Zoogeography and Faunistics ACTA ZOOLOGICA BULGARICA Research Article Acta zool. bulg., 66 (4), 2014: 477-484

Distribution of the of Montfort, 1810 (: ) in : an Overview

Vanja Marković¹, Jelena Tomović¹, Marija Ilić¹, Margareta Kračun-Kolarević¹, Boris Novaković², Momir Paunović¹, Vera Nikolić3

¹Institute for Biological Research ‘Siniša Stanković’, University of Belgrade, 142 Despota Stefana Blvd., 11060 Belgrade, Serbia; E-mail: [email protected] ²Serbian Environmental Protection Agency, Ministry of Energy, Development and Environmental Protection, Ruže Jovanovića 27a, 11160 Belgrade, Serbia 3Department of Zoology, Faculty of Biology, University of Belgrade, Studentski trg 16, 11000 Belgrade, Serbia

Abstract: The distribution of freshwater of the Theodoxus in Serbia is examined. The study comprises literature data and field surveys carried out at 84 watercourses and 5 reservoirs during 2010-2013. The oc- currence of three species is confirmed:, T. danubialis and T. transversalis. The most widespread species is T. fluviatilis (river nerite). The previously dominant nerite T. danubialis ( nerite) currently persists mainly in hilly and mountain parts of central Serbia and remains in few localities only in large rivers such as the Danube and the Sava. The endangered species T. transversalis (striped ner- ite) shows patchy distribution; the Drina River – Lim River and the Velika Morava River – Južna Morava River – Nišava River systems are revealed as important refugia for this species. Moreover, at some locali- ties at the Nišava River and the Velika Morava River, the striped nerite is found to be the dominant in the present gastropod communities. Regular studies are crucial for the timely prediction of population health and for conservation of freshwater nerite species in .

Keywords: Field survey, literature data, Theodoxus fluviatilis, T. danubialis, T. transversalis

Introduction The gastropod family Neritidae (nerites) is one of the oxygenated, calcium-rich rivers (Fr e t t e r , Gr a h a m oldest lineages of snails. It includes marine, brack- 1962, Gl ö e r 2002). ish and freshwater taxa restricted to the tropical re- There are at least 34 described species of gions and the Southern Hemisphere. The exception the genus Theodoxus (see Bu n j e 2004), although is the genus Theodoxus Montfort, 1810, a fresh- some authors consider that the real diversity is water and brackish group native to Europe and the much lower and recognise only few valid species Mediterranean region. No other freshwater nerites (Bu n j e , Li n d b e r g 2007). The species are charac- are distributed sympatrically with Theodoxus spp. terised by a pronounced phenotypic plasticity re- (Ba n d e l 2001, Bu n j e , Li n d b e r g 2007). The members sulting into substantial morphological variability of this genus are dioecious and semelparous snails (Zh a l a y et al. 2008, Fe h é r et al. 2009), which with biennial or triennial life cycle; they lay capsules makes the morphological identification difficult. In with numerous eggs that hatch a single juvenile per this regard, misidentifications may occur, as in the capsule and the remaining eggs serve as nourishment case of (C. Pfeiffer, 1828) for it (Fr e t t e r , Gr a h a m 1962, Ba n d e l 2001). The in (details in Bu n j e 2007) or T. prevostianus species are herbivores and detritivores, and usually (C. Pfeiffer, 1828) in (Sî r b u , Be n e d e k dwell on hard substrates (preferably rocks) in well- 2005).

