Biologia 67/2: 384—389, 2012 Section Zoology DOI: 10.2478/s11756-012-0020-2
The spread of non-native Cepaea nemoralis and Monacha cartusiana (Gastropoda: Pulmonata) in the Czech Republic with comments on other land snail immigrants Alena Peltanová1, Libor Dvořák2 & Lucie Juřičková3 1 Agency for Nature Conservation and Landscape Protection of the Czech Republic, Nuselská 39, CZ–14000 Praha 4-Nusle, Czech Republic; e-mail:
[email protected] 2 Municipal Museum Mariánské Lázně, Goethovo náměstí 11, CZ–35301 Mariánské Lázně, Czech Republic; e-mail:
[email protected] 3 Charles 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 species Cepaea nemoralis and Monacha cartusiana 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 European biota has experienced a substantial shift during The Quaternary. In pre-historic times, the North European Fauna and Flora ranges slide across the whole continent and Pleistocene glacial migrations were followed by the post-glacial re-colonization in Early Holocene (Taberlet et al. 1998). Historic times of wars and trade have brought moving masses of people and goods: Romans military campaign and medieval trade route significantly contributed to the present form of biota in some European countries (Evans 1972). Nowadays, a response to the recent trend of climate warming is obvious – species range significantly shifts towards the poles or altitudinal upward (Parmensan & Yohe 2003). Recent changes in Central European invertebrate fauna distributions are one of the most appropriate examples: thermophilous terrestrial species with Atlantic or Mediterranean distribution are intensively spreading northwards (Roques et al. 2009; Peltanová et al. 2011). In the Czech Republic, recent occurrence of 247 mollusc species is well-documented (Horsák et al. 2010). 169 land snails (except greenhouse gastropod) can be divided into two main groups: a) 154 native or longterm naturalized species (Ložek 1964) and b) 15 nonindigenous recent incomers (Juřičková 2006; Horsák et al. 2010).
c 2012 Institute of Zoology, Slovak Academy of Sciences
The main goals of our study are to visualise and describe the spread of two non-indigenous species: the Mediterranean Monacha cartusiana (O.F. M¨ uller, 1774) (Gastopoda: Pulmonata: Hygromiidae) and the Atlantic Cepaea nemoralis (L., 1758) (Gastropoda: Pulmonata: Helicidae), both occurring within Czech malacofauna from the end of the 19th century. Material and methods The total number of occupied quadrates (11.2 × 12 km) of KFME (Kartierung der Flora Mitteleuropas), the Central Europe grid for animal and plant mapping (Ehrendorfer & Haman 1965), was counted for two land snail species Cepaea nemoralis and Monacha cartusiana on the model area of the Czech Republic over the 110-years-period (1900–2010). The data were collected from different credible sources: publications (articles, monographs), museum and private malacological collections (unpublished findings), field excursions and web presented map reports (Peltanová & Novák 2011 a, b). Recent M. cartusiana distribution was also confronted and compared with historical data from older (e.g., Ložek 1948) and latest (Kolouch 2005) known sites. Part of C. nemoralis distributions in the western part of the Czech Republic were systematically mapped (Honěk 1995 a, b; Honěk & Martínková 2003; Dvořák & Honěk 2004). Interpretation of distribution trends is based on the number of occupied quadrates. N (%) is the ratio of recently occupied quadrates (Nr) to cumulative number of all quadrates ever occupied by the species in 1900–2010 (Nt).
Non-native land snails spreading in the Czech Republic
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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 occurrence of species in 1990–2010. If N < 33.3%, the species is consider expanding (van Swaay & Waren 1999; Konvička & Beneš 2002). N = (1 − N r/N t) × 100 ArcView 3.0 was used for grid map outputs, where changes in both species distribution are visualised. Distribution grid maps (Fig. 1) present species occurrence within three periods of time: 1900–1950, 1951–1989 and 1990–2010. The period “1900–1950” presents the result of first malacological surveys prior to the end of the Second World War. The second period “1951–1989” is characterised by decreasing temperature trend in the Central Europe and intensive land use changes (Dale 1997; Bičík et al. 2001; Rabitsch 2008) the last period “1990– 2010” dates from the opening of borders in Eastern Europe in 1989 and marks the beginning of increasing
temperature trend in Central Europe (Dobrovolný et al. 2010).
