Archiv für Molluskenkunde 150 (1) 13–29 Frankfurt am Main, 30 June 2021

Genital anatomy, jaw, and radula of the species of the genus Jeanneretia (: Cepolidae), endemic to western Cuba

Maike Hernández1, Thierry Backeljau2, 3 & Manuel A. Bauzá1

1 Institute of Ecology and Systematics, Carretera Varona Km 31/5, Capdevila, Havana, Cuba ([email protected] https://orcid.org/0000-0003- 1026-1979, [email protected] https://orcid.org/0000-0002-3712-9794). 2 Royal Belgian Institute of Natural Sciences (RBINS), Vautier- straat 29, B-1000 Brussels, Belgium ([email protected] https://orcid.org/0000-0002-9057-9727). 3 Evolutionary Ecology Group, University of Antwerp, Universiteitsplein 1, B-2610 Antwerp, Belgium. • Corresponding author: M. Hernández.

Abstract. This study provides the first data on the genital anatomy, jaw, and radula of 4Jeanneretia spe- cies (Helicoidea: Cepolidae). The auxiliary copulatory organ in this genus is highly peculiar and differs from that of other cepolids. It consists of an elongated, pedunculated mucus gland inserted apically on a muscular papilla (dart sac absent) and a structure of soft consistency, all covered by a sheath. Two ac- cessory tubular glands are inserted at the base of the atrial sac. Parts of the genitalia of Jeanneretia spp. are unusually large compared to the size of the , such as the bursa-copulatrix duct and flagellum, which can exceed 200 mm in length. Another peculiarity of the genus is the presence of a fertilization pouch–spermathecal complex with a single exposed spermatheca. The jaw is oxygnath, solid, and high arched, with a well-developed, wide median projection, and slight striae. The radula has triangular, monocuspid central and lateral teeth (central teeth smaller than the rest). The transitional teeth have ectocones, and the marginal teeth are multicuspid with a mesocone and ectocones. The features of the reproductive system of Jeanneretia deviate conspicuously from the pattern assumed to be typical of the Cepolidae. However, the atrial sac, accessory gland, and pedunculated mucus gland of the auxiliary copulatory organ of Jeanneretia are considered homologous with, respectively, the atrial sac, accessory gland, and pedunculated mucus gland of the auxiliary copulatory organ in other cepolid genera. Key words. Auxiliary copulatory organ, reproductive system, , West Indies. DOI. https://doi.org/10.1127/arch.moll/150/013-029

Introduction (Baker 1943; Moreno 1950). Yet, for other cepolid gen- era little, if any, information on genital anatomy is avail- The stylommatophoran family Cepolidae Ihering, able. With respect to the genus Jeanneretia L. Pfeiffer, 1909 is one of the least well-studied helicoid families. 1877, for example, there are hitherto only genital data for The family is distributed throughout the Greater Antilles J. parraiana (d’Orbigny, 1841) (Poey 1856–1858) and J. and some Lesser Antilles, the Bahamas, and Florida, in- dermatina (Shuttleworth, 1854) (Baker 1943, copied by cluding the Florida Keys (Miller & Naranjo-García Schileyko 2004), but not for its type species Helix multi­ 1991). striata Deshayes, 1832 (= Helix bicincta Menke, 1830). Ihering (1909) created Cepolinae to include the gen- The genus Jeanneretia comprises 5 species which era Cepolis Montfort, 1810 and Beck, 1837. are characterized by their relatively large (height: According to Nordsieck (1987, 2017), Cepolidae are 13–27 mm), globose to subglobose, yellowish to brown characterized by a reproductive system having a single shells, with or without darker spiral bands. The shell dart apparatus consisting of a dart sac with a pedun- sculpture mainly consists of weak to rather strong spiral culated mucus gland on its apical end, both covered by threads or ridges (Fig. 1). The snails are endemic to west- a sheath. At the base of the dart apparatus is a pair of ern Cuba, where they mainly occur in limestone areas accessory glands, and there is no diverticulum. However, of mountainous regions from the province of Matan- these early diagnoses were largely based on details of zas westward to the province of Pinar del Rio and on the reproductive system of only a small number of taxa, the Isla de la Juventud. The center of species diversity including Setipellis Pilsbry, 1926, Cepolis Montfort, is in the Sierra de los Órganos and at Pan de Guajaibón 1810, Polymita Beck, 1837, and Dialeuca Albers, 1850 (Clench & Aguayo 1951). Pilsbry (1893–1895) placed

© E. Schweizerbart’sche Verlagsbuchhandlung and Senckenberg Gesellschaft für Naturforschung, 2021 ISSN 1869-0963, eISSN 2367-0622