Research Article ISSN 2336-9744 (online) | ISSN 2337-0173 (print) The journal is available on line at www.biotaxa.org/em

The 46 years overlooked species Monacha oshanovae Pintér & Pintér, 1970

DILIAN GEORGIEV

Department of Ecology and Environmental Conservation, University of Plovdiv, Tzar Assen Str. 24, BG-4000 Plovdiv, Bulgaria, email: [email protected]

Received 1 June 2016 │ Accepted 14 June 2016 │ Published online 15 June 2016.

Abstract The species Monacha (Monacha) oshanovae Pintér & Pintér, 1970 (Mollusca: Gastropoda) was re-collected during the period 2007-2016 from various localities of Bulgaria, including and its type locality. New information on this very poorly known species concerning its morphology, anatomy, distribution, biology and ecology was provided. Short notes on the Bulgarian species of the genus Monacha were made.

Key words: Hygromiidae, Balkans, distribution.

Introduction

The species Monacha (Monacha) oshanovae Pintér & Pintér, 1970 (Mollusca: Gastropoda) was described from SW Bulgaria from the area of Skrat village at the north foothills of Belasitsa Mts. near the Macedonian border. It was also found in Petrich town by same authors (species collected by N. Oshanova on 23- 24.04.1969). For a long time it was surprisingly omitted from all the lists of valid species for Bulgaria or was considered as one “with unclear status” (Damyanov & Likharev, 1975; Dedov, 1998; Hubenov, 2005; Irikov, 2008; Irikov & Eröss, 2008). The same situation was and concerning the always ignored record of Monacha (Monacha) frequens (Mousson, 1859) which was proven by Pintér & Pintér (1970) from the same area in the country by anatomical study and providing of a drawing of the genitalia. The very poor knowledge on the Bulgarian Monacha species provoked my interest to revise my old collections and to initiate some new collecting trips. In this paper I provide new information on the very poorly known species Monacha oshanovae concerning its morphology, anatomy, distribution, biology and ecology.

Material and Methods

Snails were collected during the period 2007-2016 from various localities in Bulgaria. Animals were preserved in 75% ethanol, some of which were firstly over narcotized with diethyl ether. Dissections were made under binocular microscope Ziess. Some of the genital systems or their parts were observed or/and prepared in slides with glycerin. Photographs of the genital organs were made with Canon camera in glycerin, and in some cases in ethanol. Measurements were taken by caliper. For calculations of standard deviation, average values and variance Microsoft Excel was used. The material is deposited in the author`s

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collection, and the collections of Ivailo Dedov, Viktor Gashtarov, and The Regional Natural History Museum – Plovdiv. Some of the collected live specimens during 2016 were kept in small terrariums for observations on their behavior and biology, and are still living (June 2016). The taxonomic important features and the terminology follow Hausdorf (2000). Abbreviations used: D – shell diameter, H – shell height, AH – aperture height, LW – last (body) whorl width, ap – appendicula; ep – epiphallus; fl – flagellum; gm – glandulae mucosae; p – penis; sod – spermoviduct; vd – vas deferens, rs – receptaculum seminis, bc – bursa copulatrix, StdDev – standard deviation, Var – variance

