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Home , Chilostoma, Eucobresia, Eucobresia diaphana, Eucobresia nivalis, Limacidae, Malacolimax tenellus

March 2013 C.J.P.J. (Kees) Margry Mozartlaan 41 NL 5283 KB Boxtel, the Netherlands. [email protected] diaphana (Draparnaud, 1805)

Vitrinidae () and other molluscs in Parc national de la Vanoise in the French Alps. A report from a short research in July 2012.

Résumé En Juillet 2012, une étude a été menée sur la présence des dans une partie du Parc national de la Vanoise. Dans trois régions, huit sites ont été échantillonnés. On a fait des recherches sur les animaux vivants et les coquilles vides. Les coquilles vides des autres espèces ont aussi été collectées. Au total, aux côtés de certains résidus qui n’ont pas pu être identifiés, 18 espèces ont été observées, y compris deux espèces de Vitrinidae: pellucida (Semilimace commune) et Eucobresia glacialis (Semilimace recouverte). On a trouvé des E. glacialis de taille remarquable. En cherchant dans les années antérieures, il est connu que le stabilei (Semilimace des Alpages) vit également dans le Parc national de la Vanoise.

Key words: Parc national, Vanoise, Gastropoda, Eucobresia, Vitrina, Phenacolimax.

Introduction In the summer of 2012 the occurrence of Vitrinidae has been investigated in a part of Parc national de la Vanoise and the surrounding area in the French Alps. The motivation for this research is the personal interest for Vitrinidae in general and (Draparnaud, 1805) in particular (Margry & Boesveld, 2009). The Family of Vitrinidae (glass snails) is a mainly Palaearctic group of semislugs. On the European mainland this family has a center of diversity in alpine habitats (Hausdorf, 2002). Glass snails have very thin translucent shells. Most of the have such a little house, that they are not able to withdraw themselves into their shell. They are adapted to low temperatures and live in wet and cold habitats (Kerney et al., 1983). In lowland areas they are more active in winter and have a one-year lifecycle with a period of rest in summer (i.e. “summer decline”). On higher altitude they are more forced to activity in summer and have a lifecycle up to a few years (Umi!ski 1975a, 1975b, 1979, 1983) In Europe (including Turkey but except the Macaronesian Islands) 25 vitrinid species are recorded (Forcart, 1944, 1946; Kerney et al., 1983; Fechter & Falkner, 1990; de Winter, 1990; Wiese & Welter-Schultes, 1992; Turner et al., 1998; Falkner et al., 2001, 2002; Hausser, 2005; Nardi et al., 2007; Gavetti et al., 2008; Boschi, 2011; Egorov, 2011; Giusti et al., 2011; Welter-Schultes, 2012). In eleven species are recorded, but there is doubt about the occurence of semilimax (J. Férussac, 1802) and Vitrinobrachium breve (A. Férussac, 1821) (Falkner et al., 2002: 131-132; Gargominy et al., 2011). Table 1 shows which vitrinid snails have been found in, or close to, the French Alps. The French Alps are of great interest in respect to the surveys in the Valley of Susa in and the surveys in the north and northeast in Switserland (Gavetti et al., 2008; Boschi, 2011; Map 1). Only three species are recorded in both these Italien and Swiss researches.

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 1 2012. 1-12. www.parcnational-vanoise.fr

black spot = Parc national de la Vanoise red area = the Valley of Susa

Map 1. The research area in the French Alps

Falkner et al. (2002) suggested to look for Semilimax kotulae (Westerlund, 1883) in the (Haute) Savoie. One of the conditions for the authorization for this research was the compiling of a report. In this report only the research is described from the part within the borders of the National Park.

Species Record from Record from Record in Record in Distribution France Switserland, the Italien the French map for the (Falkner et close to the part from the part from region of al., 2002; French Susa Valley the Susa the Gargominy border (Gavetti et Valley Vanoise et al (2011) (Boschi, al., 2008) (Gavetti et National 2011) al., 2008) Park in Al = alpine Welter- Schultes (2012) (X) Semilimax kotulae X X Semilimax pyrenaicus X Vitrinobrachium breve (X) X Eucobresia diaphana X Al X X X Al X X Eucobresia pegorarii X Eucobresia glacialis X Al X X X X X Al X X X X X Al X X Phenacolimax stabilei X Al X X X Phenacolimax locardi X Oligolimax annularis X Al X X X X Table 1. Vitrinid snails according to literature found in or close to the French Alps.