Results and discussion Monacha cartusiana occupied 64 quadrates (9% of the CZ area) in the period “1900–2010”, while 80% of its distribution represents new occurrences in 2004– 2010. The ratio of recently occupied sites (N ) equals to 23.53%; therefore the species is considered expanding. Cepaea nemoralis occupied 144 quadrates (more than 20% of the CZ area) in the period “1900–2010”, while near 30% of its distribution represents new occurrences in 2004–2010. The ratio of recently occupied sites (N ) equals to 36.81%; therefore the species is close to be considered expanding.
386 How do land snails spread? Land snails have a weak active dispersal potency (Godan 1983), therefore the natural passive dispersal way (anemochory, hydrochory or zoochory) is one of the most effective processes for their spread (Figuerola & Green 2002), even for a long distance (Gittenberger et al. 2006; Shinichiro et al. 2012). The most of recently expanding land snails are xerotolerant and/or thermophilous species, with pre-adaptation to survive during unfavourable annual period (aestivation/hibernation) (Godan 1983). These species have potential abilities to stay alive during the transportations. They solve only one crucial problem: to board itself or to be boarded. The first technique plays an important role in the spread of land snails with the climbing behaviour (e.g., Helicellinae and some others hygromiids, Enidae, some helicids). Probably due to their necessity to reduce the body temperature during hot days in insolated habitats (Sacchi 1971; Jaremovic & Rollo 1979), they climb to horizontal objects, including the potential vectors (Aubry et al. 2006). This was also verified in arboreal land snails (Cockerell 1921; Gittenberger et al. 2006) and we could assume the same mechanism of transportation among the other vertically moving European species (e.g., Succineidae or selected taxons of Clausillidae). Results of passive dispersal research in different parts of Europe show the importance of traffic (rails, cars and other vehicles mobility) for the spread of land snails (Aubry et al. 2006). Occurrences of new incomers are increasing, especially in the areas with particularly high concentration of cargo (freight terminals, wholesale market or shopping centres) (Benke & Renker 2005; Juřičková & Kapounek 2009; Hlaváč & Peltanová 2010; Říhová & Juřičková 2011a). Not only individual observations, but also systematical research show that both expanding species use climbing behavior as successful way of distribution: dozens of C. nemoralis individuals sitting on a passing truck in Prague, Czech Republic (Juřičková, personal observ.); cargo boxes in Basel docks, Switzerland (M. cartusiana – Peltanová, personal observ.); as well as few individuals on the disused wagons on the railway station in Leipzig, Germany (M. cartusiana – Benke & Renker 2005) are the evidence of linking human activities and the spread of non-indigenous species. Non-indigenous species in the Czech Republic During historic time, alien species were gradually, yet in a lower number, spreading to the area of the Czech Republic; but since the 1990s the number of new recorded non-indigenous land snails rapidly increased (Juřičková 2006; Peltanová et al. 2011). We have recorded 15 nonnative outdoor living species (near 7% of the Czech terrestrial malacofauna), where 3 are rapidly expanding: C. nemoralis, M. cartusiana and Arion vulgaris (syn. lusitanicus), 11 species have stable populations without expanding events (Boettgerilla pallens, Cernuella neglecta, Cornu aspersum, Deroceras panormitanum, Helix lucorum, Hygromia cinctella, Lucilla scin-
