Helix (Helix) schlaeflii MOUSSON , 1859

Neubert, Eike, 2014, Revision of Helix LINNAEUS, 1758 in its eastern Mediterranean distribution area, and reassignment of Helix godetiana KOBELT, 1878 to Maltzanella HESSE, 1917 (Gastropoda, Pulmonata, Helicidae), Contributions to Natural History 26, pp. 1-200 : 136-142

Helix (Helix) schlaeflii MOUSSON, 1859 ( Figs 220–226 )

1859 Helix schläflii MOUSSON, Vierteljahrsschrift der naturforschenden Gesellschaft in Zürich 4: 266 [Iannina, Sziza ex Schläfli 1858].

1879 Helix schläfli var. praestans WESTERLUND & BLANC, AperÇu sur la faune malacologique de la Grèce: 83 [Corfu, Pantocrator 1000 m].

1906 Helix (Helicogena) pomatia var. krüperi KOBELT, Systematisches Conchylien-Cabinet VI : 258, Taf. 365, Fig. 1, 2 [Agnanta (upper reach of the Arachthos, Pindos, not too far from the sanctuary of Dodona) = Agnanda].

1970 Helix secernenda edlaueri URBANSKI , Bulletin de la Societé des Amis des Sciences et des Lettres de Poznan, Série D 11: 76 [ Albanien, Burghof in Kruje].

Type specimens:

schlaeflii : 2 syntypes ZMZ 506230, illustrated syntype: H = 42.75; D = 40.55; PH = 29.1; PD = 21.7; PrD = 6.1; praestans: syntype GMN 1754: H = 50.5; D = 48; PH = 34.8; PD = 25.3; PrD = 6.7; W = 5.25. krüperi: holotype SMF 9584 ( Zilch 1952): H = 43.3; D = 46.4; PH = 31.2; PD = 24.7; PrD = 5.7; W = 5. edlaueri: not seen .

Specimens examined:

FYROM: Jablanitsa Mts. , up to village Gorna Belitsa, alpine meadow, 41.2312 20.5241, July 2006, leg. I. Dedov, NMBE 520399 / 3; Galicica Mts. , Magaro Peak, 40.9351 20.8204, leg. I. Dedov, NMBE 520400 /2.