Methods The study area was visited between July 10th and 27th, 2012. All sampling areas within the borders of the Vanoise National Park were in the region of Peisey-Nancroix, Col d’Iséran and la Thuile, the lowest part of the National Park in the southeast of Bourg-Saint-Maurice (Region Rhône-Alpes, Departement Savoie) (Table 2, Map 2, Fig 1-8). Vitrinidae are difficult to identify on the mere basis of their appearance. For this reason authorization was obtained to collect both empty shells and living . The localities and

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 2 2012. 1-12. www.parcnational-vanoise.fr sampling sites were chosen at random. The sampling was done by hand by means of looking under rocks and dead wood and utilizing a special big pair of glasses that makes it possible to find snails as small as 1 mm. The research was not done in a standardized way. If after about a quarter of an hour nothing was found, the sampling site was abandoned without collection of any field data. If one or more shells were found the search could take over an hour. Besides Vitrinidae, empty shells of other snail species were collected as well. Field data and photos were taken to document the samples as precisely as possible. The plant species on the plots were not identified. The living Vitrinidae were initially kept in a refrigerator (winter snails!) in order to study their habits and take photos. If available the genital anatomy from at least one vitrinid specimen from each plot was studied to confirm the identification. The genital anatomy of Vitrinidae is described and illustrated in several references (Mermod, 1930; Forcart, 1944, 1946; Umi!sky, 1975a; Kerney et al., 1983; de Winter 1990; Nardi et al., 2007; Egorov, 2011; Giusti et al., 2011). All figures of Boschi (2011) are derived from Kerney et al. (1983). A figure of the genitals of Phenacolimax locardi (Pollonera, 1884) was kindly provided by Gianbattista Nardi. From a few species Umi!ski (1975a: 373) shows several stages of the changes of the genitals during maturation. Phenacolimax blanci (Pollonera, 1884) was not included in this study. It is considered to be an Italian endemic in the surroundings of Cuneo and the Imperia province close to the French border in the south (Welter-Schultes, 2012).

Map 2. The northern part of Parc national de la Vanoise with the 8 sampling plots

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 3 2012. 1-12. www.parcnational-vanoise.fr

Fig. 1. plot 1 photo Ingrid Margry Fig. 2. plot 2 photo Kees Margry

Fig. 3. plot 3 photo Kees Margry Fig. 4. plot 4 photo Ingrid Margry

Fig. 5. plot 5 photo Kees Margry Fig. 6. plot 6 photo Kees Margry

Fig. 7. plot 7 photo Kees Margry Fig. 8. plot 8 photo Kees Margry

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 4 2012. 1-12. www.parcnational-vanoise.fr

For identification of the other species Kerney et al (1983), Gittenberger & Bank (1996), Wiktor (2004), Hausser (2005), Gargominy & Ripken (2011) and Gargominy & Neubert (2011) were used. The nomenclature used, is according to Fontaine et al, 2010. (Chilostoma glaciale (A. Férussac, 1832) = Phontanilli glacialis = Delphinatia glaciale). Refer to appendix 1 for the French names. Measurements of the shells were taken with a calliper to the nearest 0.01 mm. The collected shells and alcohol samples are kept in the collection of the author.

Plot locality sampling site Coordinates Altitude Vegetation sampling nr (m) belt* nr 1 Peisey- Rocher des 45˚29’11.7’’ N – 006˚50’27.1’’ E 2151 alpine 20120716.2 Nancroix, Mindières 2 Rocher des 45˚29’24.6’’ N – 006˚50’27.8’’ E 2120 alpine 20120716.3 Mindières 3 Plan de la 45˚29’49.5’’ N – 006˚50’24.6’’ E 2095 alpine 20120716.4 Plagne 4 Refuge du Mont 45˚31’44.9’’ N – 006˚49’37.7’’ E 2380 alpine 20120722.1 Pourri 5 la Thuile about 1 km 45˚33’51.1’’ N – 006˚53’19.9’’ E 1249 montane 20120727.1 south of the village 6 Col close to Col 45˚25’24.4’’ N – 007˚02’45.2’’ E 2853 nival 20120718.1 d’Iséran Pers 7 close to Col 45˚25’24.2’’ N – 007˚02’48.6’’ E 2910 nival 20120718.2 Pers 8 Pointe des 45˚25’54.9’’ N – 007˚01’40.8’’ E 2829 nival 20120718.3 Lessières Table 2. Localities and sampling sites. * Vegetation belt according to Nagy & Grabherr (2009). montane = forest, alpine = alpine meadow, nival = Scree, rocks and snow level, scant patchy vegetation

Results Table 3 provides a list of species for each sampling site, in alphabetical order. In the upper part only Vitrinidae species are indicated, while the lower part contains al the other molluscs.

Vitrinidae Vitrinidae were recorded in six out of eight sampling sites. On three sampling sites only Vitrinidae were found. In total 16 living animals and 39 empty shells were collected.

Vitrina pellucida Semilimace commune (Fig. 9) From one sampling site (plot 3) in the valley of Plan de la Plagne, Vitrina pellucida could be identified by anatomical research (Fig. 11). From two empty shells of plot 1 it is presumed that they are from V. Pellucida as well.

