Systematics and Phylogenetics ACTA ZOOLOGICA BULGARICA Research Article Acta zool. bulg., 67 (2), 2015: 167-177

Identification Keys to the Clausiliidae J. E. Gray, 1855 (Mollusca: Stylommatophora) from Bulgaria

Atanas Irikov*, Dilian Georgiev

Department of Ecology and Nature Conservation, Faculty of Biology, University of Plovdiv “Paisii Hilendarski”, 24 Tsar Assen Street, 4000 Plovdiv, Bulgaria, E-mails: [email protected]; [email protected]

Abstract: The paper presents the currently known diversity of the family Clausiliidae J. E. Gray, 1855 on the territory of Bulgaria. All new systematic changes are noted and all new taxa described during the last years are listed. The main key characters for species identification are presented, with additional descriptions of some characters.

Keywords: identification, genera, species, subspecies, Clausiliidae, Bulgaria

Introduction The terrestrial malacofauna of Bulgaria is char- the different groups of Clausiliidae require expertise. acterised by high taxonomic diversity of the family According to the same authors, the genital system Clausiliidae Gray, 1855. The Europaen representa- structure also has an important taxonomical value in tives of this family are distributed mostly on the identifying some of the species. Balkan Peninsula. On the territory of Bulgaria, there Specific for the species and subspecies of is an extremely high diversity of species and sub- this family is that most of them have local distri- species of clausiliids. Most of these taxa are sum- butions on isolated rock complexes in Bulgaria of marised in the monograph of Da m j a n o v , Li k h a r e v the so-called ‘island-type’. Some of them occur (1975). Following the latter publication, many no- on very small territories as Macedonica pirinensis menclatural changes were made, new distributional Jaeckel, 1954, Macedonica zilchi Urbański, 1972, data were published and some additional species and Macedonica hartmuti Irikov, 2003, Macedonica subspecies were described. At present, the identifica- teodorae Irikov, 2006, Bulgarica varnensis trimont- tion characters of the taxa of the family Clausiliidae siana Irikov, 2006, Alinda biplicata karlukovoensis suggested by Da m j a n o v , Li k h a r e v (1975) are not Dedov, 2009 and other (Ja e c k e l 1954, Ur b a ń s k i functional and need to be updated. The shell struc- 1972, Ir i k o v 2003, 2006, De d o v 2009). Since many ture and the genital system were the features con- sites of ‘limestone islands’ of Bulgaria are still in- sidered with the highest taxonomic value for iden- sufficiently explored it can be highlighted that the tifying clausiliids. As a result of numerous studies clausiliid fauna of the country needs more studies of this snail group, concerning their morphology and many taxa remain unknown. In this paper, we and anatomy, some new criteria for determination of summarise all data concerning the identification the genus, species and subspecies were introduced characters of the clausiliids from Bulgaria. To sup- (No r d s i e c k 2002). Leading taxonomical value now port future studies on this group of snails we sug- has the closing apparatus of the shell (clausilium). gest identification keys to the genera, species and No r d s i e c k , Ne u b e r t (2002) point out that the key subspecies based on all the current knowledge re- characters for the determination of the species from garding this group from Bulgaria.

*Corresponding author: [email protected]

167 Irikov A., D. Georgiev

Fig. 1. Schematic illustration of the clausilial apparatus in accordance with No r d s i e c k , Ne u b e r t (2002) considered in this paper: 1 – frontal view, 2 – dorsal view (for the abbreviations see ‘Material and Methods’)

the construction of the keys, the comparative shell collection of A. Irikov was used as well as all the descriptions of taxa by Ro s s m ä s s l e r (1835, 1839), We s t e r l u n d (1884, 1892), Wa g n e r A. (1927), Ja e c k e l S. (1954), Da m j a n o v , Li k h a r e v (1975), No r d s i e c k (1963, 1969, 1972, 1973, 1974, 1977, 1982, 1985, 2002, 2008), No r d s i e c k , Ne u b e r t (2002), Ur b a ń s k i (1960, 1964, 1969, 1972, 1977), De d o v (2009, 2011, 2012), and We l t e r -Sc h u l t e s F. W. (2012). Abbreviations Plicae of the pariet-columellar side = lamella superior lamella (parietalis, sul) and spiral la- mella (spiralis, spl): inferior lamella (columellaris, il), subcolumellar lamella (subcolumellaris, scl), clausilium, consisting of stalk (cs) and plate (cp), Fig. 2. Schematic illustration of the genitalia of Clausilii- The space between superior lamella and infe- dae (according No r d s i e c k 1985). Abbreviations: ad = al- losspermiduct (of spermiduct); at = genital atrium; bb = rior lamella is named interlamellar. bursa (of bursa copulatrix); db = diverticulum (of bursa A parallel lamella (parallelis, pll) is often copulatrix); ep = epiphallus; fl = flagellum; fod = free present near to the suture. oviduct; od = oviduct (of spermoviduct); p = penis; pb = pedunculus (of bursa copulatrix); pc = penial caecum; Plicae of the palatal side = plicae (in the pro = prostate (of spermoviduct); rp = penial retractor; v strict sense): = vagina; vd = vas deferens principal plica (principalis, pri), palatal plicae (lunellar): upper palatal plica (up), if lunella present, with Material and Methods anterior part (upa) and posterior part (upp), The schematic drawings of the clausilial ap- middle palatal plicae resp. lunella (lun), paratus with main key characters are in accordance lower palatal plica (lp), if lunella present, with with No r d s i e c k (1982) and No r d s i e c k , Ne u b e r t anterior part (basalis, bas) and posterior part (sub- (2002) (Fig. 1), and the one of the genital system is claustralis, scs), in accordance with No r d s i e c k (1985) (Fig. 2). For lowest palatal plica (sulcalis, sulc),

