A N N A L E S Z O O L O G I C I (Warszawa), 2008, 58(4): 857-880 LIFE CYCLES OF CLAUSILIIDS OF POLAND – KNOWNS AND UNKNOWNS TOMASZ K. MALTZ1 and ANNA SULIKOWSKA-DROZD2 1Museum of Natural History, Wrocław University, Sienkiewicza 21, 50-335 Wrocław, Poland; e-mail: [email protected] 2Department of Invertebrate Zoology and Hydrobiology, University of Łódź, Banacha 12/16, 90-237 Łódź, Poland; e-mail: [email protected] Abstract.— Among the 24 native clausiliids, 15 were subject to laboratory observations. Eleven of them were found to be oviparous, three – egg retainers and one – ovoviviparous. Batches, containing most often one to about a dozen of partly calcified, ellipsoidal or spherical eggs, appeared usually in the spring and autumn (in non-hibernating individuals throughout the year). Probably the main factors determining the onset of reproduction are humidity and temperature while the photoperiod has no significant effect. The incubation period is ca. two weeks (room temperature), the hatching is synchronous or asynchronous. Cases of intra-batch and inter-batch cannibalism were observed. The minimum time from hatching/birth till adult size is ca. 3–9 months and after further 5–8 months the snails start producing eggs/babies. Clausiliids are iteroparous. Anatomical studies on the development of the reproductive system show that just before lip completion the reproductive system is still incompletely developed. Penis, epiphallus and spermatheca develop within the first month after growth completion (which would indicate attainment of ability to copulate), and the reproductive system becomes wholly mature only after a few months. The clausiliid development strategy is probably the following: the quickest possible growth and attainment of adult size, then development of the reproductive system and attainment of sexual maturity. ± Key words.— Terrestrial snails, Clausiliidae, life cycle, reproduction strategy, reproductive season, development, growth, maturity. INTRODUCTION by one species (4.1%) (Riedel 1988). A detailed classifi- cation is presented in Table 2. The European group Recent terrestrial malacofauna of Poland includes includes widely distributed species found in the whole 174 species (Wiktor 2004) classified in 27 families. country (8 species), and some with regional ranges: Besides the Helicidae (19% species) and the Zonitidae Clausilia bidentata (lowlands of north-western (11.5%), one of the numerously represented families Poland), C. cruciata and C. parvula (uplands and are clausiliids with their 24 species (13.8%) assigned to mountains) and Balea perversa (a few sites in the nine genera in three subfamilies (Table 1) (Kerney et Sudetes and one in the Eastern Beskidy Mts). Alpine al. 1983, Riedel 1988, Wiktor 2004). clausiliids: Cochlodina costata, Charpentieria Zoogeographically, European clausiliids constitute ornata and Macrogastra badia, have only single the biggest group (54.2%), followed by Carpathian spe- sites in the Sudetes, whereas Carpathian species occur cies (29.2%), while the Alpine component is represent- most of all in the south-eastern part of the country, ed by three (12.5%), and the boreo-montane component though the range of some of them is not confined to the PL ISSN 0003-4541 © Fundacja Natura optima dux doi: 10.3161/000345408X396783 858 T. K. MALTZ and A. SULIKOWSKA-DROZD Table 1. Systematic index of clausiliids of Poland (Riedel 1988, Wiktor 2004). Family Subfamily Genus Subgenus Species Clausiliidae Alopiinae Cochlodina Cochlodina C. costata (C. Pfeiffer, 1828) C. laminata (Montagu, 1803) Paracochlodina C. orthostoma (Menke, 1828) Charpentieria Itala Ch. ornata (Rossmässler, 1836) Clausiliinae Ruthenica R. filograna (Rossmässler, 1836) Macrogastra M. badia (C. Pfeiffer, 1828) M. latestriata (A. Schmidt, 1857) M. plicatula (Draparnaud, 1801) M. tumida (Rossmässler, 1836) M. ventricosa (Draparnaud, 1801) Clausilia C. bidentata (Ström, 1765) C. cruciata (Studer, 1820) C. dubia Draparnaud, 1805 C. parvula Férussac, 1807 C. pumila C. Pfeiffer, 1828 Baleinae Laciniaria L. plicata (Draparnaud, 1801) Alinda A. biplicata (Montagu, 1803) Balea Balea B. perversa (Linnaeus, 1758) Pseudalinda B. fallax (Rossmässler, 1836) B. stabilis (L. Pfeiffer, 1847) Vestia Vestia V. elata (Rossmässler, 1836) V. gulo (E. A. Bielz, 1859) V. turgida (Rossmässler, 1836) Bulgarica Strigilecula B. cana (Held, 1836) Carpathians: Macrogastra tumida has a few locali- habitats, 13 clausiliids (54.2%) are regarded as threat- ties in the Eastern Sudetes, and M. latestriata in- ened in Poland: seven are low risk species while six are habits also the lowlands of north-eastern Poland. The endangered, of these as many as five critically (Wiktor region holds also two European species typical of up- and Riedel 2002) (Table 3). land and mountainous areas: Cochlodina orthosto- Literature information on clausiliid life cycles is ma and Clausilia dubia (Riedel 1988, Wiktor 2004) usually fragmentary and pertains only to 11 among the (Fig. 1). 