477 Marković V., J. Tomović, M. Ilić, M. Kračun-Kolarević, B. Novaković, M. Paunović, V. Nikolić

On the territory of the central Balkans and Table 1. Scale for estimation of the relative abundance of Serbia, three species are present: Theodoxus fluviati- Theodoxus spp. in Serbian waters. lis (L., 1758), T. danubialis and T. transversalis (C. Relative Number of individu- Description Pfeiffer, 1828). abundance als per sample The Balkan region is recognised as one of the 1 present 1 European centres of biodiversity (Sa v i ć 2008). Serbia is part of this remarkable region, the central Balkans, 2 low abundance 2-5 including the silicate west part of the Dinarides and 3 moderate abundance 6-30 the carbonate east parts of the Carpathians and Stara 5 high abundance 31-60 Planina (Balkan) Mountains. Through the north- 7 very high abundance 61-100 ern part, which belongs to the Pannonian Basin, 9 mass present >100 some of the largest European rivers (Danube, Sava and Tisa) flow. The central part is occupied by the investigations in 2010-2013 provided new data. Our Velika and Južna Morava river basins, which serve four-year field campaign covered 84 rivers, streams as a sort of connection with the Vardar River and the or canals as well as 5 lakes. We collected 638 sam- Mediterranean basins. According to Il l i e s (1978), ples from 159 localities. The investigations involved four ecoregions are present on the territory of Serbia. the whole territory of Serbia, including the catchment A recent delineation of these ecoregions is given by areas the rivers Danube, Tisa, Sava, Kolubara, Velika Pa u n o v i ć (2007). Morava, Južna Morava, Zapadna Morava, Ibar, Lim, Despite the numerous investigations, there are Nišava, Mlava, Pek and Timok. The benthic samples still not sufficient data regarding the diversity and dis- were taken by a standard hand-net (semi-quantitative tribution of the freshwater gastropods in this region. technique), preserved in 70% ethanol solution or 4% The first data about the freshwater snails in Serbia formaldehyde solution and further processed in the were provided by Pf e i ff e r (1857) and Mö l l e n d o r f laboratory. The identification of the gastropods was (1873), with recognition of two Theodoxus species. performed by using appropriate keys (Pf l e g e r 1999, Many authors have given their contribution to the Gl ö e r 2002). All collected materials were deposited better knowledge of the freshwater snails in Serbia in the collection of the Benthological Section of the (see details in Ka r a m a n , Ka r a m a n 2007), either Institute for Biological Research ‘Siniša Stanković’ by studying specific groups (e.g. Hydrobiidae, see in Belgrade. Ra d o m a n 1977) or in frames of more or less exten- The relative abundance of species was evalu- sive malacological studies (e.g. He ss e 1929, Ja e c k e l ated according to the scale given in Table 1. et al. 1958, Fr a n k et al. 1990). The most detailed overview is provided by Ka r a m a n , Ka r a m a n (2007). Results and Discussion The aim in this work is to give a compre- Three species belonging to the genus Theodoxus hensive and up-to-date review of the past and recent were reported from Serbian waters. Typical habitats distribution of Theodoxus spp. in the central part of of these species in Serbia are large rivers in the case the Balkans (Serbia). We also try to draw more atten- of T. fluviatilis (Fig. 1a), and smaller rivers for T. tion to the IUCN endangered taxon T. transversalis danubialis and T. transversalis (Fig. 1b). in this part of its range. Finally, providing the list of Theodoxus fluviatilis (Linnaeus, 1758) – river nerite records is necessary for some more detailed studies such as assessment of the intraspecies and interspe- The distribution range of the most widespread cies variability. species of Theodoxus covers the area of Europe from Scandinavia (Sk o o g 1971) to Greece and Crete Island (Sc h u t t 1986, Bu n j e 2005), and from Material and Methods Ireland and the Iberian Peninsula (Lu c e y et al. 1992, The study is based on an extensive data-set, which Ze t t l e r et al. 2004) to the Sea of Azov Sea and the includes the available literature and databases ob- Baltic States (An i s t r a t e n k o et al. 1999). This spe- tained by long-term investigations (internal data of cies is also recorded in Asia, i.e. Anatolia (Yı l d ı r ı m Department of Hydroecology and Water Protection, 1999) and Iran (Gl ö e r , Pe š i ć 2012) while its reports Institute for Biological Research ‘Siniša Stanković’), from Africa are doubtful (Br o w n 2002). According regular monitoring (Annual Water Quality Reports to IUCN database, it is currently regarded as inva- 1999-2010, 2011-2012) or from mixed sources sive in the Danube River basin (Ke b a p ç ı , Va n Da m m e (BAES; Si m i ć et al. 2006). In addition, recent field 2012). The tolerance of T. fluviatilis to organic pollu-

