Abstract
Hemicycla laurijona sp. nov. and H. fulgida sp. nov. are described from the laurel forests of La Gomera and Tenerife islands, respectively (Canary Islands). Both species belong to the Helicinae group of genera sharing the “presence of a specialized twin papillae system” in the penis, the adaptive advantage of which has not hitherto been discussed. Both species present a proximal penial papilla homologous to the “penial papilla” of other Stylommatophora, and an apomorphic distal penial papilla. The arrangement of these organs is described in a specimen with the distal genital system everted, showing that the distal penial papilla is an accessory papilla whose main function may be to lengthen the male duct in the evaginated penis. The accessory papilla may also have another function; its base forms a protuberant ring in the everted penis, perhaps for anchorage during mating or to impede a too deep penetration. The dart and an undigested part of the spermatophore of H. fulgida are also described, the species status of both H. invernicata (Mousson, 1872) and H. consobrina (A. Férussac, 1822) is confirmed and some aspects of Hemicycla relationships are discussed.References
Adamo, S.A. & Chase, R. (1988) Courtship and copulation in the terrestrial snail Helix aspersa. Canadian Journal of Zoology, 66, 1446–1453.
Adamo, S.A. & Chase, R. (1990) The “love dart” of the snail Helix aspersa injects a pheromone that decreases courtship duration. Journal of Experimental Zoology, 255, 80–87.
Adamo, S.A. & Chase, R. (1996) Dart shooting in helicid snails: An ‘honest’ signal or an instrument of manipulation? Journal of theoretical Biology, 180, 77–80.
Alonso, M.R., Goodacre, S.L., Emerson, B.C., Ibáñez, M., Hutterer, R. & Groh, K. (2006a) Canarian land snail diversity: conflict between anatomical and molecular data on the phylogenetic placement of five new species of Napaeus (Gastropoda, Pulmonata, Enidae). Biological Journal of the Linnean Society, 89, 169–187.
Alonso, M.R., Henríquez, F. & Ibáñez, M. (1991) Nuevas especies de moluscos terrestres (Gastropoda, Pulmonata) de la isla de Alegranza (Archipiélago Canario). Bonner zoologische Beiträge, 42, 325–338.
Alonso, M.R., Ibáñez, M. & Ponte-Lira, C.E. (2002) Canariella (Canariella) falkneri new species (Gastropoda Pulmonata: Hygromiidae) from La Gomera (Canary Islands). In: Falkner, M., Groh, K. & Speight, M.C.D. (Eds.), Collectanea Malacologica—Festschrift für Gerhard Falkner. Conchbooks, pp. 79–87.
Alonso, M.R., Ibáñez, M. & Ponte-Lira, C.E. (2003) Canariella (Salvinia), new subgenus, and three new species of Canariella Hesse, 1918 (Gastropoda: Pulmonata: Hygromiidae). The Veliger, 46, 69–76.
Alonso, M.R., Nogales, M. & Ibáñez, M. (2006b) The use of the computer-assisted measurements utility. Journal of Conchology, 39, 41–48.
Alonso, M.R., Ponte-Lira, C.E., Castillo, C., Yanes, Y., Groh, K. & Ibáñez, M., (2006c) A new Canariella species (Gastropoda: Helicoidea: Hygromiidae) of the new subgenus Majorata, both endemic to the Jandía peninsula (Fuerteventura, Canary Islands). Zootaxa, 1316, 45–56.
Alonso, M.R., Valido, M.J., Groh, K. & Ibáñez, M. (2000) Plutonia (Canarivitrina), new subgenus, from the Canary Islands, and the phylogenetic relationships of the subfamily Plutoniinae (Gastropoda: Limacoidea: Vitrinidae). Malacologia, 42, 39–62.
Bank, R.A., Groh, K. & Ripken, T.E.J. (2002) Catalogue and bibliography of the non-marine Mollusca of Macaronesia (with colour-plates 14–26). In: Falkner, M., Groh, K. & Speight, M.C.D. (Eds.), Collectanea Malacologica—Festschrift für Gerhard Falkner. Conchbooks, Hackenheim, 89–235.
