Systematics of the Baleinae (Clausiliidae) of Europe (except Caucasian region)

Hartmut Nordsieck (IX.2016)

I. Introduction

For the first time, Baleinae have been separated by Steenberg (1914: 43) from the Clausiliinae (= IIième Groupe) as IIIième Groupe, because of the structure of their male copulatory organs. Later on, Nordsieck (1969: 248) classified them as subfamily Baleinae A. J. Wagner 1913, a name which originally was given to the Clausiliinae on the whole. The subfamily was defined by its transformed male copulatory organs which are unique within the Clausiliidae (Nordsieck 1969: fig. 4; 2007a: chapter I: fig. 10, chapter VIII: fig. 3) and are therefore evaluated as the essential autapomorphy of the group (2007a: chapter I: 15-16).
In the molecular phylogeny of the Clausiliidae, presented by Uit de Weerd & Gittenberger (2013: tree fig. 3), the Baleinae came out as a group within the Clausiliinae sensu lato, paraphyletic with respect to the Clausiliinae sensu stricto. This result, however, has only a limited validity, because the number of examined species (5) was too low (Nordsieck 2013). In other trees (mtDNA 16S, Hausdorf unpubl., written communication), which are based on a greater number of species, the Baleinae appear as a clade.
In the following, a system of the Baleinae is proposed, based on shell and genital characters. It is coordinated with the available molecular trees.

II. Systematics of non-Caucasian genera

Genera:

In the following, only genera of non-Caucasian Europe are treated. They are distributed nearly over the whole of Europe, but their distribution center is Pontian Europe (Stara planina, Rhodopes and E. Dinarids, within the political frontiers of Bulgaria, Serbia, Macedonia and Greece). The genera are defined by shell and genital characters, and their distribution and the species included are given.

Shell characters which have revealed tü be essential for genus diagnoses are the following (terms see Nordsieck 2007: Appendix 2: fig. 1):
Basal keel (prominence); upper lamellae (relation of superior lamella and spiral lamella), subcolumellar lamella (emergence), anterior palatal plicae (presence and relation to lunellar), clausilium plate (shape, Nordsieck 1973: figs. 5-16).
The anterior lower palatal plica is named basalis.

Genital characters which have revealed to be essential for genus diagnoses are the following (terms see Nordsieck 2007: Appendix 2: fig. 4):
Vagina (form); male copulatory organs (form of penis, development of proximal penis and penial ligaments, Steenberg 1914: figs. 22-25; Nordsieck 1973: figs. 17-28, 1977: figs. 5-6, 2007b: figs.1-2).
The terms proximal and distal are used as seen from the gonad. The species the genitalia of which have been examined by me, are marked by an asterisk*.

Laciniaria genus group

In this group Mentissa, Mentissella, Laciniaria, Alinda, and Pseudalinda are assembled.
Shell: Spiral lamella ending less deeply than inferior lamella (Fig. 4). Clausilium plate distally bent (except in Pseudalinda) (Fig. 1).

Mentissa H. & A. Adams 1855

Shell: Basal keel prominent; superior lamella separated from spiral lamella; subcolumellar lamella not emerging; upper palatal plica separated from lunella, elongated as anterior upper palatal plica, basalis connected with lunella; clausilium plate emarginate.
Genitalia: Vagina proximally widened; penis ± cylindrical, proximal part delimited, penial ligaments present.
Distribution: Crimea.
Species: canalifera (Rossmässler 1836)*, gracilicosta (Rossmässler 1836), velutina Baidashnikov 1990.

Mentissella H. Nordsieck 1973

Shell: Basal keel prominent; superior lamella separated from spiral lamella; subcolumellar lamella not emerging; upper palatal plica separated from lunella, elongated as anterior upper palatal plica, basalis connected with lunella; clausilium plate simple (neither emarginate nor narrowed).
Genitalia: Like Mentissa; proximal part of penis large.
Distribution: E. Stara planina.
Species: rebeli (Sturany 1897)*.

Laciniaria Hartmann 1842

Shell: Basal keel mostly prominent; often with peristome folds; superior lamella separated from spiral lamella; subcolumellar lamella mostly not emerging (if not elongated by a peristome fold); upper palatal plica connected with lunella, anterior upper palatal plica, if present, separated from upper palatal plica, basalis short to missing; clausilium plate simple (neither emarginate nor narrowed).
Genitalia: Vagina proximally widened; penis cylindrical, proximal part weakly or not delimited, penial ligaments present or missing.
Distribution (Nordsieck 2008): Carpathians, Stara planina, Rhodopes to Turkish Thracia, S. and E. Dinarids to northernmost Greece, one species (L. plicata) farly distributed in central and E. Europe.
Species: bajula (A. Schmidt 1868)*, exalta (Westerlund 1878), macilenta (Rossmässler 1842)*, plicata (Draparnaud 1801)*.