168 Identification Keys to the ClausiliidaeJ . E. Gray, 1855 (Mollusca: Stylommatophora) from Bulgaria

If subclaustralis and sulcalis are fused or indis- an exception of Carinigera schuetti Brandt, 1962, tinguishable, only the term posterior lower palatal with rounded back side of last whorl). plica should be used. 14. (25) Lunella present. A sutural plica (suturalis, sut) is often present 15. (22) Last whorl with one basal keel; lower near to the suture. palatal plica lacking. The callous thickening of the palatal wall be- 16. (19) Lunella situated laterally. hind the peristome is named palatal callus. 17. (18) Palatal end of aperture usually with short lamellae or nodules; wider side of clausilium plate strongly twisted; penis without head ..Laciniaria Results 18. (17) Shell without lamellae on aperture edge; wider side of clausilium plate only slightly Key to the genera of the family Clausiliidae from twisted; head of penis is more or less distinctly sepa- Bulgaria rated ...... Аlinda 1. (26) Spiral lamella present (with the excep- 19. (16) L lunella situated on back side. tion of Laciniaria bajula mursalicae Urbański, 1969 20. (21) Inferior lamella positioned vertically and L. b. ditrichi Nordsieck, 1977). and hardly visible in aperture; lower palatal plica in- 2. (9) Spiral lamella in contact with superior sufficiently developed; palatal callus lacking; basal lamella or both are fragmented and situated in one keel more or less clearly visible ...... Euxina line, at equal distances from collumela. 21. (20) Inferior lamella situated partly spirally 3. (6) Periphery of the aperture with many small and better visible from aperture; lower palatal plica lamellae. well developed and terminates frontally with palatal 4. (5) Whole aperture edge with many lamel- callus; basal keel distinct ...... Galeata lae and nodules, subcolumellar lamella interrupted 22. (15) Last whorl with double keel: always and composed of variously developed lamellae; dis- well developed basal and distinct or insufficiently tal part of clausilium plate inclined inwards; diver- developed dorsal keel; lower palatal plica present. ticulum of bursa copulatrix longer than bursa and its 23. (24) Lunella strongly developed and below stalk ...... Serrulina transformed into lower palatal plica, well visible in 5. (4) Only columellar end of aperture with la- aperture ...... Carinigera mellae and nodules; subcolumellar lamella not in- 24. (23) Lunella less distinctly developed, not terrupted; distal part of clausilium plate not twisted; well visible in aperture ...... Bulgarica diverticulum of bursa copulatrix shorter than bursa 25. (14) Lunella lacking ...... Mentisella and its stalk ...... Dobattia 26. (1) Spiral lamella missing or distinctly re- 6. (3) Aperture edge different. duced. 7. (8) Aperture narrow and high; last whorl with 27. (26) Clausilial apparatus strongly reduced, clear basal keel and groove; end of clausilium plate only underdeveloped superior lamella present ...... rounded and twisted with a clear angle ....Clausillia ...... Balea 8. (7) Aperture more rounded; last whorl with 28. (27) Clausilial apparatus not reduced, with weak basal keel; aperture rounded at its lower part, developed lamellae. with weak basal keel; clausilium plate with notched, 29. (32) Shell spindle-shaped, with sculpture of hooked ...... Vestia more or less developed ribs; subcolumellar lamella 9. (2) Spiral lamella does not touch superior hardly visible in aperture; additional lamellae lacking. lamella and both are not situated in one line but lo- 30. (31) Shell protruded, small (with height of cated at different distances from the columella. up to 70 mm), with fine striae and sharply pointed end 10. (13) Last whorl without keels. with rounded apex; inferior lamella not well visible 11. (12) Palatal plicae begins from back side in aperture; callus present, situated at basal sector of last whorl; callus above lower palatal plica of aperture and insufficiently developed dorsal and present; clausilium plate with two cuttings, one big basal keels; lunella situated sideward on last whorl; and one small connection, with large outgrowth short and deeply situated principal plica present; dis- and concavity in wider section of clausilium plate . tal part of vas deferens thickened; vagina and penis ...... Cochlodina very short ...... Micridylla 12. (11) Palatal plicae begins dorso-laterally, lat- 31. (30) Shell spindle-shaped, large (height > erally or deeper on last whorl; callus above lower pala- 70 mm), with sculpture of densely situated ribs and tal plica lacking; clausilium plate with one big cutting pointed apex; inferior lamella relatively well visible in connection, with outhgrowth and regularly-rounded in aperture; well-developed callus in palatal part wide side of clausilium ...... Macedonica of aperture and slight basal groove; lunella situated 13. (10) Last whorl with one or two keels (with backside on last whorl; principal plica lacking; distal

169 Irikov A., D. Georgiev part of vas deferens normal, vagina and penis rela- C. laminata partita (Westerlund, 1892) tively long ...... Idyla Shell slightly convex, slender, gradually and 32. (29) Shell tower-shaped, not densely but finely pointing along apex with 12-13 whorls. distinctly ribbed; subclaustralis relatively well-vis- Genus Macedonica O. Boettger, 1877 ible in aperture; additional plica under lower palatal There are 17 species and subspecies of this ge- plica present ...... Ruthenica nus known from Bulgaria. 1. (7) Lunella rudiment of lower palatal plica Keys to the species and subspecies of the family present. Clausiliidae from Bulgaria 2. (3) Aperture without thickened inner lip; Subfamily Serrulininae Ehrmann, 1927 subcolumellar lamella slightly to clearly visible in- Genus Serrulina Mousson, 1873 side aperture (excl. M. martae, in which lamella and There is only one known species of this genus clausilial apparatus significantly reduced); retractor from Bulgaria. of vagina inserted into pedunculus at distance from Serrulina serrulata (L. Pfeiffer, 1847) its transition into vagina; base of penis-papillae con- Shell ribbed, pale yellow; end of aperture with vex, ± long. many lamellae and nodules. 3. (6) Shell with well-developed sculpture of Genus Dobatia Nordsieck, 1973 stripes and ribs. There is only one known species of this genus 4. (5) Clausilial apparatus significantly reduced; from Bulgaria. palatal plicae short; clausilium plate not closing well Dobatia goettingi (Brandt, 1961) opening to aperture ...... M. martae Shell surface smooth, pale yellow; only colu- 5. (4) Clausilial apparatus well-developed: mellar end of aperture with lamellae and nodules. palatal plicae are long, clausilium plate completely Subfamily Alopiinae A.J. Wagner, 1913 closes opening to aperture ...... M. pinteri Genus Carinigera Moellendorff, 1873 6. (3) Shell surface with slight sculpture or al- Three subspecies and one species of this genus most smooth ...... M. frauenfeldi are known from Bulgaria. 7. (1) Lunella rudiment missing. Last whorl with two slightly twisted keels: 8. (17) Shell without white outer surface stra- more developed basal keel and less developed dorsal tum; aperture with relatively thickened inner lip; re- keel. Lunella situated dorsally or dorso-laterally on tractor of vagina inserted into pedunculus near to its last whorl, massive. transition into vagina; penis of small size, base of Carinigera buresi buresi (A. J. Wagner, 1928) penis papillae reduced (excl. M. teodorae with large Surface of upper whorls with small white papil- penis papilla). lae close to suture. Aperture with small lip. Lunella 9. (12) Sculpture of shell distinctly devel- with well-developed basalis. oped. Carinigera buresi damjanovi (Likharev, 1972) 10. (11) Shell with large, rounded ribs ...... Surface of upper whorls with big, white papillae ...... M. hartmuti and nodules close to suture. Aperture widely-open 11. (10) Shell with white, fine and sharp ribs .... and big. Lunella with well-developed basalis...... M. teodorae Carinigera buresi dramaensis (Nordsieck, 12. (9) Sculpture of shell insufficiently devel- 1977) oped. Shell without white suture, with rare small white 13. (14) Shell slender, thin, with simple aper- papillae close to suture of upper whorls. Aperture ture ...... M. brabeneci widely-open and big. Lunella with extremely long 14 (13) Shell spindle-shaped, thick, with thick and thick basalis. aperture. Carinigera schuetti Brandt, 1962 15. (16) Shell of large size and rather spindle- Shell often decollated, with small fine white shaped ...... M. marginata ribs. Aperture is widely-open and thickened. Parallel 16. (15) Shell of smaller size and rather tower- lamella present. Lunella situated dorso-laterally, shaped ...... M. dobrostanica with long and massive basalis. 17. (8) Shell with white outer surface stratum; Genus Cochlodina Férussac, 1821 aperture without separate inner lip (simple aperture One species is known from Bulgaria, represent- margin); retractor of vagina inserted into pedunculus ed with two subspecies. Shell smooth, without ribs. at distance from its transition into vagina but closer Last whorl without keel. to vagina in comparison with frauenfeldi species; C. laminata laminatа (Montagu, 1803) base of penis papillae clear. Shell distinctly convex, sharply pointed at apex 18. (19) Shell with significantly developed with 10-111/2 whorls. sculpture of ribs and lamellae ...... M. zilchi