A. Peltanová et al. tilla, Monacha cantiana, Oxychilus alliarius, O. draparnaudi and Zonitoides arboreus) and the last species Limacus flavus is strictly eusynathropic slug with a random occurrence (citations summarized in Peltanová et al. 2011). Changes in distribution of A. vulgaris in the Czech Republic were discussed in detail (Dvořák & Horsák 2003; Juřičková 2006). Our attention is focused now on the ongoing expansion of C. nemoralis and M. cartusiana. Monacha cartusiana in the Czech Republic The Cartusian snail M. cartusiana is a Mediterranean species with original distribution in Southern Europe. The shell (extremely variable shell diameter 6–18 mm) is horny yellowish with reddish margin; the aperture with a prominent white lip inside occurs in adults. Nowadays, the stable populations were found in suitable habitats (open areas in lowlands) all over Europe: Southern (European and African part of the Mediterranean coast), Western (the Atlantic coast), Northern (Denmark), Central (Austria, Czech Republic, Hungary, Germany, Poland and Slovak Republic) and Eastern, including probably only eastern parts of the Black Sea region and Western Caucasus, where species was probably introduced by man (Hausdorf 2000a, b; Georgiev 2006). Monacha cartusiana successfully spreads across the area of Central Europe (e.g., Fischer & Duda 2004; Wictor 2004; Benke & Renker 2005; Míkovcová & Juřičková 2008), where it inhabits different types of open habitats (semi-natural steppe and grassland, synathropic or ruderal localities, river banks and marches) (Kerney et al. 1983). The species was also introduced to SE England in prehistoric times and over the Atlantic Ocean in the 20th century (Michalak & Price 2010). There are no fossil records of M. cartusiana occurrence in the Czech Republic. The nearest Holocene localities in Central Europe are Nitrianský hrádok in Slovakia (Ložek 1964) and Vienna basin in Austria (Frank 2006). The first known locality of the Cartusian snail in the Czech Republic is Hustopeče surroundings (Uličný 1885) in Moravia. The first Bohemian site Choceň was observed in 1937 (collection of The National Museum in Prague, CZ). In 1943 empty shell was found in Čermníky near Žatec (Ložek 1948), but presence of this population has never been confirmed there. In the 1950s and 1960s, numerous sites were recorded in South-West Moravia (summarized by Lisický 1991). During the last twenty years, the number of new evidences has significantly increased (Míkovcová & Juřičková 2008). However, some populations on South and SouthWest Moravia extinct, we have new evidences from Northern Moravia (Maňas 2002; Novák & Novák 2009; Horsák personal comm.). In Bohemia, rapid increasing of records is concentrated especially to the warmer areas: the Labe River valley, Bohemian Karst, foothills of the Ore Mts (Krušné hory) and sporadically in urban areas (Hudec 1954; Urbánek 1968; Kolouch 1996; Juřičková 1998; Ložek 1999; Maňas 2002; Kolouch 2005;
Non-native land snails spreading in the Czech Republic Hlava & Vrabec 2008). The Cartusian snail is recently distributed mainly in the warmest areas of the Czech Republic, so called thermophyticum, along great rivers (Tolasz 2007). A few of new sites (2000 – Plzeň, M. Mergl, personal comm.; 2009 – Nové Hodějovice, Pech & Pechová 2009; 2010 – Tábor surroundings, NGO Daphne observ.) are situated in the middle-temperature areas of the Czech Republic, which could be a consequence of the increasing trend in the average annual temperature (Dobrovolný et al. 2010) and