478 Distribution of the species of Theodoxus Montfort, 1810 (Gastropoda: Neritidae) in Serbia an Overview

often with triangular whitish spots (Fig. 2a). The oper- culum is reddish, usually with red edges. The opercu- lar hinge has a broad (lamellar) ridge. (Fig. 2b) The freshwater form (T. fluviatilis) prefers cen- tral and lower parts of larger rivers. It feeds domi- nantly on periphyton, forages mostly at night and requires a hard substrate (preferably rock or wood) (Ba n d e l 2001, Gl ö e r 2002). T. fluviatilis toler- ates moderate organic pollution and very low oxy- gen contents (even below 2 mg/l) but does not en- dure too long periods of drought or ice (Ca r l ss o n Fig. 1a. Locality Golubac (Iron Gate; the Danube River): an 2000, Gl ö e r 2002). Similarly to the majority of example of typical locality of Theodoxus fluviatilis in Serbia the European nerites, T. fluviatilis has biennial life cycle. Sexual maturity is reached at the age of av. 18 months. Up to 150 eggs are laid in a yellowish spherical capsule about 1 mm in diameter, which is attached to any hard substrates, including on other snails. The egg laying occurs in the warmer period (usually from mid-April to October). Normally, only one embryo develops to a miniature adult, depend- ing on temperature (20°C – 65 days, 25°C – 30 days) (Or t o n , Si b l y 1990). Theodoxus danubialis (C. Pfeiffer, 1828) – Danube nerite Theodoxus danubialis is native to the Danube River basin. Besides, there are some documented populations in northern Italy (Bo d o n , Gi o v a n n e l l i Fig. 1b. Prosek (the Nišava River): a locality with abun- 1995). Although the species is not on the IUCN Red dant populations of Theodoxus transversalis and Theo- List, the Danube nerite is critically endangered in doxus danubialis (Ju n g b l u t h , v o n Kn o r r e 2009), (Re i s c h ü t z , Re i s c h ü t z 2007) and tion (Mo u t h o n , Ch a r v e t 1999), which is recognised (http://mollusca.sav.sk/malacology/redlist.htm). as one of major treats to the Danube River, facilitates The shell has a short semi-ellipsoid form (more its spreading. Due to the patchy distribution and phe- globular than in T. fluviatilis) and the slightly promi- notypic plasticity of the river nerite, a vast number nent accounts for about one third of its length of various morphs is present. Among them, the best (Bo d o n , Gi o v a n n e l l i 1995, Pf l e g e r 1999). The col- known are probably the freshwater (f. fluviatilis) and our pattern of the shell is predominantly with brown- brackish (f. littoralis) forms, which differ morpho- ish or purplish zigzag stripes on a yellowish or whit- logically, ecologically, reproductively and behav- ish background (Fig. 3a) but in some cases brown iourally (Gl ö e r 2002, Gl ö e r , Me i e r -Br o o k 2003). or almost black specimens were found (Fe h é r et al. Bu n j e (2005) showed on the basis of genetic studies 2009). The is usually paler than in T. flu- that the taxonomic distinctions of these forms are viatilis and without a broad lamellar ridge (Fig. 3b). insignificant, though some of them have been de- Two main forms, stragulatus and danubialis, are scribed as separate species. Therefore, even the com- distinguished on the basis of the shell morphology, mon freshwater and brackish forms are most likely i.e. more or less conspicuous keel on body is to be only ecomorphs of this highly plastic species present in stragulatus (Gl ö e r 2002). Some authors (Ze t t l e r et al. 2004, Ze t t l e r 2008). consider the populations with more sharp and pro- In general, the shell has an elongate ovoid form, nounced ridges, which are found in the Sava River with only a slightly prominent spire and usually up to drainage (mainly in ), as another valid form, 2½ whorls ( 1 whorl, and teleoconch up to and argue that the stragulatus form is a transitional 1½) (Pf l e g e r 1999, Ba n d e l 2001). The colouring of between the sharp-keeled carinatus and non-keeled is very variable and ranges from almost danubialis forms (Sc h u t t 1988). The closer relations black to various brighter with darker zigzag stripes, between these forms and their taxonomic differenti-

479 Marković V., J. Tomović, M. Ilić, M. Kračun-Kolarević, B. Novaković, M. Paunović, V. Nikolić

or yellowish-brown without bands. The operculum is usually reddish, with red margins, resembling the colouration of T. fluviatilis but the difference is the lack, similarly to T. danubialis, of a broad lamella on the big apophyse ridge (Fig. 4b). T. transversalis, according to a recent phy- a b logenetic analysis (Bu n j e , Li n d b e r g 2007), forms a monotype that is located at the very base of Fig. 2. Theodoxus fluviatilis (Golubac, the Danube River): the Theodoxus genealogy (approximately 30 mil- a) shell; b) operculum lion years old). Therefore, this relict has great sig- nificance in attempts to reveal the phylogeny of the a b whole genus. Historical data The first documented reports of Theodoxus in Serbia were published by Pf e i ff e r (1857) and Mö l l e n d o r ff (1873). Theodoxus transversalis was present in both reports (as transversalis in the Danube River near Golubac) while Mö l l e n d o r ff (1873) reported for the first time the presence of T. danubialis (T. danubialis stragulatus as Neritina Fig. 3. Theodoxus danubialis (Prosek, the Nišava River): stragulata from the Danube). a) shell; b) operculum The presence of the most common European Theodoxus species (T. fluviatilis) was not reported in th a b the Serbian waters until the late 20 and the beginning of the 21st centuries. The first documented findings of the river nerite in Serbia were in 1992 in the Danube River at Karataš, near Kladovo (Ka r a m a n 2001) and in 1993 in the Svrljiški Timok River (Si m i ć 1993). Since then, it has rapidly become the dominant ner- ite species and, along with Lithoglyphus naticoides (C. Pfeiffer, 1828), the dominant gastropod species Fig. 4. Theodoxus transversalis (Prosek, the Nišava in the Danube River and some of its main tributar- River); a) shell; b) operculum ies (Tisa). This status was confirmed by three large international investigations of the Danube River in ation as subspecies is questionable (Ne s e m a n n et al. the period from 2001 to 2007: First Joint Danube 1997, Gl ö e r 2002, Fe h é r et al. 2009). Survey (JDS1), Aquaterra Danube Survey (ADS) It should be noted that T. danubialis is closely and JDS2. Si m i ć , Si m i ć (2004) did not found T. fluvi- related to the extremely rare black nerite (T. prevos- atilis in the investigations at the Iron Gate (conduct- tianus). The latter species is listed as critically en- ed during 1994, 1995 and 2000) while it was present dangered by IUCN and forms a clade native to the and common already in 2001 (JDS1). Therefore, Pannonian basin (Bu n j e 2007, Fe h é r et al. 2009). basing on these data, it could be assumed that the beginning of the 21st century marked the beginning Theodoxus transversalis (C. Pfeiffer 1828) – of a rapid spread of the river nerite in the Serbian striped nerite Danube. Besides its tolerance to organic pollution, The striped nerite is endemic for the Danube the species is also well adapted to the conditions of River basin. Once common, it is currently an endan- increased sedimentation, such as those present at the gered species according to IUCN (Só l y m o s , Fe h é r Iron Gate in the late 20th and the beginning of the 21st 2011), with most of the remnant populations existing century. Another possibility is that T. fluviatilis has in some smaller tributaries, where the water quality been overlooked and misidentified as T. danubialis, is still suitable. at least in some cases. The morphological similarity The periostracum is grey or yellowish-grey, of these two species (T. fluviatilis and T. danubialis) with three dark longitudinal or transverse bands (Fig. could be a reason for taxonomical confusion as it 4a). Rarely, the entire shell may be dark brown, black was speculated in the case of the nerites in northern

