Alopia species from Dâmboviţa valley system, Romania (Gastropoda, Stylommatophora, Clausiliidae), an example of high diversity

Hartmut Nordsieck (XII.2015)

Introduction

In the Dâmboviţa valley system, Judeţul Argeş, Romania, the genus Alopia exhibits a high diversity which is exemplary for the whole genus. The region is characterized by several gorges (chei(le)s), which were cut by the Dâmboviţa river and its tributaries into the Jurassic limestone of the southern slope of the Piatra Craiului Mountains, and offers thus favorable habitats for Alopia species. In 1975 and 1978 I collected Alopia species at twenty localities of the region and visited it again in 2014 and 2015 to complete these collections at eighteen further localities. In this article the results of the examination of the collected material are presented.

History

The Dâmboviţa valley region which until World War I was part of the Romanian Old Kingdom (Wallachia) has not been explored by the Transylvanian malacologists of the nineteenth century, with the exception of the Dâmbovicioara valley from which Clausilia elegans and varieties were described (Bielz 1852, 1867, Rossmässler 1858, M. Kimakowicz 1894). The other parts of the region were explored by R. Kimakowicz until World War II; as result he (1962) described Alopia magnifica with two subspecies. Since the fifties of the past century new efforts were made to explore the Alopia fauna of the region (results in Grossu 1981, Szekeres 2007, Nordsieck 2008). More exact information on the distribution of the taxa found until then was given in Nordsieck (2008: 9-10). In the same paper (: 16) the only known case of syntopy of left- and right-coiled Alopia species in the region was listed. This case was the reason for the description of the first Alopia taxon based only on mtDNA analysis (Deli & Szekeres 2011). In the molecular phylogeny of Fehér et al. (2013), which actually is a phylogeny of the COI gene (comment Nordsieck 2013b), all examined Alopia taxa of the region are assembled in one and the same clade (D2, named A. glorifica).

Results

The region under consideration is inhabited by three Alopia species, with altogether six subspecies and several geographical forms. In the following the taxa are defined and their occurrences (collecting sites see map) are given. Revisory notes are added.

A. glorifica (Charpentier 1852)
Peristome always detached; body whorl before peristome distinctly ribbed.

A. g. magnifica R. Kimakowicz 1962 (Figs. 1-2)
Mostly darkly coloured; weakly sculptured, also upper whorls; superior lamella not reaching or reaching spiral lamella; inferior lamella high; palatal plicae long to short; 1-2 middle palatal plicae, occasionally more; clausilium plate leaving no or more gaps, outer lobe protruding.
Lower parts of Valea Dâmboviţei, Valea Cheii and Valea Ghîmbavului.
The samples from different gorges or parts of gorges differ by the more or less weakened clausilial apparatus.
Form clessini R. Kimakowicz 1962, Valea Cheii (Fig. 3): More strongly sculptured; outer lobe of clausilium plate rarely not protruding.
Form transitional to A. g. elegantissima, upper Valea Cheii (Fig. 4): sculpture stronger; palatal plicae short, middle palatal plica lunella-like.
Notes 1, 2.

A. g. elegantissima H. Nordsieck 1977 (Figs. 5-6)
= C. elegans Bielz 1852 [non Cantraine]
Brightly coloured; distantly ribbed; superior lamella not reaching or reaching spiral lamella; inferior lamella mostly high; palatal plicae moderately long to short, middle palatal plica mostly lunella-like; clausilium plate leaving more gaps, outer lobe protruding.
Cheile Dâmbovicioarei, Valea Crovului.
Form from Valea Crovului on average more distantly ribbed.
Form transitional to A. g. magnifica in Valea Crovului above mouth.

A. g. intercedens (A. Schmidt 1857) (Figs. 7-8)
Like A. g. elegantissima, but darkly coloured; more weakly sculptured.
Form cerasina A. Schmidt 1857 sensu Bielz (Fig. 7): Ribbed (more weakly and densely than A. g. elegantissima); palatal plicae short; outer lobe of clausilium plate in part not protruding.
Valea Dâmbovicioarei between Dâmbovicioara and below Cabana Brusturetului.
Form intercedens sensu Bielz (Fig. 8): Weakly sculptured, upper whorls rib-striated; palatal plicae on average longer.
Valea Dâmbovicioarei above mouth of Valea Muierii and immediately above Cabana Brusturetului.
Note 3.

A. g. glorifica (Fig. 9)
Darkly coloured; weakly and irregularly ribbed, upper whorls more distinctly; superior lamella not reaching spiral lamella; inferior lamella high to moderately high; palatal plicae short, middle palatal plica lunella-like or missing; clausilium plate leaving more gaps, outer lobe protruding or not.
Valea Dâmbovicioarei above Cabana Brusturetului.
Note 4.