evidence of distribution area natural shifting. Cartusian snail is an unmistakable species in the Czech malacofauna – as the rarity of our malacofauna was already observed by malacologists and interested other biologists, each of new site has been carefully published. Due to this history, we expect growing number of new sites significantly connected with increasing number of new landing; the impact of higher sampling effort is probably negligible. Cepaea nemoralis in the Czech Republic The brown lipped snail C. nemoralis is an Atlantic species with the most recent range in Western and Central Europe. The yellow, pink or brown shell (diameter 18–25 mm) typically is with a dark brown apertural lip and a light periostracum, with or without dark brown colour bands. In its original area of Western Europe, many populations live at grassy or forest habitats, while in the eastern and southern parts of its current area the species is nearly restricted to human impacted sites, only few localities are situated in floodplain forest (Dvořák & Honěk 2004; Kerney 1999). Its recent occurrence covers Europe from Ireland and Scotland (except for islands) to S Portugal and C Spain to SW Hungary [introduction to NE Hungary in 2003 (Domokos 2003)] and Bosnia, in Italy to Lucania, in the North to S Sweden, in the East to NW Poland, Latvia, Kaliningrad, Estonia (Hiiumaa island) (Welter-Schultes 2011) and also introduced overseas (Godan 1983). Although C. nemoralis is assumed to have appeared during the Central Europe Holocene interglacial (Ložek 1964) and West European population may have existed thousands years ago (Currey & Cain 1968), the first reference to its modern occurrence in the Czech Republic comes from the end of the 19th century (Uličný 1892–1895). Several authors since then have published records of C. nemoralis (e.g., Hlaváč 1940; Petrbok 1940, 1954) afterwards all the known data were reviewed by Flasar (1989). Only last thirty years of expansion is well-documented (Honěk 1995a, b; Kovanda, in Juřičková 1995; Kolouch 1997; Dvořák 1998, 1999; Flasar 1998; Juřičková 1998; Hlaváč et al. 2002; Dvořák et al. 2003; Honěk & Martínková 2003; Dvořák & Honěk 2004; Drvotová et al. 2008; Peltanová & Novák 2011b). Details of C. nemoralis spreading were summarized by Dvořák & Honěk (2004): The brown lipped snail starts to penetrate from Germany to N Bohemia, where it spreads further into C Bohemia and W Moravia. Other isolated populations in the Czech Republic could be connected (in both directions) with those in Poland
387 (N and C Moravia populations), Austria (population in S Bohemia and S Moravia) and probably also in Bavaria (W Bohemia), but no genetic study, which can clear this spreading, exist. In 2004–2010, new records are situated primarily in areas where this species has long been known: e.g., great cities surroundings (České Budějovice, Kladno, Prague, Ostrava), E and W Bohemia. In some regions (České Budějovice, W Bohemia), the higher number of new sites could be consequence of increased research activity (L. Dvořák, personal observ.). After rapid spatial spreading documented since 1990, current distribution area has been thickening without long distance expansion to the unoccupied regions. Acknowledgements This paper falls under research projects MSM 6293359101 and 0021620828. Many thanks to all our colleagues for their field data (in alphabetical order): Luboš Beran, Petr Bogush, Alena Borská, Petr Červenka, Adam Furek, Michal Holec, Alois Honěk, Michal Horsák, Jitka Horáčková, Petr Kment, Miroslav Kesl, Vojen Ložek, Michal Maňas, Miroslav