Helix (Helix) cincta O. F. MüLLER, 1758

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 : 84-91

Helix (Helix) cincta O. F. MüLLER, 1758 ( Figs 132–147)

1774 Helix cincta O. F. MÜLLER, Vermium terrestrium et fluviatilium II: 58 [no type locality given].

1838 Helix lemniscata BRUMATI, Catalogo sistematico delle conchiglie, Monfalcone: 22 [non Helix lemniscata LESSON, 1831 nec Helix lemniscata WEBB & BERTHELOT, 1833 ].

1849 Helix Pollinii DA CAMPO, Memorie dell'Accademia d'Agricoltura, Commercio ed Arti di Verona 23:113 [not seen].

1876 Helix giuliae BOURGUIGNAT , Species novissimae molluscorum: 56 [ Malta].

1891 Helix (Pomatia) cincta var. anatolica KOBELT, Iconographie der Land- und Süsswassermollusken 2 (5): 26, Taf.128, Fig. 767–769 [no type locality mentioned; the specimens figured in No. 767 and 768 come from " Hieronda , Caria " in SW Turkey (cf. ZILCH 1952)] .

1891 Helix (Pomatia) cincta var. cyprica KOBELT, Iconographie der Land- und Süsswassermollusken 2 (5): 26, Taf. 128, Fig. 770 [ Cyprus].

1893 Helix (Pomatia) cincta var. trojana KOBELT, Iconographie der Land- und Süsswassermollusken 2 (6): 55, Taf. 164, Fig 1048 [Troja].

1895 Helix (Pomatia) stenarochila KOBELT, Iconographie der Land- und Süsswassermollusken, Suppl. 1: 44, Taf. 16, Fig. 2, 4 [ Zypern].

1895 Helix (Pomatia) trixenostoma KOBELT, Iconographie der Land- und Süsswassermollusken, Suppl. 1:45, Taf. 16, Fig. 5 [auf Zypern].

1895 Helix (Pomatia) cornarae KOBELT, Iconographie der Land- und Süsswassermollusken, Suppl. 1: 46, Taf. 17, Fig. 1, 2 [auf Zypern].

1895 Helix (Pomatia) pediaea KOBELT, Iconographie der Land- und Süsswassermollusken, Suppl. 1: 47, Taf. 17, Fig 3, 4 [auf Zypern, namentlich in der Fruchtebene des Pediaeos].

1895 Helix (Pomatia) epidaphne KOBELT, Iconographie der Land- und Süsswassermollusken, Suppl. 1: 52, Taf. 21, Fig. 2–5 [Antiochia (Epidaphnes in the ancient times), today Antakia].

1903 Helix (Helicogena) cincta libanica KOBELT, Systematisches Conchylien-Cabinet VI : 115, Taf. 333, Fig. 3–7 [ Libanon].

1906 Helix (Helicogena) cincta var. elegans KOBELT, Systematisches Conchylien-Cabinet VI : 270, Taf. 368, Fig. 7, 8 [Crkvenica].

1921 Helix (Helicogena) castanostoma GERMAIN, Mollusques terrestres et fluviatiles de Syrie I: 130, Fig. 7, 8 [Tripoli de Syrie].

1921 Helix (Helicogena) pachya var. incrassata PALLARY in Germain: Mollusques terrestres et fluviatiles de Syrie I: pl. 11, fih. 13 (nomen nudum).

1928 Helix cincta melanotica RENSCH, Zoologischer Anzeiger 78: 2, Fig. 1 [Istrien, Rovigno, La Figarola Grande].

Type specimens:

anatolica: syntype SMF 9949: H = 36.7; D = 34.9; PH = 24.7; PD = 17.9; PrD = 4.8; W = 5.25. giuliae : syntype MHNG 18144: H = 36.9; D = 41.2; PH = 22.2; PD = 26.6; PrD = 4.5; W = 5.75. stenarochila : lectotype SMF 9641 (SD Zilch 1952: 151): H = 42.1; D = 41.7; PH = 29.8; PD = 20.2; PrD = 4.3; W = 5.5. trixenostoma : lectotype SMF 9794 (SD Zilch 1952: 151): H = 41.2; D = 38.6; PH = 27; PD = 18.8; PrD = 4.65; W = 5.5. cornarae : lectotype SMF 9802 (SD Zilch 1952: 151): H = 43.6; D = 42.8; PH = 30.3; PD = 21; PrD = 4.1; W = 5.5. epidaphne : lectotype SMF 9654 (SD Zilch 1952: 154): H = 42.4; D = 41.8; PH = 30.8; PD = 21.4; PrD = 5.4; W = 5. libanica: holotype SMF 9715 ( Zilch 1952: 152): H = 43.1; D = 39.4; PH = 28.2; PD = 29; PrD = 4.3; W = 5.5. castanostoma : syntype (s) MNHN-IM-2000-27766, coll. Germain (ex Férussac?): H = 40.4; D = 36.6; PH = 26.9; PD = 18.5; PrD = 3.5; W = 5.5. trojana: lectotype SMF 9946 (SD Zilch 1952: 151), H = 44.1, D = 42.6, PH = 26.2, PD = 26.7.

