A Taxonomic Overview of Protozoa of Shkodra Lake, Albania

Dr. Violeta Alushi

Department of Biology-Chemistry, University of Shkodra “Luigj Gurakuqi”,Shkoder, Albania [email protected]

Abstract During the last decade, at the Department of Biology & Chemistry of Shkodra University, the field for studying the protozoa of fresh waters, was opened. Protozoa play an important role in the stability of water ecosystems, particularly through its involvement in the decay of macrophytes, in the decreasing of the level of water saprobia and serving as a link in the transfer of food and energy in food chains of aquatic ecosystems. From the previous studies on Protozoa of Shkodra Lake, we can mention: Alushi (2004, 2005, 2006, 2011) and Alushi & Dhora (2006, 2007, 2008); Rakaj, Alushi & Dhora (2007), Mani (2010), Mani & Dhora (2011) as well as for the species found in the Montenegrin part of the Lake, we mention: Gessner (1934), Nedejlkovic (1959), Milovanovic & Zivkovic (1965), Petkovic St. (1977, 1980, 1981), Petkovic, Sm & Petkovic, St (1984) dhe Nikolic et al. (2001). This study shows the results in terms of the taxonomic aspect of Protozoa of Shkodra Lake, meaning the identification and positioning in the taxonomic system. Water samples for the study of protozoa were taken during the years 2004 – 2011 and from various sampling stations such as from the shore waters, from amidst the submerged aquatic vegetation and from plankton. The identifying issue is closely related with the variety of cultivation methodologies used to identify the potential taxonomic situation of Protozoa of the Lake. For this purpose, we’ve used the cultivation of water samples taken from the Lake in 10 different nutritious terrains. The identification is done with microscope and with the help of the contemporary bibliography. A total of 227 protozoa species of Shkodra Lake are identified and a description of their features is given. Along with 50 species previously identified by foreign scientist, from which 16 of them were also found by us, the total number of Protozoa of Shkodra Lake now is 261. All species are listed in the taxonomic system taken from S. M. Adl et al. 2005, reflecting the latest taxonomic changes. Several stances are held in terms of some taxonomic dilemmas in the amoeba group as well as in some genus and ciliophora species. The largest number of species belongs to the Ciliophora phylum, with 174 species followed by Amoebozoa phylum with 56 species. The classes with the largest number of species are: Oligohymenophorea class with 54 species. The Arcellinida order is represented with 33 species and the Sessilida order with 24 species. We’ve given the bibliography of publications about the protozoa of Shkodra Lake. Generally speaking, the Protozoa of the lakes, rivers, lagoons etc. are little known in Albania. Our study group of the Protozoa has programmed the extension of studies for the protozoa of Albania and the institutionalization of the contribution in Balkan level.

Keywords: Protozoa, Shkodra Lake, taxonomic system, identification, description, species

Introduction The Propozoa have been the least studied group of living organisms from Shkodra Lake just before the last decade. Consequently, a deep study of them has been considered necessary, especially on their taxonomic aspect.

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 1 During the last decade, for the first time the new field of study for this group was opened at the Department of Biology of Shkodra University, making it possible to fill the gap between the importance of these organisms and the lack of studies about them. Until now a considerable number of studies are done by some authors of which we can mention the doctorate of V. Alushi (2006), 3 Masters works and the valuable contribution of Prof. Dr. Dhimiter Dhora, who has always encouraged and supported the studies on Protozoa of Shkodra Lake. Protozoa of Shkodra Lake have a high diversity of species and they play an important role at the ecosystem. In this survey, a full list of Protozoa of Shkodra Lake is presented. It has been fulfilled with the data from the latest studies about this group and for the first time it’s given the bibiliography of studies about the Protozoa of the Lake.

Material and methods This study is based on all the published literature on the protozoa of the Shkodra Lake, for both the Albanian and Montenegrin part. 100% of the publications given in the bibliography are analysed in details. All species of protozoa are listed according to the taxonomy given by Adl et al 2005, also the synonymy has been eliminated. Most of the publications belong to the author ( Alushi, V) which are realized during the last decade. Most of protozoa species are found by us. Our study for the identification of protozoa of Shkodra Lake is done during different periods of time, from 2003 – 2011 and the water samples are taken in 11 stations. The stations represent different habitats of shores, submers vegetation and plankton. The selection of sampling stations is done by considering the Lake seasonal developments and the influence of various ecological factors. The stations are: S 1 - Zogaj, shore: S 2 - Zogaj, in open waters, 200 - 400m away from the shore; S 3 - Shiroka, shore, S 4 - Shiroka, among macrophytes. S 5 - Gril, among macrophytes. S 6 - Kamicë, shore. S 7 - Kamicë, among macrophytes. S 8 - Kamicë, 600-800 m from the shore, in open waters. S 9 – Shegan’s Spring, S 10 – Shegan’s Spring, shore, S 11 – Shegan’s Spring, among macrophytes. The stations are given in Fig 1. Sampling is done mainly based on the methods provided by Magazzu (1978). Different techniques were used to cultivate the samples from the Lake in order to identify species potencially present in the Lake. The species found in the cultures are identified 6 – 7 times during a period of three months after their cultivation. The study was made through a detailed microscope observation, identification is based mainly on Streble & Krauter (2002), Lee et al (2000), Foissner et al (1999), Kumar, A. & Dalby, A. P., (1998), Smirnov, A & Brown, S (2004).