Alinda H. & A. Adams 1855

Shell: Basal keel mostly prominent; superior lamella separated from spiral lamella; subcolumellar lamella not emerging; upper palatal plica connected with lunella, anterior upper palatal plica, if present, connected with upper palatal plica or separated from it, basalis short to missing; clausilium plate simple (neither emarginate nor narrowed).
Genitalia: Vagina distally widened; penis proximally widened, proximal part delimited, penial ligaments present.
Distribution (Nordsieck 2008): Stara planina, Rhodopes to N. E. Greece, Dinarids to central Albania and N. Greece, W. Carpathians, one species (A. biplicata) farly distributed in central and W. Europe.
Species:atanasovi (Urbanski 1964)*, biplicata (Montagu 1803)*, elegantissima A. J. Wagner 1914*, vratzatica (Likharev 1972), wagneri (A. J. Wagner 1911)*.

Pseudalinda O. Boettger 1877

Shell: Basal keel not prominent; superior lamella separated from spiral lamella or connected with it; subcolumellar lamella emerging; upper palatal plica connected with lunella, the latter occasionally separated and weakened, other palatal plicae missing; clausilium plate distally not bent, simple (neither emarginate nor narrowed).
Genitalia: Vagina distally widened; penis proximally widened, proximal part delimited, in part weakly, penial ligaments present.
Distribution: Carpathians, S. and E. Dinarids to northwesternmost Greece and Macedonia.
Species:fallax (Rossmässler 1836)*, golesnicensis (A. J. Wagner 1914), jugularis (Vest 1859)*, nordsiecki Dedov & Neubert 2002, stabilis (L. Pfeiffer 1847)*, viridana (Rossmässler 1836)*.


Vestia Hesse 1916

Shell: Basal keel prominent or not; superior lamella continuous with spiral lamella, in part separated from it; subcolumellar lamella mostly not emerging; upper palatal plica connected with lunella, anterior upper palatal plica missing, basalis mostly missing; clausilium plate distally much narrowed (with hook-like end); spiral lamella ending more deeply than or as deeply as inferior lamella (fig. 2).
Genitalia: Vagina distally widened; penis proximally widened, compact, proximal part delimited, penial ligaments present, short.
Distribution: Carpathians and Sudetes, S. and E. Dinarids to Macedonia, in some species (V. turgida, V. elata) Carpathian distribution continued in Czechia with adjacent Bavaria and S. Poland, one species (V. ranojevici) also in Stara planina and Rhodopes (Rila mountains).

(Vestia)

Species: elata (Rossmässler 1836)*, gulo (Bielz 1859)*, turgida (Rossmässler 1836)*.

(Brabenecia) H. Nordsieck 1974

Shell: Like V. (Vestia), but streaked; basal keel more prominent; subcolumellar lamella emerging.
Genitalia: Like V. (Vestia), but vagina proximally widened; penial ligaments of normal length.
Species: ranojevici (Pavlović 1912)*.

(Vestiella) H. Nordsieck 1977

Shell: Like V. (Vestia), but basal keel prominent; in part peristome folds present; subcolumellar lamella emerging; in part basalis present.
Genitalia: Like V. (Vestia), but proximal part of penis indistinct, penial ligaments of normal length.
Species: lazarovii Dedov 2012, roschitzi (Brancsik 1890)*.

(Pavlovicia) H. Nordsieck 1973

Shell: Like V. (Vestia), but basal keel prominent; subcolumellar lamella in part emerging; basalis separated, connected with palatal callus.
Genitalia: Like V. (Vestia), but no proximal part of penis discernible, penial ligaments of normal length.
Note: Pavlovicia is a subgroup of Vestia, not of Bulgarica, as formerly proposed (Nordsieck 1973: 192). This is proved by the following shell characters:
1. spiral lamella penetrating more deeply than inferior lamella,
2. superior lamella continuous with spiral lamella,
3. clausilium plate distally finger-like narrowed.
Species: pavlovici (H. Nordsieck 1972)*.

Bulgarica O. Boettger 1877

Shell: Basal keel prominent; superior lamella separated from spiral lamella; subcolumellar lamella emerging or not; upper palatal plica connected with lunella, rarely not, anterior upper palatal plica, if present, separated from upper palatal plica or connected with it, basalis connected with lunella or separated from it; clausilium plate distally narrowed, in part with outer corner; spiral lamella ending as deeply as or less deeply than inferior lamella (fig.3).
Genitalia: Vagina cylindrical or distally widened; penis cylindrical or proximally widened, no proximal part discernible, penial ligaments present or indistinct.
Distribution: Dinarids, Carpathians, Dobrogea, Stara planina, Rhodopes, some species (B. cana, B. vetusta) distributed in central and E. Europe, other ones (B. denticulata and relatives) in N. and central Greece, Aegean islands and westernmost Turkey.