Mergl, Jiří Novák, Pavel Pech, Hana Pechová, Milan Řezáč, Pavel Tlachač, Bohdan Zvarič, Daphne ČR – Institute of Applied Ecology and Biolib server stuff. Many thanks also to Tomáš Peltan and Daniel Franke for help with visualizations of species distribution data. References Aubry S., Labaune C., Magnin F., Roche P. & Kiss L. 2006. Active and passive dispersal of an invading land snail in Mediterranean France. J. Anim. Ecol. 75 (3): 802–813. DOI: 10.1111/j.1365-2656.2006.01100.x Benke M. & Renker C. 2005. Vorkommen von Monacha cartusiana (O.F. M¨ uller, 1774) und Cernuella neglecta (Draparnaud, 1805) im Stadtgebiet von Leipzig (Sachsen). Malakol. Abh. 23: 109–115. Bičík I., Jeleček L. & Štěpánek V. 2001. Land–use changes and their social driving forces in Czechia in the 19th and 20th centuries. Land Use Policy 18 (1): 65–73. DOI: http://dx.doi.org/10.1016/S0264-8377(00)00047-8 Cockerell T.D.A. 1921. The dispersal of snail by bird. Nature 108: 496–497. DOI: 10.1038/108496d0 Currey J.D. & Cain A.J. 1968. Studies on Cepaea IV. Climate and selection of banding morphs in Cepaea from the climatic optimum to the present day. Philosoph. Trans. Roy. Soc. London B 253 (789): 483–489. Dale V.H. 1997. The relationshuip between land–use change and climate change. Ecol. Appl. 7 (3): 753–769. DOI: http://dx.doi.org/10.1890/1051-0761(1997)007[0753:TRB LUC]2.0.CO;2 Dobrovolný P., Moberg A., Brázdil R., Pfister C., Glaser R., Wilson R., Van Engelen A., Limanówka D., Kiss A., Halíčková M., Macková J., Riemann D., Luterbacher J. & B¨ ohm R. 2010. Monthly, seasonal and annual temperature reconstructions for Central Europe derived from documentary evidence and instrumental records since AD 1500. Climatic Change 101 (1–2): 69–107 DOI: 10.1007/s10584-009-9724-x Domokos T. 2003. Behurcolt Cepaea nemoralis (Linnaeus, 1758) a B¨ ukkben [Imported Cepaea nemoralis (Linnaeus, 1758) in the B¨ ukk Mountains (N-Hungary)]. Malacological Newsletter 21: 65–67. Drvotová M., Hlaváč J.Č., Horsák M., Beran L., Dvořák L., Juřičková L. & M¨ uckstein P. 2008. Měkkýši (Mollusca) Žďárských vrchů [Molluscs (Mollusca) of the Žďárské vrchy
388 Mts]. Vyd. 1. Hlinsko, AOPK ČR, Správa CHKO Žďárské vrchy, Parnassia 3, 79 pp. ISBN: 978-80-87051-49-8 Dvořák L. 1998. Páskovka hajní (Cepaea nemoralis L.) v jižních Čechách [Cepaea nemoralis in Southern Bohemia]. Sbor. Jihočes. Muz. v Čes. Budějovicích, Přír. Vědy 38: 18. Dvořák L. 1999. Malakofauna sklepů, štol a jeskyní západních Čech a oblasti Šumavy [Molluscs of cellars, galleries and caves of West Bohemia and of the Bohemian Forest Region]. Silva Gabreta 3: 141–154. Dvořák L., Ditrich O. & Juřičková L. 2003. New records of three ruderal snail species (Mollusca. Gastropoda) in South Bohemia. Sbor. Jihočes. Muz. v Čes. Budějovicích, Přír. Vědy 43: 111–113. Dvořák L. & Honěk A. 2004. The spread of the Brown Lipped Snail, Cepaea nemoralis (Linnaeus, 1758), in the Czech Republic. Čas. Nár. Muz., Řada Přírod. Zoology173 (1–4): 97– 103. Dvořák L. & Horsák M. 2003. Současné poznatky o plzáku Arion lusitanicus (Mollusca: Pulmonata) v České republice. Čas. Slez. Muz. v Opavě 52: 67–71 Ehrendorfer F. & Hamann U. 1965. Vorschl¨ age zu einer Floristischen Kartierung von Mitteleuropa. Ber. Deutsch. Bot. Gesell. 78: 35–50. Evans J.G. 1972. Land Snails in Archeology. 