10 9 11

6 8 7

5

2 1 4 3

Fig. 1. Protozoa sampling points of Shkodra Lake

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 2 Results and discussion

1- Bibliography of Publications of Shkodra Lake Protozoa In the bibliography titles treating Protozoa of Shkodra Lake are listed 34 publications. All publications are listed in the end of the article. As it’s shown from Fig. 1, 11, publications are done before 2003, with 23 publications being done after 2003. Fig. 2 shows that 68% of publications are done by Albanian authors and 32% by others. Fig. 3 shows the number of species found by us and by foreign authors and the total number of the protozoa of Shkodra Lake.

Fig. 1. Number of publications before and after 2003

Fig. 2. Number of publications done by Albanian authors and by others

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 3

Fig. 3. Total number of protozoa species, those found by us and by others

2. The list of Protozoa of Shkodra Lake All Protozoa species are listed in the following. Fig 4 shows that the Ciliophora are represented with the highest number of species, 174 of them, followed by Amoebozoa with 61 species.

Fig. 4. Number of Protozoa species

Fig. 5 shows the number of species of Amoboezoa; Arcellinida has the largest number of species (33 of them).

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 4

Fig. 5. Number of species for Amoboezoa Fig. 6 shows the number of species of Ciliophora. Oligohymenophorea is represented with the largest number of species, 54 of them

Fig 6. The number of species for Ciliophora

List of Shkodra Lake Protozoa

PROTISTA AMOEBOZOA

Tubulinea Tubulinida Amoeba BORY DE ST. VINCENT,1822 Amoeba gorgonia PENARD 1902 Amoeba proteus (PALLAS,1766) Amoeba hylobates PENARD,1902

Trichamoeba FROMENTEL,1874 Trichamoeba villosa WALLICH,1863 Trichamoeba clava SCHAEFER, 1926 Trichamoeba pilosa (CASH, 1904) Trichamoeba sp.

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 5 Polychaos SCHAEFER, 1926 Polychaos fasciculatum (PENARD, 1902)

Metachaos SCHAEFER, 1926 Metachaos laureata PENARD, 1902 Metachaos gratum SCHAEFFER, 1926

Dinamoeba LEIDY, 1874 Dinamoeba mirabilis LEIDY,1874

Testacealobosia Arcellinida Penardochlamus DEFLANDRE, 1953 Penardochlamus arcelloides (PENARD, 1904)

Pyxidicula EHRENBERG, 1838 Pyxidicula operculata (AGARDH, 1827)

Arcella EHRENBERG, 1832 Arcella gibbosa PENARD, 1890 Arcella vulgaris EHRENBERG, 1830 Arcella discoides EHRENBERG, 1843 Arcella dentata EHRENBERG, 1830 Arcella haemisphaerica PERTY, 1852 Arcella megastoma PENARD 1902 Arcella conica (PLAYFAIR, 1917)

Difflugia LECLERC, 1815 Difflugia lobostoma LEIDY 1879 Difflugia pyriformis PERTY, 1849 Difflugia acuminata EHRENBERG, 1838 Difflugia corona WALLICH, 1864 Difflugia limnetica LEVANDER, 1900 Difflugia leidyi WAILES, 1913 Difflugia pyriformis bryophila PENARD, 1902 Difflugia urceolata CARTER, 1864 Difflugia elegans PENARD, 1890

Centropyxis STEIN, 1857 Centropyxis aculeata (EHRENBERG, 1830) Centropyxis ecornis (EHRENBERG, 1841) Centropyxis hemisphaerica (BERNARD, 1879) Centropyxis arcelloides PENARD, 1902

Paraquadrula DEFLANDRE, 1932 Paraquadrula irregularis DEFLANDRE, 1932

Lesqueruesia SCHLUMBERGER, 1845 Lesqueruesia spiralis (EHRENBERG, 1840) Lesqueruesia modesta RHUMBLER, 1896

Hyalosphenia STEIN, 1859 Hyalosphenia papilio LEIDY, 1875 Hyalosphenia elegans LEIDY, 1879

Heleopera LEIDY, 1879 Heleopera sp. aff. sphagni Heleopera petricola LEIDY, 1879

Nebela LEIDY, 1874

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 6 Nebela griseola PENARD 1890 Nebela tubulosa PENARD, 1890 Nebela lageniformis PENARD, 1890

Cryptodifflugia PENARD, 1890 Cryptodifflugia sp. aff. oviformis

Flabellinea Dactylopodida Mayorella SCHAEFER, 1926 Mayorella sp. aff cantobrigensis Mayorella bigemma SCHAEFFER, 1918