Beese, K., Beier, K. & Baur, B. (2006) Bursa tract diverticulum in the hermaphroditic land snail Arianta arbustorum (Stylommatophora: Helicidae): Morphology, function, and evolutionary implications. Journal of Morphology, 267, 940–953.
Beck, T. & Rähle, W. (2006) Description of a newly discovered extinct representative of the genus Hemicycla Swainson, 1840 (Gastropoda, Pulmonata, Helicidae) from La Gomera, Canary Islands. Basteria, 70, 53–56.
Boettger, O. (1908). Liste der Mollusken aus einem Sande im Barranco von Tegina auf Tenerife (Canaren). Zeitschrift der Deutschen geologischen Gesellschaft, 60, 246–249.
Castillo, C., Yanes, Y., Alonso, M.R. & Ibáñez, M. (2006) Napaeus lajaensis sp. nov. (Gastropoda: Pulmonata: Enidae) from a Quaternary Aeolian Deposit of Northeast Tenerife, Canary Islands. Zootaxa, 1307, 41–54.
Chase, R. & Blanchard, K.C. (2006) The snail’s love-dart delivers mucus to increase paternity. Proceedings of the Royal Society B, 273, 1471–1475.
Chung, D.J.D. (1986) Stimulation of genital eversion in the land snail Helix aspersa by extracts of the glands of the dart apparatus. Journal of Experimental Zoology, 238, 129–139.
Davison, A., Wade, C.M., Mordan, P.B. & Chiba, S. (2005) Sex and darts in slugs and snails (Mollusca: Gastropoda: Stylommatophora). Journal of Zoology, London, 267, 329–338.
Dorello, P. (1923) Sopra uno speciale organo annesso al pene nel genere Helix. Atti Reale Accademia Nazionale dei Lincei, anno 320, serie quinta, Rendiconti Classe di scienze fisiche, matematiche e naturali, 32, 257–259.
Dorello, P. (1924) Osservazioni sopra il corpo erettile del pene nel genere Helix. Atti Reale Accademia Nazionale dei Lincei, anno 321, serie quinta, Rendiconti Classe di scienze fisiche, matematiche e naturali, 33, 412–415.
Férussac, A.E.J.P.J.F. d'Audebard de (1821–1822) Tableaux systématiques des animaux Mollusques, classés en familles naturelles, dans lesquels on a établi la concordance de tous les systèmes; suivis d'un prodrome général pour tous les mollusques terrestres ou fluviatiles, vivants ou fossiles.—3 parties; pp. V–XLVII [Tableaux systématiques] and plates [1822] ; pp. 1–92 (= Folio edition), pp. 1–88 (= Quarto edition) [limaçons] [1821]. Arthur Bertrand Libr., Paris.
Giusti, F. & Andreini, S. (1988) Morphological and ethological aspects of mating in two species of the family Helicidae (Gastropoda: Pulmonata): Theba pisana (Müller) and Helix aperta Born. Monitore Zoologico Italiano (Nuova Serie), 22, 331–363.
Giusti, F. & Lepri, A. (1981) Aspetti morfologici ed etologici dell'accoppiamento in alcune specie della famiglia Helicidae (Gastropoda: Pulmonata). In: Giusti F. (Ed.), Atti del IV° Congresso della Società Malacologica Italiana (Siena 6–9 Ottobre, 1978), pp. 11–71.
Giusti, F., Manganelli, G. & Schembri, P.J. (1995) The non-marine molluscs of the Maltese Islands. Monografie Museo Regionale di Scienze Naturali (Torino), 15, 607 pp.
Grasset, A. (1857) Description de Coquilles nouvelles provenant des îles Canaries. Journal de Conchyliologie, 5, 345–348, pl. 13.
Groh, K. (1985) Landschnecken aus quartären Wirbeltierfundstellen der Kanarischen Inseln (Gastropoda). Bonner zoologische Beiträge, 36, 395–415.
Groh, K., Alonso, M.R., Ibáñez, M. & Henríquez, F. (1992) Rediscovery of Hemicycla saulcyi (d'Orbigny, 1839), a revision of its fossil allies (Gastropoda: Helicidae), and a description of a new species of Napaeus (Enidae), both from La Isleta, Gran Canaria, Canary Islands. Schriften zur Malakozoologie, 5, 1–12, pl. 1–3.