Greece: SMF 9675 Corfu, Pantokrator, coll. Kobelt; SMF 1818190/2 Corfu; ditto, SMF 18192 , coll. O. Boettger ex Reitter; ditto, SMF 18191 / 2, coll. Kobelt ex Wohlberedt ¸ SMF 18189/5, coll. O. v. Moellendorff; ditto, SMF 86146 , coll. Nägele; ditto, SMF 74400 , coll. Krüper; ditto, SMF 74401 , coll. Krüper; ditto, SMF 74399 , coll. Jetschin ex Linnaea; ditto, SMF 74398 / 2, coll. Jetschin ex Conéménos; ditto, SMF 74402 / 2, coll. C.R. Boettger; ditto, SMF 74397 / 2 Corfu (?), coll. K.L. Pfeiffer; Kerkyra, 39.62 19.92, NMBE 18480 / 2; Kerkyra, slope of Mt. Pantokrator , 39.7249 19.8776, 20.7.1975, NMBE 524235 / 1; Kerkyra, above Petalia, 39.7468 19.8425, 7.8.1995, NMBE 523281 / 1; Kerkyra, summit of Pantokrátor, radiostation, 39.7471 19.8723, 5.8.1995, NMBE 523273 / 3; Kerkyra, Mt. Loutses , 39.7722 19.8928, 2.8.1995, NMBE 523275 / 1; SMF 9674 Epirus, coll. Kobelt; ditto, SMF 18185 ; ditto, SMF 18186 ; ditto, SMF 74396 / 2, SMF 74392 / 4, coll. Jetschin ex Conéménos; ditto, SMF 74392 / 2+2, coll. Krüper; ditto, SMF 299519 , coll. Kaltenbach ex Linnaea; SMF 18188 Kamarina, 39.1356 20.6722, coll. O. Boettger ex Conéménos; SMF 18187 Goumenitsa [= Igoumenitsa ], 39.5061 20.2655, coll. O. Boettger ex Conéménos; ditto, SMF 74391 / 2, coll. Jetschin ex Conéménos; SMF 74394/2, Arta, 39.1592 20.9868, coll. K.L. Pfeiffer ex Wohlberedt; ditto, SMF 74394 / 4, coll. C.R. Boettger; SMF 74653/2, Agnanda, coll. Krüper (types of krüperi?); SMF 20222/2, Tschumerka [= old name for the mountain chain E of Agnanda, Oros Athamanon], coll. O. v. Moellendorff; ditto, SMF 74710 , Tschumerka , coll. Krüper; Amphithea close to Ioannina 200 m in direction to Longades, 31.05.1985, HAUS (3) ; Amphithea close to Ioannina 3.5 km in direction to Longades, 31.05.1985, HAUS (2) ; Perama close to Ioannina 3.5 km in direction to Metsovon, 39.6872 20.8739, 22.08.1982, HAUS (2) ; Perama 3 km in direction to Metsovon, rocky canyon, 30.05.1985, HAUS (1) ; Ioannina in direction to Driskos, rocks at the crossroads to Kastritsa, 39.6342 20.9930, 31.05.1985, HAUS (1) ; Ioannina, rocks at the crossroads to Kastritsa SE shore of the Ioannina Lake , 39.599 20.919, 20.07.1990, NMBE 524740 / 2; Ioannina, Baltuma, 6.7 km in direction to Ioannina, 30.05.1985, HAUS (3) ; rocks in Mikro Peristeri, 39.7395 21.0888, 30.05.1985, HAUS (3) ; Lakmos Oros, rock walls above church in Megalo Peristeri, 39.7234 21.0622, 30.05.1985, HAUS (1) ; Vrosina 1 km in direction to Igoumenitsa , 39.6505 20.5079, 23.08.1982, HAUS (2) ; Vrosina 10 km in direction to Igoumenitsa , 39.5981 20.4940, 01.06.1985, HAUS (1) ; Igoumenitsa 6 km in direction to Plataria, rocky canyon close to the lake, 24.08.1982, HAUS (1) ; Margariti 7 km in direction to Igoumenitsa , 24.08.1982, HAUS (1) ; Mt. Zalongon N of Prevesa, 39.1481 20.6837, 24.08.1982, HAUS (1) ; Prevesa , rocks above Dropos, 39.1530 20.7280, 03.06.1985, HAUS (1) ; Metsovon, 10.5 km in direction to Milea, 39.8247 21.1841, 29.05.1985, HAUS (2) ; Votonosion 7.5 km in direction to Metsovon, 29.05.1985, HAUS (2) ; Sitsa 3.8 km in direction to Lithinon, 01.06.1985, HAUS (1) ; Thesprotias, Paramithia 2 km in direction to Ag. Kiriaki, 39.4795 20.5199, 01.06.1985, HAUS (1) ; E of Amalthea 1.5 km in