Astramoeba VEJDOVSKY, 1881 Astramoeba radiosa (EHRENBERG, 1830)

Dactylosphaerium HERTWIG & LESSER 1874 Dactylosphaerium vitraeum HERTWIG & LESSER 1874

Hyalodiscus HERTWIG & LESSER, 1874 Hyalodiscus rubicundes HERTWIG & LESSER, 1874

Thecamoebida Thecamoeba FROMENTEL, 1874 Thecamoeba verrucosa (EHRENBERG, 1838)

Cochliopodium HERTWIG & LESSER 1874 Cochliopodium bilimbosum AUERBACH 1856 Cochliopodium granulatum PENARD, 1890

Pelomyxa GREEFF, 1874 Pelomyxa schiedti SCHAEFFER, 1918 Pelomyxa palustris GREEFF, 1874 Pelomyxa tertia (GRUBER, 1885)

Dinamoeba PASCHER 1916 Dinamoeba mirabilis LEIDY 1874

Incertae sedis AMOEBOZOA Filamoeba PAGE, 1967 Filamoeba nolandi PAGE, 1967

Allelogromia BLANC Allelogromia brunneri (BLANC) DE SAEDELEER, 1934

Lieberkuehnia CLAPAREDE & LACHMANN, 1859 Lieberkuehnia wagneri CLAPAREDE & LACHMANN, 1859

Asterocaelum CANTER, 1973 Asterocaelum sp.

Vampyrella LEIDY, 1879 Vampyrella lateritia (FRESENIUS 1856)

RHIZARIA

Cercozoa Silicofilosea Amphitrema ARCHER, 1869

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 7 Amphitrema wrightianum ARCHER, 1869

Eugliphida Euglypha DUJARDIN, 1841 Euglypha ciliata EHRENBERG 1848 Euglypha alveolata DUJARDIN, 1841 Euglypha brachiata LEIDY, 1878

Trinema DUJARDIN, 1841 Trinema enchelys (EHRENBERG 1838), LEIDY 1878

Cyphoderia SCHLUMBERGER, 1845 Cyphoderia ampulla (EHRENBERG, 1840)

Paulinella LAUTERBORN 1895 Paulinella chromatophora LAUTERBORN 1895

CHROMALVEOLATA

Actinophryidae Actinophrys EHRENBERG, 1830 Actinophrys sol EHRENBERG, 1830

Actinosphaerium STEIN, 1857 Actinosphaerium eichorni (EHRENBERG, 1840)

Alveolata Ciliophora Postciliodesmatophora Karyorelictea Trachelocerca EHRENBERG, 1833 Trachelocerca sp.

Loxodes EHRENBERG, 1833 Loxodes sp. Loxodes sp. aff vorax

Heterotrichea Stentor OKEN, 1815 Stentor polymorphus (MÜLLER, 1773) Stentor igneus EHRENBERG, 1838 Stentor amethystinus LEIDY, 1880 Stentor coeruleus (PALLAS, 1766) Stentor roeseli EHRENBERG, 1835 Stentor niger (MÜLLER, 1773) Stentor sp.

Blepharisma PERTY, 1849 Blepharisma steini KAHL, 1932

Climacostomum STEIN 1859 Climacostomum virens (EHRENBERG, 1838)

Pseudoblepharisma KAHL, 1926 Pseudoblepharisma tenue KAHL, 1926

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 8 Spirostomum EHRENBERG, 1833 Spirostomum minus ROUX 1901 Spirostomum ambiguum (MÜLLER, 1786)

Linostomella AESCHT, in FOISSNER, BERGER & SCHAUMBURG, 1999 Linostomella vorticella (EHRENBERG, 1833)

Intramacronucleata Spirotrichea Hypotrichia Aspidisca EHRENBERG, 1831 Aspidisca cicada (MÜLLER, 1786)

Euplotes EHRENBERG, 1830 Euplotes patella (MÜLLER, 1773) Euplotes muscicola KAHL 1932 Euplotes charon (MÜLLER, 1786)

Oligotrichia Rimostrombidium JANKOWSKI, 1978 Rimostrombidium humile (PENARD, 1922)

Limnostrombidium KRAINER, 1995 Limnostrombidium viride (STEIN, 1867)

Choreotrichia Codonella HAECKEL, 1873 Codonella cratera (LEIDY, 1877)

Tintinnidium KENT, 1881 Tintinnidium semiciliatum STERKI, 1879 Tintinnidium fluviatile STEIN, 1833

Strobilidium SCHEWIAKOFF, 1893 Strobilidium caudatum (FROMENTEL, 1876)

Stichotrichia Holosticha WRZESNIOWSKI, 1877 Holosticha navicularum KAHL, 1932

Urostyla EHRENBERG, 1830 Urostyla grandis EHRENBERG, 1830 Urostyla viridis STEIN, 1859

