Helix (Helix) pronuba WESTERLUND & BLANC , 1879

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 : 122-126

Helix (Helix) pronuba WESTERLUND & BLANC, 1879 ( Figs 190–197 )

1859 Helix nucula , – L. Pfeiffer, Monographia heliceorum viventium 4: 160 [Alexandria] [non Helix figulina var. nucula MOUSSON, 1854 ].

1879 Helix thiesseana var. pronuba WESTERLUND & BLANC, AperÇu sur la faune malacologique de la Grèce: 80 [Crete, Messarà].

1879 Helix thiesseana var.? nupta WESTERLUND & BLANC, AperÇu sur la faune malacologique de la Grèce: 82 [Crete, Messarà].

1904 Helix (Helicogena)? rueppelli KOBELT, Systematisches Conchylien-Cabinet VI : 122, Taf. 325, Fig. 13, 14 [Egypten].

1905 Helix (Helicogena) grothei KOBELT, Systematisches Conchylien-Cabinet VI : 201, Taf. 349, Fig. 7, 8 [Regentschaft Tripolis, Libyen].

1950 Helix nucula forma incultarum, – Kaltenbach, Archiv für Molluskenkunde 79 (1/3): 62, Taf. 10, Fig. 9 [Cyrenaika, Plateau s. der Östl. Kaserne von Derna].

1950 Helix nucula forma semidesertorum, – Kaltenbach, Archiv für Molluskenkunde 79 (1/3): 62, Taf. 10, Fig 10–11 [Cyrenaika, Dünen beim südlichen Salzsee von el-Coefia, etwa 11 km n. Bengasi].

Type specimens:

nucula : no original specimens were found in major European museums. pronuba : syntype GNM 1723, H = 26.3; D = 25.5; PH = 19.8; PD = 12.7; PrD = 4.3; W = 4.75. nupta: no specimens found in GNM. rueppelli : holotype SMF 9702, D = 26.7. grothei : holotype SMF 9836, not measured.

Note: In contrast to the commonly accepted use throughout the last 150 years, Pfeiffer did not describe a new species in 1859 but simply tried to redefine H. nucula Mousson using the specimen(s?) he had received from Parreyss. The quotation of Mousson in the synonymy of this species shows that he was aware of the true authorship of this name. Unfortunately, this was in fact the dark-mouthed North African Helix , and not the true white-mouthed H. nucula from Turkey. For this reason, the next available name has to be used, i.e. pronuba WESTERLUND & BLANC, 1879 .

Specimens examined:

Greece, Crete: Arkasa, Karpathos , 35.4782 27.1196, 9.7.1982, NMBE522643 / 1; Lendas , Gortys , Kreta, 34.9308 24.9243, 29.12.1984, NMBE503511 /9, SMF/20; Palaiochora, Kreta, 35.2268 23.6811, 27.12.1980, NMBE 503510 /3; Hagios Deka, Gortys, Kreta, 35.06 24.95, 2.5.1994, NMBE 18457 / 1. Dodekanissos: Chalki Island , 21.5.1983, SMF/8, NMBE 528722 /3.

