Helix (Helix) borealis 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 : 98-104

Helix (Helix) borealis MOUSSON, 1859 ( Figs 156–163 )

1859 Helix ambigua MOUSSON, Vierteljahrsschrift der naturforschenden Gesellschaft in Zürich 4: 15 [non Helix ambigua LINNAEUS, 1758 ] [De la Grèce et de la Thessalie].

1859 Helix ambigua var. borealis MOUSSON, Vierteljahrsschrift der naturforschenden Gesellschaft in Zürich 4: 16 [Corfou, dans les broussailles des rochers de la citadelle].

1860 Helix cyrtolena BOURGUIGNAT, Revue et Magasin de Zoologie (2e série) 12 (1): 165 [separatum: Aménités Malacologiques II: 183] [nom. nov. pro Helix ambigua MOUSSON, 1859 ].

1878 Helix (Pomatia) Thiesseana KOBELT, Jahrbücher der deutschen Malakozoologischen Gesellschaft 5: 320 [ Greece, Evvia, Chalkis].

1889 H. ambigua clathrata WESTERLUND, Fauna der in der Paläarctischen Region lebenden Binnenconchylien, II. Helix : 459 [Kerkira und Epirus] .

1892 Helix (Pomatia) ambigua var. aetolica KOBELT, Iconographie der Land- und Süsswassermollusken 2 (5): 106, Taf. 146, Fig. 936, 937 [ Greece, Etoliko, Vrachori].

Type specimens:

borealis : probable syntype ZMZ 506307: H = 39.4, D = 35.65, PH = 26.6, PD = 18.9, PrD = 3.6, W = 5.25; thiesseana : lectotype SMF 9627 (SD ZILCH 1952: 151): H = 37.9, D = 38.2, PH = 26.4, PD = 19.6, PrD = 3.9, W = 5.5; clathrata: lectotype GNM 1718 (lectotype herewith designated): H = 34.4, D = 33.1, PH = 24.2, PD = 16.8, PrD = 3.35, W = 5.5; aetolica: syntype SMF 24313a: H = 41.4, D = 39.9, PH = 31.1, PD = 20.7, PrD = 3.85, W = 5.15. ambigua : syntype ZMZ 548103: H = 33.2, D = 30.2, PH = 20.8, PD = 20.1.

Remarks on type specimens: borealis : The collection of Mousson contains six lots labelled as Helix borealis , but only two of them were collected by Schläfli and originate from Sagiada, a place NW of Igoumenitsa. The type specimens of H. ambigua var. borealis from the citadel of Kerkira, which were collected by Schläfli as well, are missing. A third lot contains four specimens from Argostoli on Kefallinia, which were collected by Mousson himself in 1858. The label of this lot (ZMZ 506307) reads " H. cyrtolena Brgt. var. borealis Mss., Argostoli (M. 58)" which proves that it was written after 1860, the publication date of Helix cyrtolena Bourguignat. Mousson does not mention Argostoli in the description of H. ambigua var. borealis , but he had them already at his disposal, so these specimens can only be considered possible syntypes. clathrata: The type lot contained two specimens. The somewhat larger specimen is selected, because it shows the sculpture and bands of the upper whorls as described by Westerlund very clearly. aetolica: As the original specimens for figures Iconographie der Land- und Süsswassermollusken 936 and 937 are not in the SMF ( Zilch 1952: 150), no lectotype was designated. A syntype specimen from SMF 24313 coll. Kobelt ex BrÖmme (not SMF 42313 in Zilch!) is figured.

Specimens examined:

Greece: Corfu: SMF 9848 Corfu , 39.6242 19.9216, coll. Kobelt; ditto, SMF 74133 / 2, coll. C.R. Boettger; SMF 214799 Corfu, Pantokrator, 39.7383 19.8723, coll. S. H. Jaeckel ex Martens ex Brenske. Lefkada : SMF 74062, Island of St. Maura (= Lefkas ), coll. Krüper.; Karia , 2 Km in dir. Lefkada , 38.7756 20. 6554, 440 m alt., 5.8.1975, NMBE 524297 / 1; Nidri , hill slope S of village, 38.6915 20.6979, 20 m alt., 3.8.1975, NMBE 524294 / 1; Sivros , 38.6668 20. 6433, 170 m alt., 3.8.1975, NMBE 524311 / 4; Sivros, Mt. Stavrotas , 38.6704 20.6373, 200 m alt., 3.8.1975, NMBE 524318 / 6; Kaligoni , 38.8148 20.7124, 31.7.1975, NMBE 524310 / 2. Kefalonia: Poros , canyon NW of village, 38.1506 20.771, 30 m alt., 6.5.2005, NMBE 522334 / 8; Argostoli , 1.5 Km SE of village, 38.163 20.5105, 10 m alt., 7.5.2005, NMBE 522331 / 1; Mt. Énos, 7.2 Km from crossroads Argostoli-Sami, 38.1539 20.6311, 1200 m alt., 7.5.2005, NMBE 522336 / 1. Sterea Ellada: SMF 299540 Epirus, coll. C. Bosch ex W. Schlüter; ditto, SMF 74114 / 4, coll. Jetschin ex Conéménos; ditto, SMF 299546 , coll. C. Bosch ex Schlüter; Igoumenitsa , 39.4858 20.2818, 40 m alt., 21.7.1975, NMBE 524298 / 1; ditto, SMF 74112 / 2+1, coll. Jetschin ex Conéménos; Igoumenitsa , 5 Km E of city, 39.4882 20. 3274, 370 m alt., 22.7.1975, NMBE 524296 / 1; SMF 74113, Epirus, Prevesa , 38.9592 20.7517, coll. Jetschin ex Conéménos; ditto, SMF 24308 , coll. O. Boettger ex Conéménos; ditto, SMF 74132 / 3, coll. C.R. Boettger; SMF 24313/4 Vrachori (= Agrinion), 38.6244 21.4095, coll. Kobelt ex BrÖmme (syntypes aetolica Kobelt ); SMF 24305 Vonitsa , Gulf of Amvrakikos , 38.9190 20.8889, coll. O. Boettger ex Conéménos ; Vonitsa , 38.92 20.9089, 50 m alt., 14.4.1988, NMBE 524305 / 1; SMF 74070 Acarnania, Varassovo , coll. Krüper; SMF 74116/5 Acarnania, coll. Krüper; SMF 86163 Acarnania, Messolóngi, 38.3686 21.4304, coll. Nägele ex Krüper; ditto, SMF 74066 / 2, coll. Jetschin ex Krüper; ditto, SMF 214800 , coll. S. H. Jaeckel; SMF 74069/3+1 Acarnania, Aetolico, coll. Krüper; ditto, SMF 74067 , coll. Jetschin ex Krüper; SMF 24307, Aetolia, coll. O. v. Moellendorff; SMF 74122 Kalyakuda (= mountain S of Karpenission), 38.7958 21.7598, coll. Krüper; Zalongo, 500 m from crossroad in dir. Igoumenítsa, 39.1248 20.6568, 90 m alt., 6.8.1975, NMBE 524295 / 1; road Filiates-Igoumenitsa , cave chapel, 39.6434 20. 2135, 150 m alt., 19.7.1990, NMBE 524330 / 1; Kartéri , 39.3953 20. 3702, 170 m alt., 20.7.1990, NMBE 524326 / 3; ditto, 15.5.1995, NMBE 524304 / 1; Kotsika , 39.6592 20.2471, 300 m alt., 13.5.1995, NMBE 524307 / 2; Igoumenitsa, crossroads to Agia Marina , 39.4771 20. 