1st Ed. Seminar Press, London and New York, 436 pp. ISBN: 0128295503, 978-0128295502 Figuerola J. & Green A. J. 2002. Dispersal of aquatic organism by waterbirds – a review of past research and priorities for future studies. Freshwater Biol. 47(3): 483–494. DOI: 10.1046/j.1365-2427.2002.00829.x Fischer W. & Duda M. 2004. Beitr¨ age zur Kenntnis der Mol¨ luskenfauna Osterreichs VII. Cernuella virgata (da Costa 1778), neu f¨ ur die Molluskenfauna Wiens, sowie Bemerkungen zur Ausbreitung von Monacha cantiana (Montagu 1803), Cernuella neglecta (Draparnaud 1805), Hygromia cinctella (Draparnaud 1801) und Cornu aspersus (O.F.M¨ uller 1774) in Nieder¨ osterreich und Wien (Mollusca. Gastropoda). Nachrichtenblatt der Ersten Vorarlberger Malakologischen Gesellschaft 12: 10–14. Flasar I. 1989. Die Verbreitung der Schwarzm¨ undigen B¨ anderschnecke Cepaea (Cepaea) nemoralis (L., 1758) in der Tschechoslowakei (Gastropoda, Stylommatophora. Helicidae). Malakol. Abh. Staatl. Mus. Tierkd. Dresden 14,13: 111–117. ISSN: 0070-7260 Flasar I. 1998. Die Gastropoden Nordwestb¨ ohmens und ihre Verbreitung. Heldia 3 (Sonderheft 4), M¨ unchen, 210 pp. ISSN: 0932-4143 ¨ Frank C. 2006. Plio–pleistoz¨ ane und holoz¨ ane Mollusken Osterreichs. Austrian Academy of Sciences, Vienna, 860 pp. ISBN: 3700136749, 978-3700136743 Fr¨ omming E. 1954. Biologie der mitteleurop¨ aischen Landgastropoden. Duncker und Humblot, Berlin, 404 pp. ISBN: 9783-428-00454-6 Georgiev D. 2006. A contribution to the knowledge of the malacofauna of Sveti Iliiski Heights (South-Eastern Bulgaria). Scientific Studies of the University of Plovdiv, Biology, Animalia 42: 13–20. http://web.uni-plovdiv.bg/dilian/002 Georgiev (2006).pdf Gittenberger E., Groenenberg S.J., Kokshoorn B. & Preece R.C. 2006. Molecular trails from hitch–hiking snails. Nature 439: 409. DOI: 10.1038/439409a Godan D. 1983. Pest Slugs and Snails, Biology and Control. Springer–Verlag, Berlin, Heidelberg, 445 pp. ISBN: 3540118942, 9783540118947 Hausdorf B. 2000a. The genus Monacha in Turkey (Gastropoda: Pulmonata: Hygromiidae). Arch. Molluskenk. 128 (1/2): 61– 151. Hausdorf B. 2000b. The genus Monacha in the Western Caucasus (Gastropoda: Hygromiidae). J. Nat. Hist. 34 (8): 1575–1594. DOI: 10.1080/00222930050117495 Hewitt G.M. 1999. Post–glacial re–colonization of European Biota. Biol. J. Linn. Soc. 68 (1–2): 87–112. DOI: 10.1111/j. 1095-8312.1999.tb01160.x Hlava J. & Vrabec V. 2008. Malacozoological research of the Bílina mine territory near Mariánské Radčice (North–West
A. Peltanová et al. Bohemia, Czech Republic), pp. 88–89. In: Kubík Š. & Barták M. (eds), Proceedings of the „Workshop of animal biodiversity”, 3.–4. July 2008, Jevany. ISBN: 978-80-213-1865-6 Hlaváč J.Č., Horsák M., Beran L., Dvořák L., Juřičková L. & Vrabec V. 2002. Měkkýši Českého lesa – I. Vybrané lokality v severní části (západní Čechy) [Molluscs of the Český Les Mts – I. Selected localities in the northern part (Western Bohemia)]. Silva Gabreta 8: 205–228. Hlaváč J.Č. & Peltanová A. 2010. First occurrence of the Kentish Snail Monacha cantiana (Mollusca. Gastropoda. Hygromiidae) in the Czech Republic. Malacologica Bohemoslovaca 9: 11–15. Hlaváč V.F. 1940. Nový výskyt druhu Tachea nemoralis L. v zemi České [New finding of Tachea nemoralis L. in Bohemia]. Čas. Nár. Muz., Řada Přír. 114: 228–229. Honěk A. 1995a. Geographic distribution and shell colour and banding polymorphism in marginal population in Cepaea nemoralis (Gastropoda. Helicidae). Malacologia 37 (1): 111– 122. Honěk A. 1995b. Distribution and shell colour and banding polymorphism of the Cepea species (Gastropoda. Helicelidae) in Bohemia. Act. Soc. Zool. Bohem. 