Egypt (if not mentioned otherwise, all samples from coll. Kaltenbach): Alexandria (Al Iskandariyah), 31.2 29.92, NMBE 22856 /3; ditto, 1854, NMBE 22853 /2; ditto, SMF 24427 /1, coll. Rossmässler; ditto, SMF 24439 /1, ex Knobbe; ditto, SMF 74104 , coll. C. R. Boettger; Djebel Bakhevia, Upper Nil, NMBE18493 /8; Port Said / Bur Said, 31.27 32.3, 2.1913, NMBE18479 /1; SMF 172486/3, El Alamein, 30.8199 28.9504; SMF 172476/2, 83 km W Alexandria, 30.816 29.1599; SMF 172447/1, 32 km E Sidi Barani, 31.5312 26.2473; SMF 172478/1, 9.4 km W Sidi Hamid, 30.6206 30.8095; SMF 172479/ 2, 108 km W Marniet (?); SMF 172480/1, 10 Alexandria in dir. Kairo, 31.1739 30.0766; SMF 172481/ 1, 102 km W Marsa Marniet; SMF 172482, Alexandria, S of 1. dam over the Mariut Lake, 31.1712 29.9058; SMF 172483/1, 85 km E Sollum [Sidi Barami], 31.6012 25.9387; SMF 172484/1, Bakis (= Bakisch, Bakus?) W Alexandria, 31.2379 29.9671; SMF 172485/2, valley 183.7 kmW Alexandria, 31.0670 28.1787; SMF 172487/1, Mariut Lake 3 km from Alexandria in dir. Kairo, 31.1655 29.9242; SMF 172492/8, Alexandria, garden of the Borromaeum; SMF 172493/4, 10 km E Sollum, 31.5140 25.2022; SMF 172494/7, km 344 E Sollum , 31.0283 28.4991; SMF 172488/2, between the 2 branches oft he Mariut Lake , 31.1439 29.8656; SMF 172489/1, Marsa Matruh, harbour, 31.3615 27.2456; SMF 172490/1, 20 km from Sidi Barami to Marsa Matruh, 31.5522 26.1262; SMF 172491, few kilometers E Sid Abd el Bachwan; SMF 172495/7, Alexandria, hill SW of 1. dam over the Mariut Lake ; ditto, 172496/5,; SMF 172497, Alexandria, hills W of Mariut Lake; ditto, 172498/4; SMF 172499/10, Abu Mena SW Alexandria, 30.841 29.663; SMF 172502/2, 14 km E Beleig [Balig Hasaf?] (= 8 km W crossroads Alexandria – Kairo), 31.0802 30.3187; SMF 172503/2, 3.5 km W Matruh, 31.3539 27.1876; SMF 172500/13, 43 km from Alexandria to Kairo, 31.0735 30.3447; SMF 172508/1, 3 km E Debba; SMF 172509/8, road Alexandria to Sollum , somewhat W Behig, 30.9458 29.5106; SMF 172510/12, Abu Mena, 40 km SW Alexandria, 30.841 29.663; SMF 172504/ 2, 107 km W Marsa Matruh, 31.5333 26.2367; SMF 172505/1, 56 km W Alexandria, 30.9099 29.4355; SMF 172506/1, 10 km E Marsa Matruh, 31.3416 27.3408; SMF 172507/2, Behig, 30.9322 29.6066; SMF 172511/17, quarry N Behig, 30.9501 29.5801; SMF 172512/29, 11 km W road Alexandria to Kairo, 31.1647 30.085, coll. Kaltenbach; SMF 24438/2, Ramleh close to Alexandria, 31.228 29.9762; ditto, SMF 24434 /4, ex Loebbecke; ditto, SMF 74099 /2, coll. Jetschin; ditto, SMF 74105 /3, coll. C. R. Boettger; SMF 9686/2, unknown, Orig. MCH 325, fig. 10, 11 (reduced granulation); SMF 24225/2, Kairo, Palais of the Khediva; SMF 1411/2, Heliopolis, 30.0917 31.3224, ex Rolle; SMF 24436/3, El Mecks close Alexandria, 31.1445 29.8367, coll. O. Boettger; SMF 24444/21, Mariut Alexandria, 31.1712 29.9058, ex Schweinfurth .

