Delimini (Gastropoda, Stylommatophora, Clausiliidae) from central and southern Apennine Italy with adjacent Sicily (except genus Papillifera Hartmann)

Hartmut Nordsieck

Introduction

Delimini species of Apennine Italy (with adjacent Sicily) with normal clausilial apparatus (= CA) are widely distributed and especially in southern Italy relatively diverse, but have never been revised. The species concerned cannot be determined easily, because clear diagnoses do not exist. Also, until now, exact informations on the distribution have not been given.
The last revision is that of O. Boettger (1879: 91-100, 103-104, list 150-151) in continuation of Rossmässler's Iconography, based on shell characters. It contains descriptions and illustrations of several forms, but no clear diagnoses of the species. Since then, only systematic lists without descriptions have been published. A. J. Wagner (1924: 121-123, 1925: 54-57) has given a list with information on distribution and description of some new taxa. Forcart (1965: 116-120) has published a list of the species from southern Apennine Peninsula, with information on the literature and distribution. The list of Manganelli et al. (1995: 24-25, 48) gives only insufficient distributional data, my own list (Nordsieck 2002: 32, 35) some revising remarks at least. What concerns genital morphology, only some figures have been published, without information on species differences (sketches by A. J. Wagner 1925: fig. 27 (Siciliaria stigmatica Rossmässler), fig. 29 (S. gibbula Rossmässler); more accurate figures by Forcart 1965: fig. 5 (S. paestana Philippi, not S. kobeltiana Küster!), Nordsieck 1969: fig. 9 (S. gibbula) and Giusti 1973: fig. 28 D, F (S. vulcanica Paulucci)).
The material on which the present paper is founded is from the collection of the Forschungsinstitut Senckenberg (SMF) and my own collection. The latter has mainly been collected by myself during four journeys in Apennine Italy in 2009 and 2010. For the localities of examined material see map 1. The species of which I have examined the genital morphology have already been listed in a former paper (Nordsieck 2002: 27-28).

Results

Induced by the results of the DNA analysis of several Delimini species (Hausdorf unpubl.) I have checked once more the relationship of the genus taxa Charpentieria Stabile and Siciliaria Vest (1.) and the systematic position of Delima piceata Rossmässler (2.).
1. Now as before, I have found only two differences of the genitalia between Charpentieria and Siciliaria, as already discussed in a former paper (Nordsieck 2002: 33):
Charpentieria: Epiphallar thickening at the transition epiphallus-penis missing; delimitation of proximal part of penis distinct.
Siciliaria: Epiphallar thickening at the transition epiphallus-penis present; delimitation of proximal part of penis indistinct.
If Siciliaria is regarded as an independent genus as in the following, it can, at present, morphologically only be defined by the named characters. But it should not be concealed that these characters are in some species weakly or even not expressed. Therefore, they alone will not be sufficient for the generic separation.
To the genus Siciliaria belongs also Mauritanica O. Boettger, because it exhibits the same characters.
2. D. piceata has been affiliated to the genus Delima mainly based on one genital character (Nordsieck 1972: 39, 2002: 32). It has a distinct penial papilla base like the Delima species. On the other side, it has an indistinct delimitated proximal part of penis like the Siciliaria species. A repeated comparison of the shell and other genital characters has shown that D. piceata corresponds more with S. paestana than with externally similar Delima species like D. amoena L. Pfeiffer or D. blanda Rossmässler. Such shell characters are the missing lamella inserta (A. Schmidt 1868: 45) and the inferior lamella which tends to end with two branches at the peristome. Not only the delimitation of the proximal penis, but also the measurements of the genitalia, especially the long penial papilla, are like those of S. paestana. The ecological preferences (living in woods) and the distribution on the Apennine peninsula speak also in favour of a relationship to S. paestana.
The classification of D. piceata was obviously wrong, caused by a wrong evaluation of the distinct penial papilla base (which is possibly a plesiomorphic character). The species is therefore transferred to the genus Siciliaria as defined in 1. The subgenus name Delima (Piceata) O. Boettger becomes a synonym of Siciliaria (Stigmatica) O. Boettger.