3137, 210 m alt., 25.7.1976, NMBE 524301 / 1; Tsara 200 m in dir. to Prevesa, rocky slope, 02.06.1985, HAUS (3) ; Prevesa 0.5 km in dir. to Arta, 38.9986 20.7389, 03.06.1985, HAUS (1) ; Domiani , 6.5 km in dir. to Agia Trias, 39.0084 21.7938, 16.06.1985, HAUS (1) ; Kato Retsina , 38.4268 21.4313, 08.06.1985, HAUS (1) ; northern part of Klissoura , 25.08.1982, HAUS (1) ; Astakos , 38.5454 21.0902, 25 m alt., 8.5.2005, NMBE 522335 / 3, NMBE 522330 / 3; Astakos , 1 Km E of village, 38.5323 21.0998, 19.5.1991, NMBE 524317 / 16; Astakos, SE slope of Mt. Veloutsa , 38.537 21.0692, 185 m alt., 19.5.1991, NMBE 524328 / 3; Petrona , 38.9258 21.4066, 500 m alt., 20.7.1990, NMBE 524322 / 1; Kefalovrisso , 38.4598 21.367, 30 m alt., 30.9.1989, NMBE 524315 / 7; Ancient Kalydon, 2 Km from bridge over Evinos River , 38.3725 21.5332, 1.10.1989, NMBE 524314 / 3; Karpenissi, 7.5 Km WNW of village, 38.9245/21.7454, 1250 m alt., 9.4.1986, NMBE 524323 / 1; Thessaly: Loutropigi 2 km in dir. to Karditsa , 39.1521 22.0455, 18.06.1985, HAUS (1) ; limestone rocks above Loutropigi S of Karditsa, 39.1172 22.0415, 27.08.1982, HAUS (1) ; ditto, 16.06.1985, HAUS (1) ; Pilion, Platania on the road in dir. to Rodia, 39.1534 23.2827, 15.10.1986, LIEB (2) ; Dafnospilia , S of village, cave, 39.2274 21. 9427, 260 m alt., 12.5.1997, NMBE 522288 / 2; Evvia : Evvia, 1876, NMBE 18449 / 3; ditto, SMF 24304 , coll. Reinhardt ex Heldreich; ditto, SMF 74106 , coll. K.L. Pfeiffer ex Staudinger; ditto, SMF 299543 , coll. C. Bosch ex Rolle ex Schlüter; ditto, SMF 74110 / 5, coll. Krüper; SMF 74109 N Evvia, coll. Jetschin ex Krüper; ditto, SMF 299544 , coll. O. Boettger ex Thiesse ; SMF 9628, Chalkis, 38.4645 23.605, coll. Kobelt ex Thiesse ex Godet (paralectotype thiesseana KOBELT ); ditto, SMF 9826 / 3, coll. O. v. Moellendorff ex Thiesse; ditto, SMF 24309 / 3, coll. O. Boettger ex Thiesse; ditto, SMF 74111 / 2, coll. C.R. Boettger; ditto, 21.5.1983, NMBE 503512 / 6; quarry 2 km E Ellinika , 38.9957 23.3488, 18. 06. 1994, NMBE 512598 / 1; Ellinika, 1.7 Km in dir. Vassilika, 38.9979 23.3488, 35 m alt., 30.4.1991, NMBE 524320 / 2; Kourbatsion (= Artemision ), 39.02 23.23, 58 m alt., 1884, NMBE 18451 / 26/3, NMBE 18455 / 4, NMBE 18454 / 3, NMBE 18453 / 2, NMBE 18452 / 3; Koulouros , 38.86 23.24, 1876, NMBE 18450 / 3. Northern Sporades: SMF 74122 Giura [Alonissos], 39.3953 24.1736, coll. Krüper; ditto, 25.05.1979, LIEB (2) ; SMF 74063/2 Skopelos , coll. Krüper; Skopelos, on the road to Stathmos, 39.0968 23.7219, 1.11.1981, LIEB (1) ; Piperi , 39.3496 24.3252, 11.06.1981, LIEB (2) . Peloponnessos : SMF 24306 Morea (= Peloponnesos), coll. O. Boettger ex E. Brenske; SMF 74107 Achaia in Morea, coll. Jetschin ex Conéménos; SMF 74064/2 Patras, 38.2466 21.7345, coll. Jetschin ex