Specimens examined:

Italy: Castle of Arco, Rovereto , Lombardia, 45.922 10. 8853, 150 m alt., 30.4.1997, NMBE 513052 / 3; Desenzano del Garda / Brescia / Lombardia, 45.47 10.53, 1847, NMBE 22823 / 6; ditto, NMBE 22825 / 3; ditto NMBE 22824 / 4; Lake Garda, Tremosine, above Pieve to Mezzama , 45.7687 10. 5737, 510 m alt., 15.5.1994, NMBE 513041 / 4; Milano , Lombardia, 45.5 9.5, NMBE 22827 / 2; Riva del Garda, Trento , 45.88 10.83, 4.1997, NMBE 505535 / 2; ditto, 1847, NMBE 22826 / 4; Verona , Veneto, 45.45 10.9937, NMBE 18485 /8.

Croatia: Island Pag (= Isola di Pago ), 44.45 15.0527, 7.1985, NMBE 22830 / 4; Mali Losinj , Istria, 44.53 14.4731, 8.1980, NMBE 512950 / 10; Obrovac, Velebit , 44.2 15.68, 200 m alt., NMBE 513042 / 5; Supetarska Draga, island Rab , 44.8121 14.7165, 20 m alt., 25.7.2002, NMBE 522628 / 20; Vransko Jezero N Pakostane , 43.9407 15.51925, 9.1985, NMBE 513040 / 2; Zadar , 44.12 15.24, NMBE 18484 / 2; Pag road between Novalja and Pag, 8.1978, NMBE 524289 / 1; Pag , SE of Dinjiska, 44.3563 15.1923, 12.6.1970, NMBE 524288 / 6; Paklenica National Park, Starigrad , 44.3002 15.4699, 17.6.1970, NMBE 524287 / 1; Rab, Supetarska Draga , 44.8121 14.7165, 20 m alt., 25.7.2002, NMBE 522628 / 20; Karlobag , 44.5258 15.0785, 20 m alt., 30.7.2002, NMBE 523336 /2.

Greece, Dodekanes : Tilos: Tilos, 36.4230 27.3733, 25.05.1996, LIEB (6) . Leros : Alinda in direction to Platamos, 37.1609 26.8257, 04.07.1997, LIEB (4) . Lipsi : Lipsi, 37.2953 26.7685, 25.–30.06.1997, LIEB (36) . Lesbos: Skala Eresos, Lesbos, 39.15 25.9, 17.1996, NMBE 22836 /1.

Cyprus: Cape Andreas, Karpas Peninsula , 35.7 34.5823, 7.1977, NMBE 18483 / 2; Kato Pyrgos , 35.1817 32.6851, 10 m alt., 3.3.1990, NMBE 512592 / 1; Limassol, 34.68 33.03, NMBE 18488 / 3; ditto, NMBE 18487 / 3; Petra tou Romiou ca. 30 km W Limassol, 34.664 32.62724, 24.2.1990, NMBE 512591 / 1; on the road from Dipkarpes in direction to Yeni ErenkÖy , 35.5734 34.2932, 01.01.1986, LIEB (24) ; Larnaca, on the market, 24.10.1981, LIEB (5) ; on the road from Kourion in direction to Episkopi , 34.6794 32.7155, 14.10.1981, LIEB (1) ; Famagusta, Rizokarpaso , 35.5972 34.3792, 5.1984, BANK/6 .