Results and Discussion

Monacha (Monacha) oshanovae Pintér & Pintér, 1970

Material examined: Sashtinska Sredna Gora Mts.: 3 shells, 03.11.2007, near the railway station of Strelcha town, UTM KH80, N42 29 34.0 E24 19 53.0, 399 m a.s.l.; 3 shells, 21.11.2007, N site of Hisarya town, UTM LH11; 2 spec. in ethanol, 24.11.2007, banks of Luda Yana River, W of Dyulevo vill., UTM KH80, N42 26 55.5 E24 20 02.1, 362 m a.s.l.; all D. Georgiev, S. Stoycheva leg.; (all published as Monacha sp. in Georgiev & Stoycheva, 2009); 1 live spec., 28.04.2016, near the river E of Mihiltsi vill., N42 31 29.2 E24 48 43.6, 307 m a.s.l., D. Georgiev leg.; 15 shells, 28.04.2016, near small flood E of Hisarya town, N42 31 14.2 E24 43 55.7, 440 m a.s.l.; Sarnena Gora Mts.: 1 spec. in ethanol, 2 live spec., 30.04.2016, near small stream E of Shanovo vill., N42 33 42.2 E25 38 56.3, 297 m a.s.l.; 3 shells, 1 live spec., 30.04.2016, near small stream N of Edrevo vill., N42 35 51.3 E25 48 42.4, 267 m a.s.l., all. D. Georgiev leg.; East Rhodopes Mts.: 1 spec. in ethanol, 11 dry shells, 03.04.2016, Zimzelen village surroundings, N41 39 00.4 E25 23 50.7, 278 m a.s.l.; 4 spec. in ethanol, 11.04.2016, same locality; 1 spec. in ethanol, 5 shells, 23.04.2016, near water source below the quarry of Beli Plast vill., N41 46 52.3 E25 26 03.6, 458 m a.s.l., all D. Georgiev leg.; Below Kozhuh Mt., near the hot springs: 1 live specimen observed, not collected, 21.02.2016; more than 20 dry shells, 06.05.2016, D. Georgiev, V. Gashatov, V. Ivanova leg.; Type locality: 3 shells, 06.05.2016, near the river in E part of Skrat vill., D. Georgiev leg.; 2 spec. in ethanol, 1 live spec., 1 dry shell, wet meadow W of Skrat vill., N41 22 34.5 E22 58 00.3, 280 m a.s.l., D. Georgiev, V. Ivanova leg.

Figure 1. Shell of Monacha oshanovae: fresh shell of dissected specimen from Beli Plast village area, E Rhodopes Mts. (D = 7.3 mm).

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THE 46 YEARS OVERLOOKED SPECIES MONACHA OSHANOVAE

Figure 2. Exterior view of Monacha oshanovae: typical, brownish one with more globular shell from Mihiltsi vill. area, sole and back side view (left), and specimen with almost entire black mantle and depressed conical shell from the area of Zimzelen vill. area (right).

Table 1. Shell measurements and proportions of M. oshanovae from various localities. For abbreviations see Material and Methods.

Locality H AH D LW H/D AH/H LW/D Mihltsi, 28.04.16 6.9 4.71 8.56 2.63 0.81 0.68 0.31 Shanovo, 30.04.16 5.24 3.60 7.54 2.66 0.69 0.69 0.35 Shanovo, 30.04.16 5.27 3.40 7.48 3.08 0.70 0.65 0.41 Skrat, 06.05.2016 6.73 4.45 9.24 2.53 0.73 0.66 0.27 Skrat, 06.05.2016 7.38 4.94 10.10 3.98 0.73 0.67 0.39 Luda Yana, 24.11.2007 5.00 3.30 6.80 2.20 0.74 0.66 0.32 Hisarya, 28.04.2016 5.00 3.10 6.10 2.50 0.82 0.62 0.41 Hisarya, 28.04.2016 4.80 3.20 7.40 2.10 0.65 0.67 0.28 Edrevo, 30.04.2016 5.20 3.20 7.30 2.10 0.71 0.62 0.29 Zimzelen, 11.04.2016 4.80 3.20 7.10 1.80 0.68 0.67 0.25 Zimzelen, 11.04.2016 4.90 3.70 7.60 1.80 0.64 0.76 0.24 Beli Plast, 23.04.2016 5.70 3.30 7.30 1.60 0.78 0.58 0.22 Average 5.35 3.49 7.46 2.26 0.72 0.66 0.31 StdDev 0.87 0.61 1.16 0.75 0.06 0.04 0.07 Var 0.76 0.37 1.35 0.56 0.00 0.00 0.00

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Shell: The shell is small, fragile, transparent, brown-reddish-corneous (rarely brown-greenish), depressed conical-globular to depressed conical (more rarely), with 4.5-5.5 convex whorls. The protoconch is smooth, brown-reddish or white. Teleoconch is with well distinct spiral striae sometimes forming smooth irregular ribs, and has scattered hair scars. Body whorl is rounded or slightly angular with slightly visible white band at the periphery, narrow, slightly widening at the end. Aperture is almost circular, upper insertion of the peristome usually distincly descending, peristome sharp, white with well visible, wide red margin. Umbilicus is narrow but open, partly covered by the reflected columellar edge (Fig. 1). Shell measurements and proportions (Table 1): D = 6.1-10.1 mm, H = 4.8-7.4 mm, AH = 3.1-4.9 mm, LW = 1.6-3.1 mm, H/D = 0.6- 0.8, AH/H = 0.6-0.8, LW/D = 0.2-0.4.