170 Identification Keys to the ClausiliidaeJ . E. Gray, 1855 (Mollusca: Stylommatophora) from Bulgaria

19. (18) Shell with slightly developed sculp- tal plicae present between principal plica and lower ture ...... M. pirinensis palatal plica, ending with palatal callus in front part The taxa of the genus Macedonica are subdi- of аperture. Basalis large with one lunella rudiment. vided into three groups (No r d s i e c k 1974, 1977). Subspecies with relatively large penial papilla Marginata species group exception in frauenfeldi group, usually not very long Macedonica marginata marginata (Rossmässler, but convex. 1835) – (Syn. Clausilia auriformis Mousson, 1859) Macedonica frauenfeldi riedeli Urbański, Shell surface smooth, with regular, small, fine 1977 stripes on upper and lower whorls. Shell with white suture between all whorls. Macedonica marginata major (Rossmässler, Sculpture well visible, with fine lines and small ribs. 1839) Aperture periphery slightly thickened, indistinctly Shell of large size and with well-developed separated in aperture. Principal plica long, well vis- clausilial apparatus. Palatal plicae usually four. ible in aperture, going deep inside last whorl. Lower Macedonica marginata frivaldskyana palatal plica visible in inclined view into aperture, (Rossmässler, 1839) with slightly developed lunella rudiment. Male part Shell convex, of relatively small size, with big of genital organs short and convex. ribs at distance from each other, and patches of white Macedonica frauenfeldi tau Nordsieck, 1977 touches at suture. Aperture periphery well separated in aperture. Macedonica marginata balcanica (A. Wagner, Lunella deep inside last whorl. Principal plica well 1927) developed frontally in aperture. Shell of relatively small size. Clausilial appara- Macedonica pinteri Sajó, 1968 tus rather reduced. Shell slender, with weak stripes and large Macedonica brabeneci brabeneci Nordsieck, rounded ribs. Aperture periphery with thin and part- 1977 ly separated lip in aperture. Subcolumellar lamella Shell slender, gradually shaped, thinner at top. large and well visible in aperture. Palatal plicae well Aperture of elongate-oval shape. Principal plica rel- developed. One lunella rudiment. atively short. Macedonica martae Sajó, 1968 Macedonica brabeneci prismatica Dedov, Shell with whitish surface, convex, with regular 2012 and widely spaced ribs. Aperture periphery with thin Shell rounded at its middle part, sharply and partly separated lip in aperture. Subcolumellar shaped, thin at top. Aperture of oval-auricular shape. lamella missing. Clausilial apparatus significantly Principal plica long. reduced. All palatal plicae short, sometimes with Macedonica hartmuti Irikov, 2003 shape of callus. Shell small, tower-shaped, with massive, round- Macedonica species group ed ribs, at distance from each other.Macedonica teo- Macedonica pirinensis Jaeckel, 1954 dorae Irikov, 2006 Shell with white surface stratum, slight sculp- Shell large, with sparse regular thin ribs, with ture excluding neck part, insufficiently developed white touches at suture. Only species of marginata lamella and closing apparatus. Clausilium plate with group with distinctly developed and long penis pa- clearly visible cutting. pilla. Macedonica zilchi Urbański, 1972 Macedonica dobrostanica Irikov, 2012 Differs from other forms of the genus Shell small, smooth. Genital system of small Macedonica mostly by its large size, convex shell, size, in distal part of vagina with longitudinal, seg- well-developed bluish-white shell surface stratum mented fold, so far unknown in other species of this and clear sculpture of ribs and clausilial apparatus. genus. Subcolumellar lamella hardly visible inside aper- Frauenfeldi species group ture. Lower palatal plica strongly developed and Macedonica frauenfeldi sigma (Westerlund, visible inside aperture. Clausilium plate with dou- 1884) ble cutting, similar to сochlodina species (with one Shell almost smooth, with fine ribs at top, on larger and one smaller, widely rounded cutting). middle and lower whorls clearly visible at suture. Male section of genital system of large size aperture with thickened periphery. Principal plica compared to marginata species. Penis papilla large, very long, ending inside. Basalis with lunella rudi- long, and situated along whole length of penis reach- ment, not well visible in аperture. ing almost to genital atrium. Macedonica frauenfeldi regia Nordsieck ,1974 Subfamily Mentissoideinae Lindholm, 1924 Sculpture of shell of fine stripes. Aperture with Genus Idyla Н. & А. Adams, 1855 thin periphery. Principal plica long. Additional pala- One subspecies of this genus is known from