480 Distribution of the species of Theodoxus Montfort, 1810 (Gastropoda: Neritidae) in Serbia an Overview

Italy. Bo d o n , Gi o v a n n e l l i (1995) hypothesised that Theodoxus species was found; that was most often the Italian populations of T. danubialis represent a T. fluviatilis. The rare striped nerite species, T. trans- recent (historical) invasion of populations from the versalis, was found as the only Theodoxus species eastern tributaries of the Adriatic Sea. However, it is at Markovac Bridge (the Velika Morava River). On also possible that much older samples of T. danubia- the other hand, at only five sampling sites, all three lis in Italy had simply been overlooked or misidenti- species were found: Bagrdan (the Velika Morava fied as T. fluviatilis( Bu n j e 2007). River), Niš (the Nišava River), Lešnica (the Jadar The literature data reveal that T. danubialis was River), and Priboj and Prijepolje (the Lim River). the most common nerite in Serbia in the past. Some However, all three species were not found together of these old records are quite interesting, as is the case in any of the samples. with Krupačko Vrelo, a karst well near the Nišava The distribution of T. fluviatiliscovers the north River in east Serbia. Two specimens of the Danube part, i.e. the large Pannonian Rivers (the Danube nerite were collected by L. Dokić in the late 19th cen- River, the Tisa River and the Sava River) but it is tury (deposited in the Museum of Natural History in also spreading to the south, through the larger wa- Belgrade, see Ka r a m a n , Ka r a m a n 2007), and after tercourses (the Velika Morava River – Nišava River that, this population was vanished. T. danubialis, in system as the main corridor). On excluding the re- contrast to the results of recent investigations, was ports from sampling sites on the tributaries (the present in the Ibar River, the Svrljiški Timok River Kolubara and the Nera Rivers) situated in proximity and the Crnica River. The species was also found in to the above-mentioned large rivers, some isolated the Karaš canal and in the Vrujci well. findings at the rivers Jadar and Lim are noteworthy. The past distribution of the rare striped nerite, As these were a single sample and specimen find- similarly to that of the mentioned Danube nerite in ings at localities with relatively stable populations the Danube River, is much wider. It was found at of T. danubialis, they may well be accidental. The several sites along the Danube River, from Novi Sad, frequency of occurrence and the relative abundance through the Iron Gate to the Timok River conflu- of T. fluviatilis in Theodoxus samples varied during ence. The striped nerite was also reported for some the investigated period (Figs. 6 and 7). other rivers such as the Sava River and the Moravica The Danube nerite, on the other hand, is bare- River. The more recent reports were from the Iron ly present in the Danube River, absent in the Tisa Gate, where the species was assessed as common River, while in the Sava River coexists with the (Si m i ć , Si m i ć 2004). With regard to this species we should point to the specimen found in the Jelenački stream, Fruška Gora Mountain (Ži v i ć et al. 2000). This finding could indicate the potential presence of an isolated population of that rare . Furthermore, such streams could be considered as possible refugia for some rare taxa in the polluted environment of the Pannonian Plane. Recent investigation The recent field investigations (2010-2013) confirmed the presence of all three Theodoxus spp. and showed their wide distribution. The snails were found in 112 samples at 51 localities in 17 rivers and one lake, which were in the expected range of dis- tribution of this genus in Serbia (Pa u n o v i ć 2007). The localities of those recent records are provided in Fig. 5. The majority of sites were located in the Danube River (16). The river nerite (T. fluviatilis) was present at 33 localities, the Danube nerite (T. danubialis) was registered at 25, and the striped ner- ite (T. transversalis) at 15 localities. At 9 localities, mixed occurrence of different Theodoxus spp. were found. The juvenile specimens at those sites were Fig. 5. Distribution of Theodoxus species in Serbia (the identified as Theodoxus sp. At 17 sites only one last decade)