59: 63–77. Honěk A. & Martínková Z. 2003. Persistence and shell band morph frequencies in urban populations of Cepaea. Symposium on Slugs and Snails. Agricultural, Veterinary and Environmental Perspectives 80: 165–170. ISBN: 1 901396 80 0 ISSN: 0306-3941: No 80 Hudec V. 1954. Rozšíření hlemýždě Theba cartusiana na jižní Moravě [Distribution of snail Theba cartusiana in Southern Moravia]. Vesmír 33 (9): 314–315. Jaremovic R. & Rollo D. 1979. Tree climbing by the snail Cepaea nemoralis (L.). a possible method for regulation temperature and hydratation. Can. J. Zool. 57 (5): 1010–1014. DOI: 10.1139/z79-128 Juřičková L. 1995. Měkkýší fauna Velké Prahy a její vývoj pod vlivem urbanizace [Molluscan fauna in the territory of Prague agglomeration and its development in urban influence]. Natura Pragensis 12: 212 pp. ISSN: 0862-366X Juřičková L. 1998. Měkkýši Hradce Králové [Mollusca of Hradec Králové, East Bohemia, Czech Republic]. Acta Mus. Reginahradecensis, Ser. A 26: 101–172. ISBN: 80-85031-22-1 Juřičková L. 2006. Mollusca – Suchozemští plži. [Mollusca – Terrestrial gastropods], pp. 214–215. In: Mlíkovský J. & Stýblo P. (eds), Nepůvodní druhy ve fauně a flóře České republiky [Alien species of fauna and flora of the Czech Republic], ČSOP, Praha. ISBN: 80-86770-17-6 Juřičková L. & Kapounek F. 2009. Helix (Cornu) aspersa (O.F. M¨ uller, 1774) (Gastropoda. Helicidae) in the Czech Republic. Malacol. Bohemoslov. 8: 53–55. Kerney M.P., Cameron R.A.D. & Jungbluth J.H. 1983. Die Landschnecken Nord- und Mitteleuropas. Verlag Paul Parey, Hamburg und Berlin, 384 pp. ISBN: 3-490-17918-8 Kolouch L.R. 1997. Měkkýši (Mollusca) lokality „Na Plachtě“ v Hradci Králové [Molluscs in the locality “Na Plachtě”, Hradec Králové]. Acta Mus. Reginaehradecensis, Ser. A 25: 69–74. ISSN: 0231-9616 Kolouch L.R. 2005. Další lokality tmavoretky bělavé [New evidence of the occurrence of the Carthusian snail]. Živa 53 (3): 123. Konvička M. & Beneš J. 2002. Metodika mapování [Distribution Atlas. Survey Methods]. In: Beneš J., Konvička M., Dvořák J., Fric Z., Havelda Z., Pavlíčko A., Vrabec V. & Weidenhoffer Z. (eds), Motýli České republiky. Rozšíření a ochrana I, II [Butterflies of the Czech Republic. Distribution and conservation I, II], SOM, Praha, 478 + 379 pp. ISBN: I.&II.: 8090321208 Ložek V. 1948. Prodromus českých měkkýšů [Prodrome of the Czech Molluscs]. Matice česká, Orbis, Praha, 191 pp. Ložek V. 1956. Klíč československých měkkýšů [Key of Czechoslovak Mollusca]. Vydavatelstvo Slovenskej Akademie Vied, Bratislava, 437 pp. Ložek V. 1964. Quart¨ armollusken der Tschechoslowakei. Nakladatelství ČSAV Praha sv. 31, 374 pp., 32 tab., 4 supplements.
Non-native land snails spreading in the Czech Republic Ložek V. 1999. Jihoevropský plž Monacha cartusiana v Českém Krasu [South European Gastropod Carthusian Snail in the Bohemian Karst]. Živa 47 (4): 175. Maňas M. 2002. Měkkýši (Gastropoda, Bivalvia) černovírského slatiniště u Olomouce [Molluscs (Gastropoda, Bivalvia) of Černovír fens near Olomouc]. Ms. Depon. In: OkÚ Olomouc Michalak P.S. & Price T. (eds) 2010. New Pest Response Guidelines – Temperate Terrestrial Gastropods. United States Department of Agriculture, Animal and Plant Health Inspection Service, Cooperating State Departments of Agriculture, 118 pp. Míkovcová A. & Juřičková L. 2008. Hledá se tmavoretka bělavá [The Carthusian snail wanted]. Živa 56 (2): 73. Novák J. & Novák M. 2009. Dvě nové lokality tmavoretky bělavé Monacha cartusiana (O.F. M¨ uller, 1774) na Moravě [Two new localities of Monacha cartusiana (O.F. M¨ uller, 1774) in Moravia, Czech Republic]. Malacol. Bohemoslov. 8: 29–30. Parmesan C. & Yohe G. 2003. A globally coherent fingerprint of climate change impacts across natural system. Nature 421: 37–42. DOI: 10.1038/nature01286 Pech P. & Pechová H. 2009. Monacha cartusiana (Gastropoda. Hygromiidae) in South Bohemia. Malacol. Bohemoslov. 