A. lischkeana (Charpentier 1852)
Right-coiled, otherwise like A. glorifica.

A. l. lischkeana (deceptans Deli & Szekeres 2011) (Fig. 10)
Right-coiled, otherwise like A. g. glorifica (= enantiomorph of g. glorifica). On average more densely ribbed than the syntopically occurring enantiomorph.
Valea Dâmbovicioarei above Cabana Brusturetului.
Form from immediately above Cabana Brusturetului: More weakly sculptured; palatal plicae on average longer; clausilium plate less leaving gaps, outer lobe mostly protruding (= enantiomorph of g. intercedens sensu Bielz).
Notes 5, 6.

A. plumbea (Rossmässler 1839)
Peristome in part adnate; body whorl before peristome densely ribbed.

A. p. plumbea (Rudăriţa form) (Fig. 11)
Darkly coloured; weakly sculptured, also upper whorls; superior lamella mostly not reaching spiral lamella; inferior lamella mostly high; palatal plicae moderately long, 1-3 middle palatal plicae; clausilium plate leaving more gaps, outer lobe mostly protruding.
Valea Rudăriţa.
Note 7.

Notes

1. A. magnifica clessini sensu R. Kimakowicz (1962: 513) contains two different subspecies. Therefore, in Nordsieck 1977 (: 75) the name clessini was restricted to the magnifica form from Cheile Cheiţa Cheii (Fig. 3) which R. Kimakowicz had figured (1962: fig. 2). The other form from Valea Crovului (Fig. 6) belongs to A. g. elegantissima. It is of interest that R. Kimakowicz had formerly the intention to describe both forms as separate subspecies (on labels in SMF, material deposited in 1956).

2. Clausilia regalis var. valachiensis „Bielz“ O. Boettger (1879: 58, pl. 168, fig. 1691) has been identified by Szekeres (2007: 10, figs. 5-6) with a form of A. g. magnifica from upper Valea Cheii opposite to the mouth of Valea Crovului (see map). The examination of the type series (SMF 59233), however, had the result that this identification is erroneous, because C. regalis var. valachiensis is near to A. g. glorifica (see diagnoses). The type locality is „Koltzu Teleje“, not Valea Teleajen, as formerly given by Szekeres (1969: 30). Thus, the name A. magnifica is not threatened by the older name A. valachiensis.

3. A. Schmidt (1857: 78) described Clausilia elegans var. cerasina (defined by „subtilius striata“) and C. intercedens (diagnosis in comparison with C. regalis). Rossmässler (1858: 124-125) degraded C. intercedens (following a written communication of A. Schmidt) to a variety of C. elegans (: pl. 86, fig. 956) and mentioned a brighter coloured transitional form as cerasina. Both forms were said to occur in „Dumbrowitzoare“. The examination of both (intercedens SMF 59318, cerasina 59297) revealed that they do not correspond to the descriptions of these varieties given by Bielz (1867: 117). C. e. var. intercedens sensu Rossmässler is a weakly sculptered form of C. e. var. cerasina sensu Bielz, while C. e. var. cerasina sensu Rossmässler is not much different from the type form of C. elegans. Besides the weakly sculptured specimens the original sample of C. intercedens sensu Rossmässler contains also a ribbed specimen = true cerasina.
As my collections have shown, the varieties as described by Bielz occur in different parts of the Valea Dâmbovicioarei between Dâmbovicioara and immediately above Cabana Brusturetului (see map), the ribbed cerasina sensu Bielz (Fig. 7) in nearly the whole valley and the larger smooth intercedens sensu Bielz (Fig. 8) in a part of the upper valley. Above the mouth of Valea Muierii, they occur in immediate neighbourhood, as far as known, without transitions. Both are so different (see diagnoses and figures) that they should be treated as subspecies, but because of the nomenclatural problems it is difficult to name them correctly.

4. The glorifica form of the Valea Dâmbovicioarei farther above Cabana Brusturetului (Fig. 9) is not A. g. intercedens, as determined by Deli & Szekeres (2011) and Fehér et al. (2013), but A. g. glorifica. This was already realized by Grossu (1981: 67-68), but his judgement was brought to nothing by his faulty figure of the distribution (: fig. 31, „ g. intercedens“ = g. glorifica, „g. elegantissima“ = g. intercedens). A. g. glorifica does not only occur in the Valea Dâmbovicioarei „farther upstream“ from A. l. lischkeana (deceptans), as asserted by Deli & Szekeres (2011), but in the whole valley from somewhat above Cabana Brusturetului; at some localities it occurs syntopically with the enantiomorph A. l. lischkeana. (Nordsieck 2008: 16, Deli & Szekeres 2011: 19; see map).

