Check-list of Recent Clausiliidae
Subspecies concept
Faunen Europas
Neniinae
Siciliaria Sicily
Dextral Clausiliidae
Clausiliidae Vietnam
Albinaria Crete
Papillifera bidens
Charpentieria itala
Cochlodina diversity
Systematics Clausilia
Alopia Dambovita
Alopia System
Revisoral remarks Phaedusinae
Systematics Baleinae
Plio-Pleistozän
Welter-Schultes 2012
Feher & al. 2013
Welter-Schultes 2010

Charpentieria itala: subspecies, species, superspecies

Hartmut Nordsieck (XI.2011)

Introduction

In the revision of the genus Delima (later on Charpentieria) from the southern Alps (Nordsieck 1963: 174-190) I tried to describe the intraspecific diversity of Charpentieria itala (G. Martens 1824) by a subdivision of the species of that region into several subspecies. These are (from W to E) C. i. albopustulata (Cristofori & Jan 1832), C. i. latestriata (Küster 1850), C. i. rubiginea (Rossmässler 1836), and C. i. serravalensis (H. Nordsieck 1963), besides, as transitional forms within C.i. albopustulata, but treated by following authors as subspecies, C. i. braunii (Rossmässler 1836) and C. i. baldensis (Charpentier 1852) [already described by Strobel (1851: 25), therefore corrected to C. i. baldensis (Strobel 1851)]. A complex of forms which in shell morphology and ecological preferences are similar to the related species C. stenzii (Rossmässler 1836), but in genital morphology more similar to C. itala and connected with it by transitional forms, was added to the latter as "stenzioid subspecies" of C. itala (Nordsieck 1963: 171-173, 181-187): C. i. clavata (Rossmässler 1836), C. i. variscoi (Pini 1883), C. i. balsamoi (Strobel 1850), C.i. lorinae (Gredler 1869). On practical grounds, that complex has been separated later on (Nordsieck 1979: 259) as species C. clavata.
In the 1963 paper the subspecies of C. itala from beyond the southern Alps (Monti Berici and Colli Euganei, Provence, Maritime and Ligurian Alps, northern and central Apennines) are not considered. The related species C. dyodon (Studer 1820) from Piedmont is not treated either, because at this time its close relationship to C. itala was not known.
Zilch (1972), in a type catalogue of the Delimini, has also listed subspecies from beyond the southern Alps: C. i. itala from M. Berici and C. Euganei, and C. i. nigra (Issel 1866) and C.i. punctata (Michaud 1831) from the remaining distributional range.
The following Italian authors called in question the proposed subdivision of C. itala into subspecies, with the following arguments:
Giusti & Mazzini (1971: 294): "Sulla scorta delle sue descrizioni [Nordsieck 1963] abbiamo provato a determinare alcuni materiali dell' Italia settentrionale, tuttavia, senza riuscirci mai con sicurezza. Tanto variabili sono i caratteri conchiliologici..."
Boato et al. (1985: 312): C. itala cfr. punctata: ".. ., sia questa, che le numerose altre che normalmente si includono nella C. itala sono spesso variabili e poco definite".
Manganelli et al. (1995: 47): "Molte delle entità sottospecifiche convalidato da Nordsieck (1963 ...) e da Zilch (1972 ...) qui riportate appaiano di validità dubbia".
These statements, however, are only opinions, apparently not based on further revision.
Therefore, I thought it necessary to check the shell characters once more and to supplement the revision of 1963 by an inclusion of the forms of C. itala from beyond the southern Alps and of C. dyodon from Piedmont. The "stenzioid subspecies" of C. itala = C. clavata are not considered this time, because new results cannot be presented. For the revision, the examined shell material of C. itala has been increased by a third (from 350 samples for the 1963 paper to 460 for this article), and 25 samples of C. dyodon have been included. The material is deposited in the collection of the Forschungsinstitut Senckenberg (SMF) and in my own collection.
The characters used in 1963 have been checked in the added material, and it resulted that the most important characters, those of lunellar and clausilium plate, are not at all too variable and are therefore suited for the definition of subspecies. Obviously, these characters were not considered or not understood by the named authors. Therefore, the statements cited above must be contradicted. The characters of lunellar and clausilium plate are once more described and figured in this paper; hopefully they will be better understood.

