A Conspectus of the Black Sea fauna of benthic ciliates Текст научной статьи по специальности «Биологические науки»

Protistology 3 (2), 72-91 (2003)

Protistology

A conspectus of the Black Sea fauna of benthic ciliates

Andrey I. Azovsky1 and Yuri A. Mazei2

1 Department ofHydrobiology, Biology Faculty, Moscow State University, Moscow, Russia

2 Department of Ecology, Penza State Pedagogical University, Penza, Russia

Summary

Species list of the Black Sea free-living benthic ciliates was compiled using most of the published sources available as well as original data. The total list presented contains 476 species from 147 genera. The Romanian and Bulgarian coasts were the richest ones (299 and 253 species), then followed the northeastern part of the sea (144 species) and the northwestern part (83 species). Statistical analysis using species/genera ratio, average taxonomic diversity and its variance showed that the latter two regions had significantly low taxonomic diversity due to disproportional under- or overrepresentation of some taxa. The Black Sea benthic ciliofauna appears to be the richest one of all the seas studied.

Key words: ciliata, protozoa, benthos, Black Sea, diversity

Introduction

Protozoological studies at the Black Sea have a long history, especially at Romanian and Bulgarian coasts and in the Crimea (Merezhkovsky, 1880; Andrussova, 1886; Perejaslawzewa, 1886; Lepsi, 1926a, 1926b; Jeliaskova-Paspalewa, 1933; Valkanov, 1934, 1935, 1936). These regions had been repeatedly studied during the second half of the XXth century. Nevertheless, there are no inventories that summarize and review the data accumulated up-to-date. The estimates reported by some authors were based upon fragmented and obviously incomplete data and yielded rather low figures (Tuco-lesco, 1961; Kovaleva and Golemansky, 1979; Djur-tubaeva and Mokhammed, 1995). Since these bio-

diversity figures were noticeably lower than those for some other well-studied seas, the Black Sea has long been recognized as relatively poorly populated by ciliates (Agamaliev, 1983; Burkovsky, 1984).

However, even a cursory overview of the publications suggests that the diversity of benthic ciliates of the region may be rather high. Therefore the objectives of this study have been as follows: (i) to summarize and unify (as far as possible) the literature data and to compile a total species list of the Black Sea benthic ciliates; (ii) to estimate taxonomic diversity of the sea and its parts; (iii) to compare the parts as to their species richness and composition.

We have not attempted to undertake any comprehensive taxonomic revision of the Black Sea fauna,

© 2003 by Russia, Protistology

so only the most evident changes and corrections were made, necessary for the unification of the species lists compiled from different sources (see Material and methods).

Material and methods

Data sources. Compiling the species list, we used the data from the following sources: Perejaslawzewa,

1886; Lepsi, 1926a, 1926b; Jeliaskova-Paspalewa, 1933;

Valkanov, 1934, 1935, 1936, 1954, 1957; Czapik, 1952a,

1952b; Tucolesco, 1961, 1962; Kovaleva, 1966; Bacescu et al., 1967; Detcheva 1973, 1977, 1980, 1981, 1983;

Groliere and Detcheva, 1974, 1979; Groliere et al.,

1974; Petran, 1963, 1968, 1976; 1997; Djurtubaev,

1977; Kovaleva and Golemansky, 1979; Gulin et al.,

1986; Djurtubaeva and Mokhammed, 1995; Azovsky,

2002; Azovsky and Mazei, 2003.

The data were grouped according to the four regions where the investigations had been carried out (Fig. 1): a) Bulgarian coasts (BC); b) Romanian coasts (RC); c) the northwestern part (the Gulf of Odessa and the Crimea, NW); and d) the northeastern part (Russian shore sector from Adler to Anapa, NE).

Taxonomic remarks. The species list compilation was based on the review by Carey (1991). In the study we mainly followed the rather conservative system of Corliss (1979), which is accepted in the above review.

The main results of recent revisions of some groups were also taken into account (Esteban et al., 1995; Foissner,

1995, 1996 1997a, 1997b; 1998; Foissner and Al-

Fig. 1. Map of the Black Sea showing approximate location of the regions considered (see text for the denotations).

Rasheid, 1999a, 1999b; Foissner and Dragesco, 1996a, 1996b).