The most important characters for species diagnoses are those of the clausilial apparatus = CA, especially of the lunellar and the clausilium plate (figs. 1-5). For the palatal plicae see fig. 1, for the clausilium plate and its relations to the plicae see figs. 2-5.

Abbreviations: alp = anterior lower palatal plica (basalis), aup = anterior upper palatal plica, cp = clausilium plate, il = inferior lamella, lu = lunella, plp = posterior lower palatal plica, pr = principal plica, pup = (posterior) upper palatal plica, sc = subcolumellar lamella, sl = superior lamella.


The shell diagnoses of the Delimini species with normal CA from central and southern Apennine Italy and adjacent Sicily are as follows:

Siciliaria Vest 1867

S. (Gibbularia) Monterosato 1908

S. (G.) gibbula (Rossmässler 1836):
Lunella not or shortly interrupted above; posterior lower palatal plica separated from lunella; anterior lower palatal plica = basalis fully developed; anterior upper palatal plica mostly present, connected with or separated from upper palatal plica; clausilium plate slightly emarginate, distal part groove-like, outer corner more or less pointed and bent up (fig. 2).
The species is subdivided into two main subspecies: S. g. gibbula sensu lato, S. g. honii (O. Boettger 1879).
S. g. honii differs from S. g. gibbula s.l. by the following characters: Inferior lamella ending with two branches; anterior upper palatal plica on an average shorter; outer corner of clausilium plate less upbent.
Distribution: Central and southern peninsular Italy (Tyrrhenian coast and islands, Penisola Salentina, Aspromonte?).
S. g. gibbula s. l. contains beside the nominotypical subspecies with normally sculptured shell, which is farly distributed, some minor subspecies with restricted occurrences.
Distribution: Central and southern peninsular Italy (from Emilia-Romagna to western Puglia and northern Basilicata, probably introduced in the northern Adriatic regions).
Synonyms: C. laurae Adami 1886, C. gibbula var. solidula Monterosato in Cecconi 1908 [non L. Pfeiffer].
The minor subspecies are four ribbed subspecies and one normally sculptured one:
S. g. niethammeri (Rensch 1934) from Foresta Umbra, Monte Gargano, Puglia, and S. g. multiplex (Westerlund 1884) (= vana Westerlund 1884 = vaga Westerlund 1884) from Trani, Puglia, both with detached aperture and anterior upper palatal plica separated from upper palatal plica, S. g. multiplex besides with outer corner of clausilium plate less upbent;
S. g. pelagosana (O. Boettger 1877) from Palagruža Islands, Croatia, and S. g. selecta (Monterosato in Cecconi 1908) from Capraia and Cretaccio, Tremiti Islands, Puglia, both with non-emerging subcolumellar lamella, S. g. selecta besides with outer corner of clausilium plate not upbent;
S. g. sanctangeli (A. J. Wagner 1925), a normally sculptured subspecies from Monte Gargano, Puglia, with larger shell than in the nominotypical subspecies and outer corner of clausilium plate less upbent.
The latter has been collected only in the western part of Monte Gargano. An occurrence near Monte S. Angelo, as asserted by A. J. Wagner, could not be stated; in that locality only S. g. gibbula has been found.

S. (Stigmatica) O. Boettger 1877

S. (S.) stigmatica (Rossmässler 1836):
Lunella ± reduced above; posterior lower palatal plica ± separated from lunella; basalis fully developed; anterior upper palatal plica in part present, displaced from upper palatal plica; clausilium plate emarginate (fig. 3).
In Italy only S. stigmatica sturmii (L. Pfeiffer 1848) (= S. s. maritima sensu auct.) occurs; it is restricted to the Penisola Salentina, Puglia (Nordsieck 2002: 35) (see map 1). In comparison with the nominotypical subspecies it has a more distinct dorsal keel and an emerging subcolumellar lamella. The species might not be introduced from the Balkan peninsula, as assumed by former authors, because it has been found in natural habitats in several localities of the peninsula.