171 Irikov A., D. Georgiev

Bulgaria. Idyla castalia boschi Nordsieck, 1973. tal plica long, inclined, such that on inside almost Shell with fine, dense ribs. innferior lamella divergating with principal plica, and in its front part hardly visible in aperture. Neck part of last whorl ending with callus. Lunella short, massive. with two keels: basal keel and weak dorsal keel. Subfamily Clausiliinae J.E. Gray, 1855 With one massive lunella and palatal callus near ap- Genus Ruthenica Lindholm, 1924 ertural edge. Palatal plicae missing: basalis poorly Monotypic genus. developed. Ruthenica filograna (Rossmässler, 1836) Genus Euxina Boettger, 1877 Shell of very small size, slender with tower- Euxina circumdata (L. Pfeiffer, 1848) like shape, entirely covered with regular, sharp ribs. Shell slender, gradually growing thiner to Aperture small, with slightly open apertural end. apex, densely covered with fine ribs and with white Keels missing, neck side of last whorl with big dor- touches at suture. Aperture ovally-elipsoid. Inferior sal bulge. Inferior lamella slightly visible in aper- lamella hardly visible in aperture. Last whorl with ture, front part ending with thin lamella at periphery clear basal keel and slight dorsal bulge. Principal pli- of aperture edge. One short and thin lamella under ca long, situated high, near suture. Lunella massive, inferior lamella, at end of aperture. Subcolumellar with lower end twisted inwards. Any other palatal lamella partly visible in aperture. Principal plica plicae missing. Upper side of clausilium plate regu- short, situated deep in last whorl on inside, penetrat- larly rounded, without any cuttings. Diverticulum ing behind lunella, cannot be seen in aperture from of bursa copulatrix significantly shorter than bursa frontal view. Lunella well developed. Any other of bursa copulatrix. Penis long, convex at its back palatal plicae lacking. part. Diverticulum of bursa copulatrix twice as long Euxina persica paulhessei (Lindholm, 1925) as bursa of bursa copulatrix. Penis short. Penial re- Shell strongly convex in its middle part, sharp- tractor big and connected with proximal end of pe- ly growing thinner at apex, with fine stripes and no nis. white touches. Aperture large, rounded. Inferior la- Genus Micridyla Nordsieck, 1973 mella not well visible in aperture. Last whorl with There is one known species of this genus from clear basal keel and slight dorsal bulge. Principal Bulgaria. plica very long, longer than lunella inside. With pal- Micridyla pinteri (Nordsieck, 1973). atal callus near aperture edge and massive lunella. Shell small (smallest species of the fam- Clausilium plate with parallel sides and not widen- ily Clausiliidae in Bulgaria), slightly convex, fast ing at its upper side which rounded. Bursa and di- tapering upwards, with obtuse apex. Sculpture verticulum of bursa copulatrix equal in length. Penis consists of very fine, regular and dense small ribs. long, with ligament and with penial retractors, fold- End of aperture separated. Last whorl with neck ing it and dividing it in two sections. bulge, slight basal keel and unclear basal groove. Euxina pontica borisi (P. Hesse, 1912) Inferior lamella situated vertically and almost Shell slender, gradually growing thinner at its invisible in aperture. Subcolumellar lamella partly apex, with fine ribs with tufts of white touches at visible in front part of aperture. Principal plica and suture. Aperture brown and large with round-ellip- lower palatal plica insufficiently developed, short, tical shape, and thin pale lip. Last whorl with clear not visible in front part of aperture, situated behind basal keel and slight dorsal bulge. Principal plica lunella on inside. Lunella well developed, with small long, situated high, near suture. Lunella distinctly subclaustralis. Palatal callus in lower part, in front developed. Clausilium plate widened at front side part of aperture, penetrating inside. Any other palatal and with few small cuttings. Biggest cutting forming plicae missing. Clausilium plate narrow with tongue- growth with thickened sharp top. like shape. Genus Galeata O. Boettger, 1877 Vagina and penis very short. One known species of this genus from Genus Clausilia Draparnaud, 1805 Bulgaria. There is one known species of this genus from Galeata schwerzenbachii (L. Pfeiffer, 1848) Bulgaria. Shell of small size, with slender to tower-like Clausilia (Clausilia) pumila pumila C. Pfeiffer, shape, whole surface covered with dense and fine 1828 ribs with white touches. Aperture small with well- Shell at lower whorls strongly convex, fast separated apertural edge. Inferior lamella hardly tapering upwards to apex. Shell surface intensely visible in aperture, transformed into thin lamella at and regularly ribbed with bundles of white touches front part, ending at apertural edge. Clear basal keel on ribs. End of aperture open, thickened, clearly and wide rounded dorsal keel. Principal plica long, separated, and white in color. Superior lamella penetrating inside from behind lunella. Upper pala- merges with spiral lamella. Inferior lamella almost