Groh, K., Hutterer, R. & Voggenreiter, V. (1996) On the identity of Helix digna Mousson, 1872, and description of another extinct helicid from La Gomera, Canary Islands. Basteria, 59, 115–125.
Gude G.K. (1896) Description of a new species of Vitrina, and new forms of Helicidae, with a list of the helicoid shells hitherto found in the Canary Islands. Proceedings of the Malacological Society of London, 2, 15–22.
Hesse, P. (1912) Die Anatomie einiger Arten des Genus Hemicycla Swainson. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 31, 76–82, pl. 3.
Hesse, P. (1918) Die Subfamilie Helicodontinae. Nachrichtenblatt der Deutschen malakozoologischen Gesellschaft, 50, 99–120.
Hesse, P. (1920) Subfamilia Helicinae. In: E. A. Rossmässler (forgesezt von W. Kobelt), Iconographie der Land- und Süßwasser-Mollusken mit vorzüglicher Berücksichtigung der Euroipäischen nocht nicht abgebildeten Arten, N.F., 23, 153–262, pl. 651–660.
Ibáñez, M. & Alonso, M.R. (2001) La extinción reciente de la malacofauna endémica en Tenerife y Gran Canaria. Estudios Canarios, 45, 9–22.
Ibáñez, M. & Alonso, M.R. (in press) A tale of two snails: Le Pouchet, from Adanson (Mollusca, Gastropoda, Helicoidea, Helicidae). Zoosystema.
Ibáñez, M., Alonso, M.R. Groh, K. & Hutterer, R. (2003) The Genus Obelus Hartmann, 1842 (Gastropoda, Pulmonata, Helicoidea) and its phylogenetic relationships. Zoologischer Anzeiger, 242, 157–167.
Ibáñez, M., Alonso, M.R. Henríquez, F. & Valido, M.J. (1997) Distribution of land snails (Mollusca, Gastropoda, Pulmonata) on the island of Gran Canaria (Canary Islands) in relation to protected natural areas. Biodiversity and Conservation, 6, 627–632.
Ibáñez, M., Alonso, M.R. & Ponte-Lira, C.E. (2002) A new subgenus and two new species of Canariella Hesse 1918 (Gastropoda: Pulmonata: Hygromiidae). American Malacological Bulletin, 17, 85–90.
Ibáñez, M., Alonso, M.R. & Valido, M.J. (2001) Plutonia solemi new species (Gastropoda: Vitrinidae: Plutoniinae) from La Palma (Canary Islands). Journal of Conchology, 37, 149–157.
Ibáñez, M., Alonso, M.R., Yanes, Y., Castillo, C.& Groh, K. (in press) Presence of the genus Napaeus (Gastropoda: Pulmonata: Enidae) living in all the islands of the Canarian archipelago: Napaeus lichenicola sp. nov. from Fuerteventura island. Journal of Conchology.
Ibáñez, M., Barquín, J., Cavero, E. & Alonso, M.R. (1988a) La variabilidad de Hemicycla bidentalis (Gastropoda: Helicidae). Malacologia, 28, 105–117.
Ibáñez, M., Groh, K., Alonso, M.R. & Cavero, E. (1988b) Revision of the genus Hemicycla Swainson, 1840 (Mollusca, Helicidae) from Tenerife: Adiverticula n. subgen., and description of 3 new taxa. Bulletin du Muséum national d'Histoire naturelle, série 4, section A, Zoologie, 10, 309–326.
Ibáñez, M., Groh, K., Alonso, M.R., Castillo, C., Yanes, Y., (2006a) The subgenus Monilearia (Lyrula) Wollaston, 1878 (Gastropoda: Helicoidea: Cochlicellidae), from Lanzarote and Fuerteventura (Canary Islands), with the description of Monilearia (Lyrula) tubaeformis sp. nov. Zootaxa, 1320, 29–41.
Ibáñez, M., Groh, K., Cavero, E. & Alonso, M.R. (1987) Revision of the genus Hemicycla Swainson 1840 on Tenerife: The group of H. plicaria (Lamarck, 1816) (Mollusca: Helicidae). Archiv für Molluskenkunde, 118, 77–103.