481 Marković V., J. Tomović, M. Ilić, M. Kračun-Kolarević, B. Novaković, M. Paunović, V. Nikolić

genated. Yet, according to our field investigations, the gastropod community there is composed only of Ancylus fluviatilis Müller, 1774 and Amphimelania holandrii (C. Pfeiffer, 1828) (unpublished data). The absence of Theodoxus species in the Ibar River is intriguing and could be related to the intense anthro- pogenic pressures in the near past, mostly related to the intensive mining and resulting heavy metal pol- lution (details in Mi l a d i n o v i ć et al. 2012). Fig. 6 .Average frequencies of Theodoxus taxa in positive The striped nerite, T. transversalis, was found samples (period 2010-2013) in the Danube River in only one sample (Stari Slankamen, 2012) and was assessed as low abun- dant. This finding is of special interest considering that this species was reported as extinct in the up- per part of the Danube River (Fr a n k 1984, Fr a n k et al. 1990), and extremely rare in the middle and lower parts (Ci o b o i u 2010). In addition, it was not recorded during the large international investiga- tions of the entire Danube River course (JDS1, ADS and JDS2). Pa v l o v a et al. (2013) reported recent- ly the presence of this taxon on two islands in the lower Danube River and supposed that such unpol- Fig. 7. Average relative abundances of Theodoxus taxa in luted river islands might be its preferred habitat. As positive samples (period 2010-2013) this particular site (Stari Slankamen) is located just upstream of the Tisa mouth and is relatively well river nerite and according to our findings, the lat- preserved, with a few river islands nearby, it could ter species has become dominant. However, due to be considered as one of the few remaining refu- its relatively wide distribution in the southern, hilly gia for T. transversalis in this part of the Danube and mountainous part of Serbia (Fig. 5), the Danube River. There were some rare findings in the Tisa nerite is not considered as endangered on the nation- River (site Martonoš) and Crni Timok River (site al level; yet, continuous monitoring is needed. The Zaječar). Therefore, the main refugia for the striped most abundant populations were found at the rivers nerite were found to be the Drina River – Lim River Drina, Lim, Velika Morava, Južna Morava, Nišava and Velika Morava River – Južna Morava River – and Beli Timok. Populations with lower abundanc- Nišava River systems, especially the last one. The es were registered at the rivers Kolubara, Mlava, most representative locality is Prosek, at the Nišava Zapadna Morava, Toplica and some other smaller River (downstream of the Sićevo gorge), in which watercourses. The Nera River and Ram – Banatska mass abundance of the striped nerite population was Palanka (Danube sites) could be singled out because assessed (sample taken during 2013). While at the of the presence of moderately abundant co-occur- mentioned locality two Theodoxus species (T. trans- rence of T. danubialis and T. fluviatilis. The Ram – versalis and T. danubialis) coexist, only the species Banatska Palanka site is characterised by diversity Theodoxus transversalis was found at the Markovac of habitats, unusual for this part of the Danube River Bridge site in the Velika Morava River. The species (Pa u n o v i ć et al. 2005) and by a rich gastropod com- was assessed as moderately abundant during the in- munity. The Nera River is a mountainous river with vestigated period. gravel substrate and relatively unpolluted water. The In conclusion, the data about the distribution case of the Ibar River, a relatively large river flowing and ecological preferences of the species are dynam- across hilly region of central Serbia, could also be ic variables that require regular monitoring. The in- mentioned. With regard to the structure of the habi- creased pollution and degradation of habitats results tats in comparison with some similar watercourses in the overall freshwater diversity loss. The most re- in the region (e.g., the Zapadna Morava and Nišava sistant and tolerable species (T. fluviatilis), expands Rivers), the Ibar River fulfills the primary conditions its range from the large rivers of the Pannonian plain for maintaining communities of grazing prosobranch to mountainous watercourses, suppressing and re- snails. The riverbed includes a lot of hard substrates placing the natives. As this mountainous part of and plenty periphyton, and the water is well oxy- Serbia acts as the main refugia for the other two spe-