Eucobresia glacialis Semilimace recouverte (Fig. 10, 15) From two sampling sites close to Col Pers Eucobresia glacialis could be identified by anatomical research (Fig. 12). On plot 6 en 7 a total of 15 living specimens and 25 empty shells were found. All those empty shells are most likely derived from E. glacialis. Most of them show the typical conchological characters of this species. In Figure 14 the measurements of the shells found on plot 6 and 7 are given.

The only two empty shells found on plot 5 were damaged in such extent, that identification was impossible. Al other empty shells from plot 1 and 2 are obviously not from Oligolimax annularis (Studer 1820) and probably neither from Vitrina pellucida. Broken parts of shells were found as well. Most of those broken parts are not suitable for further research. In the National Park close to la Thuile, no Vitrinidae were found.

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 5 2012. 1-12. www.parcnational-vanoise.fr

Other molluscs On five out of eight sampling sites molluscs other than Vitrinidae were found. On two sampling sites only other mollusc species were recorded. The subspecies of Chilostoma zonatum could be identified as Chilostoma zonatum flavovirens (Dumont & Mortillet, 1852). This subspecies is known from the triangle Bourg-Saint-Maurice (Col de la Seigne), Col du Mont Cenis and Savines-le-Lac (Haute Alpes). This area is not indicated on the distribution maps of Chilostoma in Formenti (2012). In one sample (plot 2), 14 empty shells from Cochlicopa were found. In the Cochlicopa in the Alps at least three species can be distinguished. Four specimens are juveniles or have a broken shell. Five specimens with a length of less than 5 mm have a width of respectively 2.05, 2.06, 2.08, 2.09 and 2.31 mm. Five specimens with a length of more than 5 mm have a width of respectively 2.07, 2.15, 2.37, 2.39 and 2.46 mm. Those measurements do not provide enough support for identification of the second species C. lubrica (O.F. Müller, 1774). All the shells are listed as Cochlicopa cf lubricella (Rossmässler, 1834). A fresh dead could be identified as tenellus (O.F. Müller, 1774) (Fig. 13). From another slug (plot 3) only the internal shell was found.

Peisey-Nancroix La Col d’Iséran Thuile Species 1 2 3 4 5 6 7 8 16.2 16.3 16.4 22.1 27.1 18.1 18.2 18.3 Vitrinidae

Eucobresia glacialis 1g 1g 12a 1a Vitrina pellucida cf 2d 1g Vitrinidae indet 2d + r 8d +r - 2d - 14d +r 11d - Other molluscs

Aegopinella pura X Arianta arbustorum forma X alpina Chilostoma glaciale X Chilostoma zonatum X X flavovirens Columella columella X Cochlicopa cf lubricella X Cochlodina laminata X Discus rotundatus X Ena montana X Euconulus fulvus X X X Macrograstra plicatula X X X X Nesovitrea petronella X X X Punctum pygmaeum X pusilla X Vitrea subrimata X Hygromiidae indet X X / X Total number of species 2 10 6 1 10 1 1 1 Table 3. Number of idividuals of vitrinid species and records of other molluscs (X) on each sampling site. a = alive without anatomical research g = anatomical research of genitals d = dead, empty shells +r = broken empty shells, no measurements possible

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 6 2012. 1-12. www.parcnational-vanoise.fr

Fig. 9. Vitrina pellucida photo Ingrid Margry

Fig. 10. Eucobresia glacialis photo Ingrid Margry

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 7 2012. 1-12. www.parcnational-vanoise.fr

Fig. 11. Vitrina pellucida Fig. 12. Eucobresia glacialis Fig. 13. Malacolimax tenellus

Fig. 11-13. Genitals from three species (Fig. Kees Margry). A = atrium, BC = bursa copulatrix, H = hermaphrodite gland, P = penis, PR = penial retractor muscle, SO = spermoviduct, V= vagina.

Figure 14. Measurements of shells from Eucobresia close to Col Pers (plot 6 and 7). red = Eucobresia glacialis, identified according to genitals

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 8 2012. 1-12. www.parcnational-vanoise.fr

Discussion

Vitrinidae No specimens of Eucobresia diaphana were found. In France records of E. diaphana are known from the northeast (Bichain, 2003; inpn.mnhn.fr/espece/cd_nom/163168/tab/sources, viewed 10-ii-2013). According to records in this species could be expected in the French Alps. More research has to be done. E. diaphana resembles E. glacialis. Old records of E. diaphana on higher altitudes probably concerns the species E. glacialis or E. nivalis (Dumont & Mortillet, 1854).