Conéménos; ditto, 74108/2, coll. Jetschin ex Conéménos; ditto, 74065/2, coll. Jetschin ex Schneider; ditto, SMF 24310 Patras , coll. O. Boettger ex Conéménos; ditto, SMF 74068 , coll. Krüper; Kaiafas NW of Zaharo, thermal springs, 37.5193 21.6022, 7.4.1986, NMBE 524308 / 2; SMF 74120, Olympia , 37.6447 21.6254, coll. Krüper; ditto, SMF 299545 / 3, coll. C. Bosch ex Schlüter; ditto, SMF 74119 / 2, coll. K.L. Pfeiffer; SMF 24311, Santameri, 37.9877 21.5724, coll. O. Boettger ex Conéménos; SMF 74118/2, Sparta, 37.0744 22.4302, coll. K.L. Pfeiffer; ditto, SMF 74115 / 5, coll. Krüper; SMF 74123 Taygetos , coll. Krüper; Githio in dir. Sparti, hill slope, 36.7594 22.5562, 30 m alt., 18.2.1981, NMBE 524316 / 7; Skala, 13 Km in dir. Githio , 36.7959 22.5957, 50 m alt., 14.7.1979, NMBE 524303 / 1; Githio, 10 Km in dir. Areopoli , canyon, 36.7299 22.5055, 40 m alt., 18.2.1981, NMBE 524302 / 1; 4 Km N crossroads to Molai , 36.8456 22. 8448, 250 m alt., 14.7.1979, NMBE 524293 / 1; SMF 24312/2 Pylos ( Navarino ), 36.91 21.7, coll. Kobelt; ditto, NMBE 18458 / 4; Eleohori (= Janitsa ), 37.0408 22. 1817, 545 m alt., 10.5.1991, NMBE 524329 / 1; Paleokastro S of Petrohori, 36.9582 21. 6568, 120 m alt., 18.4.1988, NMBE 524325 / 2; Gargaliani, 1 Km in dir. Kiparissia , 37.0667 21.6199, 200 m alt., 10.7.1979, NMBE 524321 / 2; castle of Kiparissias , 37.2506 21.6783, 100 m alt., 29.06.1987, NMBE 512670 / 1; ditto, 10.7.1979, NMBE 524309 / 2; mountain SW Hrisokellaria , 36.7864 21.8801, 400 m alt., 11.5.1991, NMBE 524306 / 1; Pigi, 15 Km in dir. Kalamata , 36.8592 22.2564, 25 m alt., 13.7.1979, NMBE 524300 / 1; Skala 6.5 km in dir. Githion , rocky slope, 36.8053 22.6183, 01.10.1986, HAUS (1) ; crossroads W of Gerakion 5 km in dir. Kosmas, 37.0248 22.7078, 05.10.1986, HAUS (1) ; Kalivia 0.5 km in dir. to Skutarion, 37.3134 22.1389, 04.10.1986, HAUS (1) ; Mistras , 37.0745 22.3666, 05.07.1987, NMBE 512671 / 1; plateau above Gerolimenas , 36.486 22.387, 100 m alt., 7.7.1987, NMBE 512672 / 1; Rizes 1 km to Agios Petros at the crossroads to Astros, 37.4405 22.4880, 06.10.1986, HAUS (3) ; Vouno, Takka Lake , 37.4265 22.413, 680 m alt., 20.4.1988, NMBE 524327 / 7; crossroads to Agriakona , 37.3217 22. 2701, 705 m alt., 8.5.1991, NMBE 524312 / 1; Klitoria, 2 Km S in dir. Tripoli , 37.8808 22.1355, 500 m alt., 9.7.1979, NMBE 524324 / 1; Kerpini, crossroads Kalavrita-Diakofto to Rogi , 38.0534 22.1384, 700 m alt., 13.5.1991, NMBE 524319 / 2; Agionori, 2 Km in dir. Limnes , 37.7439 22.8739, 700 m alt., 4.4.1986, NMBE 524313 / 1. Kythira : SMF 74117/3, Keramoto, 36.1998 22.9731, coll. Krüper; ditto, SMF 86162 , coll. Nägele ex Krüper ex Leonis 1902; ditto, SMF 86161 /2, coll. Nägele ex Krüper.