172 Identification Keys to the ClausiliidaeJ . E. Gray, 1855 (Mollusca: Stylommatophora) from Bulgaria vertical, slightly visible in aperture. Lunella with ture end. Neck part of last whorl with a slight basal curved inner ends. Principal plica long, penetrating keel. Principal plica well visible in front part of ap- behind lunella. Palatal callus from which lower pala- erture, penetrating deep inside from behind lunella. tal plica separated. Wide part of clausilium curved Anterior part of upper palatal plica ditinctly inclined with clear angle. to principal plica and ends before lunella. Subfamily Baleinae A.J. Wagner, 1913 Laciniaria macilenta (Rossmässler, 1842) Genus Mentissella Nordsieck, 1973 Shell tower-like shaped with a sharp apex, of There is one known species of this genus from small size (smallest Laciniaria species, 10-11 сm Bulgaria. shell height), with fine, dense, regular ribs. Aperture Mentissella rebeli (Sturany, 1897). with a few small lamellae, ending inside near prin- Shell slender, with tower-like shape and sharp cipal plica and upper palatal plica. Last whorl at apex, and fine, dense ribs with bundles of white neck with one basal keel, with a deep basal groove touches. Aperture with elliptical-oval shape, apertural at front. Principal plica and upper palatal plica long, end slightly open. Inferior lamella almost vertical well visible in front of aperture, prolonged inside be- and hardly visible in aperture. Last whorl with well- hind lunella. Lunella well developed. developed basal keel and deep basal groove at front. Laciniaria bajula bajula A. Schmidt, 1968 Three palatal plicae: long principal plica, short upper Shell spindle-shaped, with regular ribs on its palatal plica and very massive lower palatal plica. whole surface. Columellar part of aperture with few Ends of principal plica and of lower palatal plica lamellae and nodules, situated rather inside than at visible at front part of aperture. One palatal callus end of aperture. Inferior lamella vertical and hardly in aperture, as part of upper palatal plica. Lunella visible in aperture. One interlamellar lamella situat- missing. Clausilium plate with a wide front end. ed at front of inferior lamella. Basal groove shallow, Genus Laciniaria Hartmann, 1842 not reaching apertural end. Neck part of last whorl According to No r d s i e c k (2008) there are convex, with a slight basal keel. Palatal plicae, with eight valid species and subspecies of this genus in exception of principal plica, reduced. Principal plica Bulgaria. Laciniaria potochensis Dedov, Neubert not visible in front part of aperture. Lunella well (2006) a synonym of Alinda (Alinda) atanasovi developed. atanasovi (Urbański, 1964; Nordsieck 2008). Laciniaria bajula mursalicae (Urbański, 1969) Lower palatal plica missing. Palatal end of ap- Shell with thin ribs, mostly on middle and up- erture usually with lamella and nodules. Sometimes per whorls. Spiral lamella missing. Basal groove in- such lamellae present also at parietal end of aper- sufficiently developed. Last whorl convex at its neck ture. part, without keel. Principal plica short, almost com- Laciniaria plicata plicata (Draparnaud, 1801) pletely reduced and not visible in aperture. Lunella Shell spindle-shaped, with thin, dense, regu- and clausilium missing. lar ribs with patches of bundles of white touches. Laciniaria bajula lunella Nordsieck, 1973 Aperture end in palatal and more rarely in colume- Shell surface of middle and upper whorls with llar part, with plicae and nodules, situated further dense, thin ribs. Aperture end with few lamellae and inside apertural edge (sometimes with a missing la- nodules on its palatal part, situated rather inside ap- mella at columellar part). Last whorl on neck with a erture edge. Basal groove inscufficiently developed. well-developed basal keel, in front part there a deep Last whorl at its neck part convex without a keel. basal groove. Principal plica and upper palatal plicae Principal plica long, hardly visible in front part of ap- long and almost parallel to each other. Posterior part erture, penetrating deep inside from behind lunella. of upper palatal plica divergates with principal plica Lunella massive, rainbow-curved with partly formed close to lunella. Lunella well developed. subclaustralis. Laciniaria plicata kueprijae Nordsieck, 1973 Laciniaria bajula ditrichi Nordsieck, 1977 Shell spindle-shaped. sculpture consists of Shell surface of middle and upper whorls with clear, widely spaced, whitish ribs. Lamellae missing fine ribs, enlarged at aperture end. Clausilial appara- at aperture end. Anterior part of upper palatal plicae tus distinctly reduced. partly reduced. Genus Alinda Н. & А. Adams, 1855 Laciniaria plicata rhodopensis Nordsieck, There are 13 species and subspecies known 2008 from this genus from Bulgaria. Shell with widely spaced, regular and sharp Shell ribbed, with white touches on suture. ribs, with patches of white touches on them. clausilial apparatus developed to a different extent. Between superior lamella and inferior lamella some Alinda (Alinda) biplicata bilicata (Montagu, intermediate nodules (interlamellar and mostly 1803) subinterlamellar). Basal groove not reaching aper- Shell spindle-shaped, thin and densely ribed,

173 Irikov A., D. Georgiev with white touches. apeture end thickened, sepa- erture. Without interlamellar plicae. Basal groove rated by open aperture edge. Inferior lamella well reaches end of aperture. Neck part of last whorl with visible in aperture. At its parietal part aperture ends insufficiently developed basal keel. Lunellae dorso- with one or two not very large lamellae or nodules. lateralis, principal plica not well visible in front part Sometimes between superior lamella and inferior of aperture, and penetrating inside to upper end of lamella there are one or two interlamellar lamellae. lunella. Anterior part of upper palatal plica small, Neck part of last whorl with an unclear basal keel. short or missing, most often separated from upper Principal plica begining rather inside lunella. Upper palatal plica. Lunella insufficiently developed. palatal plica situated at an angle to principal plica, Alinda (A.) biplicata karlukovoensis Dedov, begining from upper end of lunella and ending near 2009 end of aperture. Lunella well developed and situated Shell spindle-shaped, with thin and dense ribs, at right side of last whorl. Clausilium with distinctly and patches of white touches. On upper whorls with curved upper part. groups of small nodules at suture. Aperture with Alinda (A.) biplicata michaudiana (L. Pfeiffer, an open lip. Rarely with interlamellar plicae. Basal 1848) groove shallow, reaching almost to end of aperture at Shell elongated, spindle-shaped, with dense front. Neck part of last whorl with a well-developed ribs, and patches of white touches near suture. basal keel and dorsal bulge. Principal plica and up- Inferior lamella relatively big, well visible in aper- per palatal plica long, well visible in front part of ture. Basal groove reaches almost to end of aperture. aperture. Principal plica penetrates deeply behind Neck part of last whorl with clear basal keel and dor- lunella. Anterior part of upper palatal plica situated sal bulge. Palatal plicae and clausilim partly reduced. at a wide angle to principal plica and ending before Principal plica and upper palatal plicae penetrate lunella. Often with additional anterior upper palatal inside behind lunella, very thin almost rudimental, plica. Lunella long with distinctly developed upper in front part of aperture slightly visible. Anterior part end. of upper palatal plicae short. Often upper palatal pli- Alinda (A.) biplicata alibotushensis Dedov, cae completely missing. Lunella rudimental or com- 2009 pletely missing. Shell widely ribbed, with white touches. Alinda (A.) biplicata euptychia (Ehrmann (in Superior lamella situated near spiral lamella. Anterior Ur b a ń s k i 1960)) part of upper palatal plica separated, owing to higher Shell elongate and spindle-shaped, with white altitude where this subspecies lives. touches on ribs near suture. Shell surface with fine Alinda (Alinda) atanasovi atanasovi (Urbański, and dense small ribs. Inferior lamella big, well vis- 1964) ible in aperture. Neck part of whorl with clear ba- Shell tower-like shaped, with widely situated sal keel. Principal plica long and penetrating inside whitish ribs and white touches at suture. Inferior la- behind lunella. Anterior part of upper palatal plica mella vertical, hardly visible in aperture. Between situated at wide angle to pricipaland separated from superior lamella and inferior lamella most often with it, on inside ending in front of lunella, well visible in two interlamellar lamellae. Neck part of last whorl aperture. Sometimes with short pseudo-palatal plica with slight basal keel. Principal plica long, penetrat- situated under upper palatal plica. Lunella massive, ing inside from behind lunella, in front paret hardly with distinctly curved upper end parallel to principal visible in aperture. Upper palatal plica situated at a plica. clear angle to principal plica and reaching lunella. Alinda (A.) biplicata orientalis Nordsieck, Lunella long with slightly curved upper end. 2008 Alinda (Alinda) atanasovi kremenensis (Dedov, Shell convex, sharply tapering at its apex, 2009) widely ribbed, with white touches near suture. Shell widely ribbed with white touches. Inferior lamella with well visible aperture. Neck Alinda (Alinda) wagneri wagneri (A. Wagner, part of last whorl with short basal keel and dorsal 1911) bulge. Principal plica long, on inside it prolongs Shell distinctly convex, massive. Sculpture con- behind lunella. Anterior part of upper palatal plica sists of very fine, dense stripes. Inferior lamella and short, situated at wide angle to pricipalis. Sometimes basal groove hardly visible in aperture. Neck part of with short pseudo-palatal plica situated under upper last whorl with small basal keel and big dorsal bulge. palatal plica. Lunella short with upper end strongly Spiral lamella, palatal plicae and clausilium partly curved inwards, parallel to principal plica. reduced. Alinda (A.) biplicata irikovi Nordsieck, 2008 Alinda (A.) wagneri petrohanica (Urbański, Shell convex, widely ribbed, with patches of 1969) white touches. Inferior lamella hardly visible in ap- Clausilial apparatus distinctly reduced. Shell