5. The lischkeana form of the Valea Dâmbovicioarei farther above Cabana Brusturetului (Fig. 10) has been described by Deli & Szekeres (2011: 19, figs. 1) as a new subspecies of A. glorifica, A. g. deceptans, because its COI gene is identical with that of the enantiomorph A. g. intercedens sensu Deli & Szekeres = A. g. glorifica. (Fehér et al. 2013: 266). Doing so, it was not considered that the identity could be caused by hybridization, which can be expected because of the syntopy of both species (see Nordsieck 2012). Because the lischkeana form from Valea Dâmbovicioarei does not differ from
A. l. lischkeana from other parts of the Piatra Craiului Mountains, it is not separated from it as a subspecies (the COI identity with the enantiomorph
glorifica is indicated in brackets as deceptans). A. l. lischkeana occurs only in the lower part of the Valea Dâmbovicioarei above Cabana Brusturetului, at some localities syntopically with the enantiomorph A. g. glorifica (see note 4 and map), while the upper part of the valley, as far as I could state, is inhabited by A. g. glorifica alone.

6. Immediately above Cabana Brusturetului a lischkeana form has been collected, which is the enantiomorph of A. g. intercedens sensu Bielz, which also occurs there (see map). Until further notice, it is included in A. l. lischkeana.

7. In a tributary valley of Valea Cheii, Valea Rudăriţa, I discovered a form of A. plumbea (Rudăriţa form, Fig. 11) which was unknown until now (it is not mentioned by any of the former authors). It is similar to the neighbouring nominate subspecies of A. plumbea, but differs from it by the more weakened closing apparatus (inferior lamella in part moderately high; palatal plicae moderately long; clausilium plate leaving more gaps). Further examination will clear up the status within the species.

Recommendation

In both last years I could state that compared with the seventies of the past century at several localities Alopia species had diminished in number; in some localities, especially in those far from brooks, I found several specimens fresh dead by drought. This may be due to the climate change, as already discussed for other clausiliid groups. Because many Alopia forms have restricted occurrences, it is urgently recommended not to collect such a large amount of specimens as usual until now (also by myself in the seventies, but especially by other collectors in the past years, see the respective literature). As already recommended in a former paper (Nordsieck 2013a), only few specimens should be collected and only for scientific purposes, not for supplementing collections or even for selling.


Figs. 1-11. Alopia species from Dâmboviţa valley system.
Frontal and body whorl dorsal; phot. S. Hof.
Abbreviations: collection Nordsieck = N, collection of Forschungsinstitut Senckenberg = SMF, shell width = D (mm), shell height = H (mm). 
Fig. 1. Alopia glorifica magnifica, Valea Dâmboviţei, Cheile Plaiul Mare 3 km towards Săticu de Jos (right side), ex SMF 345851; H/D 18.1 / 5.0.
Fig. 2. Alopia g. magnifica, Valea Ghâmbavului (big spring), ex SMF 155737, H/D 19.1 / 4.6.
Fig. 3. Alopia g. magnifica (clessini), Valea Cheii, Cheile Cheiţa Cheii (right side), ex SMF 345850, H/D 17.5 / 4.45.
Fig. 4. Alopia g. magnifica >< elegantissima, Valea Cheii, upper part above mouth of Valea Crovului (right side), ex SMF 345831, H/D 16.9 / 4.1.  
Fig. 5. Alopia glorifica elegantissima, Cheile Dâmbovicioarei (upper part, left side), ex SMF 330863, H/D 14.8 / 3.9.
Fig. 6. Alopia g. elegantissima, Valea Crovului (right side), ex SMF 345847, H/D 15.55 / 3.8.
Fig. 7. Alopia glorifica intercedens (cerasina), Valea Brusturetului 1 km above Peştera Dâmbovicioara (right side), ex SMF 345818, H/D 15.2 / 3.8.
Fig. 8. Alopia g. intercedens (intercedens), Valea Brusturetului 2 km above Peştera Dâmbovicioara (right side), ex SMF 345805, H/D 20.1 / 4.7.  
Fig. 9. Alopia g. glorifica, Valea Brusturetului above Cabana Brusturet (Valea Seacă a pietrelor, right side, ca. 1050 m), ex SMF 345811, H/D 15.1 / 4.2.
Fig. 10. Alopia l. lischkeana (deceptans), Valea Brusturetului above Cabana Brusturet (Valea Seacă a pietrelor, right side, ca. 1100 m), ex SMF 345814, H/D 15.6 / 4.0.
Fig. 11. Alopia p. plumbea (Rudăriţa form), Valea Rudăriţa (right side, ca. 1000 m), ex SMF 345829, H/D 19.7 / 5.3.  