Fig. 15. Eucobresia glacialis, juvenile with shell from 4.39 mm (photo alc. Specimen 20120718.1c)

The measurements of the shells of Eucobresia glacialis suggest that there are at least two cohorts present during the same time period. More strikingly is the large size of some shells, up to almost 8 mm. In literature the lenght is given as 4.5-6 mm width (Kerney et al., 1983; Boschi, 2012; Welter-Schultes, 2012). These shells are probably from the cohort that died last winter. E. glacialis is known from the nival zone. Gavetti et al (2008) found E. glacialis on Monte Rocciamelone at 3400 m asl. Vitrina pellucida is a common species that can also be found in much drier places. It is recorded all over Europe and not restricted to places of higher altitude. In Parc national de la Vanoise at least one more vitrinid species is recorded. Two years ago also Semilimace des alpages Phenacolimax stabilei (Lessona, 1880) was found within de boundaries of the National Park. More research will clarify which Vitrinidae live in this area. Especially for this group the climat change can cause significant problems as Müller et al. (2009) showed for Semilimax kotulae.

Other species On the sampling site close to la Thuile (plot 5), some empty shells of Aegopinella pura (Alder, 1830) and Euconulus fulvus (O.F. Müller, 1774) were collected. These specimens still have their periostracum but do not have any colour. Albinism in shells is found in several species, especially on specimens in wet and shady places (Clessin, 1873: 48; Schlesch, 1927: 38). In the lower part of the National Park close to la Thuile probably more species can be found. Because of an upcoming heavy thunderstorm this plot was left before the collecting was finished. Some Hygromiidae could not be identified. According to different references this group of snails contains a variable number of species, some of which are often covered with hairs. DNA- research on a few species showed that the distinction between species is rather complicated (Dépraz et al., 2009).

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 9 2012. 1-12. www.parcnational-vanoise.fr

Acknowledgements The author would like to thank Michael Delorme for the authorization for the research in the National Park, Bernd Shalmann (Haus der Natur Cismar) and Folco Giusti for provision of literature, Gianbattista Nardi for his figure of the genitals of Phenacolimax locardi, Joop Eikenboom for the identification of Chilostoma zonatum flavovirens, Edi Gittenberger, Wim Maassen and Ton de Winter (Naturalis) for their help with literature and sharing of their knowledge and expertise. Thanks to Bram Margry, Françoise Chan and Kobus Margry for their help with figures and the English and French language and, last but not least, my wife Ingrid for her help with the fieldwork and all the photos she took.

References

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Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 11 2012. 1-12. www.parcnational-vanoise.fr

Appendix 1. French names of the molluscs in this report.

Vitrinidae Semilimax semilimax (J. Férussac, 1802) Semilimace des montagnes Semilimax kotulae (Westerlund, 1883) Semilimace alpine Semilimax pyrenaicus (A. Férussac, 1821) Semilimace atlantique Vitrinobrachium breve (A. Férussac, 1821) Semilimace germanique Eucobresia diaphana (Draparnaud, 1805) Semilimace aplatie Eucobresia nivalis (Dumont & Mortillet, 1854) Semilimace des neiges Eucobresia pegorarii (Pollonera, 1884) (not available) Eucobresia glacialis (Forbes, 1837) Semilimace recouverte Vitrina pellucida (O.F. Müller, 1774) Semilimace commune Phenacolimax major (A. Férussac, 1807) Semilimace des plaines Phenacolimax stabilei (Lessona, 1880) Semilimace des alpages Phenacolimax locardi (Pollonera, 1884) (not available) Oligolimax annularis (S. Studer, 1820) Semilimace globuleuse

other molluscs Aegopinella pura (Alder, 1830) Petite luisantine Arianta arbustorum alpicola (A. Férussac, 1821) Hélice des Alpes Chilostoma zonatum flavovirens (Dumont & Mortillet, 1852) Hélicon des granites Columella columella (G. von Martens, 1830) Columelle alpine Cochlicopa cf lubricella (Rossmässler, 1834) cf Petite brillante Cochlodina laminata (Montagu, 1803) Fuseau commun Chilostoma glaciale (A. Férussac, 1832) Hélicon des Alpes Discus rotundatus (O.F. Müller, 1774) Bouton commun Ena montana (Draparnaud, 1801) Bulime montagnard Euconulus fulvus (O.F. Müller, 1774) Conule des bois Macrograstra plicatula (Draparnaud, 1801) Massue costulée Malacolimax tenellus (O.F. Müller, 1774) Limace jaune Nesovitrea petronella (L. Pfeiffer, 1853) Luisantine brune Punctum pygmaeum (Draparnaud, 1801) Escargotin miniscule (Vallot, 1801) Pyramidule commun Vitrea subrimata (Reinhardt, 1871) Cristalline méridionale

Entrance of Parc national de la Vanoise close to la Thuile

Margry, C.J.P.J., 2013. Vitrinidae and other molluscs in Parc national de la Vanoise. A report from a short research in July 12 2012. 1-12. www.parcnational-vanoise.fr