Results

1. Character analysis:
The lunellar of C. itala (fig. 1) consists of (posterior) upper palatal plica, lunella and posterior lower palatal plica. An anterior upper palatal plica and an anterior lower palatal plica (= basalis) are not present. The posterior lower palatal plica is identical with the subclaustralis; it is connected with the lunella by an obtuse or even right angle. A lowest palatal plica, the sulcalis, is in part weakly developed; as a rule, it is not connected with the subclaustralis. The outer edge of the clausilium plate rests against the lunella and the subclaustralis in the closed position. There are two possible shapes of lunellar (Nordsieck 1963: 175; see figs. 2-9). In one type of lunellar the upper palatal plica is connected with the lunella by an angle; the lunella is about perpendicular compared with the shell axis (figs. 5, 6). In the other type the upper palatal plica is connected with the lunella by a curve; the lunella is more oblique compared with the shell axis (figs. 2-4, 7-9). There are also two possibilities how the clausilium plate rests against the subclaustralis; this can be observed by an oblique view into the aperture (figs. 6b-9b: arrow!). The outer edge of the clausilium plate rests against the subclaustralis up to its distal end and does not overlap this plica (figs. 6b-7b), or the outer edge of the clausilium plate overlaps that plica (at least in most specimens of the sample) at its distal end (figs. 8b-9b). C. i. serravalensis, C. i. rubiginea and C. i. punctata (see 3.) show the first type of clausilium plate, C. i. itala (see 2.), C.i. albopustulata, C. i. latestriata, C. i. braunii and C. i. baldensis the second one.
Another shell character used for defining subspecies of C. itala is the length of the principal plica. The remaining shell characters, size, shape, sculpture, development of peristome, inferior lamella and palatal callus are of minor importance; only the three first characters are used.


Fig. 1. Lunellar of Charpentieria itala (scheme).  
Abbreviations:
lu = lunella; plp = posterior lower palatal plica (subclaustralis); pr = principal plica; pup = (posterior) upper palatal plica; sc = subcolumellar lamella; sul = lowest palatal plica (sulcalis).  
Figs. 2.-5. C. itala. Actual shell height (mm) = H.
Body whorl dorsal, view on lunellar.  
2. C. i. itala, Mira (Villa Ducale) (ex SMF 193974), H = 18.9.
3. C. i. itala, Vicenza (syntype of. C. i. var. vicentina, ex SMF 196209), H = 21.3.
4. C. i. albopustulata, Varone waterfall near Riva (ex SMF 174367), H = 18.3.
5. C. i. serravalensis, Vittorio Veneto (ex SMF 265953), H = 18.4.  
Figs. 6-9. C. itala. Actual shell height (mm) = H. 
a. Body whorl dorsal, view on lunellar.
b. Body whorl frontal, oblique view into aperture. Different position of clausilium plate with respect to the subclaustralis marked by an arrow.
6. C. i. rubiginea, Bozen (ex coll. Bosch), H = 17.1.
7. C. i. punctata, Fivizzano (ex SMF 186744), H = 16.7 (apical part broken).
8. C. i. albopustulata, Lugano (ex coll. Bosch), H = 18.4.
9. C. i. braunii, Heidelberg (castle) (ex coll. Bosch), H = 15.9.  

2. C. i. itala:
G. von Martens (1824: 442, pl. 3, fig. 1) described C. itala from Miravecchia ("... Garten des aelterlichen Hauses ...", according to E. von Martens 1857: 129 between Mira and Dolo). Zilch (1972: 247) regarded the forms from M. Berici and C. Euganei (with Mira) as representing the nominotypical subspecies and listed the specimens collected by me in Mira (SMF 193974, see Nordsieck 2007: pl. 13, fig. 6) as topotypes. The type form from Mira is relatively small and similar to large forms of C. i. albopustulata (eastern form, see Nordsieck 1963: 177). The samples from M. Berici (e. g. Arcugnano) and Vicenza (C. i. var. vicentina A. Schmidt 1868, syntypes SMF 196209) have a larger and more ventricose shell. The samples from C. Euganei (Abano Terme, Teolo) (C. i. var. vicentina f. tridentina O. Boettger 1879, types SMF 167941) have in part a larger shell , but of the same shape as the type form. In contrast to the other samples, in those from Teolo the clausilium plate does not overlap the subclaustralis.
Among the samples classified by Zilch (: 247) with C. i. itala, however, some from Padova are wrongly affiliated to this subspecies. The sample from Padova (botanical garden) (SMF 156968, including the representative specimen figured by Zilch 1960: fig. 1532) belongs to C. i. braunii.
"Fern" (Vicenza), in the internet forum Natura Mediterraneo (V. 2008), who had, in order to identify C.i. itala, collected specimens between Mira and Dolo, stated that they look like C. i. serravalensis ("Corrispondono ... anche alla serravalensis"), but did also not exclude a close relationship to C. i. albopustulata ("Non posso ... escludere la possibilità che albopustulata si insinui nella pianura veneta.") For his comparison, however, he did not consider the differences between C. i. serravalensis and C. i. itala in the clausilium plate.
C. i. itala does not much differ from C. i. albopustulata (eastern form), but is maintained as subspecies because of its shell size (on average larger) and its isolated occurrence (see map). The vicentina form differs from C. i. albopustulata also by its shell shape (more ventricose); thus the type form of C. i. itala is transitional between the vicentina form and C. i. albopustulata. The form from Teolo resembles C. i. serravalensis.