S. (S.) incerta (Küster 1860):
Lunella not or less reduced above; posterior lower palatal plica connected with lunella; basalis relatively short; anterior upper palatal plica missing; clausilium plate not or nearly not emarginate.
Distribution: N. E. Sicily (Madonie, Nebrodi), S. Calabria.
Synonym: C. monterosati Benoit 1882.

S. (S.) kobeltiana (Küster 1876):
Lunella not or less reduced above; posterior lower palatal plica ± separated from lunella; basalis mostly short; mostly without anterior upper palatal plica; clausilium plate not or nearly not emarginate.
Distribution: Calabria.
Synonyms: C. calabrica Westerlund 1878, C. k. var. furcata Paulucci 1878, C. k. var contorta Paulucci 1878.

S. (S.) vulcanica (Paulucci 1878):
Lunella ± reduced above; posterior lower palatal plica ± separated from lunella; basalis long; upper palatal plica frequently elongated by an anterior upper palatal plica; clausilium plate emarginate (fig. 4).
Distribution: Calabria, N. E. Sicily, Eolie Islands.
In the type form of the latter from N. E. Sicily inferior lamella mostly ending with only one branch.
Because of its external similarity with S. g. honii, S. vulcanica has been regarded by O. Boettger and following authors as a subspecies of S. gibbula. Its clausilium plate, however, is emarginate, outer corner not pointed and upbent.

S. (S.) paestana (Philippi 1836):
Lunella not reduced above; posterior lower palatal plica connected with lunella; basalis mostly missing; anterior upper palatal plica missing; clausilium plate entire (fig. 5).
Distribution: Central and southern peninsular Italy (from Toscana (Monte Argentario) and S. Abruzzo to N. Calabria).
S. p. paestana: Basalis indistinct or missing.
Synonyms: C. neumeyeri Küster 1850, C. paestana var. semisculpta Paulucci 1878, C. p. var. tenuisculpta O. Boettger 1879, D. piceata sangroensis A. J. Wagner 1925.
S. p. intustructa (Westerlund 1883): Basalis ± well-developed.
The latter subspecies (Nordsieck 2002: 35, fig. 7), which is not mentioned in the list of Manganelli et al. (1995), occurs in Basilicata (see map 1). A form transitional to the nominotypical subspecies has been collected in E. Campania (Savignano Irpino).

S. (S.) ernae Fauer 1978:
Like S. paestana, but peristome detached; lunella less arched; posterior lower palatal plica tending to be separated; basalis partly present.
Distribution: Campania (Monti Alburni).

S. (S.) piceata (Rossmässler 1836):
Lunella not reduced above; posterior lower palatal plica in part separated from lunella; basalis short or missing; anterior upper palatal plica in part present, separated from upper palatal plica; clausilium plate entire.
In contrast to S. paestana the shell of S. piceata is yellowish-brown, with indistinct sutural papillae.
Distribution: Central peninsular Italy (from Marche to N. Lazio and Abruzzo).
Synonyms: C. adaucta A. & G. Villa 1841, C. piceata f. gularis Küster 1876.