References

Bielz, E. A. (1852): Zwei neue Schliessmundschnecken. – Verhandlungen und Mittheilungen des siebenbürgischen Vereins für Naturwissenschaften in Hermannstadt, 3: 31-32.

Bielz, E. A. (1867): Fauna der Land- und Süsswasser-Mollusken Siebenbürgens. 2. Aufl. – VIII + 216 pp. Hermannstadt.

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

Deli, T. & Szekeres, M. (2011): Two new dextral subspecies of Alopia H. & A. Adams 1855 (Gastropoda: Clausiliidae). – Malakológiai Tájékoztató [Malacological Newsletter], 29: 19-23.

Fehér, Z., Németh, L., Nicoară, A. & Szekeres, M. (2013): Molecular phylogeny of the land snail genus Alopia (Gastropoda: Clausiliidae) reveals multiple inversions of chirality. – Zoological Journal of the Linnean Society, 167: 259-272.

Grossu, A. V. (1981): Gastropoda Romaniae Ordo Stylommatophora 3 Suprafamiliile Clausiliacea şi Achatinacea: 269 pp. Universitatea din Bucureşti Facultatea de Biologie. Bucureşti.

Kimakowicz, M. von (1894): Prodromus zu einer Monographie des Clausilia-Subgenus Alopia H. & A. Adams. Beitrag zur Mollusken-Fauna Siebenbürgens, III. Nachtrag. – Verhandlungen und Mittheilungen des siebenbürgischen Vereins für Naturwissenschaften in Hermannstadt, 43: 19-58.

Kimakowicz, R. von (1962): Zwei neue Alopien aus Rumänien. – Travaux du Muséum d`Histoire Naturelle „Grigore Antipa“, 3: 513-515.

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

Nordsieck, H. (2008): The system of the genus Alopia H. & A. Adams 1855 (Gastropoda: Stylommatophora: Clausiliidae). – Mitteilungen der Deutschen Malakozoologischen Gesellschaft, 79/80: 7-18.

Nordsieck, H. (2012): Interspecies hybridization in the genus Alopia (Gastropoda, Stylommatophora, Clausiliidae). – www.hnords.de.

Nordsieck, H. (2013a): A new species of the genus Alopia H. & A. Adams (Gastropoda, Pulmonata, Clausiliidae) from Lotru valley, southern Carpathians, Romania. – Mitteilungen der Deutschen Malakozoologischen Gesellschaft, 89: 1-6.

Nordsieck, H. (2013b): Comment on Fehér et al. (2013): Molecular phylogeny of Alopia. – www.hnords.de.

Rossmässler, E. A. (1835-1858): Iconographie der Land- und Süsswasser-Mollusken, mit vorzüglicher Berücksichtigung der europäischen noch nicht abgebildeten Arten. – (1) 3 (5/6): I-VIII, 81-140, pls. 81-90 [1858]. Leipzig (Costenoble).

Schmidt, A. (1857): Ueber die Baleen und baleaartigen Clausilien Siebenbürgens. – Giebel`s Zeitschrift für Naturwissenschaften, 8: 407-413.

Szekeres, M. (1969): Neue Angaben zur Kenntnis der Alopien (Clausiliidae). – Archiv für Molluskenkunde, 99 (1/2): 27-33.

Szekeres, M. (2007): Four new subspecies of Alopia H. & A. Adams 1855 (Gastropoda, Pulmonata: Clausiliidae). – Schriften zur Malakozoologie, 23: 7-18, 4 pls.


Figs. 12-15. Habitats of Alopia species from Dâmboviţa valley system. 
Fig. 12. Cheile Dâmbovicioarei, view from entrance upstream.  

Fig. 13. Valea Brusturetului 1 km above Peştera Dâmbovicioara, below mouth of Valea Muierii, view downstream.  

Fig. 14. Valea Crovului, right side above mouth.  

Fig, 15. Alopia p. plumbea (Rudăriţa form), Valea Rudăriţa (right side, ca. 1000 m).  

Map. Collecting sites of Alopia species from Dâmboviţa valley system.
Alopia subspecies of sites are indicated by letters; figure numbers are added in circles. If sites cannot be localized exactly, only letters are given.
Explanation of letters: c = g. intercedens (cerasina), e = g. elegantissima, g = g. glorifica, i = g. intercedens (intercedens), l = l. lischkeana, m = g. magnifica, m (c) = g. magnifica (clessini), m/e = g. magnifica >< elegantissima, p = p. plumbea (Rudăriţa form).
Sites of the seventies localized by the help of the locality desciptions, sites of 2014 and 2015 by GPS data.  

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