3. C. i. punctata:
The various forms of C. itala from Provence, Maritime and Ligurian Alps and northern and central Apennines (see map) are characterized by a stronger sculpture and a clausilium plate which does not overlap the subclaustralis. The form from the Provence (C. punctata Michaud 1831) and those from the Apennines (C. alboguttulata var. obesa Issel 1866 (non L. Pfeiffer 1861) from Lucchese, C. a. var. nigra Issel 1866 from Volterra, and C. a. var. elegans Gentiluomo 1868 (non Cantraine 1835) from Vallombrosa) do not much differ and are therefore not separated as subspecies (in contrast to Giusti & Mazzini 1971: 295, Zilch 1972: 251). The whole subspecies has to bear the oldest name C. i. punctata. This was already proposed by Boato et al. 1985: 312, but as cfr. punctata, and Manganelli et al. 1995: 24, but with the statement (: 47): "L' attribuzione del materiale della Liguria e dell' Appennino a C. i. punctata permane incerta."
The alleged syntypes of C. punctata from the Rossmässler collection (SMF 196374) are much smaller and more ventricose than the type specimens (Michaud 1831: 55, pl. 15, fig. 23); they belong to C. i. braunii. This has been noticed neither by the previous authors (Rossmässler, O. Boettger) nor by Zilch (1972: 251), who listed them as syntypes.
It should be checked, if C. i. punctata is distributed in Italy beyond the southern border indicated in the map, e. g. as far as the Colli Albani, where it is said to occur by E. von Martens (1857: 136).

4. C. i. "braunii" from Meran:
The C. itala form from Meran and surroundings, South Tyrol, differs much from C. i. braunii (type form fig. 10) to which it was affiliated in the 1963 paper (: 177). Unlike C. i. braunii, it has a long principal plica and a clausilium plate not overlapping the subclaustralis. It thus resembles C. i. serravalensis (the only subspecies in which a long principal plica is frequently present) and is therefore classified with that subspecies.

5. C. dyodon:
C. dyodon, which was formerly placed with another group (Dilataria Vest) in another subfamily (Zilch 1960: 401), has been recognized by me by an examination of the genitalia (Nordsieck 1972: 38) as closely related to C. itala. The shell differs from that of C. itala by the complete or nearly complete reduction of the sutural papillae and a lunellar tending to reduction (lunella weakly developed, posterior lower palatal plica missing, clausilium plate distally rounded). Until now, forms transitional to C. itala have not been found. Therefore, in spite of its genital similarity to C. itala, C. dyodon is regarded as an independent species. Its distributional range in Piedmont (see map) fills partly the gap between that of the C. itala subspecies from the southern Alps and that of C. i. punctata. Therefore I assume that C. dyodon has originated from an isolate of the common stem form in that region. C. itala (with C. clavata) and C. dyodon might belong to a monophyletic group of closely related species, the superspecies of C. itala.
What concerns the subspecies division of C: dyodon, see Nordsieck (2002: 36).