Determination key

1. Peristome detached and protruding: S. ernae.
2. Peristome adnate:
....1. Anterior lower palatal plica (basalis) short or missing:
........1. Shell with indistinct sutural papillae; posterior lower palatal plica in part separated from lunella: S. piceata.
........2. Shell with distinct sutural papillae:
............1. Posterior lower palatal plica not separated from lunella: S. p. paestana.
............2. Posterior lower palatal plica ± separated from lunella: S. kobeltiana.
....2. Anterior lower palatal plica (basalis) ± long:
........1. Clausilium plate distally ± groove-like, outer corner ± upbent:
............1. Inferior lamella ending with two branches: S. gibbula honii.
............2. Inferior lamella ending with only one branch: S. g. gibbula s. l..
........2. Clausilium plate distally not groove-like, outer corner not upbent:
............1. Lunella not reduced above; clausilium plate not emarginate:
................1. Subcolumellar lamella not emerging: S. incerta.
................2. Subcolumellar lamella mostly emerging: S. paestana intustructa.
............2. Lunella ± reduced above; clausilium plate ± emarginate:
................1. Anterior upper palatal plica, if present, elongation of upper palatal plica: S. vulcanica.
................2. Anterior upper palatal plica, if present, displaced from upper palatal plica: S. stigmatica sturmii.

Distributional relations (see map 1)

The Siciliaria species of central and southern Apennine peninsula and adjacent Sicily have, as a rule, allopatric ranges, as can be expected for closely related species. Only S. (S.) piceata and S. (G.) gibbula have common ranges with other species. The range of S. (S.) piceata overlaps that of S. (S.) paestana in central Lazio (e. g. Tivoli, SMF). S. (G.) gibbula and S. (S.) piceata have a common range in Marche and Abruzzo. The ranges of S. (G.) gibbula and S. (S.) paestana overlap in Basilicata (Forcart 1965, SMF), that of S. (G.) gibbula (honii) and S. (S.) stigmatica in Penisola Salentina (own collection). S. (G.) gibbula (honii) and S. (S.) paestana occur together at M. Circeo, Lazio (own collection) and on the island of Ischia (SMF). S. (G.) gibbula and S. (S.) paestana have been collected in Montella, Campania, S. (G.) gibbula and S. (S.) kobeltiana in Cosenza, Calabria (according to Degner 1927), but in both cases S. (G.) gibbula has not been collected in the surroundings of these localities..

References

Boettger, O. (1879): Gattung Clausilia Drap. – In: Rossmässler, Icon. Land- und Süsswasser-Mollusken, (1) 6 (4/6): 52-153, pls. 167-178. Wiesbaden.

Degner, E. (1927): Zur Molluskenfauna Unteritaliens. – Mitt. Zool. Staatsinst. Zool. Mus. Hamburg, 43: 39-124.

Forcart, L. (1965): Rezente Land- und Süsswassermollusken der süditalienischen Landschaften Apulien, Basilicata und Calabrien. – Verh. naturf. Ges. Basel, 78 (1): 59-184.

Giusti, F. (1973): Notulae Malacologicae. XVIII. I Molluschi terrestri e salmastri delle Isole Eolie. – Lav. Soc. Ital. Biogeogr., (NS) 3: 113-306, 16 pls.

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.

Nordsieck, H. (1969): Zur Anatomie und Systematik der Clausilien, VI. Genitalsystem und Systematik der Clausiliidae, besonders der Unterfamilie Alopiinae. – Arch. Molluskenkunde, 99 (5/6): 247-265.

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

– – – (2002): Contributions to the knowledge of the Delimini (Gastropoda: Stylommatophora: Clausiliidae). – Mitt. dtsch. malakozool. Ges., 67: 27-39.

Schmidt, A. (1868): System der europäischen Clausilien und ihrer nächsten Verwandten: 175 pp., 1 tab. Cassel.

Wagner, A. J. (1924): Systematisches Verzeichnis der mir heute bekannten Arten und Formen der Clausiliidae. III. – Ann. zool. Mus. Polon. Hist. nat., 3 (3/4): 99-126.

– – – (1925): Studien über die Systematik, Stammesgeschichte und geographische Verbreitung des Genus Delima (Hartmann) A. J. Wagner. – Ann. zool. Mus. Polon. Hist. nat., 4 (1): 1-73, 16 pls.

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