direction to Seatozi, 23.08.1982, HAUS (1) ; Karditsa, rocks W of Morfovounion , 39.3556 21.7372, 18.06.1985, HAUS (1) ; Karditsa, rocks at Musaki 1.5 km in direction to Krioneri, 19.06.1985, HAUS (1) ; Krionéri, SW slope of Mt. Varassova , 38.3477 21.5973, 1.10.1989, NMBE 524210 / 2; Lefkada, Frini in dir. Tsoukalades, 38.8236 20.68, 5.8.1975, NMBE 524236 / 2; Loutropigi, 39.1173 22.0424, 18.5.1995, NMBE 523267 / 2; Louros valley , Agios Georgios, 39.2723 20.8446, 28.9.1989, NMBE 524248 / 2; Gliki, road to Koukoulii, 39.3322 20.603, 15.5.1995, NMBE 523285 / 1; Morfovouni, 39.3567 21.7464, 23.4.2003, NMBE 523270 / 11; road Ioanina – Arta, crossroads to Klissoura, 39.3607 20.888, 14.5.2011, NMBE 523253 / 1; Kartéri, 1 Km N village, 39.3929 20.3718, 14.5.1997, NMBE 523286 / 1; ditto, 21.4.2009, NMBE 523256 / 12; Kartéri, 1 Km E village, road Margariti – Igoumenitsa , 39.3953 20.3702, 20.7.1990, NMBE 524253 / 10; ditto, 15.5.1995, NMBE 523279 / 1; Louros valley , N exit of canyon, 39.3991 20.838, 20.7.1990, NMBE 524238 / 1; ditto, 28.9.1989, NMBE 524232 / 2; ditto, 14.5.2011, NMBE 523240 / 3; Theodoriana, 6.3 Km S Ort, Athamanon Geb., 39.4021 21.2068, 15.5.2008, NMBE 523255 / 1, NMBE 523233 / 1; crossroads to Porti, 1.6 Km S Mouzaki, 39.4113 21.663, 23.4.2003, NMBE 524746 / 1; Platanoussa, 39.43 21.0117, 16.5.2008, NMBE 523239 / 2; source of Louros River , 39.4327 20.8413, 28.9.1989, NMBE 524244 / 1; Plataria, hills W of the bay, 39.4408 20.271, 18.5.2008, NMBE 523262 / 2; Plataria, above Camping «Kalami Beach», 39.474 20.2493, 9.5.2011, NMBE 523261 / 3; Frossini 39.4496 20.5951, 15.5.1995, NMBE 523276 / 2; Kataraktis, above the waterfalls, 39.4627 21.1126, 13.5.1997, NMBE 523274 / 4; Igoumenitsa , pass 35 km E Igoumenitsa, 39.491 20.478, 11.9.1991, NMBE 524739 / 2; Parapotamos, 2 Km in dir. Ioanina, 39.5297 20.3426, 23.7.1975, NMBE 524228 / 4; Ano Vereniki , 39.5534 20.5487, 13.5.1995, NMBE 523277 / 2; Dovla, canyon of small creek, 39.5679 20.5481, 13.5.1995, NMBE 523280 / 2; Vrossina, 16 Km in dir. Igoumenitsa , 39.5777 20.4766, 14.4.1988, NMBE 524237 / 1; Vrossina, 13 Km in dir. Igoumenitsa , 39.5944 20.4918, 11.5.2011, NMBE 523235 / 1; Ioannina, rocks at the crossroads to Kastritsa, 39.599 20.919, 20.7.1990, NMBE 524740 / 2; Dafni, 39.6356 20.3898, 10.5.2011, NMBE 523238 / 1; Kastritsa, 39.6393 20.9145, 24.7.1975, NMBE 524234 / 10; road Filiates – Igoumenitsa , cave chapel, 39.6434 20.2135, 19.7.1990, NMBE 524259 / 2; ditto, 15.5.1997, NMBE 523272 / 5; Rizo, crossroads Filiates – Keramitsa 39.6435 20.4164, 19.7.1990, NMBE 524200 / 2; Tsamantas, SE of village, waterfall, 39.669 20.3214, 17.7.1990, NMBE 524246 / 3, ditto, 11.5.2011, NMBE 523265 / 1; Mazia, 39.6705 20.9485, 11.4.1988, NMBE 524252 / 2; Amfithea, 2 Km E village, 39.6802 20.8871, 24.7.1975, NMBE 524233 / 6; Amfithea, 39.686 20.8727, 14.5.2011, NMBE 523266 / 2; Ioannina, cave of Perama, 39.695 20.846, 20.7.1990, NMBE 512577 / 1; Pérama, 39.6986 20.8482, 28.9.1989, NMBE 524209 / 24; ditto, 12.5.1995, NMBE 523284 / 1; mountain slope E Méga Peristéri, Lakmos Mts. , 39.7099 21.0941, 21.7.1990, NMBE 524203 / 5; road Kalambaka – Ioanina, 39.7326 21.0589, 28.7.1975, NMBE 524229 / 1; ditto, 29.7.1976, NMBE 524226 / 