Soft body: The whole animal is dark pigmented. Head is grey-brown, the tentacles are dark grey. The mantle is black, dark grey or brownish with grey or white dots and spots, or having reticulation or/and spots of white, yellow or paler brown-reddish color. The very dark specimens, with more depressed shells look like Trochulus hispidus (Linnaeus, 1758). Always the mantle has light whitish periphery. The sole has white- yellow medial zone and darker, greyish-brown lateral parts. Sometimes the backside of the sole could be white-yellow and at the tip (Fig. 2, 3).

Figure 3. Soft body and genitalia of specimens from E Rhodopes Mts.: 1 – mantle, dorsal and ventral view, 2 – sole view, 3 – penis view of the genitalia of specimen from Zimzelen vill., 4 – genitalia and eggs of specimen from Beli Plast vill.

Genitalia: The atrium is thick and short. The penial papilla is cylindrical. The penis is thicker than the epiphallus and well delimited. It is connected to the epiphallus by weak muscle bands. The epiphallus is 1.3- 1.6 times as long as the flagellum. The appendicula inserts at the proximal section of the vagina. Its long basal section could vary in width in one and a same population. It can be two main types: only indistinctly

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THE 46 YEARS OVERLOOKED SPECIES MONACHA OSHANOVAE delimited against the terminal section and to looks almost cylindrical or to be well delimited against the terminal section but transitional shapes between them were also observed. Both sections are about same length. The two slightly ramified glandulae mucosae insert at the distal section of the vagina. The bursa of the bursa copulatrix is elongated oval (Fig. 3, 4). The right ommatophoral retractor passes to the left of the genitalia. Measurements (3 specimens): 1. (23.04.16, Beli Plast, D = 7.3 mm): p = 1.8 mm, ep = 2.8 mm, fl = 2.2 mm, p/ep = 0.6, fl/ep = 1.3; 2. (14.11.2007, Luda Yana, shell not measeured): p = 1.8 mm, ep = 2.7 mm, fl = 1.8 mm, p/ep = 0.7, fl/ep = 1.5; 3. (06.05.2016, Skrat, type loc., D = 10.1 mm): p = 2.5 mm, ep = 3.2 mm, fl = 2.0 mm, p/ep = 0.8, fl/ep = 1.6.

Fig. 4. Genitalia of Monacha oshanovae, specimen from surroundings of Luda Yana River, W of Dyulevo village, collected on 24.11.2007 (photographed in glycerin).

Eggs: Eggs are oval, whitish, with diameter of 1.75 mm, laid in clutches of about 10-20 (Fig. 5).

Habitat: The species is hygrophilic but relatively draught resistant. It tend to live at moisture grassy areas along rivers, creeks and water sources, low to the ground at the base of the vegetation (Fig. 6, 7). It inhabits open, grassy terrains occupied by tall wet loving species such as Mentha sp., Rannunculus sp., Scirpus sp., Carex sp., Persicaria sp., Dactylorhiza sp., Trifolium sp., Plantago sp., Taraxacum sp., Urtica sp., and various Poaceae species (Fig. 6, 7). It is more rarely climbing on the grass vegetation compared to simpatric living species from the Monacha cartusiana – complex, and could be found on the ground, among the grass, sand and detritic parts. It seems that M. oshanovae is not connected to limestone areas as it was found on zeolite, riolite, and silicate terrains. It was registered at hilly areas at altitudes of 200-500 m a. s. l. The species was associated mainly with following terrestrial mollusc species: Deroceras sturanyi, Oxychilus glaber, Monacha cartusiana – complex, Helix lucorum, Cepaea vindobonensis, Oxyloma elegans, and Zonitoides nitidus.