482 Distribution of the species of Theodoxus Montfort, 1810 (Gastropoda: Neritidae) in Serbia an Overview cies, among which is T. transversalis listed as endan- river (with increased organic pollution). However, gered by IUCN, the range expansion of T. fluviatilis unless the ecological state of these rivers improves, surely raises some concerns. However, our investi- a possibility of extinction of these Theodoxus popu- gation also revealed the possibility of an increased lations may arise. Therefore, as mentioned above, adaptability of the native species. In support of this regular monitoring of these rivers, as well as of the it may be noted that the most abundant populations entire Serbian surface-water network, is crucial for of T. transversalis were found at the Velika Morava timely prediction of the population health and for River –Južna Morava River – Nišava River system, conservation of rare species. which is under an intense anthropogenic influence. According to the recent ecological status assess- Acknowledgments: This study was supported by the Ministry ments (Ma r k o v i ć et al. 2011, Ko l a r e v i ć et al. 2012), of Education, Science and Technological Development of the the Velika Morava River was a moderately polluted Republic of Serbia, Projects TR 37009 and OI 17173025.

References

An i s t r a t e n k o O. Y., St a r o b o g a t o v Y. I. and V. V. An i s t r a t e n k o Neritidae)–a result of ancestral or recent effects? – North- 1999. Mollusks of the genus Theodoxus (Gastropoda, Pec- Western Journal of Zoology, 8 (2): 300-307. tinibranchia, Neritidae) from the Black and the Azov seas Fe h é r Z., M. Ze t t l e r and K. Sz a b o 2009. An attempt to reveal the basin. – Vestnik Zoologii, 33: 11-19. systematic relationship between Theodoxus prevostianus An n u a l Wa t e r Qu a l i t y Re p o r t (1999-2010). Republic Hydro- (C. Pfeiffer, 1828) and Theodoxus danubialis (C. Pfeiffer, meteorological Service of Serbia. 1828) (, Gastropoda, Neritidae). – Mollusca, 27 An n u a l Wa t e r Qu a l i t y Re p o r t (2011-2012). Serbian Environ- (2): 95-107. mental Protection Agency, Ministry of Energy Develop- Fr a n k C. 1984. Beiträge zur Molluskenfauna Ungarns I: Die ment and Environmental Protection. Donau bei Visegrad. – Zeitschrift für Angewandte Zoologie, Ba n d e l K. 2001. The history of Theodoxus and Neritina con- 71: 29-69. nected with description and systematic evaluation of Fr a n k C., J. H. Ju n g b l u t h and A. Ri c h n o vsz k y 1990. Die Mol- related (Gastropoda). – Mitteilungen aus lusken der Donau vom Schwarzwald bis zum Schwarzen dem geologisch-paläontologischen Institut der Universität Meer: eine monographische Darstellung. Vácrátót, 142 p. Hamburg, 85: 65-164. Fr e t t e r V. and A. Gr a h a m 1962. British prosobranch molluscs. Bo d o n M., M. M. Gi o v a n e l l i 1995. Sulla sistematica e dis- Their functional anatomy and ecology. Ray Society, Lon- tribuzione di Theodoxus danubialis (Pf e i ff e r , 1828) in don, 755 p. Italia. – Bollettino-Museo Regionale di Scienze Naturali, Gl ö e r P. 2002. Süsswassergastropoden Nord-und Mitteleuropas: 13: 493-544. Bestimmungsschlüssel, Lebensweise, Verbreitung, Vol 73. Br o w n D. S. 2002. Freshwater snails of Africa and their medical ConchBooks, Hackenheim, 327 p. importance. CRC Press. 673 p. Gl ö e r P. and C. Me i e r -Br o o k 2003. Süsswassermollusken: Bu n j e P. M. 2004. Diversification and comparative phylogeogra- Ein Bestimmungsschlüssel für die Bundesrepublik phy of neritid gastropods. Unpublished PhD Dissertation, Deutschland. DJN, Hamburg, 138 p. University of California, Berkeley, 411 p. Gl ö e r P. and V. Pe š i ć 2012. The freshwater snails (Gastropoda) Bu n j e P. M. 2005. Pan-European phylogeography of the aquatic of Iran, with descriptions of two new genera and eight new snail Theodoxus fluviatilis (Gastropoda: Neritidae). – Mo- species. – ZooKeys, 219: 11-61. lecular Ecology, 14 (14): 4323-4340. He ss e P. 1929. Schnecken aus dem nördlichen Serbien. – Archiv Bu n j e P. M. 2007. Fluvial range expansion, allopatry, and par- fur Molluskenkunde, 61 (1): 230-240. allel evolution in a Danubian snail lineage (Neritidae: Il l i e s J. 1978. Limnofauna europaea. Gustav Fischer Verlag, Theodoxus). – Biological Journal of the Linnean Society, Stuttgart, 532 p. 90 (4): 603-617. Ja e c k e l S.V., Kl e m m W. and W. M e i s e 1958. Die Land- und Bu n j e P. M., D. R. Li n d b e r g 2007. Lineage divergence of a fresh- Süsswasser-Mollusken der nördlichen Balkanhalbinsel. – water snail clade associated with post-Tethys marine basin Abhandlungen und Berichte aus dem staatlichen Museum development. – Molecular Phylogenetics and Evolution, für Tierkunde Dresden, 23 (2): 141-205. 42 (2): 373-387. Ju n g b l u t h J. H., D. v o n Kn o r r e 2009. Rote Liste der Binnenmol- Ca r l ss o n R. 2000. The distribution of the gastropods Theodoxus lusken [Schnecken (Gastropoda) und Muscheln (Bivalvia)] fluviatilis (L.) and Potamopyrgus antipodarum (Gray) in in Deutschland. 6. revidierte und erweiterte Fassung lakes on the Aland Islands, southwestern Finland. – Boreal 2008. – Mitteilungen der Deutschen Malakozoologischen environment research, 5 (3): 187-196. Gesellschaft, 81: 1-28. Ci o b o i u O. 2010. Gastropoda in the Danube–Carpathian hydro- Ka r a m a n B 2001. Contribution to the knowledge of freshwater graphic space: possible impact of climate change. In Pro- snails (Gastropoda, Mollusca) in Yugoslavian part of the ceedings of the 38th IAD Conference, Dresden (pp. 1-6). Danube. Proceedings Ecological Truth, IX International Fe h é r Z., C. Al b r e c h t , A. Ma j o r , S. Se r e d a and V. Kr i zs i k 2012. Scientific and Professional Meeting., Donji Milanovac, Extremely low genetic diversity in the endangered striped 304-309. (In Serbian). nerite, Theodoxus transversalis (Mollusca, Gastropoda, Ka r a m a n B., G. Ka r a m a n 2007. Catalogus of the freshwater snails