1; crossroads from road Ioanina – Konitsa, 39.7345 20.7737, 29.7.1975, NMBE 524201 / 3; Lia, 5.9 Km in dir Lista, 39.7355 20.4581, 17.7.1990, NMBE 523289 / 1; Monastir Pateron (= W of Zitsa ), 39.7642 20.6167, 22.5.1991, NMBE 524250 / 1; Monastir Pateron, in the valley 1 Km NE, 39.7729 20.6146, 13.5.1997, NMBE 523278 / 3; Dikorifo, Mitsikéli Mts. , 200 m S, 39.7828 20.8041, 23.5.1991, NMBE 524245 / 1; Kepéssovo, 7.2 Km S of village, old bridge, 39.862 20.7749, 23.5.1991, NMBE 524242 / 1; Kipi, 200 m W, old bridge, 39.8647 20.7897, 12.5.1995, NMBE 523287 / 1; Kalpaki, ca. 50 m S of Velas Monastery , 39.8705 20.6253, 13.4.1988, NMBE 524241 / 2; ditto, 16.7.1990, NMBE 523290 / 1; road Kalpaki – Monodéndri, 39.8753 20.6951, 30.7.1975, NMBE 524227 / 1; Monodendri, 5 km above village towrds Farangiou Vikou, 39.89 20.738, 19.7.1990, NMBE 524744 / 1; Vikos canyon, hiking path to Aristi and Papigo, 39.879 20.7365, 11.4.1988, NMBE 524257 / 3; Vikos canyon, ca. 200 m from monastery Ag. Paraskevi, 39.888 20.7539, 27.9.1989, NMBE 524258 / 2; Vikos canyon, N exit at Klidonia, 39.9677 20.6634, 30.7.1975, NMBE 524225 / 1; road Kerassovo – Lavdani, 39.8799 20.4671, 17.7.1990, NMBE 524255 / 3; Skamnéli, in dir. Tsepélovo, 39.9089 20.825, 22.5.1991, NMBE 524205 / 6; Skamnéli, 1 Km E village, 39.9131 20.8738, 23.5.1991, NMBE 524249 / 3; Papigo, 3h Fussweg E Katafigi, Timfi-Gebirge, 39.9773 20.7644, 24.5.1991, NMBE 524208 / 9; Porto Koufos, easternmost mountain, Sithonia, Chalkidiki, 39.942 23.944, 2.9.1991, NMBE 524743 / 7; Toroni, Chalkidiki, 39.98 23.9, 29.8.1991, NMBE 524742 / 6; 17 km S Sarti, Sithonia, Chalkidiki , 39.993 23.959, 8.9.1991, NMBE 524745 / 2; crossroads from road Konitsa – Molivdoskepastis, 40.0038 20.5617, 16.5.1997, NMBE 523288 / 1; Konitsa, 600 m from old bridge, 40.0331 20.748, 26.9.1989, NMBE 524239 / 1; Pades, ca. 28 Km E Konitsa, 40.0408 20.9054, 16.7.1990, NMBE 524202 / 5; Kakolakos, Mt. Dousko , 40.0411 20.4757, 12.5.1995, NMBE 523271 / 9; road Konitsa – Molivdoskepastis to Pogonisko, 40.0438 20.5617, 16.7.1990, NMBE 524247 / 3; ditto, 26.9.1989, NMBE 524240 / 2; Konitsa, Topolca creek , tributary to the Aoos, 40.0706 20.7608, 26.9.1989, NMBE 524207 / 13; Aetia, 2.2 Km E in dir. Grevena, 40.0739 21.2015, 24.4.2003, NMBE 523251 / 3; crossroads from road Konitsa – Neapoli, 40.0836 20.7422, 12.4.1988, NMBE 524256 / 6; Sarantaporos canyon between Exohi and Agia Varvara , 40.1101 20.7203, 11.5.1995, NMBE 523283 / 1; Sarantaporos canyon, 40.1114 20.7218, 16.7.1990, NMBE 524260 / 3; Sarantaporos River , 3 Km N Drossopigi, 40.2281 20.9133, 15.7.1990, NMBE 524243 / 1; Agia Paraskevi, 40.1438 20.8965, 25.9.1989, NMBE 524230 / 2; Loutra N of Amarantos , 40.1676 20.7321, 25.9.1989, NMBE 524206 / 6; Loutra, summit of mountain, 40.1735 20.731, 12.4.1988, NMBE 524254 / 3; Loutra, W slope of mountain, 40.1745 20.7297, 12.4.1988, NMBE 524251 / 1; Kastania, Kozani, 40.1741 22.0232, 4.8.1991, NMBE 524741 / 2; mountain between Epano Aréna and Kato Aréna, E slope, 40.2948 20.9381, 11.5.1995, NMBE 523282 / 1; Epano Aréna, E slope of mountain, Grammos Mts. , 40.3447 20.8845, 15.7.1990, NMBE 524204 / 2; Pérama, 24.7.1975, NMBE 524231 / 6; Epirus, NMBE 18490 / 1; Epirus, NMBE 18481 /2.

