1971) and Iranopsis Bank & Neubert, 1998 (with the a columellaris but with an indistinct thickening at the subgenus Mordaniella Bank & Neubert, 2016 = a height of the palatalis superior), and Pseudojaminia nomen novum for Mordania Bank & Neubert, 1988 gen. nov. (two parallel V-shaped lobes that are par - [non Dworakowska, 1979]). tially attached to the penis wall; penial papilla absent; The subfamily Eninae can be subdivided into three shell with or without a parietalis, columellaris and/or tribes, viz. the Chondrulini Wenz, 1923, the Multiden - palatalis superior ). These four groups are considered tulini Schileyko, 1978 , and the Enini. The Chondrulini to have genus status; the latter two are described as are characterized by a well developed penial papilla new to science. with an open groove and by the penis wall that is dec - orated with small papillae. In the Multidentulini the Key words: Gastropoda, Pulmonata, Enidae, Anatolya , Pseudo - penis wall is smooth, and the penial papilla is perfo - jaminia , anatomy, Turkey. rated (i.e. with an open canal). Finally, in the Eninae the penis wall is smooth or decorated with longitudi - Basteria 80 (4-6): 177-192 (2016) B2016-21-Gargely:Basteria-2015 11/19/2016 8:40 PM Page 178 nal rolls, and the penial papilla is absent or, when assigned to Imparietula and Pseudochondrula . Much to present, not perforated. our surprise it turned out that both genera have to be The Chondrulini consists, when one follows the redefined, and that it is necessary to establish two new above given diagnostic criteria, the following genera: genera in order to accommodate taxa that have previ - Thoanteus Lindholm, 1925; Peristoma Krynicki, 1833, ously been classified under Imparietula or Pseudochon - Caucasicola P. Hesse, 1917 (synonyms: Medea O. drula . Boettger, 1883 [non Eschscholtz, 1825] ; Helle P. Hesse, All species that are anatomically examined by us 1916 [non Osten-Sacken, 1896] ); Zebrina Held, 1838 share the following traits: penis appendix well-devel - (synonym : Zebrinus Westerlund, 1887); Georginapaeus oped, long, its retractor inserts on the distal end of the Schileyko, 1998; Rhabdoena Kobelt & Moellendorff, organ around the boundary of A1 and A2; the retrac - 1902; Chondrus Cuvier, 1817 (synonym: Antichondrus tor muscle of the penial appendix and the penis attach Lindholm, 1925); Brephulopsis Lindholm, 1925 (syn - on the diaphragm next to each other; penis moder - onym: Ramusculus Lindholm, 1925) ; Ayna Páll- ately long, the insertion of the penial retractor is on Gergely, 2009; Leucomastus A.J. Wagner, 1928 the penis; epiphallus long, usually cylindrical, usually (synonym: Aschera Pallary, 1939 [nomenclaturally in - with a small epiphallar caecum; the distal portion of valid name]); Chondrula Beck, 1837 (synonyms: Gon - the epiphallus (= the portion situated closer to the odon Held, 1838; Eucore Charpentier, 1837; Chondrulus penis) is usually more slender than the proximal part; Westerlund, 1887; Dentistomus M. von Kimakowicz, the proximal, thicker epiphallar portion is internally 1890) ; Mastus Beck, 1837 (synonyms : Pseudomastus O. decorated with transversal slit-like pockets, which Boettger, 1889; Amphitrorsus M. von Kimakowicz, probably play a role in forming the spikes of the sper - 1890); Meijeriella Bank, 1985 (synonym: Borlumastus matophore; flagellum very short or absent ; bursa cop - Örstan & Yıldırım, 2004) and Eubrephulus A.J. Wagner, ulatrix without diverticulum. In the following, we 1928. refrain from detailed descriptions of the genitalia, but We assign to the Multidentulini, following the highlight the observed diagnostic features, with spe - above given diagnostic criteria, the following genera: cial focus on the inner structure of the penis . Euchondrus O. Boettger, 1883 (synonym: Multidentinia Lindholm, 