24 species (Table 4). Complete data based on field Most clausiliids (91.7%) live in natural forest habi- observations exist only for Vestia elata (Piechocki tats, some (among others Laciniaria plicata, Alinda 1982), the life cycle of Balea perversa has been part- biplicata) are found also in forest-like habitats (den- ly studied (Baur 1990, Baur and Baur 1992). In the re- se, shaded shrubs, neglected parks etc.). Only Char- maining cases the data pertain either to a selected pentieria ornata and Balea perversa prefer more aspect of clausiliid biology, e.g. copulation (Nordsieck open habitats. With respect to humidity requirements 2005), or are fragmentary and/or problematic (Fröm- only Vestia gulo is a typical higrophile while the re- ming 1954 – literature review, Likharev 1962, Hudec maining ones live in mesic habitats with moderate 1963, Tompa 1979a – literature review, Bulman 1996, humidity fluctuations during the vegetation season Kuźnik-Kowalska 1998), and thus require verification. (mesophile species). Cochlodina costata, Charpen- The objective of this paper is to present and discuss tieria ornata and Clausilia parvula require a high laboratory-based data on 15 species of clausiliids, and calcium content in the substratum. The last two spe- pertaining, among other things, to the mode and period cies and Balea perversa are petrophiles (Maltz and of reproduction (oviparity: time of egg-laying, number Pokryszko 2007) (Fig. 2). of eggs per batch, hatching, cannibalism; egg reten- Because of the largely man-induced environment- tion, and ovoviviparity; reproductive season in the lab- al changes, and the resulting disappearance of natural oratory), growth and maturation, which are crucial to LIFE CYCLES OF CLAUSILIIDS OF POLAND 859 Table 2. Zoogeographic composition of clausiliids of Poland (Riedel 1988). General division Detailed division Species Boreo-Alpine Boreo-Alpine Clausilia cruciata Alpine s. lato E Alpine-Dinaric Cochlodina costata E Alpine Charpentieria ornata Macrogastra badia Carpathian s. lato Carpathian s. stricto Macrogastra latestriata Macrogastra tumida Balea stabilis Vestia gulo Vestia turgida E Carpathian Vestia elata Carpathian-Balkan Balea fallax European s. lato European s. stricto Cochlodina laminata E European Ruthenica filograna C European Macrogastra plicatula Macrogastra ventricosa Clausilia dubia Clausilia parvula Alinda biplicata E and C European Clausilia pumila Laciniaria plicata C and E European Cochlodina orthostoma W European Balea perversa N and NW European Clausilia bidentata E and SE European Bulgarica cana protection of endangered species and at the same time Tilia sp, Fagus sp.), fragments of limestone and tufts make it possible to rectify problematic information pre- of mosses (Ceratodon sp., Atrichum sp., Funaria sp., sented in earlier publications. Hypnum sp., Dicranum sp., Trichostomum sp.). Calcium was provided in the form of writing chalk or dolomite tablets. The snails were fed leaves of lettuce, MATERIAL AND METHODS Pekingese lettuce, sometimes Italian or headed cab- bage, pieces of carrot, cucumber and mushroom The studies were carried out in two institutions: (champignon, chanterelle). Museum of Natural History, Wrocław University [1] Humidity in the containers was practically constant and the Department of Invertebrate Zoology and Hy- and close to 100% as a result of regular spraying of drobiology, University of Łódź [2]. Fifteen species the tissue, stones, bark and moss. The culture was observed in the laboratory were subject to the studies checked every 2–3 days (when necessary daily), while (Table 5). total cleaning (exchange of tissue paper, container Adults were kept in plastic containers of 300, 500, cleaning) took place twice a month. In Wrocław the 700 and 1200 cm3 volume, single snails or pairs were culture containers were kept at room tempera- kept in smaller containers, while other containers held ture (18–22°C) and natural light regime throughout groups of 5–10 individuals. Apart from the species the year; in Łódź the snails hibernated in a cool which were introduced into the culture later (C. ortho- room: preparing for hibernation (half of October till stoma, C. pumila, C. dubia), detailed observations half of November – 8–12°C), hibernation (half of No- involved five pairs of each C. laminata and L. plicata vember till the end of February – 3°C) and prepar- and 10–15 pairs of each of the remaining species. ing for activity (March – 8–12°C). During that time Each tissue-lined box contained pieces of deciduous the photoperiod