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Figure 5. Eggs and juvenile of Monacha oshanovae: 1 – eggs laid in terrarium, 2 – hatchling in terrarium (Shanovo vill.), 3 – eggs found in nature (Skrat vill.).

Diet: Eating green and decaying leafs and detritus. The specimens kept in terrarium accepted to eat: Lactuca sativa, Rannunculus sp. (only decaying leafs), Mentha sp., Trifolium sp., decaying leafs of Corrylus sp., cucumber (Cucumis sativus), and aquarium fish food.

Activity: Mostly active during night and in the morning till about 8.00-9.00h. Most of the terrarium raised animals stopped activity after 7.30h. when attached to the walls, plant leafs or dig into the sand on the bottom.

Figure 6. Habitat view of the type locality of M. oshanovae – wet meadow, near the road, W of Skrat vill., foothills of Belasitsa Mts., near the Macedonian border (06.05.2016).

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THE 46 YEARS OVERLOOKED SPECIES MONACHA OSHANOVAE

Figure 7. Typical habitat of Monacha oshanovae (near Shanovo vill.), and specimens as they were found in the nature: brown-greenish specimen, Shanovo (left) and brown-reddish one, Mihiltsi (right) villages.

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Biology: In the albumen gland of one of the dissected specimens from Zimzelen village area, collected on 03.04.2016, 5 eggs were found (Fig. 5). Almost developed eggs were found and in one of the specimens from Skrat, collected on 06.05.2016. On same date and locality a laid clutch of about 20 eggs was found among roots of hygrophyte plants and detritus (Fig. 5). Animal from Shanovo vill. kept in terrarium laid eggs on 04.05.2016, possibly during night (observed at 7.20h in the morning, and adults were already not active), air temperature was 17°C. Two clutches with 10 eggs each were observed. One was deposited into dry leaf of Corrylus, and another, few millimeters below surface of wet sand at the bottom of the terrarium (Fig. 5). The eggs laid on the Corrylus leaf were accidently dried for about two days and were shriveled. After moisture they quickly (for about two minutes) regained their original shape, absorbing water. All of them survived, and began to hatch on 18.05.2016 in the morning at 18.8°C air temperature. It can be supposed that species has two years cycle, lay eggs during spring mainly in the period of April-May, and youngsters are hatched after about two weeks.

Distribution: The hilly areas and mountain foothill slopes of S Bulgaria between 200-500 m a. s. l. Recorded at the foothills of Belasitsa Mts., Kozhuh Mt., Sashtinska and Sarnena Sredna Gora Mts., and East Rhodopes Mts (Fig. 8). Could be expected and in wider area of similar conditions, also in Greece and Republic of Macedonia. It can only be speculated that being hygrophilic, M. oshanovae is relatively resistant to drowning and could successfully disperse by running waters.

Figure 8. Known localities of M. oshanovae in Bulgaria: 1 – Skrat vill. (Type locality); 2 – Petrich town and the hot springs below Kozhuh Mt., 3 – Strelcha town and Dyulevo village, 4 – Hysaria town and Mihiltsi village, 5 – Shanovo village, 6 – Edrevo village, 7 – Zimzelen and Beli Plast villages.

Discussion

The ecological preferences of M. oshanovae being hygrophilic species are rare in the genus Monacha which is consisted mainly by xerophilic species. The other species strongly connected to wet marshy meadows is Monacha (Paratheba) pusilla Hausdorf, 2000, known from the alpine areas of Anatolia in Turkey (Hausdorf, 2000). Morphologically M. oshanovae resembling with Monacha pilosa Pintér, 1968 or/and Monacha (Monacha) ovularis (Bourguignat, 1855), from which it differs mainly by the more depressed shell, not so

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THE 46 YEARS OVERLOOKED SPECIES MONACHA OSHANOVAE thick shell wall with not so dense, and well visible hair scars. While the genitalia of M. ovularis is not known the one of M. pilosa was studied by Körnig (1983), and Irikov (2008). It clearly differs by very short penis and flagellum compared to the epiphallus, complex structure of the appendicula, and strongly oval bursa copulatrix. M. oshanovae is similar and with M. solidior from which it differs by its smaller and darker body and shell with not so many hair scars, shorter atrium, more proximal insertion of the appendicula, narrower penis, and shorter flagellum according the penis and the epiphallus.