Hypotrichidium ILOWAISKY, 1921 Hypotrichidium conicum ILOWAISKY, 1921

Spiretella BORROR, 1972 Spiretella plancticola (GELEI, 1933)

Strongylidium STERKI 1878 Strongylidium crassum STERKI 1878

Paraurostyla BORROR, 1972 Paraurostyla weissei STEIN, 1859

Balladyna KOWALESKI, 1882 Balladyna fusiformis KAHL, 1932

Halteria DUJARDIN, 1841

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 9 Halteria grandinella (MÜLLER, 1773)

Pelagohalteria FOISSNER, SKOGSTAD & PRATT, 1988 Pelagohalteria cirrifera (KAHL, 1932)

Tachysoma STOKES, 1887 Tachysoma pellionellum (MÜLLER, 1773)

Tetmemena EIGNER, 1999 Tetmemena pustulata EIGNER, 1999

Stylonychia EHRENBERG, 1830 Stylonychia mytilus (MÜLLER, 1773)

Urosoma KOWALEWSKIEGA, 1882 Urosoma cienkowskii KOWALEWSKIEGA, 1882

Oxytricha BORY DE SAINT-VINCENT 1824 Oxytricha fallax STEIN, 1859 Oxytricha saprobia KAHL, 1932

Armophorea Brachonella JANKOWSKI, 1964 Brachonella spiralis (SMITH, 1894) Brachonella sp. aff. spiralis

Tropidoatractus LEVANDER, 1894 Tropidoatractus acuminatus LEVANDER, 1894

Metopus CLAPAREDE & LACHMANN, 1858 Metopus es (MÜLLER, 1776) Metopus laminarius KAHL, 1927 Metopus sp.

Caenomorpha PERTY, 1852 Caenomorpha medusula PERTY, 1852

Discomorphella CORLISS, 1960 Discomorphella pectinata (LEVANDER, 1894)

Epalxella CORLISS, 1960 Epalxella exigua PENARD, 1922 Epalxella mirabilis (ROUX, 1899)

Saprodinium LAUTERBORNN, 1908 Saprodinium dentatum LAUTERBORNN, 1908

Litostomatea Didinium POCHE, 1913 Didinium nasutum (MÜLLER, 1773)

Monodinium FABRE-DOMERGUE, 1888 Monodinium chllorelligerum KRAINER, 1995 Monodinium balbianii balbianii FABRE-DOMERGUE, 1888 Monodinium balbianii breviproboscis FOISSNER, BERGER & SCHAUMBURG, 1999

Cyclotrichium MEUNIER, 1910 Cyclotrichium viride GAJEWSKAJA, 1933

Actinobolina STRAND, 1928

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 10 Actinobolina radians (STEIN, 1867) Actinobolina smalli HOLT, LYNN & CORLISS, 1973

Teuthophrys CHATTON & BEAUCHAMP, 1923 Teuthophrys trisulca CHATTON & BEAUCHAMP, 1923

Paradileptus WENRICH, 1929 Paradileptus elephantinus (ŠVEC, 1897)

Pelagodileptus FOISSNER, BERGER & SCHAUMBURG, 1999 Pelagodileptus trachelioides (ZACHARIAS, 1894)

Trachelius SCHRANK, 1803 Trachelius ovum (EHRENBERG, 1831)

Dileptus DUJARDIN, 1840 Dileptus cygnus (CLAPAREDE & LACHMANN, 1859) Dileptus anser (MÜLLER, 1786) Dileptus sp.

Lacrymaria BORY DE ST. VINCENT, 1926 Lacrymaria olor (MÜLLER, 1786)

Homalozoon STOKES, 1890 Homalozoon vermiculare (STOKES, 1887)

Balantidion EBERHARD, 1862 Balantidion pellucidum EBERHARD, 1862

Spathidium DUJARDIN, 1841 Spathidium stammeri WENZEL, 1955 Spathidium pectinatum KAHL, 1926 Spathidium opimum KAHL, 1930

Lepidotrachelophyllum NICHOLLS & LYNN, 1984 Lepidotrachelophyllum lineare (LEPSI, 1960)

Lagynophrya KAHL, 1927 Lagynophrya rostrata KAHL, 1927

Trachelophyllum CLAPAREDE & LACHMANN, 1859 Trachelophyllum sigmoides KAHL, 1926

Pelagovascicola JANKOWSKI, 1980 Pelagovascicola cinctum (VOIGT, 1901)

Lilimorpha GAJEWSKAJA, 1930 Lilimorpha viridis GAJEWSKAJA, 1928

Amphileptus EHRENBERG, 1830 Amphileptus sp. EHRENBERG, 1830 Amphileptus claparedi STEIN, 1867 Amphileptus pleurosigma (STOKES, 1884) Amphileptus trachelioides (ZACHARIAS, 1893)

Loxophyllum DUJARDIN, 1841 Loxophyllum meleagris (MÜLLER, 1773)

Litonotus WRESNIOWSKI, 1870

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 11 Litonotus sp. WRESNIOWSKI, 1870 Litonotus cygnus (MÜLLER, 1773) Litonotus lamella (MÜLLER, 1773)