Diagnosis: shell large, large protoconch broadly submammillate, upper spiral bands often fused, aperture brown, umbilicus closed.

Description: shell large to very large, shell form subspherical broad to conical; basic shell colour whitish grey, with the bright spiral band No. 2 and No. 3 almost always fused, sometimes even fused with No. 1, median area of last whorl usually with a sharply pronounced white zone, band No. 4 and No. 5 narrow and always free; protoconch submammillate, usually with a shallow suture and consisting of 2.5 whorls, its diameter ranging from 6–7 mm; teleoconch of 3–4 whorls; covered by fine but rugose and somewhat irregularly arranged riblets or strongly pronounced growth lines; grey axial streaks frequently occur giving a cloudy appearance to the shell; aperture obliquely subquadrate, peristome wide, its rim only weakly calloused; columella and columellar triangle brown, the peristomial rim usually light brown; parietal area usually with a reduced brown colour marking; umbilicus closed in adult shells.

Genital organs ( Fig. 225 ): penis elongate club-shaped, epiphallus considerably longer than penis; flagellum long, three times longer than epiphallus+penis; mrp attaching in a proximal position on the epiphallus; internally, both papillae of equal size, penial chamber very short, smooth inside, both papillae conical with a central perforation; atrial stimulator a triangular knob; female system with a short vaginal tube, dart sac and glandulae mucosae strongly developed, the glandulae with a thick basic stem, multi-branched with a few glandular tubules, longer than dart sac; pedunculus stem short, with a thick and short diverticulum, bursa copulatrix longer than pedunculus stem, vesicle rounded and bent downwards.

Distribution ( Fig. 226 ): This species inhabits the Epirus and the southwestern Rhodope Mts.; it has been reported by Dhora & Welter-Schultes (1996: Figs 216 (" 7 km SE of Ballsh"), 217 (Gjirokaster, castle) and 220–221 (" 12 km SE Korce )) at least from southern Albania, it probably can be found in Central Albania as well (see also the remark section under H. secernenda ). These three localities are included in the map here.