Short notes on the Monacha records in Bulgaria It can be summarized that 9 species from the genus Monacha are known till now from Bulgaria: Monacha carascaloides (Bourguignat, 1855), Monacha cartusiana (O. F. Müller, 1774), Monacha claustralis (Menke, 1828), Monacha ocellata (L. Pintér, 1968), Monacha oshanovae I. Pintér et L. Pintér, 1970, Monacha solidior (Mousson, 1873), Monacha venusta (L. Pintér, 1968), Monacha frequens (Mousson, 1859), and Monacha pilosa Pintér, 1968. Monacha (Monacha) claustralis (Menke, 1828) and Monacha (Monacha) carascaloides (Bourguignat, 1855). From literature survey it is evident that most widely distributed species in Bulgaria are M. claustralis and M. carascaloides. From both M. carascaloides is much rare and lacking from many anthropogenic sites, especially in some large areas of towns and cities. M. claustralis possibly occupy almost the whole territory of Bulgaria from sea level up to about 2000 m alt. Monacha (Monacha) cartusiana (O. F. Müller, 1774). Only two sure records of this species in Bulgaria are known: the Stara Zagora city (Upper Thracia), and in and around Hisarya town (Sashtinska Sredna Gora Mts.) (Georgiev, 2008; Georgiev & Stoycheva, 2009). Even that it looks like M. claustralis is much more widely distributed than M. cartusiana in the country conclusions concerning both species distribution has to be made with caution. Monacha (Monacha) solidior (Mousson, 1873). M. solidior was recorded by anatomical studies only in four localities of S Bulgaria: Zlatna Ribka Resort, Arkutino and Tsarevo, at the Black Sea coast (Irikov & Mollov, 2015) and in Svetiiliiski Heights of Upper Thracian Lowland (Georgiev, 2006). I consider that without any anatomical investigations all old records of M. carascaloides or M. solidior are not sure but most probably they refer to the first species, which large form (D>16 mm) is well distinguished. Revising my materials, contrary to some previous studies, I consider M. carascaloides as more widely distributed species in Bulgaria from both. Monacha (Monacha) frequens (Mousson, 1859). The species was recorded for Skrat and Boboshevo villages in W Bulgaria by Pintér & Pintér (1970) but later without any convincing arguments was omitted in all species lists for Bulgaria. It is very frequent in neighboring areas of SW Balkans (Welter-Schultes, 2012), and occur in the nearby Greece in Epirus (Reischütz & Sattman, 1990). I recollected M. frequens on 06.05.2016 from Skrat village. Monacha (Monacha) ovularis (Bourguignat, 1855) – species complex, Monacha pilosa Pintér, 1968. Pintér & Pintér (1970) stated that the taxonomic statute of Monacha pilosa Pintér, 1968 is unclear. Till now it is not proven anatomically that this species is conspecific with previously described Monacha (Monacha) ovularis (Bourguignat, 1855) from the area of Eski-Baba, Turkey. According Hausdorf (2000) there are no constant conchological differences between many small Monacha species in Turkey, or they just slightly differ each other by shell morphology: M. crenophila (L. Pfeiffer), all species from the group of M. hemitricha (Hesse), M. ascania (Hausdorf), M. bythinica (Hausdorf), M. margarita (Hausdorf), and also the small M. stipulifera (Hausdorf) are also similar. So, till anatomy of M. ovularis from Eski-Baba is unknown, it cannot be stated that M. pilosa is not a true species. Anatomically similar specimens which genital system morphology fits with this one described for M. pilosa from Ropotamo River are reported by Irikov (2008) from some areas at the Black Sea Coast. Specimens from Sakar Mts. recorded by Georgiev (2005) were determined only as empty shells, and it is more correctly to be noted as belonging to “Monacha (Monacha) ovularis (Bourguignat, 1855) – species complex”. Both M. ocellata and M. venusta occur possibly along the coastal area of the Bulgarian Black Sea. Future studies are needed to clear their actual distribution in the country. The diversity of Monacha species in Bulgaria is possibly much higher than considered before. The new finds of Monacha oshanovae after 46 years from its description by Pintér & Pintér (1970) stress that detailed studies are needed to accept validity or reject the occurrence of a particular species in an area. It also can be speculated that some new species could be found in the country during future investigations.