(Strigilecula) Kennard & Woodward 1923

Shell: No peristome folds; subcolumellar lamella elongated along basal furrow; clausilium plate relatively narrow, outer margin distally less or not upbent.
Genitalia: Penial ligaments of normal length.
Species: cana (Held 1836)*, pindica H. Nordsieck 1974, vetusta (Rossmässler 1836)*.

(Bulgarica)

Shell: In part with peristome folds; subcolumellar lamella truncate, not elongated along basal furrow (if not by a peristome fold); clausilium plate distally narrowed, outer margin more or less upbent, in part with outer corner.
Genitalia: Penial ligaments mostly short, very short or indistinct.
Species: bulgariensis (L. Pfeiffer 1848), denticulata (Olivier 1801)*, erberi (Frauenfeld 1867), fraudigera (Rossmässler 1839), fritillaria (Frivaldsky 1835)*, hemmenorum H. Nordsieck 2015, hiltrudae H. Nordsieck 1974, iniucunda (Brandt 1962)*, pseudofraudigera H. Nordsieck 1973, rugicollis (Rossmässler 1836)*, serbica (Moellendorff 1873)*, stolii (L. Pfeiffer 1859), urbanskii H. Nordsieck 1973*, varnensis (L. Pfeiffer 1848)*.

Balea Gray 1824

Shell: Basal keel not prominent; closing apparatus = CA reduced: superior lamella rudimentary, no other lamellae or plicae, no clausilium plate.
Genitalia: Vagina cylindrical; penis proximally widened, with penial papilla, penial ligaments present.
Distribution: Stara planina, Rhodopes (n. subgen.), one species (B. perversa) farly distributed in Europe (except E. and southernmost Europe), other ones (other species of B. (Balea)) coastal westernmost Europe, Atlantic islands, Tristan d’Acunha and Gough Island (Tristania species).
Note: The shell characters of Balea are without value for genus systematics, because the CA is reduced. Species taxa with reduced CA which are similar to Balea are also present in other genera of the Baleinae (e. g., Alinda, Nordsieck 2008: 139).

(Balea)

Species: nitida Mousson 1858, perversa (Linné 1758)*, sarsii (L. Pfeiffer 1847)*, Tristania species (see Preece & Gittenberger 2003).

(n. subgen.)

Shell: Like B. (Balea), but peristome detached; rudiment of superior lamella well-developed.
Genitalia: Like B. (Balea), but bursa copulatrix longer; penis longer, with larger penial papilla.
Species: eninskoensis Irikov 2006*, kaeufeli (Brandt 1962)*, pancici (Pavlović 1912)?


Figs. 1-5. Shells of Baleinae species, opened to show inner parts of closing apparatus; phot. S. Hof.
Specimens from collection Nordsieck. Actual shell height = H (mm).
Fig. 1. Alinda biplicata (body whorl, palatal wall removed); H = 16.5.
Abbreviations: cp = clausilium plate, cs = clausilium stalk, il = inferior lamella, sc = subcolumellar lamella, sl = superior lamella, sp = spiral lamella.
Figs. 2-4. Baleinae species (from above to below shell frontal, body whorl dorsal, opened to show inner lamellae endings, view from the side and from below):
Fig. 2. Vestia elata, sp ending more deeply than il; H = 16.0.
Fig. 3. Bulgarica vetusta, sp ending as deeply as or less deeply than il; H = 15.5.
Fig. 4. Laciniaria plicata, sp ending less deeply than il; H = 15.2.  

In this article, Caucasian Baleinae are not treated. For comparison with the non-Caucasian genera, however, some important shell characters are given.

Caucasian genera:

Spiral lamella ending more deeply than inferior lamella.

Quadriplicata O. Boettger 1878, Mucronaria O. Boettger 1877: Superior lamella continuous with or separated from spiral lamella; instead of lunella palatal plicae present.

Micropontica O. Boettger 1881: Superior lamella continuous with spiral lamella (if present); lunella present (occasionally reduced).

Mode of reproduction of some non-Caucasian Baleinae (Sulikowska et al. 2014):

Laciniaria. plicata, Vestia ranojevici, Pseudalinda stabilis are oviparous;
Vestia elata, V. gulo, Pseudalinda fallax have egg retention;
Alinda biplicata, Vestia turgida, Balea perversa are (ovo)viviparous.