1925); Multidentula Lindholm, 1925 (syn - Systematic part 178 onyms: Bollingeria Forcart, 1940; Tokatia Hudec, 1972; Improvisa Schileyko, 1978; Senaridenta Schileyko, 1978); Family Enidae B.B. Woodward, 1903 (1880) Chondrulopsina Lindholm, 1925; Siraphoroides Schi - Subfamily Eninae B.B. Woodward, 1903 (1880) leyko, 1977; Pentadentula Suvorov, 2006 and Merdigera Tribus Enini B.B. Woodward, 1903 (1880) Held, 1838. The Enini exhibit a multitude of genera, some of Imparietula Lindholm, 1925 them being very rich in species, ranging from the Ca - nary Islands, Europe, North Africa, and a large part of Imparietula Lindholm, 1925: 30, 39. Type species (by monotypy): Asia (ranging from Turkey/Caucasus to Japan and Bulimus leucodon L. Pfeiffer, 1846. from Kazakhstan to Indonesia). A considerable num - ber of these taxa is poorly known. Here we will dis - The history regarding the identity of leucodon , the type cuss some Turkish taxa that have previously been species of Imparietula , is complex. Forcart (1940 : 206) assigned to the genera Imparietula Lindholm , 1925, synonymized Buliminus (Brephulus ) lasistanicus Lind - and Pseudochondrula P. Hesse , 1933. holm , 1914 with leucodon , although he did not possess The type species of Imparietula and Pseudochondrula shells of either lasistanicus or leucodon . The anatomy is Bulimus leucodon L. Pfeiffer , 1846, and Buliminus flo - (jaw, radula, outer morphology of the genital system) renskii Rosen , 1914, respectively. Imparietula leucodon is of lasistanicus was described by Hesse (1933: 169-170, a species known from a restricted area in the north - fig. 11) based on syntypes provided by Lindholm; this eastern part of Turkey (Vilayets Trabzon and Rize), description was placed by Forcart under leucodon . For - whereas Pseudochondrula armeniaca florenskii lives in a cart redefined Imparietula based on this synonymy, small area around Ardanuç in the Vilayet Artvin. The and placed several taxa under Imparietula , namely the outer morphology of the genital system of florenskii monotypic taxa leucodon , adjarica Retowski , 1914, has been described by Hesse (1933: 171, fig. 13), but tetrodon Mortillet , 1853, and brevior Mousson, 1876, as nothing has been reported on the important internal well as the polytypic taxa blanda L. Pfeiffer, 1853, se - structure of the penis. A short remark has been given ductilis Rossmässler, 1837, and armeniaca Mortillet , on the genital system of leucodon , including the inter - 1854. As a consequence, Pseudochondrula was consid - nal structure of the penis, by Hausdorf (1999 : 153). We ered a synonym of Imparietula by Forcart. Gittenberger have investigated several species taxa that have been (1967: 130-137) followed the opinion of Forcart, and Basteria 80(4-6), 2016 B2016-21-Gargely:Basteria-2015 11/19/2016 8:40 PM Page 179 added the monotypic altenai Gittenberger , 1967 and agnosed as follows : the genital system lacks a diver - pelidne Biggs , 1946 to Imparietula . Akramowski (1976: ticulum, the spermathecal reservoir is somewhat to 154-158) also followed Forcart, and included in addi - markedly swollen, the epiphallar caecum is located in tion pupoides Krynicki , 1833, and sieversi Mousson , the middle of the epiphallus or slightly subterminal, a 1873, into Imparietula . The latter two species belong to normal penis appendix is present, the penis is rela - Multidentula Lindholm (tribus Multidentulini) and tively slender, a penial papilla is absent, and the inner Ljudmilena Schileyko , 1984 (tribus Enini), respectively. surface of the penis is decorated with a large lobe that Schileyko (1984: 288, 306) separated Pseudochondrula is divided in the middle. The large to middle-sized from Imparietula , and described (1984: 307-308, fig. shell is dextral
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