Biologia 67/2: 384—389, 2012 Section Zoology DOI: 10.2478/s11756-012-0020-2

Thespreadofnon-nativeCepaea nemoralis and cartusiana (: ) in the with comments on other immigrants

Alena Peltanová1,LiborDvořák2 &LucieJuřičková3

1Agency for Nature Conservation and Landscape Protection of the Czech Republic, Nuselská 39,CZ–14000 Praha 4-Nusle, Czech Republic; e-mail: [email protected] 2Municipal Museum Mariánské Lázně, Goethovo náměstí 11,CZ–35301 Mariánské Lázně, Czech Republic; e-mail: [email protected] 3Charles University, Department of Zoology, Viničná 7,CZ-12844 Praha 2, Czech Republic; e-mail: [email protected]

Abstract: The aim of our study is to describe and visualise the spread of two non-indigenous land snail Cepaea nemoralis and in the Czech Republic during more than 100 years period. Several factors play an important role in changes of the distribution of these species: ecological (climate change), ethological (passive dispersal potencial) and economic (increasing traffic as a vector of spreading). The spreading of M. cartusiana has a rapidly increasing trend. More than half sites in the Czech Republic were colonised by this species in 2000–2010. While the spread of C. nemoralis has been continuous during the last century, the rapid range extension was recorded in the last two decades. Key words: Cepaea nemoralis; Monacha cartusiana; passive dispersal; range extension; grid map; distribution trends

Introduction The main goals of our study are to visualise and describe the spread of two non-indigenous species: the The European biota has experienced a substantial shift Mediterranean Monacha cartusiana (O.F. M¨uller, 1774) during The Quaternary. In pre-historic times, the North (Gastopoda: Pulmonata: ) and the At- European Fauna and Flora ranges slide across the lantic Cepaea nemoralis (L., 1758) (Gastropoda: Pul- whole continent and Pleistocene glacial migrations were monata: ), both occurring within Czech mala- followed by the post-glacial re-colonization in Early cofauna from the end of the 19th century. Holocene (Taberlet et al. 1998). Historic times of wars and trade have brought moving masses of people and Material and methods goods: Romans military campaign and medieval trade route significantly contributed to the present form of The total number of occupied quadrates (11.2 × 12 km) of biota in some European countries (Evans 1972). Nowa- KFME (Kartierung der Flora Mitteleuropas), the Central days, a response to the recent trend of climate warm- grid for and plant mapping (Ehrendorfer & ing is obvious – species range significantly shifts to- Haman 1965), was counted for two land snail species Cepaea nemoralis and Monacha cartusiana on the model area of the wards the poles or altitudinal upward (Parmensan & Czech Republic over the 110-years-period (1900–2010). The Yohe 2003). Recent changes in Central European inver- data were collected from different credible sources: publi- tebrate fauna distributions are one of the most appro- cations (articles, monographs), museum and private mala- priate examples: thermophilous terrestrial species with cological collections (unpublished findings), field excursions Atlantic or Mediterranean distribution are intensively and web presented map reports (Peltanová & Novák 2011 a, spreading northwards (Roques et al. 2009; Peltanová et b). Recent M. cartusiana distribution was also confronted al. 2011). and compared with historical data from older (e.g., Ložek In the Czech Republic, recent occurrence of 247 1948) and latest (Kolouch 2005) known sites. Part of C. mollusc species is well-documented (Horsák et al. 2010). nemoralis distributions in the western part of the Czech Re- public were systematically mapped (Honěk 1995 a, b; Honěk 169 land snails (except greenhouse gastropod) can be & Martínková 2003; Dvořák & Honěk 2004). divided into two main groups: a) 154 native or long- Interpretation of distribution trends is based on the term naturalized species (Ložek 1964) and b) 15 non- number of occupied quadrates. N (%) is the ratio of re- indigenous recent incomers (Juřičková 2006; Horsák et cently occupied quadrates (Nr) to cumulative number of all al. 2010). quadrates ever occupied by the species in 1900–2010 (Nt).

c 2012 Institute of Zoology, Slovak Academy of Sciences Non-native land snails spreading in the Czech Republic 385

Fig. 1. Monacha cartusiana and Cepaea nemoralis distribution changes in the Czech Republic within three main periods (1900–1950, 1951–1989, 1990–2010). While the cartusian snail is changing its distribution in time (older sites without new findings), brown lipped snail has gradual expansion (the older sites still inhabited today).