Ibáñez, M., Ponte-Lira, C.E. & Alonso, M.R. (1995) El género Canariella Hesse, 1918, y su posición en la familia Hygromiidae (Gastropoda, Pulmonata, Helicoidea). Malacologia, 36, 111–137.
Ibáñez, M., Siverio, F., Alonso, M.R. & Ponte-Lira, C.E. (2006b) Two Canariella species (Gastropoda: Helicodea: Hygromiidae) endemic to the Northwest Tenerife (Canary Islands). Zootaxa, 1258, 33–45.
Kerney, M.P. & Cameron, R.A.D. (1979) A field guide to the land snails of Britain and North-West Europe. William Collins Son & Co., Ltd., London, 288 pp.
Koene, J.M. & Chase, R. (1998) Changes in the reproductive system of the snail Helix aspersa caused by mucus from the love dart. Journal of Experimental Biology, 201, 2313–2319.
Koene, J.M. & Chiba, S. (2006) The way of the Samurai snail. The American Naturalist, 168, 553–555.
Koene, J.M. & Ter Maat, A. (2001) "Allohormones": A class of bioactive substances favoured by sexual selection. Journal of Comparative Physiology A—Neuroethology sensory neural and behavioral Physiology, 187, 323–326.
Koene, J.M. & Ter Maat, A. (2002) The distinction between pheromones and allohormones—Reply. Journal of Comparative Physiology A—Neuroethology sensory neural and behavioral Physiology, 188, 163–164.
Landolfa, M.A., Green, D.M. & Chase, R. (2001) Dart shooting influences paternal reproductive success in the snail Helix aspersa (Pulmonata, Stylommatophora). Behavioral Ecology, 12, 773–777.
Lamarck, J.B. de (1816) Encyclopédie méthodique. Tableau encyclopédique et méthodique des trois règnes de la nature. Vingt-troisième partie. II. Mollusques et polypes divers. V. Agasse, Paris, 1–16, pl. 96–314.
Lamarck, J.B. de (1822) Histoire naturelle des animaux sans vertèbres, presentant les caractères généraux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres et la citation des principales espèces qui s'y rapportent; precedée d'une Introduction offrant la détermination des caractères essentiels de l'Animal, sa distinction du vegetal et des autres corps naturels, enfin, l'Exposition des Principes fondamentaux de la Zoologie. J.B. Báilliére Libr., Paris, 6 (2), 232 pp.
Lowe, R.T. (1861) Diagnoses of New Canarian Land-Mollusca. The Annals and Magazine of Natural History, Zoology, Botany and Geology, series 3, 7, 104–112.
Mabille, J. (1882) Molluscornm [sic!] novorum diagnoses succintae. Bulletin de la Société philomatique de Paris, (7) 6, 132–147.
Mabille, J. (1883a) Sur quelques espèces de mollusques terrestres. Bulletin de la Société philomatique de Paris, (7) 7, 39–53.
Mabille, J. (1883b) Diagnoses testarum novarum. Bulletin de la Société philomatique de Paris, (7) 7, 115–132.
Mabille, J. (1885) Matériaux pour une faune malacologique des Iles Canaries. I. Nouvelles Archives du Muséum d'Histoire naturelle, (2) 8 (1), 17–182.
Mabille, J. (1898) Notitiae malacologicae. Bulletin de la Société philomatique de Paris, (8) 9 (2), 78–102.
Meisenheimer, J. (1907) Biologie, Morphologie und Physiologie des Begattungsvorganges und der Eiablage von Helix pomatia. Zoologischer Jahrbücher (Abteilung für Systematik), 25, 461–502.
Meisenheimer, J. (1912) Die Weinbergschnecke Helix pomatia L. In: Ziegler, H.E. & Woltereck, R. (Eds.), Monographien einheimischer Tiere, 4. Verlag von Dr. Werner Klinkhardt, Leipzig, pp. 1–140.
Mermod, G. (1951) Les types de la collection Lamarck au Muséum de Genève. Mollusques vivants. II. Revue Suisse de Zoologie, 58, 693–753.
Morelet, A. (1864) Descriptions de Coquilles inédites. Journal de Conchyliologie, 12, 155–159.