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Acknowledgements I am grateful to my colleagues Barna Páll-Gergely, Zoltán Fehér, Zoltán Erőss, Ulrich Schneppat, Ivailo Dedov, and Atanas Irikov for the useful discussions, and literature sources. I also wish to thank my wife Veselina Ivanova for the help during some collections and the nice trips during spring of 2016.

References

Damyanov, S. & Likharev, I. (1975) Terrestrial snails (Gastropoda terrestria). Fauna Bulgarica 5., Marin Drinov Publ., Sofia, 425 pp. (in Bulgarian) Dedov, I. (1998) Annotated check-list of the Bulgarian terrestrial snails (Mollusca, Gastropoda). Linzer Biologische Beiträge, 30(2), 745–765. Georgiev, D. (2005) The mollusks (Mollusca: Gastropoda et Bivalvia) of Sakar Mountain (Southern Bulgaria): A Faunal Research. Scientific Studies of the University of Plovdiv, Biology, Animalia, 41, 5–12. Georgiev, D. (2006) A Contribution to the Knowledge of the Malacofauna of Sveti Iliiski Heights (South- Eastern Bulgaria). Scientific Studies of the University of Plovdiv, Biology, Animalia, 42, 13–20. Georgiev, D. (2008) Habitat Distribution of the Land Snails in One Village Area of the Upper Thracian Valley (Bulgaria). In: Velcheva I., A. Tsekov (Eds.). Anniversary Scientific Conference of Ecology, Proceedings, 1 November 2008, Plovdiv, 147–151. Georgiev, D. & Stoycheva, S. (2009) The molluscs and their habitats in Sashtinska Sredna Gora Mts. (Southern Bulgaria). Malacologica Bohemoslovaca, 8, 1–8. Irikov, A. (2008) Genus Monacha Fitzinger 1833 in Bulgaria (Gastropoda, Pulmonata, Hygromiidae). Linzer Biologische Beitrage, 40, 1, 785–811. Irikov, A. & Eröss Z. (2008) An updated and annotated checklist of Bulgarian terrestrial gastropods (Mollusca: Gastropoda). Folia Malacologica, 16(4), 199–207. Irikov, A. & Mollov, I. (2015) Terrestrial gastropods (Mollusca, Gastropoda) of Strandzha Mountain and the Black Sea coast (Bulgaria and Turkey). Historia Naturalis Bulgarica, 21, 13–48. Hausdorf, B. (2000) The genus Monacha in Turkey (Gastropoda: Pulmonata: Hygromiidae). Archiv fur Molluskenkunde, 128(1/2), 61–151. Hubenov, Z. (2005) Malacofaunistic diversity of Bulgaria. In: Petrova A. (Ed.), Current state of Bulgarian biodiversity – problems and perspectives, Bulgarian Bioplatform, Sofia, 199–246. (in Bulgarian). Körnig, G. (1983) Beitrag zur Ökologie und Zoogeographie bulgarischer Landgastropoden. Malakologische Abhandlungen Staatliches Museum für Tierkunde in Dresden, 9(5), 31–52. Reischütz, P. & Sattman, H. (1990) Beitrage zur Molluskenfauna des Epirus, II. Annalen des Naturhistorischen Museums in Wien, 91B, 253–272. Welter-Schultes, F. (2012) European non-marine molluscs, a guide for species identification. – Planet Poster Editions, Göttingen, 674 pp.

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