174 Identification Keys to the ClausiliidaeJ . E. Gray, 1855 (Mollusca: Stylommatophora) from Bulgaria rather slender in comparison with nominate sub- end of aperture ending with two separate lamellae. species. End of aperture not so separated and open. Subcolumellar lamella not well visible in aperture. Inferior lamella hardly visible in aperture. Principal Insignificant basal keel and groove. Long principal plica signifcantly reduced. Without lunella. plica (not well visible in aperture from front and Alinda (Alinda) vratzatica (Likharev, 1972) prolonged behind lunella) and short lunella (upper Shell tower-like shaped, small, densely ribbed, end distinctly curved inwards), situated dorsally on with thin and dense stripes, some of them white in whorl. Clausilium plate toungue-shaped and slightly color. Aperture edge white with well separated open pointed and hooked at its end. Vagina located dis- lip. Neck part of last whorl with a distinctly protruded tally and not widened proximally. basal keel. Lunellae laterally situated. Principal plica Vestia (Vestiella) roschitzi trigonostoma long and penetrating behind lunella; its front part not (Pavlovic, 1912) visible in aperture. Anterior upper palatal plica short, Shell with well visible sculpture, widely ribbed, separated, situated at a wide angle to principal plica, convex at its lower side, aperture of triangular shape. inside part ending before lunella, front part hardly With insignificant palatal and well visible basal callus visible in aperture. Lunella long, massive, distinctly inside base of aperture. Two small lamellae in front curved to inside upper end parallel to principal plica. of inferior lamella, at end of aperture. Subcolumellar Genus Balea Gray, 1824 lamella not well visible. Basal keel and basal groove There are two known species of this genus from poorly developed. Bulgaria. Vestia (Vestiella) roschitzi nordsieckiana Balea kaeufeli (Brandt, 1961) (Urbański, 1979) Shell densely ribbed with fine stripes, white in Shell of small size (average shell height of 11.2 colour at suture. Basal keel only implicit. Lunellae mm, and width of 2.7 mm), with dense fine sculpture and plicae missing except for superior lamella and and no callus at aperture end. Interlamellar space insufficiently developed inferior lamella. covered with few small lamellae. Last whorl at neck Balea eninskoensis (Irikov, 2006) part rounded. Clausilium plate tounge-shaped, its Shell surface almost smooth; no white coloration end pointed and hooked. at suture. Last whorl at its neck part convex and Vestia (Vestiella) roschitzi neubertiana Dedov, protruded, without a keel. With exception of superior 2010 lamella, no other lunellae and plicae. Shell with clear sculpture of widely situated Genus Pseudalinda O. Boettger, 1877 ribs. End of aperture with thick callus. Without in- There are two known species of this genus from terlamellar lamellae. Basal keel and groove well de- Bulgaria. veloped. With well visible basal callus further inside Pseudalinda fallax (Rossmässler, 1836) base of aperture. Shell brown, solid, finely ribbed. Superior la- Genus Bulgarica Boettger, 1877 mella long and not connected to spiral lamella, no There are 14 known species and subspecies of folds between superior and inferior lamella, inferior this genus from Bulgaria. lamella deep inside, slightly concave at lower side 1. (16) Aperture end without lamellae or with and forked inside. No lower palatal plicae, lunella single lamellae, only on parietal and columellar end dorsal and incomplete, subcolumellar lamella visible of aperture; lausilium without exterior angle, respec- from a perpendicular view, next to it with basal tively cant, near ± such. furrow. 2. (11) Last whorl with two keels: basalis and Pseudalinda golesnicensis A. J. Wagner, 1914 dorsalis. Very similar to Pseudalinda falax, but with 3. (6) False upper palatal plica transformed into rather slender shell, yellowish or greenish-brown, callus or missing, falsely connected to lunella. with shorter superior lamella. 4. (5) Shell with dense ribs, pale to horn-brown Genus Vestia Hesse, 1916 ...... B. bulgariensis There are two species and four subspecies of 5. (4) Shell except for uppermost and last whorls this genus from Bulgaria. almost smooth, horn to violet-brown ....B. hiltrudae Vestia (Brabenecia) ranojevici ranojevici 6. (3) False upper palatal plica well developed, (Pavlovic, 1912) always connected with lunella and well visible in Shell convex with a blunt apex and fine dense aperture. small ribs with white touches near suture. Superior 7. (10) Basal and dorsal keel well developed. lamella merging with spiral lamella. Lip clearly 8. (9) False upper palatal plica long ...... separated and thickened. Interlamellar space cov- ...... B. fraudigera ered with few small lamellae. Aperture rounded; 9. (8) False upper palatal plica partly developed inferior lamella deeply situated, hardly visible, at ...... B. pseudofraudigera