Mousson, A. (1872) Révision de la faune malacologique des Canaries. Neue Denkschriften der allgemeinen schweizerischen Gesellschaft für die gesammten Naturwissenschaften, 25 (1), I–V, 1–176 pp., 6 pl.
Neubert, E. & Bank, R.A. (2006) Notes on the species Caucasotachea C. Boettger 1909 and Lindholmia P. Hesse 1919, with annotations to the Helicidae (Gastropoda: Stylommatophora: Helicidae). Archiv für Molluskenkunde, 135, 101–132.
Neubert, E. & Gosteli, M. (2003) The molluscan species described by Robert James Shuttleworth. I. Gastropoda: Pulmonata. Contributions to natural history, Bern, 1, 1–123.
Odhner, N.H. (1932) Beiträge zur Malakozoologie der Kanarischen Inseln. Lamellibranchien, Cephalopoden, Gastropoden. Arkiv för Zoologi, 23 A, 1–116, pl. 1–2.
Orbigny, A. d' (1836–1842) Mollusques, Échinodermes, Foraminifères et Polypiers, recueillis aux Îles Canaries par MM. Webb et Berthelot. Mollusques. In: P.B. Webb & S. Berthelot (eds.), Histoire Naturelle des Îles Canaries. Tome II. Partie 2. Zoologie. Paris, 152 pp, 14 pls.
Ortiz, J.E., Torres, T., Yanes, Y., Castillo, C., de la Nuez, J., Ibáñez, M. & Alonso, M.R. (2006) Climatic cycles inferred from the aminostratigraphy and aminochronology of Quaternary dunes and paleosols from the eastern islands of the Canary Archipelago. Journal of Quaternary Science, 21, 287–306.
Pfeiffer, L. (1859) Nachträge zum zweiten Supplemente meiner Monographia Heliceorum. Malakozoologische Blätter, 6, 19–48.
Pilsbry, H.A. (1939) Land Mollusca of North America (North of Mexico). Academy of Natural Sciences of Philadelphia, Monographs, 3, volume I, part 1, XVII + 573 pp + IX (index).
Reeve, L.A. (1852) Conchologia Iconica: or, illustrations of the shells of molluscous animals. Vol. VII. Monograph of the genus Helix. Reeve, London, pl. 63–146.
Rogers, D. & Chase, R. (2002) Determinants of paternity in the garden snail Helix aspersa. Behavioral Ecology and Sociobiology, 52, 289–295.
Schileyko, A.A. (2006) Treatise on recent terrestrial pulmonate molluscs. Part 13. Helicidae, Pleurodontidae, Polygyridae, Ammonitellidae, Oreohelicidae, Thysanophoridae. Ruthenica, Supplement. 2, 1765–1906.
Shuttleworth, R.J. (1852a) Diagnosen einiger neuer Mollusken aus den Canarischen Inseln. Mittheilungen der naturforschenden Gesellschaft in Bern, 241/242, 137–146.
Shuttleworth, R.J. (1852b) Diagnosen neuer Mollusken. Mittheilungen der naturforschenden Gesellschaft in Bern, 260/261, 289–304.
Shuttleworth, R.J. (1975) Tabulae ineditae Molluscorum Insularum Canariensium. Backhuys, W. (Ed.), Goecke & Evers, Krefeld, 43 pp., 8 pl.
van Osselaer, C & Tursch, B. (2000) Variability of the genital system of Helix pomatia L., 1758 and H. lucorum L., 1758 (Gastropoda: Stylommatophora). Journal of Molluscan Studies, 66, 499–515.
Webb, P.B. & Berthelot, S. (1833) Synopsis molluscorum terrestrium et fluviatilium quas in itineribus per insulas Canarias, observarunt. Annales des Sciences naturelles, 28, 307–326.
Wollaston, T.V. (1878) Testacea Atlantica or the land and freshwater shells of the Azores, Madeiras, Salvages, Canaries, Cape Verdes and Saint Helena. L. Reeve, London, xi + 588 pp.
Wood, W. (1828) Supplement to the Index Testaceologicus; or a catalogue of shells, British and foreign. W. Wood, London, IV + 59 pp., 8 pl.