The apomorphic modes of reproduction (egg retention, ovoviviparity) have obviously evolved convergently. Thus, they cannot be used for genus systematics.

Consideration of DNA analyses and phylogenetic remarks:

In the trees of Van Moorsel (2001: chapter 3: figs. 4-5) Balea = Alinda and Bulgarica come out as only distantly related; this confirms the evaluation of Bulgarica as a genus of its own (Nordsieck 1973: 191). In the tree of Uit de Weerd & Gittenberger (2013: fig. 3) Vestia and Balea form a clade different from the clade including Laciniaria and Alinda. This means that the first two genera are only distantly related to the two latter ones which are more closely related. The separation of Balea and Alinda as independent genera (in contrast to Nordsieck 1977: 98-99) is thus confirmed (Nordsieck 2007b). In the trees of Hausdorf (unpubl.) these relationships are visible again. Vestia, Bulgarica and Balea appear as only distantly related to the genera of the Laciniaria group (Mentissa, Laciniaria, Alinda, Pseudalinda) which are assembled in a clade. Thus, also the separation of Vestia and Pseudalinda as independent genera (Nordsieck 1973: 188-189) is confirmed.
In the following the resulting system is given:
Vestia;
Bulgarica;
Balea;
Laciniaria group:
Mentissa (+ Mentissella?);
Laciniaria;
Alinda;
Pseudalinda.
Vestia, Bulgarica and Balea have an isolated position each; they are characterized by special morphological characters (Vestia shell, Bulgarica and Balea genitalia, see diagnoses). They are basal to the genera of the Laciniaria group, which have apomorphies of the shell (see diagnosis) and may represent a monophyletic group. Vestia may have the most basal position, because in contrast to the other genera it has the more deeply ending spiral lamella. This may be a plesiomorphic character, because it is also found in the Caucasian Baleinae (see diagnoses).
The distributional patterns of the European Baleinae are not helpful for the reconstruction of phylogeny, because all groups (except Mentissa) have their centres of distribution in the southern and southeastern part of S. E. Europe (Stara planina, Rhodopes, eastern part of S. Dinarids, see distributions). Only the centres of distribution of Pseudalinda and Vestia are situated somewhat more northerly (Carpathians, E. Dinarids). The only case of vicariance is that of Mentissa (Crimea) and remaining genera of the Laciniaria group (Stara planina, Rhodopes).


References

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

Nordsieck, H. (1973): Zur Anatomie und Systematik der Clausilien, XIII. Neue Balkan-Formen der Mentissoideinae und Baleinae (mit taxonomischer Revision der zugehörigen Gruppen). – Archiv für Molluskenkunde, 103 (4/6): 179-208, Taf. 6-7, 7a.

Nordsieck, H. (1977): Zur Anatomie und Systematik der Clausilien, XVIII. Neue Taxa rezenter Clausilien. – Archiv für Molluskenkunde, 108 (1/3): 73-107, Taf. 3-5.

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

Nordsieck, H. (2007b): Balea Gray 1824 and Alinda H. & A. Adams 1855 are separated as genera (Gastropoda: Stylommatophora: Clausiliidae). – Mitteilungen der deutschen malakozoologischen Gesellschaft, 77/78: 27-30.

Nordsieck, H. (2008): Alinda biplicata (Montagu) and Laciniaria plicata (Draparnaud), diversity in comparison, with the description of new subspecies (Gastropoda: Stylommatophora: Clausiliidae). – Archiv für Molluskenkunde, 137 (2): 133-157, 5 pls..

Nordsieck, H. (2013): Comment on Uit de Weerd & Gittenberger (2013): Phylogeny of Clausiliidae. – www.hnords.de.

Preece, R. C. & Gittenberger, E. (2003): Systematics, distribution and ecology of Balea (= Tristania) (Pulmonata: Clausiliidae) in the islands of the Tristan-Gough group. – Journal of Molluscan Studies, 69: 329-348.

Steenberg, C. M. (1914): Anatomie des Clausilies danoises. I. Les organes génitaux. – Mindeskrift for Japetus Steenstrup, 29: 46 pp., 1 pl.

Sulikowska-Drozd, A., Walczak, H. & Binkowski, M. (2014): Evolution of shell apertural barriers in viviparous land snails (Gastropoda: Pulmonata: Clausiliidae). – Canadian Journal of Zoology, 92: 205-213.

Uit de Weerd, D. R. & Gittenberger, E. (2013): Phylogeny of the land snail family Clausiliidae (Gastropoda: Pulmonata). – Molecular Phylogenetics and Evolution, 67: 201-216.

Van Moorsel, C. H. M. (2001): Molecular phylogenetics of a speciose group: Albinaria and the search for homology. – Thesis Leiden University: 120 pp.


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