Recent distribution Nr is represented by the positive occur- temperature trend in Central Europe (Dobrovolný et al. rence of species in 1990–2010. If N < 33.3%, the species is 2010). consider expanding (van Swaay & Waren 1999; Konvička & Beneš 2002). Results and discussion N =(1− Nr/Nt) × 100 Monacha cartusiana occupied 64 quadrates (9% of the ArcView 3.0 was used for grid map outputs, where CZ area) in the period “1900–2010”, while 80% of changes in both species distribution are visualised. Dis- its distribution represents new occurrences in 2004– tribution grid maps (Fig. 1) present species occurrence 2010. The ratio of recently occupied sites (N )equals within three periods of time: 1900–1950, 1951–1989 and to 23.53%; therefore the species is considered expand- 1990–2010. The period “1900–1950” presents the result of ing. first malacological surveys prior to the end of the Sec- Cepaea nemoralis occupied 144 quadrates (more ond World War. The second period “1951–1989” is char- than 20% of the CZ area) in the period “1900–2010”, acterised by decreasing temperature trend in the Cen- tral Europe and intensive land use changes (Dale 1997; while near 30% of its distribution represents new oc- Bičík et al. 2001; Rabitsch 2008) the last period “1990– currences in 2004–2010. The ratio of recently occupied 2010” dates from the opening of borders in Eastern sites (N ) equals to 36.81%; therefore the species is close Europe in 1989 and marks the beginning of increasing to be considered expanding. 386 A. Peltanová et al.

How do land snails spread? tilla, , Oxychilus alliarius, O. dra- Land snails have a weak active dispersal potency (Go- parnaudi and Zonitoides arboreus) and the last species dan 1983), therefore the natural passive dispersal way Limacus flavus is strictly eusynathropic with a ran- (anemochory, hydrochory or zoochory) is one of the dom occurrence (citations summarized in Peltanová et most effective processes for their spread (Figuerola al. 2011). & Green 2002), even for a long distance (Gitten- Changes in distribution of A. vulgaris in the Czech berger et al. 2006; Shinichiro et al. 2012). The most Republic were discussed in detail (Dvořák & Horsák of recently expanding land snails are xerotolerant 2003; Juřičková 2006). Our attention is focused now on and/or thermophilous species, with pre-adaptation to the ongoing expansion of C. nemoralis and M. cartu- survive during unfavourable annual period (aestiva- siana. tion/hibernation) (Godan 1983). These species have po- tential abilities to stay alive during the transportations. Monacha cartusiana in the Czech Republic They solve only one crucial problem: to board itself or The Cartusian snail M. cartusiana is a Mediterranean to be boarded. The first technique plays an important species with original distribution in Southern Europe. role in the spread of land snails with the climbing be- The shell (extremely variable shell diameter 6–18 mm) haviour (e.g., Helicellinae and some others hygromiids, is horny yellowish with reddish margin; the aperture Enidae, some helicids). Probably due to their necessity with a prominent white lip inside occurs in adults. to reduce the body temperature during hot days in in- Nowadays, the stable populations were found in solated habitats (Sacchi 1971; Jaremovic & Rollo 1979), suitable habitats (open areas in lowlands) all over Eu- they climb to horizontal objects, including the poten- rope: Southern (European and African part of the tial vectors (Aubry et al. 2006). This was also verified Mediterranean coast), Western (the Atlantic coast), in arboreal land snails (Cockerell 1921; Gittenberger et Northern (Denmark), Central (, Czech Repub- al. 2006) and we could assume the same mechanism of lic, , , Poland and Slovak Republic) transportation among the other vertically moving Eu- and Eastern, including probably only eastern parts of ropean species (e.g., Succineidae or selected taxons of the Black Sea region and Western Caucasus, where Clausillidae). species was probably introduced by man (Hausdorf Results of passive dispersal research in different 2000a, b; Georgiev 2006). parts of Europe show the importance of traffic (rails, Monacha cartusiana successfully spreads across the cars and other vehicles mobility) for the spread of land area of Central Europe (e.g., Fischer & Duda 2004; Wic- snails (Aubry et al. 2006). Occurrences of new incomers tor 2004; Benke & Renker 2005; Míkovcová & Juřičková are increasing, especially in the areas with particularly 2008), where it inhabits different types of open habitats high concentration of cargo (freight terminals, whole- (semi-natural steppe and grassland, synathropic or rud- sale market or shopping centres) (Benke & Renker 2005; eral localities, river banks and marches) (Kerney et al. Juřičková & Kapounek 2009; Hlaváč & Peltanová 2010; 1983). The species was also introduced to SE England Říhová & Juřičková 2011a). in prehistoric times and over the Atlantic Ocean in the Not only individual observations, but also system- 20th century (Michalak & Price 2010). atical research show that both expanding species use There are no fossil records of M. cartusiana occur- climbing behavior as successful way of distribution: rence in the Czech Republic. The nearest Holocene lo- dozens of C. nemoralis individuals sitting on a passing calities in Central Europe are Nitrianský hrádok in Slo- truck in , Czech Republic (Juřičková, personal vakia (Ložek 1964) and Vienna basin in Austria (Frank observ.); cargo boxes in Basel docks, (M. 2006). The first known locality of the Cartusian snail in cartusiana – Peltanová, personal observ.); as well as the Czech Republic is Hustopeče surroundings (Uličný few individuals on the disused wagons on the railway 1885) in . The first Bohemian site Choceň was stationinLeipzig,Germany(M. cartusiana –Benke& observed in 1937 (collection of The National Museum Renker 2005) are the evidence of linking human activi- in Prague, CZ). In 1943 empty shell was found in ties and the spread of non-indigenous species. Čermníky near Žatec (Ložek 1948), but presence of this population has never been confirmed there. In the 1950s Non-indigenous species in the Czech Republic and 1960s, numerous sites were recorded in South-West During historic time, alien species were gradually, yet in Moravia (summarized by Lisický 1991). During the last a lower number, spreading to the area of the Czech Re- twenty years, the number of new evidences has signifi- public; but since the 1990s the number of new recorded cantly increased (Míkovcová & Juřičková 2008). non-indigenous land snails rapidly increased (Juřičková However, some populations on South and South- 2006; Peltanová et al. 2011). We have recorded 15 non- West Moravia extinct, we have new evidences from native outdoor living species (near 7% of the Czech Northern Moravia (Maňas 2002; Novák & Novák 2009; terrestrial malacofauna), where 3 are rapidly expand- Horsák personal comm.). In , rapid increasing ing: C. nemoralis, M. cartusiana and Arion vulgaris of records is concentrated especially to the warmer ar- (syn. lusitanicus), 11 species have stable populations eas: the Labe River valley, Bohemian Karst, foothills without expanding events (Boettgerilla pallens, Cer- of the Ore Mts (Krušné hory) and sporadically in ur- nuella neglecta, Cornu aspersum, Deroceras panormi- ban areas (Hudec 1954; Urbánek 1968; Kolouch 1996; tanum, Helix lucorum, cinctella, Lucilla scin- Juřičková 1998; Ložek 1999; Maňas 2002; Kolouch 2005; Non-native land snails spreading in the Czech Republic 387