Libya (if not mentioned else, all from coll. Brandt or Kaltenbach). Cyrenaica: SMF 158511/24, Cyrene, Roman bath, 32.8234 21.8528; SMF 86132, Derna, 32.7667 22.6334, ex Stürmer; ditto, 16.12.1956, NMBE 18499 /3; SMF 158526/7, Tokra pass; SMF 158573/12, Tokra, castle, 32.5341 20.5666; SMF 158525/8, 6 km S Agedabia [Adschabiya], 30.7087 20.2742, ditto SMF 1558538/7; SMF 37027/1, Tobruk, 32.0772 23.9608, ex Wohlberedt; SMF 280587/5, Cyrenaika, Apollonia, 32.8997 21.9666; SMF 158519/6, Wadi Garhar, 12 km S Derna (Gahham?), 32.6644 22.5829; SMF 158515/17, steppe below Benina pass; SMF 158507/20, Cami close to El Kish, Bengasi; SMF 158523/7, Wadi Bu Msaper [Wadi Bu Musafir?], close Derna, 32.7351 22.5899; SMF 158518/5, Wadi Bu Msefer S Derna = [Wadi Bu Musafir?]; SMF 158506/19, Driana [Daryanah], E of lighthouse, 32.345 20.3031; SMF 158520/9, Wadi El Gattara [Wadi al Qattarah], 32.0543 20.0331; SMF 158524/12, Zuetina [Az Zuwaytinah], 30.9522 20.1203; SMF 161222/5, Wadi Scalium close to El Labiat; SMF 158522/11, Derna, 6 km S Fort Marabuta, 32.7043 22.6108; SMF 158514/5, Rues Mlella; SMF 158532/6, Wadi Sakal w Tobruk, 32.0762 23.9611; SMF 161450/5, Marsa Breza [Marsa Brega (al-Buraiqa)], 30.4092 19.5708; SMF 158529/5, Wadi Fil ; SMF 158521/7, 13 km S Magrun [Al Maqrun], 31.3342 20.1991; SMF 158531, Hosc es Zuama SE Bengasi [Hawsh, Ḩawsh Shatwān?], 31.9659 20.1622; SMF 158540/4, El Kisch [Dar el Kish], 32.0936 20.07, ditto, 8.10.1956, NMBE 18494 /3; SMF 158517/4, Ain Mara, 32.7511 22.3855; Wadi Mradem [Wadi el Maradim], Fiorita, 32.8713 22.3067, 12.4.1959, NMBE 18498 /4; Wadi el Maalegh, Martuba [Wādī al Mu'allaq], 32.3795 23.0886, 4.12.1955, NMBE 18497 /4; ditto, NMBE 18496 /4; Haua Zeiana, Benghasi [Ayn Zayyānah?], 32.2134 20.161, 3.3.1956, NMBE 18495 /5. Sirte: SMF 172840/80, 9 km E Sirte city, 31.2082 16.7062; SMF 172835/6, 21 km S Mesrata along the coast, 32.216 15.3151; SMF 1172836/9, 99 km W Agedabia from Sirte, 30.2847 19.4083; SMF 172837/12, 7 km W Agedabia from Sirte, 30.6911 20.1807; SMF 172838/14, 18 km E Sirte city, 31.1925 16.7802; SMF 172839/11, E Buerat [Bwayrat al Ḩasūn], 84 km W Sirte, 31.3989 15.7216; SMF 172834/12, 32 km E Buerat, 31.2719 16.0292; SMF 172833/12, 3 km E crossroads (from coastal road?) to Nafilia [An-Nufalija], 30.8446 17.9132; SMF 172819/1, 58.5 km W crossroads to Nafilia , 31.0396 17.3222. Tripolitanien : SMF 172900/77, Wadi Goam bei Houm Genit (?); SMF 172898/25, Sidi ben Aden [Suani Beni Aden], 25 km S Tripolis, 32.7172 13.077; SMF 172899/51, 16 km S Tauorga [Tawurgha] close Misrata, 31.8301 15.0407; SMF 172897/30, 11 km N Mirda ( Mizda ?), 31.5484 12.9919; SMF 172896/16, between Bir Gneim and Jeffren, Garian Mts. [Bi'r al Ghanam and Jafran], 32.1668 12.4918; SMF 172895/18, 7 km E Zneria [Zuwara], 103 km W Tripolis, 32.8933 12.1426; SMF 172876/ 4, 116 km W Buerat, 32.0185 15.0415; SMF 172877/2, 4 km W Jeffren, 32.0803 12.5042; SMF 172880/1, 29 km E Zuevia, 32.7941 12.3445; SMF 1722889/1, 6 km N Mirda, 31.5035 12.9903; SMF 172879/2, 5 km S Mirda, 31.4076 13.0029; SMF 172882, 100 km E Tripolis, 32.6305 14.2075; SMF 172883/1, 9 km E Misurata, 32.3673 15.1932; SMF 172887/1, 45 km N Mirda, 31.8489 12.9840; SMF 172889/4, 93 km W Tripolis, 32.8476 12.2337; SMF 172894/12, 59 km N Mirda in dir. Garian , 31.9733 12.9841 .