Turkey: Antalya: ruines of Lymira 7 km N Finike, 36.3620 30.1515, 15.7.1990, MENK (12+alk.1); Sahilkent E of Finike, 36.3024 30.1502, 13.1.1997, NMBE 525615 /8, Hemm (alk. 3); Finike , 36.304 30.1401, 13.1.1997, NMBE 505540 /3; valley above Olympos, ca. 60 km SSW Antalya, 01.10.1986, leg. Rähle, RÄH (1) .

Uncertain localities: SMF 299547/6 Greece, coll. Krüper; SMF 299541/2 Greece, coll. Krüper; SMF 299542/2, Greece, coll. Jetschin; SMF 24303 Egypt, coll. Gysser.

Diagnosis: shell of medium size, thin, protoconch small, aperture very large, dark brown, palatal area with expanded dark brown coloration, umbilicus closed.

Description: shell of medium size to large, thin, elongate spherical, spire high conical; basic shell colour greyish white with brown spirals, sometimes with five distinct brown spiral bands or spiral No. 2 fused with No. 3; protoconch considerably small in comparison to shell, its diameter ranging from 3–4 mm, smooth, white whorls; teleoconch of 3–4 whorls sculptured by fine axial riblets; last whorl only slightly descending below the periphery; aperture suboval to rounded, very large if compared to the total shell height; apertural rim narrow, labial callus weak, dark brown coloured, palatal area usually with an expanded dark brown coloration; columellar triangle small; umbilicus closed in adult shells.

Genital organs ( Fig. 162 ): penis elongate club-shaped, epiphallus as long as penis; flagellum long, exceeding the length of epiphallus+penis at least twice; mrp attaching in a central position on the epiphallus; internally, both pp1 smaller than pp2, penial chamber long, filled with a few strong elongate folds, both papillae conical with a central perforation; atrial stimulator an elongate pilaster terminating in 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, much shorter than dart sac; pedunculus stem broad, diverticulum short and thick, branching off in the proximal third of the pedunculus, remaining bursa copulatrix stem long, bursa copulatrix vesicle rounded and bent downwards.

Distribution ( Fig. 163 ): This species lives in the coastal areas of western Central Greece, the Ionian Island, the western and southern Peloponnesus, and the eastern part of Crete ( Psonis & al. 2014). The specimens from the Northern Sporades need to be carefully compared to H. pelagonesica , they probably belong to this species instead to H. borealis . The specimens from the northern and central part of the island Evvoia were once separated by Kobelt under the name thiesseana because of minor difference in the shell colour. In fact, the specimens from Evvoia are less strongly banded to completely white. The author was able to observe a living specimen on a steep limestone slope at the northern Dirfis Mountain in Evvoia where the species lives on the open rock faces. Here, the white colour perfectly helped to mask the animal. Dhora & Welter-Schultes (1996) list some records from Albania (obviously literature records) under H. cincta , repeating misidentifications of former authors but note that these require confirmation.

Remarks: The separation of the dark-mouthed Helix specimens from Central Greece from H. cincta is under debate ( Psonis & al. 2014). Comparing the shell characters, H. borealis has a somewhat thinner and larger shell, and a larger aperture than H. cincta . This also holds true if compared to H. melanostoma , which is smaller and even more spherical. Psonis & al. (2014) showed that there is a cluster of species originating from central Greece and western Crete, and which the authors identify as " H. cincta ". Another cluster originates from the East Aegean and Cyprus, which is named " H. cincta anatolica ". Both groups are widely separate genetically, supporting their status as independent species. This western " H. cincta " group is what is here considered to be H. borealis , while the " H. cincta anatolica " group represents the true H. cincta . Unfortunately, no specimens from the Greek mainland or the type locality Corfu were available for dissection, so the morphology of the genital organs of the typical population is unknown.

In a very small area in southwestern Turkey, there is a black-mouthed Helix species known, which is here attributed to H. borealis , because its shell is thinner and more globular than that of H. cincta , and is usually almost white. Conchologically, it very much resembles H. thiesseana from Chalkis (Evvoia), which is here considered a synonym of H. borealis . Interestingly, it only occurs in or close to ancient Greek settlements, so it seems highly probable that it is introduced there by human activities.

Figures

Figs 156–161. Helix (Helix) borealis MOUSSON, 1859. Fig. 156 probable syntype borealis: ZMZ 506307, Argostoli, D = 35.65; Fig. 157 lectotype thiesseana: SMF 9627, Chalkis, D = 38.2; Fig. 158 lectotype clathrata: GNM 1718, Kerkira und Epirus, D = 33.1; Fig. 159 syntype aetolica: SMF 24313a, Vrachori, D = 39.9; Fig. 160 syntype ambigua: ZMZ 548103, Ionische Inseln, D = 30.2; Fig. 161 NMBE 505540, Turkey, Finike, D = 30.58. — All photos Neubert, original size.

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

Fig. 163. Helix (Helix) borealis MOUSSON, 1859. Distribution.