483 Marković V., J. Tomović, M. Ilić, M. Kračun-Kolarević, B. Novaković, M. Paunović, V. Nikolić

(Mollusca, Gastropoda) of Serbia. – Crnogorska Akademija historical development and present status. – Advances in Nauka i Umjetnosti, Glasnik Odjeljenja prirodnih nauka, Arachnology and Developmental Biology, 12: 57-78. 17: 167-222. Sc h u t t H. 1986. The dissemination of Theodoxus species in Ke b a p ç ı U., D. Va n Da m m e 2012. Theodoxus fluviatilis. In: Greece including the islands (Moll. Prosobranch.). – Bio- IUCN 2013. IUCN Red List of Threatened Species. Ver- logia Gallohellenica, 12: 283-290. sion 2013.2. Available via www.iucnredlist. org. Cited 03 Sc h u t t H. 1988. Über eine gekielte Variante von Theodoxus December 2013. danubialis (C. Pfeiffer). – Mitt. dtsch. malakozool. Ges., Ko l a r e v i ć S., J. Kn e ž e v i ć -Vu k č e v i ć , M. Pa u n o v i ć , B. Va s i l j e v i ć , 42: 25-27. M. Kr a č u n , Z. Ga č i ć and B. Vu k o v i ć -Ga č i ć 2012. Seasonal Si m i ć V. 1993. Saprobiological evaluation of the Svrljiški and variations of microbiological parameters of water quality of Trgoviški Timok rivers, based on the macrozoobenthos the Velika Morava River, Serbia. – Archives of Biological communities. Belgrade, Faculty of Biology, MSc Thesis, Sciences, 64 (3): 1017-1027. 263 p. (In Serbian). Lu c e y J., M. L. Mc g a r r i g l e and K. J. Cl a bb y 1992. The distribu- Si m i ć V. M., S. B. Si m i ć 2004. Macroinvertebrates and fishes in the tion of Theodoxus fluviatilis (L.) in Ireland. – Journal of part of the Danube flowing through the Iron Gate National Conchology, 34: 91–101. Park and possibilities of their protection under in situ and Ma r k o v i ć V., A. At a n a c k o v i ć , B. Tu b i ć , B. Va s i l j e v i ć , V. Si m i ć , ex situ conditions. – Archives of Biological Sciences, 56 J. To m o v i ć , V. Ni k o l i ć and M. Pa u n o v i ć 2011. Indicative (1-2): 53-57. status assessment of the Velika Morava River based on Si m i ć , V., S. Si m i ć , A. Pe t r o v i ć , M. Pa u n o v i ć , V. Šo r i ć and V. aquatic macroinvertebrates. – Water Research and Man- Dimitrijević 2006. Biodiversity in aquatic ecosystems in agement, 1 (3): 47-53. Serbia, ex situ conservation (BAES ex situ; http://baes. Mi l a d i n o v i ć B., P. Pa p i ć and M. Ma n d i ć 2012. Groundwater pol- pmf.kg.ac.rs lution with heavy metals in the Ibar alluvium near Raška Sî r b u I., A. M. Be n e d e k 2005. The genus Theodoxus Monfort, (Serbia). – Geoloski anali Balkanskoga poluostrva, 73: 1810 (Mollusca, Gastropoda, Neritidae) in the Romanian 125-130. Inner Carpathian Basin. – Scientific Annals of the Danube Mö l l e n d o r ff O. v o n 1873. Zur Molluskenfauna von Serbien. – Delta Institute, 11: 92-98. Malakozoologische Blätter, 21: 129-149. Sk o o g G. 1971. Variations in the distribution of Theodoxus Mo u t h o n J., S. Ch a r v e t 1999. Compared sensitivity of species, fiuviatilis on stony localities in the northern Baltic proper. genera and families of Molluscs to biodegradable pollu- – Thalassia Jugoslavica, 7: 363-372. tion. – In Annales de Limnologie International Journal of Sk o o g G. 1978. Influence of natural food items on growth and Limnology, 35 (1): 31-39. egg production in populations of Lymnea Ne s e m a n n H., W. Gr a f and A. Sc h m i d t -Kl o i b e r 1997. Be- peregra and Theodoxus fluviatilis (Mollusca). – Oikos, 31 merkenswerte Gastropodenfunde aus dem österreichisch- (3): 340-348. ungarischen Draugebiet. – Nachrichtenbl. Vorarlberger So l y m o s P., Z. Fe h e r 2011. Theodoxus transversalis. In: IUCN malakol. Ges., 5: 28-32. 