Tunisia: Djerba Island , 33.88 10.85, ex Pallary, NMBE 22855 / 2; Djerba , Plage Sidi Maharez, 33.8595 10.9834, 20.08.1993, NMBE 512634 /1.

Diagnosis: shell small to very small, spherical, white or with up to 5 spirals, teleoconch with fine and densely spaced riblets and a prominent granulation, aperture deep chocolate brown.

Description: shell of small to very small size, thin, spherical, spire conical; basic shell colour greyish white with chestnut brown spirals, often the upper spirals fused to a single brown band, completely white shells abundant; protoconch large in comparison to shell, its diameter ranging from 4–5 mm, smooth, white whorls; teleoconch of 3–4 whorls, sculptured by fine and densely spaced riblets, crossed by deeply incised spiral furrows producing a prominent pattern of granules; last whorl slightly descending below the periphery; aperture rounded, apertural rim narrow and weak, labial callus weak, deep chocolate brown, columellar triangle small; umbilicus closed in adult shells.

Genital organs: There were no specimens available for dissection.

Distribution ( Fig. 197 ): This species is known from Crete and some East Aegean islands (own records; Psonis & al. 2014, under H. nucula ), and from the northern African coast from the Nile delta westwards to Tunisia. In the material paragraph only a selection of localities for H. pronuba is recorded, for more information on Libyan records see Kaltenbach (1950).

Remarks: The taxa named by Kaltenbach in 1950 were described as ecological forms (incultarum, semidesertorum) and thus have no status. At least the forma semidesertorum is illustrated to show the variability of this interesting Helix species living along the border of the Sahara.

There is an interesting shift of character sets from east to west. In Egypt, the typical form ( Helix nucula L. Pfeiffer auct.) occurs, with spirally arranged granulation, and clear brown spiral bands; this pattern continues westwards into the Cyrenaica. However, in the Syrte area, the granulation becomes almost obsolete, although still present in some specimens. Here, shells tend to be very small, almost smooth and whitish to cream without spiral bands; in western Libya, shells tend to become larger again, and also the granulation becomes stronger again, while the ribs remain small. For differentiation to H. melanostoma refer to this species.

Figures

Figs 190–196. Helix (Helix) pronuba WESTERLUND & BLANC, 1879. Fig. 190 pronuba: syntype GNM 1723, Messarà, D = 25.5; Fig. 191 rueppelli: holotype SMF 9702, Egypten, D = 26.7; Fig. 192 SMF 172512, Egypt, 11 km W road Alexandria to Kairo, D = 30.1; Fig. 193 SMF 172492, Egypt, Alexandria, garden of the Borromaeum, D = 20.7; Fig. 194 SMF 214791, Libya, El Kish close to Barka, D = 24.1; Fig. 195 semidesertorum, smallest specimen from W Agheila, D = 16.7; Fig. 196 SMF 172900, Libya, Tripolitania, Wadi Goam close Houm Genit, D = 26.7. — All photos Neubert, original size.

Fig. 197. Helix (Helix) pronuba WESTERLUND & BLANC, 1879. Distribution.

Figs 10–19. Helix (Helix) pomatia LINNAEUS, 1758. Fig. 10 claudiensis, holotype SMF 9543, Klausenburg, D = 39.5; Fig. 11 dobrudschae, holotype SMF 9541, Rumania, Dobrudscha, Comana Vlasca, D = 46.2; Fig. 12 costellata, holotype SMF 9631, Markt in Bukarest, D = 39.7; Fig. 13 elsae, lectotype SMF 9529, nahe Bukarest, D = 34.9; Fig. 14 transsylvanica, holotype SMF 9634, Markt in Bukarest, D = 40.1; Fig. 15 christinae, lectotype SMF 9542, Runku über Bukarest, D = 44.5; Fig. 16 kapellae, holotype SMF 9544, Kapella Gebirge, D = 44.2; Fig. 17 serbica, holotype SMF 9540, Maidanek, D = 53.9; Fig. 18 thessalica, lectotype SMF 9568, Makrinitza bei Volos, D = 45.4; Fig. 19 rhodopensis, syntype SMF 9599/2a, Philippoppel, D = 41.0. — All photos Neubert, original size.

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.