8: 28. Peltanová A. & Novák J. 2010a. Mapa rozšíření Monacha cartusiana v České republice. In. Zicha O. (ed.), Biological Library – BioLib. http://www.biolib.cz/cz/taxonmap/id191/ (accessed 1.6.2011) Peltanová A. & Novák J. 2010b. Mapa rozšíření Cepaea nemoralis v České republice. In. Zicha O. (ed.), Biological Library – BioLib. http://www.biolib.cz/cz/taxonmap/id186/ (accessed 1.6.2011) Peltanová A., Petrusek A., Kment P. & Juřičková L. 2011. A fast snail’s pace: colonization of Central Europe by Mediterranean gastropods. Biological Invasions (18 October 2011), pp. 1–6 DOI: 10.1007/s10530-011-0121-9 Petrbok J. 1940. Monografická studie českých měkkýšů rodu Tachea a Eulota [Monograph of the Czech molluscs of the genera Tachea and Eulota]. Rozpravy II. Třídy České Akademie 50 (13): 1–30. Petrbok J. 1954. Tachea nemoralis Linné v Podmoklech [Tachea nemoralis Linne in Podmokly]. Čas. Nár. Muz., Řada Přír. 123: 225. Rabitsch W. 2008. The times they are a–changin’. Driving forces of recent additions to the Heteroptera fauna of Austria, pp. 309–326. In: Grozeva S. & Simov N. (eds), Advances in Heteroptera Research, Festschrift in Honour of 80th Anniversary of Michail Josifov. PenSoft, Moscow, Sofia, 422 pp. ISBN: 9546423114, 9789546423115
389 Roques A., Rabitsch W., Rasplus J.-Y., Lopez–Vaamonde C., Nentwing W. & Kenis M. 2009. Alien Terrestrial Invertebrates of Europe, pp. 63–79. In: Drake J.A (ed.), Handbook of Alien Species in Europe, Invading Nature, Springer Series in Invasion Ecology, Vol. 3, Springer, ISBN: 978-1-4020-8279-5 e-ISBN: 978-1-4020-8280-1 Říhová D. & Juřičková L. 2011. The Girdled Snail Hygromia cinctella (Draparnaud, 1801) new to the Czech Republic. Malacol. Bohemoslov. 10: 35–37. Říhová D., Peltanová A. & Juřičková L. 2011. Šíří se suchobytka přehlížená, Cernuella neglecta (Draparnaud, 1805), v České republice? [Is the Dune Snail Cernuella neglecta (Draparnaud, 1805) spreading in the Czech Republic?]. Malacol. Bohemoslov. 10: 45–47. Sacchi C.F. 1971. Ecologie comparée des Gastropodes Pulmonés des dunes méditerranéennes et atlatiques. Natura, Milano 62: 277–358. Shinichiro W., Kazuto K. & Satoshi C. 2012. Snails can survive passage through a bird’s digestive system. J. Biogeograph. 39 (1): 69–73. DOI: 10.1111/j.1365-2699.2011.02559.x van Swaay C.A.M. & Warren M.S. 1999. Red Data Book of European Butterflies (Rhopalocera). Society (1999) Volume: 99, Issue: 99, Publisher: Council of Europe, Strasbourg, 1–260 pp. DOI: 10.1007/s10841-005-7548-1 Taberlet P., Fumagalli L. & Wust–Saucy A.-G. 1998. Comparative phylogeography and postgalcial recolonization routes in Europe. Mol. Ecol. 7 (4): 453–464. DOI: 10.1046/j.1365294x.1998.00289.x Tolasz R. et al. 2007. Atlas podnebí Česka. Český hydrometeorologický ústav, Univerzita Palackého v Olomouci, 255 pp. ISBN: 8086690261, 9788086690261 Uličný J. 1885. Beitrag zur Kenntniss der Molluskenfauna von M¨ ahren. Verh Naturforsch. Verein. Br¨ unn 23: 155–172. Uličný J. 1892–1895. Měkkýši čeští [Czech Molluscs]. F. Šimáček, Praha, 208 pp. Uličný J. 1896. Příspěvek ku poznání rozlohy plžů na Moravě. Program c.k. Státního gymnazia v Třebíči 19: 3–24. Urbánek L. 1968. Hlemýžď Monacha cartusiana (O.F. M¨ uller) v Čechách [Snail Monacha cartusiana (O.F. M¨ uller) in Bohemia]. Čas. Nár. Muz., Odd. Přír. 137 (3/4): 114. Welter-Schultes F. 2011. Species summary for Cepaea nemoralis. http://www.animalbase.uni-goettingen.de/zooweb/servlet/ AnimalBase/home/species?id=1370 (accessed 15.04.2011) Received September 6, 2011 Accepted January 4, 2012