175 Irikov A., D. Georgiev

10. (7) Dorsal keel less developed ...... Bulgarica (B.) varnensis gabrovnitcana Irikov, ...... B. urbanskii 2006 11. (2) Last whorl with one basal and only Shell clearly ribbed with white, rare ribs. last rudiments of second dorsal keel. whorl with rather distinctly formed double keel. 12. (13) Subcolumellar lamella extended to Bulgarica (B.) varnensis trimontsiana Irikov, edge of aperture, lower palatal plica insufficiantly 2006 developed and transformed into a callus, clausilium Shell insufficiently ribbed with dense small without external angle, respectively external cant, stripes. Dorsal keel less developed than in previous penis with long ligaments ...... B. vetusta subspecies. Upper palatal plica short, lower palatal 13. (12) Subcolumellar lamella not extending plica well developed. to edge of aperture, no palatal callus, lower palatal Bulgarica (B.) urbanskii urbanskii Nordsieck, plica well developed, clausilium with ± well visible 1973 external angle, respectively external cant, penis Shell with rarely situated ± rounded ribs. shortens by ± well-formed ligament. Without neck bulge. Principal plica long, upper 14. (15) Shell convex, with blunt apex and palatal plica small and interrupted, in front part with insufficiently-developed double keel, false upper remnant of plica similar to palatal callus. In some palatal plica insufficiently developed, short, shells with interlamellar plicae. sometimes completely missing ...... B. fritillaria Bulgarica (B.) urbanskii paganella Nordsieck, 15. (14) Shell less convex, with sharper apex 1974 and well-developed double keel, false upper palatal Shell with ± thin and sharp ribs. Neck bulge plica well developed and rarely missing ...... partly formed. Basalis partly separated from lunella...... B varnensis Upper palatal plica almost always missing. 16. (1) Aperture end with many lamellae, Bulgarica (B.) bulgariensis bulgariensis (L. clausilium with external angle, respectively external Pfeiffer, 1848) cant ...... B. denticulata thessalonica Shell small, distinctly ribbed with dense whit- Bulgarica (Strigilecula) vetusta (Rossmässler, ish ribs. Two keels well visible. Number of ribs on 2 1836) mm from penultimate whorl most often 10-18. Shell with thin, dense ribs with white touches. Bulgarica (B.) bulgariensis intricata (Mousson, Aperture end thickened, separated. Principal plica 1859) long. Upper palatal plica insufficiently developed, Shell of medium size. Average number of ribs lower palatal plica missing. With undeveloped pala- on 2 mm on penultimate whorl on average 19. tal callus. Bulgarica (B.) bulgariensis osmanica Bulgarica (Bulgarica) hiltrudae Nordsieck, (Westerlund, 1884) 1974 Shell of a relatively large size. Average rib Basal and dorsal keel well developed. With number on 2 mm from penultimate whorl most one- five clear interlamellar lamellae. often 29. Bulgarica (B.) fraudigera (Rossmässler, 1839) Bulgarica (B.) denticulatа thessalonica Shell massive, distinctly ribbed with widely sit- (Rossmässler, 1839) uated whitish ribs. Aperture end thick, clearly sepa- Shell convex with dense, fine ribs, with white rated. Inferior lamella horizontally situated, well touches. Dorsal keel insufficiently developed, basal visible in aperture. Well-developed double keel and keel narrow and deep, laterally situated. Three pala- long false upper palatal plica. tal plicae well developed. Bulgarica (B.) pseudofraudigera Nordsieck ,1973 Shell big with fine dense stripes. Well-developed Discussion double keel and short false upper and lower palatal Bulgaria is rich in species of the family Clausiliidae. plicae. In Da m j a n o v , Li k h a r e v (1975) there are 15 genera, Bulgarica (B.) fritillaria (Frivaldsky, 1835) 24 species and 20 subspecies listed. Some of these Shell convex, with thin and dense small stripes. taxa become invalid systematically or only in the Dorsal and basal keel partly formed. Basal keel situ- ated laterally at base of aperture and forming a nar- fauna of Bulgaria, while others are with changed row, small groove. Upper palatal plica short. nomenclature. After Da m j a n o v , Li k h a r e v (1975), Bulgarica (B.) varnensis varnensis (L. Pfeiffer, many new species and subspecies have been found, 1848) repoted and described and many distributional Shell with well-developed principal plica, up- data has been obtained. As a result of the recently per and lower palatal plicae. acquired knowledge on the family Clausiliidae in

176 Identification Keys to the ClausiliidaeJ . E. Gray, 1855 (Mollusca: Stylommatophora) from Bulgaria

Bulgaria, obtained during the last decades, it can species radiation of these genera. We reckon that the be summarised that 76 species and subspecies (40 family diversity is still poorly known. Many species species and 36 subspecies) have been recorded from have local distributions from the ‘island-type’ and the country till now. Most of the taxa belong to the have been discovered in different regions over the genera Macedonica (17) and Bulgarica (14), which last few years. We thus suppose that in future research characterises the Bulgarian territory as a centre of many more new taxa will be found in Bulgaria.