Remarks: There is no type designated for praestans, because the single specimen does not match the dimensions as given by Westerlund & Blanc in the original description. The authors say (1879: 83) "la hauteur et le diamétre de la coquille, sont égaux et mesurent 53 mm … l'ouverture est de 39 mm de hauteur sur 30 mm de diamétre". These numbers differ considerably from what could be taken from the specimen GMZ 1754. The description of the banding patterns is (more or less) close, but the figured specimen differs in shape of the aperture, coloration of the columella and parietal area and the fact that the lower two bands are missing. Thus, the status of the specimen GMN 1754 is somewhat vague, but it is accepted as a syntype as it obviously forms part of the original lot. The allocation of H. secernenda edlaueri to H. schlaeflii is more problematic. The type locality of edlaueri is situated in the presumed overlap area between the two species, but still, the correct distribution pattern of these Helix species in Albania is not satisfyingly clarified. Urbanski (1970) describes the shell to have rough riblets on its surface, which usually does not occur in H. secernenda , but sometimes can be seen in H. schlaeflii . Probably this is a hybrid population between the two species, but currently there is only a single preserved topotype specimen available for molecular research, which is not enough to resolve this problem (Korábek, pers. comm.).

The separation of Helix (Helix) schlaeflii as a species distinct from H. secernenda is based on the following character states: The shells of H. secernenda have the spiral bands Nos. 1, 2, and 3 usually separate (usually fused in H. schlaeflii ), the shell surface of H. secernenda is usually smooth- er than that of H. schlaeflii , which has fine axial riblets, the epiphallus in H. secernenda is shorter than in H. schlaeflii , and the glandulae mucosae are much more developed in H. secernenda than in H. schlaeflii .

Figures

Figs 220–224. Helix (Helix) schlaeflii MOUSSON, 1859. Fig. 220 syntype schlaeflii ZMZ 506230/a, Iannina, Sziza, D = 39.8; Fig. 221 syntype praestans GMN 1754, Corfu, Pantocrator, D = 48.0; Fig. 222 holotype krüperi SMF 9584, Agnanta, Pindos, D = 46.4; Fig. 223 NMBE 520399, Jablanitsa Mts., D = 40.24; Fig. 224 NMBE 524743, Porto Koufos, D = 42.05. — All photos Neubert, original size.

Fig. 225. Helix (Helix) schlaeflii MOUSSON, 1859. Genital organs. NMBE 520399, Jablanitsa Mts., situs 52.76.

Fig. 226. Helix (Helix) schlaeflii MOUSSON, 1859. Distribution.

Figs 1–9. Helix (Helix) pomatia LINNAEUS, 1758. Fig. 1 compacta, syntype SMF 9573, D = 50.07, Hungary "Gestrüppe der Berglehnen"; Fig. 2 pulskyana, syntype SMF 9525/2a, Hungary "Inseln der Donau", D = 36.6; Fig. 3 hajnaldiana, syntype SMF 9658, Hungary "Budapest, im botanischen Garten", D = 34.1; Fig. 4 solitaria, syntype SMF 74708/2a, D = 31.1; Fig. 5 sabulosa, syntype SMF 9524, D = 40.5, Fig. 6 expansilabris, holotype SMF 9538, D = 45.5; Fig. 7 luteola, holotype SMF 9527, D = 34.6; Fig. 8 pannonica, lectotype SMF 9535, D = 40.5, Fig. 9 lednicensis, syntype SMF 9534/2a, D = 38.3. — All photos Neubert, original size.

Figs 210–218. Helix (Helix) dormitoris KOBELT, 1898. Fig. 210 lectotype dormitoris SMF 9755, Durmitor, D = 32.75; Fig. 211 lectotype lubnicensis SMF 9788, Ljubicna, D = 33.2; Fig. 212 lectotype kolaschinensis SMF 9721, Kolašin, D = 33.0; Fig. 213 holotype pivensis SMF 9727, Durmitor gegen das Pivathal, D = 37.05; Fig. 214 holotype stolacensis SMF 9732, Berg Stolac, D = 42.55; Fig. 215 lectotype bosnica SMF 9757, Bjelasnica bei Serajewo, D = 30.0; Fig. 216 holotype sturanyi SMF 9772, Gacko, D = 31.09; Fig. 217 paratype hajlensis SMF 60085, Hajila Berg unweit Péc, D = 33.7; Fig. 218 paratype arnautorum SMF 60084, Aleksandrova vis, D = 31.65. — All photos Neubert, original size.