Phyllopharyngea Cyrtophoria Phascolodon STEIN, 1859 Phascolodon vorticella STEIN, 1859

Chilodonella STRAND, 1928 Chilodonella uncinata (EHRENBERG, 1838) Chilodonella cucullus (MÜLLER, 1786)

Suctoria Parapodophrya KAHL, 1931 Parapodophrya soliformis (LAUTERBORNN, 1908)

Podophrya EHRENBERG, 1838 Podophrya fixa (MÜLLER, 1786)

Metacineta BÜTSCHLI, 1889 Metacineta mystacina (EHRENBERG, 1831)

Staurophrya ZACHARIAS, 1893 Staurophrya elegans ZACHARIAS, 1893

Prodiscophrya KORMOS, 1935 Prodiscophrya collini (ROOT, 1914) KORMOS, 1935

Nassophorea Nassula EHRENBERG, 1833 Nassula gracilis KAHL, 1931

Obertrumia FOISSNER & ADAM, 1981 Obertrumia aurea (EHRENBERG, 1833)

Microthorax ENGELMANN, 1862 Microthorax sp. aff. pusillus

Leptopharunx MERMOD, 1914 Leptopharynx costatus MERMOD, 1914

Colpodea Colpoda MÜLLER, 1773 Colpoda cucullus (MÜLLER, 1773) Colpoda steinii MAUPAS, 1883

Bursaria MÜLLER, 1773 Bursaria truncatella MÜLLER, 1773

Bursaridium LAUTERBORN, 1894 Bursaridium pseudobursaria (FAURE-FREMIET, 1924)

Bryometopus KAHL, 1932 Bryometopus pseudochilodon KAHL, 1932

Prostomatea Vasicola TATEM, 1869 Vasicola lutea KAHL, 1930

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 12 Urotricha CLAPAREDE & LACHMANN, 1859 Urotricha sp. CLAPAREDE & LACHMANN, 1859 Urotricha castalia MUŇOZ, TELLEZ & FERNANDO-GALIANO, 1987 Urotricha saprophila KAHL, 1930

Longifragma in FOISSNER, BERGER & SCHUM., 1999 Longifragma obliqua (KAHL, 1926)

Placus COHN, 1866 Placus luciae KAHL, 1926

Coleps NITZSCH,1827 Coleps hirtus hirtus (MÜLLER, 1786) Coleps hirtus viridis EHRENBERG, 1831 Coleps nolandi KAHL, 1930

Lagynus QUENNERSTEDT, 1867 Lagynus elegans (ENGELMANN, 1862)

Pseudoprorodon KAHL, 1930 Pseudoprorodon sulcatus KAHL, 1927

Prorodon EHRENBERG, 1833 Prorodon niveus EHRENBERG, 1833 Prorodon viridis EHRENBERG, 1833

Holophrya EHRENBERG, 1831 Holophrya teres EHRENBERG, 1833

Bursellopsis CORLISS, 1960 Bursellopsis nigricans (LAUTERBORN, 1894)

Plagiopylea Plagiopyla STEIN, 1860 Plagiopyla nasuta STEIN, 1860

Oligohymenophorea Peniculia Lembadion PERTY, 1849 Lembadion bullinum (MÜLLER, 1786)

Marituja GAJEWSKAJA, 1928 Marituja sp. aff. caudata

Stokesia, WENRICH, 1929 Stokesia vernalis WENRICH, 1929

Disematostoma LAUTERBORN, 1894 Disematostoma tetraedricum (FAURE-FREMIET, 1924)

Frontonia EHRENBERG,1838 Frontonia acuminata (EHRENBERG,1833) Frontonia leucas (EHRENBERG, 1833)

Urocentrum NITZSCH, 1827 Urocentrum turbo (MÜLLER, 1786)

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 13 Paramecium MÜLLER, 1773 Paramecium putrinum CLAPAREDE & LACHMANN, 1858 Paramecium bursaria (EHRENBERG, 1831) Paramecium aurelia (EHRENBERG 1838) complex Paramecium caudatum EHRENBERG, 1838

Scuticociliatia Platynematum KAHL, 1935 Platynematum sociale (PENARD, 1922)

Pseudocohnilembus EVANS & THOMPSON, 1964 Pseudocohnilembus pusillus (QUENNERSTEDT, 1869)

Uronema DUJARDIN, 1841 Uronema marinum DUJARDIN, 1841

Histiobalantium STOKES, 1886 Histiobalantium bodamicum KRAINER & MÜLLER, 1995

Pleuronema DUJARDIN 1836 Pleuronema crassum DUJARDIN 1841

Cyclidium MÜLLER, 1786 Cyclidium citrullus COHN, 1865 Cyclidium glaucoma MÜLLER, 1773

Hymenostomatia Trichospira ROUX, 1899 Trichospira inversa (CLAPAREDE & LACHMANN, 1858)