Turkey: Izmir: SelÇuk , in rock crevices 20 km N SelÇuk, 38.0933 27.4061, 28.7.1984, MENK (4) ; Didim, in rock crevices 1 km S of AkkÖy at Didim , 37.4845 27.2604, 29.7.1984, MENK (2) ; Izmir (= Smyrna), 38.41 27.15, 1846, NMBE 22837 / 3, ditto 1846 NMBE 22838 / 2, ditto NMBE 18489 / 3, ditto NMBE 18486 / 2; rock crevices on Kazikbeli GeÇidi 20 km SE Denizli, 8.1984, 37.6308 29.2168, MENK (17) ; Valley of the Kücükmenderes , rocks close the road 12 km W Tire, 38.13 27.64, 11.6.1998, NMBE 512585 / 10; Çesme , 10 km S Mordogan, 38.4656 26.5504, 50 m alt., 15.6.1998, NMBE 512586 / 1; Çesme, 5 km W Abzweig Balikliova nach Çesme , 38.44 26. 54, 170 m alt., 15.6.1998, NMBE 512587 /1; Muğla: on the road approx. 20 km S of Denizli , 37.66 29.23, 16.4.1993, NMBE 505487 / 1 (alk.), ditto NMBE 512580 / 6; Ören, steep rocky slope 2 km before Ören close Keramos , 37.0314 27.8966, 10 m alt., 8.6.1998, NMBE 512583 / 4; Ören, 3 km NE Ören in direction Altikaya to KirankÖy , 37.058 27.986, 50 m alt., 8.6.1998, NMBE 512584 / 10; Ören, 2 km E MazikÖy in direction to Ören , 37.0271 27. 7564, 370 m alt., 8.6.1998, NMBE 512582 / 1; Milas , 37.3158 27.7797, 9.10.1997, LIEB ; Denizli, 3 km E of ÇubukdağI, 40 km NW Denizli , 37.9523 28.7782, 30.7.1984, MENK (30) ; Pamukkale , amphitheater, 37.927 29. 13, 399 m alt., 30.7.1980, MENK (19) , ditto 17.4.1993, NMBE 512581 / 2; Pamukkale , 02.10.1997, LIEB (19) ; Pamukkale , leg. Probst, 28.08.1964, RÄH (2); Pamukkale, Aphrodisias close to Geyre, 37.7093 28. 7234, 520 m alt., 5.5.1992, NMBE 512588 / 1; Pamukkale, ruins of Hierapolis , 37.9206 29. 1262, 330 m alt., 2.10.1997, LIEB, NMBE 503514 / 4, ditto 5.5.1992 NMBE 512589 / 17; Inlice , rock grave, 36.7482 28.9987, 07.10.1996, LIEB (6) ; Dalaman to Fethyie , GÖcek, 36.7568 28.9445, 30.09.1996, LIEB (5) ; KÖycegiz, rocks at the bath of Sultaniye , 36.873 28.601, 29.4.1991, NMBE 520392 / 1 (alk.), ditto NMBE 512579 / 2; Basin of KÖycegiz, Ekincik , 36.83 28.559, 26.4.1991, NMBE 519951 / 1; Edremit, AltIn Oluk (Balikesir), on the road to Sahindere, 39.5824 26.7576, 10.6.1989, LIEB (6) , ditto, olive platations, 22.09.2001, LIEB (2) ; excavations of Karain N Antalya (holocene), 37.0777 30.5708, RÄH (3); Priene, 37.6597 27.2977, 28.4.1992 Tannert, ex coll. Groh, NMBE 512590 / 1; GÖkova, Akyaka , rockgrave approx. 1 km S Akyaka, 37.0555 28.3264, 29.10.1997, LIEB ; GÖkova, mountain slope N Akyaka , 37.0646 28.3279, 6.10.1997, LIEB ; ruins of Stratonikeia close Yatagan , 37.3134 28.058, 7.10.1997, LIEB ; Demre , 36.2203 29.9457, 7.6.1987, NMBE 524292 / 7; Finike , 36.304 30.1401, 50 m alt., 8.6.1987, NMBE 524291 / 5; Demre , ca. 2 Km NW of sea shore, 36.2377 29.9182, 60 m alt., 6.6.1987, NMBE 524290 / 3; Alanya, Dim valley, 5 Km from coastal road, 36.5419 32.1069, 50 m alt., 7.10.2001, NMBE 523335 / 4; Antiochia , 36.2029 36.1603, coll. Kobelt ex Rolle (paralectotype epidaphne ) , SMF 9655, SMF 9942/2; ditto, coll. O. v. Moellendorff (paralectotypes epidaphne ) , SMF 9943/2; ditto, coll. C.R. Boettger (paralectotypes epidaphne ?) , SMF 74152/2; Tell am Orontes above Antakya, 20.9.1982 leg. Kinzelbach, NMBE 520414 /2; Hatay (= Antakya = Antiochia )/ Hatay, 36.2 36.16, 20.9.1982, NMBE 520414 /7.