2013. IUCN Red List of Threatened Species. Version Or t o n R. A., R.M. Si b l y 1990. Egg size and growth rate in Theo- 2013.2. Available via www.iucnredlist. org. Cited 03 doxus fluviatilis (L). – Functional Ecology, 4 (1): 91-94. December 2013. Pa u n o v i ć M. 2007. Composition of macro-invertebrate com- Yı l d ı r ı m M. Z. 1999. The Prosobranchia (Gastropoda: Mollusca) munities as indicator of running waters types in Serbia. species of Turkey and their zoogeographic distribution: 1. PhD Thesis, Faculty of Biology, University of Belgrade, Fresh and brackish water. – Turkish Journal of Zoology, Serbia, 184 p. 23 (3): 877-900. (In Turkish). Pa u n o v i ć M. M., V. M. Si m i ć , D. Ja k o v č e v -To d o r o v i ć and B. Ze t t l e r M. L., J. Fr a n k o w s k i , R. Bo c h e r t and M. Rö h n e r 2004. St o j a n o v i ć 2005. Results of investigating the macroin- Morphological and ecological features of Theodoxus flu- vertebrate community of the Danube River on the sector viatilis (Linnaeus, 1758) from Baltic brackish water and upstream from the Iron Gate (km 1083-1071). – Archives German freshwater populations. – Journal of Conchology, of Biological Sciences, 57 (1): 57-63. 38 (3): 305-316. Pa v l o v a M., M. Ih t i m a n s k a , I. De d o v , V. Bi s e r k o v , Y. Uz u n o v Ze t t l e r M. L. 2008. Zur Taxonomie und Verbreitung der Gattung and L. Pe h l i v a n o v 2013. New Localities of Theodoxus Theodoxus Montfort, 1810 in Deutschland. Darstellung transversalis (C. Pfeiffer, 1828) within European Natura historischer und rezenter Daten einschließlich einer Bibli- 2000 Network on the Islands of the Lower Danube River. ografie. – Mollusca, 26 (1): 13-72. – Acta Zoologica Bulgarica, 65 (1): 121-123. Zh a l a y E. I.., S. V. Me z h z h e r i n , Y. V. Sh u b r a t and A. V. Ga r b e r Pf e i ff e r L. (1857). Bericht über weitere Mittheilungen des Herrn 2008. On species composition of aquatic snail Theodoxus Zelebor. – Malakozoologische Blätter, 3: 175-186. genus (Gastropoda, Neritidae) in the lower Danube: solu- Pf l e g e r V. 1999. A field guide in colour to Molluscs. Blitz Edi- tion of the problem with allozymes analysis. – Scientific tions, Leicester, United Kingdom, 216 p. Bulletin of the Uzhgorod University. Series Biology, 23: Ra d o m a n P. 1977. Hydrobiidae auf der Balkanhalbinsel und Klei- 205-208. (In Ukrainian, English summary). nasien. – Archiv für Molluskenkunde, 107: 203-223. Ži v i ć , I., Z. Ma r k o v i ć and M. Br a j k o v i ć 2000. The change of the Re i s c h ü t z A., P.L. Re i s c h ü t z 2007. Rote Liste der Weichtiere structure of macrozoobenthos in the Jelenački stream under (Mollusca) Österreichs. the influence of pollution. –Ekologija , 35: 105-114. Sa v i ć I. 2008. Diversification of the Balkan fauna: its origine, Received: 29.01.2014 Accepted: 17.07.2014

484