Tetrahymena FURGASON,1940 Tetrahymena pyriformis (EHRENBERG, 1830) complex

Dexiostoma JANKOWSKII, 1967 Dexiostoma campylum (STOKES 1886)

Colpidium STEIN, 1860 Colpidium colpoda (LOSANA 1829)

Ophryoglena EHRENBERG, 1831 Ophryoglena atra LIEBERKUHN, 1856

Peritrichia Hastatella ERLANGER, 1890 Hastatella radians ERLANGER, 1890

Astylozoon ENGELMANN, 1862 Astylozoon fallax ENGELMANN, 1862

Opisthonecta FAURE-FREMIET, 1906 Opisthonecta henneguyi (FAURE-FREMIET, 1906)

Vaginicola LAMARCK, 1816 Vaginicola sp.

Vorticella LINNAEUS, 1767 Vorticella campanula EHRENBERG 1831 Vorticella convallaria LINNAEUS, 1758 complex Vorticella similis STOKES 1887

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 14 Vorticella microstoma EHRENBERG, 1830 complex Vorticella chlorellata STILLER, 1940 Vorticella aquadulcis FOISSNER et al., 1992 complex Vorticella vernalis STOKES 1887 Vorticella sp. aff. vernalis Vorticella sp. aff. picta Vorticella sp.

Pelagovorticella JANKOWSKI, 1980 Pelagovorticella natans (FAURE-FREMIET, 1924)

Carchesium EHRENBERG, 1830 Carchesium polypinum LINNAEUS, 1758

Epicarchesium JANKOWSKI, 1985 Epicarchesium pectinatum (ZACHARIAS, 1897)

Zoothamnium BORY DE ST. VINCENT, 1826 Zoothamnium ramosissimum SOMMER, 1951 Zoothamnium arbuscula EHRENBERG, 1839

Campanella GOLDFUSS, 1820 Campanella umbellaria LINNAEUS, 1767

Epistylis EHRENBERG, 1830 Epistylis pygmaeum (EHRENBERG, 1838) Epistylis variabilis STILLER, 1935 Epistylis procumbens ZACHARIAS, 1897 Epistylis plicatilis EHRENBERG, 1838

Trichodina EHRENBERG, 1830 Trichodina pedicula (EHRENBERG, 1831) Trichodina domerguei (WALLENGREN, 1897) Trichodina fultoni DAVIS, 1947

Trichodinella SRAMEK-HUSEK, 1953 Trichodinella epizootica (RAABE, 1950)

Astomatia Steinella CÉPÈDE, 1910 Steinella uncinata (SCHULTZE, 1851)

EXCAVATA Heterolobosea Vahlkampfia CHATTON & LALUNG-BONNAIRE, 1912 Vahlkampfia limax DUJARDIN 1841 Vahlkampfia guttula DUJARDIN 1841 Vahlkampfia tachypodia GLAESER 1912 Vahlkampfia mira SCHAEFFER 1926 Vahlkampfia vahlkampfii (CHATTON, 1910)

Incertae sedis PROTISTA

Centrohelida Acanthocystis CARTER, 1863 Acanthocystis sp. aff. mimetica Acanthocystis aculeata HERTWIG & LESSER 1874 Acanthocystys mimetica PENARD, 1904 Acanthocystis radiosa ( in STREBLE & KRAUTER, 2002)

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 15 Chlamydaster RAINER, 1968 Chlamydaster sterni STERN, 1924