6. Genus Charpentieria:
Charpentieria sensu stricto (without the Siciliaria groups, see website article Delimini Italy) is regarded as a genus and includes besides the superspecies of C. itala two further species, C. ornata (Rossmässler 1836) and C. stenzii (Rossmässler 1836). C. ornata differs from C. itala by shell characters more than the subspecies of the latter from each other (Nordsieck 1963: 198). C. stenzii differs from C. itala in shell morphology to about the same extent as C. dyodon, but in genital morphology more (Nordsieck 1963: 191, 201). A subgenus Itala O. Boettger, in which C. itala, C. ornata and C. stenzii are united, is not monophyletic and has therefore been rejected (Nordsieck 2002: 36). By further investigations, e. g. more extensive examinations of the genitalia and DNA analyses, it must be found out if C. dyodon, as assumed by the superspecies concept (see 5.), is really more closely related to C. itala than C. ornata and (or) C. stenzii.


Map. Distribution of C. itala subspecies and C. dyodon ("stenzioid subspecies" of C. itala = C. clavata not considered).
Marked ranges cover the regions in which samples of the respective taxa have either been collected or are mentioned in the literature. 
Fig. 10. C. i. braunii (type form), on a wall near the castle of Weinheim, Germany. The type locality of C. i. braunii ("Auf dem Odenwalde") is restricted to Weinheim (see Gysser 1863: 18). 

References

Boato, A., Bodon, M. & Giusti, F. (1985): Molluschi terrestri e d'acqua dolce delle Alpi Liguri. – Lavori della Società Italiana di Biogeografia, (NS) 9 [1982]: 237-371.

Giusti, F. & Mazzini, M. (1971): Notulae Malacologicae XIV. I molluschi delle Alpi Apuane. Elenco delle specie viventi con descrizione di una nuova specie: Vitrinobrachium baccettii n. sp. – Lavori della Società Italiana di Biogeografia, (NS) 1[1970]: 202-335, 9 pls.

Gysser, A. (1863): Die Mollusken-Fauna Baden's. Mit besonderer Berücksichtigung des oberen Rheinthales zwischen Basel und Mannheim. – 32 pp. Heidelberg (Wolff).

Manganelli, G., Bodon, M., Favilli, L. & Giusti, F. (1995): Gastropoda Pulmonata. In: Minelli, A., Ruffo, S. & La Posta, S. (Eds.): Checklist delle specie della fauna italiana, 16: 1-60. Bologna (Calderini).

Martens, E. von (1857): Reisebemerkungen über einige Binnenschnecken Italiens. – Malakozoologische Blätter, 4: 120-155.

Martens, G. von (1824): Reise nach Venedig, 2: VI + 664 pp., pls. Ulm.

Michaud, A.-L.-G. (1831): Complément de l'Histoire des mollusques terrestres et fluviatiles de la France, de J. P. R. Draparnaud. – VI+116+12 pp., pls. 14-16. Verdun.

Nordsieck, H. (1963): Zur Anatomie und Systematik der Clausilien, II. Die Formenbildung des Genus Delima in den Südalpen. – Archiv für Molluskenkunde, 92 (5/6): 169-203.

– – – (1972): Zur Anatomie und Systematik der Clausilien, XI. Neue Formen und taxonomische Revision einiger Gruppen der Alopiinae. – Archiv für Molluskenkunde, 102 (1/3): 1-51, Taf. 1-5.

– – – (1979): Zur Anatomie und Systematik der Clausilien, XXI. Das System der Clausilien, II: Die rezenten europäischen Clausilien. – Archiv für Molluskenkunde, 109 (4/6): 249-275.

– – – (2002): Contributions to the knowledge of the Delimini (Gastropoda: Stylommatophora:: Clausiliidae). – Mitteilungen der deutschen malakozoologischen Gesellschaft, 67: 27-39.

– – – (2007): Worldwide Door Snails (Clausiliidae), recent and fossil. – 214 pp., 20 pls. Hackenheim (ConchBooks).

Strobel, P. (1851): Notizie malacostatiche sul Trentino. – 114 pp. Pavia (Fusi).

Zilch, A.(1959-1960): Gastropoda. Teil 2. Euthyneura. – In: Schindewolf, O. H. (ed.), Handbuch der Paläozoologie, 6 (2): I-XII, 1-834. Berlin-Nikolassee (Borntraeger).

– – – (1972): Die Typen und Typoide des Natur-Museums Senckenberg, 50: Mollusca: Clausiliidae (2): Alopiinae: Delimini (1) . – Archiv für Molluskenkunde, 102 (4/6): 247-259, pl. 11.


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