Syria: Ruins of Qalaat al-Mudiq ( Apameia ), 35.4154 36.3938, 04.04.1996, leg. Rähle, RÄH (1); Orontes, mountains of the Alawits, 05.10.1089, RÖttger leg., RÄH (1); Latakieh (= Lattaquié), 35.5 35.7723, 3.2007, NMBE 517757 /1.

Lebanon: Sable de Beyrouth (specimen figured by Germain 1921: pl. XI, fig. 13 ex

Pallary), MNHN (1).

Diagnosis: shell of medium size, thick, protoconch small, spiral bands Nos. 1 to 3 usually fused, columellar side of aperture relatively straight, aperture reddish-brown coloured, umbilicus closed; flagellum extremely long, pedunculus stem long, usually without diverticulum.

Description: shell of medium size, thick, spherical, with the spire high conical; basic shell colour greyish white with chestnut brown spirals with spiral bands Nos. 1 to 3 usually fused to a large brown subsutural area, or band No. 1 a separated small brown band, periphery white to greyish, spiral bands Nos. 4 and 5 usually separate, often indistinct; protoconch small in comparison to shell, its diameter ranging from 3.5– 5 mm, smooth, white whorls; teleoconch of 3–4 whorls, teleoconch surface usually smooth, seldom with some patches of granulation, often with axial riblets; last whorl only slightly descending below the periphery; aperture subquadrate to rounded, apertural rim moderately broad, labial callus weak, reddish-brown coloured, columellar triangle small; columellar side of aperture relatively straight if compared to other black-mouthed species; umbilicus closed in adult shells.

Genital organs ( Figs 142–144, 145–146): penis club-shaped, epiphallus as long as penis; flagellum extremely long, exceeding the length of epiphallus+ penis at least fourfold; mrp attaching in a central position on the epiphallus; internally, both papillae of equal size, penial chamber long, filled with a few small elongate folds, both papillae conical with a central perforation; atrial stimulator a rounded knob; female system with a short vaginal tube, dart sac large, glandulae mucosae weakly developed, with a short basic stem and a few short digitiform tubules, seldom exceeding dart sac in length; pedunculus stem long, usually without diverticulum, bursa copulatrix vesicle rounded and bent downwards.

Distribution ( Fig. 147 ): This species has two separate distribution ranges. On the one hand, it is known from Lombardy in Italy and the northern Dalmatian coast. The second distribution range covers some islands in the eastern Aegean Sea and the southwestern part of Turkey. It also lives on Cyprus, the Hatay area in Turkey where it connects to Syria and probably Lebanon. It does not live in central and southern Lebanon.

Remarks: According to the genetic analysis (Korábek & al. submitted), the differences between the Italian and Dalmatian and the East Mediterranean populations are low, and they cannot be separated at species level. It is very likely that this disjunct distribution area is the result of human introduction of the species to Italy rather than a fragmented larger distribution. This hypothesis is also supported by the fact that part of the gap is inhabited by the conchologically similar H. borealis , which, however, differs in the structure of its genital organs ( Fig. 162 ) as well as in the genetic analysis ( Psonis & al. 2014; Korábek & al. submitted). Moreover, the H. cincta in Italy and Dalmatia is often found in the neighbourhood of ancient Roman settlements or heavily populated places, while in the East Mediterranean it is often found in remote places under natural conditions. It has been recorded from Malta (as giuliae ), but has never been found back there ( Giusti & al. 1995). This is probably an early confusion by Bourguignat himself or his shell provider; Malta is far beyond the distribution range of H. cincta .

For the relationship of this species to H. antiochiensis , H. valentini and H. anctostoma refer to these species.

Figures

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.

Fig. 147. Helix (Helix) cincta O. F. MüLLER, 1758. Distribution.

Fig. 162. Helix (Helix) borealis MOUSSON, 1859. Genital organs. NMBE 505540, Turkey, Finike, situs 38.33.