Heterophrys ARCHER 1869 Heterophrys myriapoda ARCHER 1869

Raphydocystis PENARD, 1904 Raphydocystis infestans WETZEL 1925

Raphydiophrys ARCHER 1867 Raphydiophrys pallida SCHULTZE, 1874

Bibliography of the publications of Protozoa of Shkodra Lake

Alushi, V. 2003: Rezultate paraprake nga studimi i Protozoarëve (Protozoa) të Liqenit të Shkodrës. Mikrotezë. SH. P. U. Tiranë Alushi, V. 2003: Një listë e protozoarëve të gjetur në Liqenin e Shkodrës. USh “Luigj Gurakuqi”, Botim i Sektorit BIO- EKO të Liqenit të Shkodrës, Nr 3, 2003, fq. 21-24. Alushi, V. 2004: Protozoarë të ujrave litorale-sipërfaqësore të pjesës shqiptare të Liqenit të Shkodrës. Studime Biologjike Nr. 9: 67-80. Tiranë. Alushi, V. 2004: Protozoarët e gjetur në Syrin e Sheganit dhe ujrat e Liqenit pranë tij. Konferenca mbi studimet në shkencat e natyrës dhe didaktikën e tyre. Shkodër. Alushi, V. 2005: Mbi protozooplanktonin pelagjik të Liqenit të Shkodrës., Bul. Shk. USh “Luigj Gurakuqi”, Nr. 55 / Ser. Shk. Nat. fq. 103-113. Alushi, V. 2011: Overview of Shkodra Lake ciliophora (Protozoa: Ciliophora). Intern. Conference on Carstic Water Bodies Alblakes 2. Elbasan, Albania, 09-11 June, Procedings books. ISBN: 99956-16-49-6, page 26-34 Alushi, V. 2012: “Contribution for the recognition of Amoeboid Protists of Shkodra Lake”. Vth International Scientific Conference on Water, Climate and Environment. BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012. (in process) Alushi, V. 2012: "A Taxonomic Overview of Protozoa of Shkodra Lake" . V-th International Scientific Conference on Water, Climate and Environment. BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012. (in proccess) Alushi, V. & Beqiri, E. 2004: Listë e protozoarëve të gjetur në ujrat e brigjeve të Liqenit të Shkodrës. Konferenca mbi studimet në shkencat e natyrës dhe didaktikën e tyre. Shkodër. Alushi, V. & Dhora, DH. 2004: Listë paraprake e protozoarëve të gjetur në Liqenin e Shkodrës. Buletini Shkencor USh “Luigj Gurakuqi”, Seria e Shkencave të Natyrës, Nr. 54: 107- 117. Shkodër. Alushi, V. & Dhora, DH. 2006: Protozoaret e Liqenit te Shkodres. Specie te reja, vleresime te pergjithshme. Buletini Shkencor USh “Luigj Gurakuqi”, Seria e Shkencave të Natyrës, Nr. 56: 107-128. Shkodër. Alushi, V. & Dhora, DH. 2007: Vleresime ekologjike mbi protozoaret e Liqenit te Shkodres. Buletini Shkencor USh “Luigj Gurakuqi”, Seria e Shkencave të Natyrës, Nr. 57: 120-149. Shkodër. Alushi, V. & Dhora, DH. 2008: Vlerësimi i cilësisë së ujërave të Liqenit të Shkodrës nëpërmjet protozoarëve indikatorë. Buletini Shkencor USH “Luigj Gurakuqi”, Seria e Shkencave të Natyrës, 2008. Nr 58: 180-192 Alushi, V. & Mani. E 2011: The potencials of Protozoa diversity (Protozoa) of Shkodra Lake. International Conference on Carstic Water Bodies Alblakes’ 2. Elbasan, Albania, 09- 11 June , Procedings books. ISBN: 99956-16-49-6, page 10-16 Alushi, V. & Mani. E 2011: The Use of Cultivation Technology for a More Complete Knowledge of Protozoa and Ecology of Shkodra Lake. International Conference “Biotechnological Developments”.Academy of Sciencies of Albania and University of Tirana. Tirane , 20- 21 November, 2011 Procedings books (ne botim)

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 16 Alushi,V. & Rakaj, M. 2011: Overview of Protists of Shkodra Lake. ICE, International Conference on Ecosystems, Procedings books, Essays on Ecosystem and Environmental Research, ICE_2011, ISBN: 978-9928-4068-1-1, June 4-6, Tirana, Albania. page 97 - 102 Alushi,V. & Rakaj, M. 2011: Overview of Protists of Shkodra Lake. International Journal of Ecosystems and Ecology Science (IJEES). Volume 1/ 2011, ” Special Issue, ICE 2011” , Tirana, Albania”page 1-6, ISSN 2224-4980 Dhora, Dh., Mani, E., Bekteshi, A. & Ulqini, D. 2009 : Euglena gracillis Klebs 1883 ne dy terrene ushqimore interesante. Bul.Shk, USh. « Luigj Gurakuqi », Nr 59, Seria e Shkencave te natyres, fq 81 -89 Gessner, F. 1934: Limnologische Untersuchungen am Skadar (Skutari) See Glas.Bot. zavoda i bašte Univ. u Beogradu, 3-1/2. Karaman, G & Beeton, M. 1981: Monitoring program for the protection of Lake Skadar. In: KARAMAN & BEETON: The biota and limnology of lake of Skadar. p.425 - 428. Titograd. Karaman, G & Beeton, M. 1981: The biota and limnology of Lake Skadar. General summary. p. 429 – 436. Titograd. Karaman, G & Nedic, D. 1981: Zoobentos of Lake Skadar. The biota and Limnology of Lake Skadar. Univerzitet “Veljko Vlahovic”, Institut za biološka i medicinska istra ivanja u SRCG, Biološki zavod Titograd, Smithsonian Institution Washington, University of Wisconsin, 222-246. Karini, S. & Dhora, Dh. 2012 : Zhvillimi i protozoarëve (Protozoa) gjatë dekompozimit të makrofiteve në breg të Liqenit të Shkodrës. Bul. Shk. USh. “Luigj Gurakuqi”, (in publication) Mani (Beqiri), E. (2010) Kulturat e protozoareve te Liqenit te Shkodres ne terrene te ndryshme ushqimore. Master ne Biologji. Tirane Mani (Beqiri), E. & Dhora Dh. 2010: Suksesionet protozoarëve të Liqenit të Shkodrës në terrene të ndryshme ushqimore. Bul. Shk. USh. “Luigj Gurakuqi”, Nr. 60, Seria e Shkencave të Natyrës, fq 70-81. Mani, E. & Dhora. Dh. 2011: Rezultate nga kultivimi i protozoarëve të Liqenit të Shkodrës. Bul. Shken. i Ush “Luigj Gurakuqi”, Nr 61, Seria e Shk. Nat., Shkodër. Milovanovic, D. & Živkovic, A. 1965: Plankton Skadarskog Jezera (1957-1958)]. Zbornik radova Biol.inst. 8(4):1-36. Nedeljkovic, R. 1959: Skadarsko jezero. Studija organske produkcije u jednom karstnom jezeru. Biol. Inst. Posebno izdanje. Knj. 4. Nikolic, V., Simonovic, P., Mrdak, D. & Radojicic, J. 2001: Trichodina fultoni Davis 1947 (Trichodinidae, Ciliophora, Protista) from Lake Skadar _ a new species for yougoslav fish parasite fauna. Prirodni potencijali kopna, kontinentalnih voda i mora Crne Gore i njihova zastita. Naucnih Skup. Nov. 103. Zabljak Petkovic, St.1977: Nove vrste u fauni Skadarskog Jezera. Glas. Republ. Zavoda Zast. Prirode- Prirodnjačkog Muzeja. Nr. 10: 31 - 36. Titograd. Petkovic, St.1980: A synthetic survay of investigations in a karstic mediterranean lake ecosystem – Protozoa. Poljopriveda i sumarstvo, XXVI, 1: 107 – 111. Titograd. Petkovic, St.1981: Zooplankton. In: KARAMAN, G. & BEETON, A. M. Biota and limnology of Lake Skadar Petkovic, Sm., & Petkovic, St. 1984: Limnoflora I limnofauna reke Morace I nekih reka u njenom slivu. Glas. Repub. Zav. Prirode – Prir. Muzeja, Titograd, 17: 47 – 68. Rakaj, M., Alushi, V. & Dhora, DH. 2006: Lista e protistëve të Liqenit të Shkodrës. CP. Shkodër, 94 fq.