Reference matophora: Clausiliidae). – Archiv für Molluskenkunde, Da m j a n o v S., I. Li k h a r e v 1975. Fauna Bulgarica IV. Gastropoda 137, 2: 133-157. terrestria. Sofia, Bulgarian Academy of Sciences, 425 p. (In Bulgarian). Ur b a ń s k i J. 1960. Bemerkenswerte Clausiliiden (Moll., Pulm.) aus Bulgarien. (Systematische, zoogeographische und öko- De d o v I. 2009. Three New Subspecies of Alinda H. & A. Adams logische Studien über die Mollusken der Balkan-Halbinsel. 1855 (Gastropoda: Stylommatophora: Clausiliidae) from VI.). – Bulletin de la Societe amis des Sciences et des Lettres Bulgaria. – Acta Zoologica Bulgarica, 61 (3): 229-234. de Poznań, ser. D, I: 113-149. De d o v I., T. Mi t е v (2011). Molluscs Fauna (Mollusca: Gas- Ur b a ń s k i J. 1964. Beiträge zur Kenntnis balkanischen Stylomm- tropoda: Bivalvia) of Mountain Osogovo. – Acta Zoologica atophoren. (Systematische, zoogeographische und ökolo- Bulgarica, 63, 1: 37-46. gische Studien über die Mollusken der Balkan-Halbinsel. De d o v I. 2012. Description of New Subspecies of the Rare Bul- VII.). – Bulletin de la Societe amis des Sciences et des garian Endemic Macedonica brabeneci Nordsieck (1977) Lettres de Poznań, ser. D, IV: 19-56. (Pulmonata: Clausiliidae). – Acta Zoologica Bulgarica, Ur b a ń s k i J. 1969. Bemerkenswerte Balkanische Stylommato- 64 (3): 325-326. phoren (Systematische, zoogeographische und ökologische Ja e c k e l S. 1954. Zur Systematik und Faunistik der Mollusken Studien über die Mollusken der Balkan-Halbinsel. IX). – der nördlichen Balkanhalbinsel. – Mittilungen Aus dem Bulletin de la Societe amis des Sciences et des Lettres de Zoologischen Museum der Berlin, 30: 54-95. Poznań, ser. D, 9: 225-262. Mo u s s o n A. 1859. Coquilles terrestres et fluviatiles requeillis Ur b a ń s k i J. 1972. Bemerkungen zur Kentnis balknischer Alopii- dans l’Orient par. M. le. Dr. Alexandre Schläfli. V. La nae (Moll., Pulm.) (Systematische, zoogeographische Bulgarie. – Vierteljahresscher Naturalis Gesselschaft und ökologische Studien über die Mollusken der Balkan- Zürich, 6: 283-297. Halbinsel. XI). – Bulletin de la Societe amis des Sciences No r d s i e c k H. 1963. Zur Anatomie und Systematik der Clausilien, et des Lettres de Poznań, D, 12/13: 251-260. I. – Archiv für Molluskenkunde, 92, 3/4: 81-115. Ur b a ń s k i J. 1977. Bemerkenswerte Clausiliiden (Moll., Pulm.) No r d s i e c k H. 1969. Zur Anatomie und Systematik der Clausilien, der Nordlichen Balkan-Halbinsel (Systematische, zoogeog- VI. – Archiv für Molluskenkunde, 99, 5/6: 247-265. raphische und ökologische Studien über die Mollusken der No r d s i e c k H. 1972. Zur Anatomie und Systematik der Clausil- Ballkan-Halbinsel. XV). – Bulletin de la Societe amis des ien, XI. Neue Formen und taxonomische Revision einiger Sciences et des Lettres de Poznań, ser. D, 17: 235-251. Gruppen der Alopiinae. – Archiv für Molluskenkunde, Ro s s m ä s s l e r E. A. 1835. Iconographie der Land- und Süßwasser- 102, 1/3: 1-51. Mollusken, mit vorzüglicher Berücksichtigung der eu- No r d s i e c k H. 1973. Zur Anatomie und Systematik der Clausilien, ropäischen noch nicht abgebildeten Arten. Band 1. Heft 2. XIII. Neue Balkan-Formen der Mentissoideinae und Balei- Drezden, Leipzig und Wiesbaden. nae (mit taxonomischer Revision der zugehörigen Grup- Ro s s m ä s s l e r E. A. 1839. Iconographie der Land- und Süßwasser- pen). – Archiv für Molluskenkunde, 103, 4/6: 179-208. Mollusken, mit vorzüglicher Berücksichtigung der eu- No r d s i e c k H. 1974. Zur Anatomie und Systematik der Clausilien, ropäischen noch nicht abgebildeten Arten. Band 2. Heft 9. XV. Neue Clausilien der Balkan-Halbinsel. – Archiv für Drezden, Leipzig und Wiesbaden. Molluskenkunde, 104, 4/6: 123-170. Wa g n e r A. 1927. Studien zur Molluskenfauna der Balkanhalbinsel No r d s i e c k H. 1977. Zur Anatomie und Systematik der Clausilien, mit besonderer Berücksichtigung Bulgariens und Traziens, XVIII. Neue Taxa rezenter Clausilien. – Archiv für Mol- nebst monographischer Bearbeitung einzelner Gruppen. – luskenkunde, 108, 1/3: 73-107. Annales zoologici Musei Polonici, 6, 4: 263-399. No r d s i e c k H. 1982. Die Evolution des Verschluβapparats der We l t e r -Sc h u l t e s F. W. 2012. European Non-Marine Molluscs, Schlieβmundschnecken (Gastropoda: Clausiliidae). – Ar- a Guide for Species Identification. Planet Poster Editions, chiv für Molluskenkunde, 112 (1981), 1/6: 27-43. Göttingen. 679 p. No r d s i e c k H. 1985. The System of the Stylommatophora (Gas- We s t e r l u n d C. A. 1884. Fauna der in der paläarctischen Region tropoda), with Special Regard to the Systematyc Position (Europa, Kaukasien, Sibirien, Turan, Persien, Kurdistan, of the Clausiliidae, I. – Archiv für Molluskenkunde, 116, Armenien, Mesopotamien, Kleinasien, Syrien, Arabien, 1/3: 1-24. Egypten, Tripolis, Tunesien, Algerien und Marocco) leb- No r d s i e c k H. 2002. How to Determine Clausiliid Species, I: enden Binnenconchylien. IV. Gen. Balea Prid. & Clausilia Widespread Species of Clausilia Draparnaud (Gastropoda: Dr. – 1-212, 1-18, 1-16. Karlskrona. (Länsboktryckeriet). Stylommatophora: Clausiliidae). – Mitteilungen Deutschen We s t e r l u n d C.A. 1892. Neue Binnenconchilien in der Paläark- Malakozoologischen Gesellschaft, 68: 31-36. tischen Region. 25-48. No r d s i e c k H. 2008. Alinda biplicata (Montagu) and Laciniaria plicata (Draparnaud), Diversity in Comparison, with the Received: 31.05.2014 Description of New Subspecies (Gastropoda: Stylom- Accepted: 18.12.2014

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