Hlava & Vrabec 2008). The Cartusian snail is recently (N and C Moravia populations), Austria (population in distributed mainly in the warmest areas of the Czech S Bohemia and S Moravia) and probably also in Republic, so called thermophyticum, along great rivers (W Bohemia), but no genetic study, which can clear this (Tolasz 2007). A few of new sites (2000 – Plzeň, M. spreading, exist. Mergl, personal comm.; 2009 – Nové Hodějovice, Pech In 2004–2010, new records are situated primarily & Pechová 2009; 2010 – Tábor surroundings, NGO in areas where this species has long been known: e.g., Daphne observ.) are situated in the middle-temperature great cities surroundings (České Budějovice, Kladno, areas of the Czech Republic, which could be a conse- Prague, Ostrava), E and W Bohemia. In some regions quence of the increasing trend in the average annual (České Budějovice, W Bohemia), the higher number of temperature (Dobrovolný et al. 2010) and evidence of new sites could be consequence of increased research ac- distribution area natural shifting. tivity (L. Dvořák, personal observ.). After rapid spatial Cartusian snail is an unmistakable species in the spreading documented since 1990, current distribution Czech malacofauna – as the rarity of our malacofauna area has been thickening without long distance expan- was already observed by malacologists and interested sion to the unoccupied regions. other biologists, each of new site has been carefully pub- lished. Due to this history, we expect growing number of new sites significantly connected with increasing num- Acknowledgements ber of new landing; the impact of higher sampling effort This paper falls under research projects MSM 6293359101 is probably negligible. and 0021620828. Many thanks to all our colleagues for their field data (in alphabetical order): Luboš Beran, Cepaea nemoralis in the Czech Republic Petr Bogush, Alena Borská, Petr Červenka, Adam Furek, The brown lipped snail C. nemoralis is an Atlantic Michal Holec, Alois Honěk, Michal Horsák, Jitka Horáčková, species with the most recent range in Western and Cen- Petr Kment, Miroslav Kesl, Vojen Ložek, Michal Maňas, tral Europe. The yellow, pink or brown shell (diame- Miroslav Mergl, Jiří Novák, Pavel Pech, Hana Pechová, Mi- ter 18–25 mm) typically is with a dark brown apertu- lan Řezáč, Pavel Tlachač, Bohdan Zvarič, Daphne ČR – ral lip and a light periostracum, with or without dark Institute of Applied Ecology and Biolib server stuff. Many thanks also to Tomáš Peltan and Daniel Franke for help brown colour bands. 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