Conclusions • A full list of Shkodra Lake Protozoa is given in this study and the latest data about the study of this group being reflected. • For the first time, the publication bibliography about the Protozoa of Shkodra Lake is presented, both Albanian and Montenegrin parts of the Lake being included. • The studies of the Protozoa are necessary to be thorough especially for their taxonomic aspect, the insertion of modern methods for their identification and the finding of rare species, mainly those in specific microhabitats of the Lake, as well as in other environments as in lagoons and rivers.

BALWOIS 2012 - Ohrid, Republic of Macedonia - 28 May, 2 June 2012 17 • For a further progress of this study, a wider working group is needed, as well as the ensuring of co-operation with qualified European institutions. An institutional collaboration in the aspect of Balkan studies is more than necessary.

Other references

(1) Adl et al,. 2005. The new higher level classification of eukaryotes with emphasis on the taxonomy of protists. J. Euk. Microbiol. 52:399-451.

(2) Dhora, DH. (2005) Liqeni i Shkodrës. Shtëpia botuese “Camaj-Pipaj”. Shkodër.

(3) Finlay B. J. (1988): A beginner’s guide to the collection, isolation, cultivation and identification of freshwater protozoa. Published: Freshwater Biological Associacion.

(4) Foissner, W., Berger, H. & Schaumburg, J. (1999) Identification and ecology of limnetic plancton ciliates. Bavarian State Office for Water Management (Editor and Publisher). Mynich.

(5) Hellawell, M. (1989): Biological indicators of freshwater pollution and environmental management. Pollution monitoring series. Advisory editor: K. Mellany. Elsevier applied sciense.

(6) Kumar, A. & Dalby, A. P.1998. Identification key for Holocene lacustrine arcellacean taxa. Palaeontologiaelectronica, palaeo-electronica.org/1998_1/dalby/issue1.htm

(7) Lee, J. J., Leedale, G. F. & Bradbury, Ph., 2000: An illustrated guide to the Protozoa. Second edition. Vol. I & II. Society of Protozoologists. Printed by Allen Press Inc., Lawrence, KS 66044 USA.

(8) Magazzu, G. (1978) Metodi per lo studio del plancton della produzione primaria societa di biologia marina. Edizione GM, p. 61 - 68.

(9) Pavllovski E. N. & Lepneva S. G. (1948): Oçerki iz zhizni pesnovodnih zhivotnih . Sovjetskaja Nauka. C. 400 – 425. Leningrad.

(10) Smirnov, A & Brown, S (2004): Guide to the methods of study and identification of soil gymnamoebae. Protistology 3 (3), 148_190

(11) Streble, H. & Krauter, D. (2002) Atlante dei microrganismi acquatici. La vita una goccia d'aqua. Franco muzzio editore. Padova.

(12) Zelikman, L. A. (1965) Praktikum po zoologi bespozvonoçnih. Izdatelstvo “ Visshaja shkolla 309 – 315. Moskva.

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