In particular, Trachelocercidae family is now being most radically revised. As a result, some species have been re-described, some new genera and families have been established, one genus was abolished, and some species were transferred to other genera or even classes. However, a few species of this family have not yet been re-investigated, and most species cannot be presently identified in the context ofthe new system. This presents significant difficulties for faunistic and ecological studies, in particular, for comparison of formerly published lists. The species re-described and the synonymy arisen are given below in accordance with the new system: Trachelocerca sagitta (Syn.: Trache-loraphis striata Raikov 1962), Trachelocerca incaudata (Syn.: Tracheloraphis incaudatus (Kahl 1933) Dragesco 1960), Tracheloraphisphoenicopterus, T. prenanti (it is rather close to T. phoenicopterus and they probably form one species complex) (Foissner and Dragesco 1996)), Tracheloraphis longicollis (Syn.: Trachelonema longi-collis Dragesco 1960), Tracheloraphis oligostriata (Syn.: Trachelonema oligostriata Raikov 1962), Kovalevia sulcata (Syn.: Trachelonema sulcata Kovaleva 1966), Tracheloraphis kahli. Trachelonema entzi Kahl 1927 was re-described as Trachelotractus entzi (Kahl 1927) Foissner 1997 in the order Prostomatida. For other trachelocercid species we used the old nomenclature, including the abolished genera Trachelonema, but with nomenclature emendations from Foissner and Dra-gesco (1996a).

In his review of the genus Kentrophoros, Foissner (1995) re-described K. fistulosus Faure-Fremiet 1950 and included into the genus two species: K. longissimus Dragesco 1954 h K. tubiformis Raikov et Kovaleva 1966.

Esteban et al. (1995) have recently revised the genus Metopus. Seventy-six nominal species described by that time were reduced to 22 morphotypes. As a result, the number of Metopus species earlier reported for the Black Sea was reduced significantly.

Most of other species names are given following Carey (1991). Forms unidentified to the species level were included only if they were the single representatives of the genus for the given sea region.

Statistical analysis. The total number of species and genera and species/genera ratio (S/G) were calculated for the whole sea and for each region under consideration.

In addition, the following diversity indices recently developed were used in order to compare the ciliate taxonomic diversity of the regions: a) average taxonomic distinctness (AvTD) as defined by Clarke and Warwick (1998): A+ = 2 (SSK. w.) /[s (s - 1)], where w.. is the taxonomic path length between species i and j, and s is the number of species in the list; and b) variation in taxonomic distinctness (VarTD) as defined by Clarke and Warwick (2001): A+ = 2 (SS<j w.2 ) /[s (s - 1)] - (D+)2. These indices can show not only species richness but also the taxonomic relatedness of species in the regional inventory. So, an assemblage comprising a group of closely related species must be regarded as less «taxonomically diverse» (and having lower AvTD value) than an assemblage of the same number of more distantly related species belonging to different taxa. Another aspect of diversity measured by VarTD is the «evenness» of the taxonomic structure. High VarTD value shows that some taxa are over-represented and others under-represented in a given regional list as compared with the total species pool, either because of specific reaction of some taxa or because of subjective selectivity of registration. Both indices do not depend on the degree of sampling effort involved in the data collection. This is of particular importance for comparison of historic species lists compiled from different sources, with different and uncontrolled sampling efforts (Clarke and Warwick, 1999). Unity step lengths were used between each taxonomic level from species to classes, giving path-length weights w=1 for pairs of species in the same genus, w=2 for species in the same family but different genera, etc.

All these statistics (S/G-ratio, AvTD and VarTD), calculated from the actual species lists, can be tested

for departure from the expectation by constructing a simulated distribution (funnel) from random subsets of species from the total species inventory (Azovsky, 1996; Clarke and Warwick, 2001). For each regional list, 1000 random subsets were performed repeatedly. Actual values of the statistics falling within the 95% probability funnel mean that the ciliate assemblage has the same taxonomic structure as a random selection from the total species pool.

To estimate similarity between the regions by their species composition, we used two indices: the Czekanowski-S0rensen index (Jongman et al., 1995): SCS = 2a / (2a + b + c); and the Simpson’s index (Simpson, 1960): SS = a / (a + min (b, c)); where a is number of species common for the both regions (species lists) compared, b and c are numbers of species found in the first or the second region only. Both indices vary from 0 (no common species) to 1 (completely coincident lists). The difference between these indices is that Simpson’s index is insensitive to the disparity in the total number of species and is thus more appropriate for comparison between unequal or incompletely studied faunas (Simpson, 1960). Significance of similarities found was tested by comparing the values of similarity observed with a probability distribution of the values obtained from random occurrence model (Randomly Arranged Matrices procedure realized in ECOS package, Burkovsky et al., 1995). The rationale behind this approach is that if any groups of species tend to be conjointly present (or absent) in some regions, the similarity values observed would significantly differ from the values expected from the situation where all taxa occur randomly and independently over all species lists. In the presence/absence matrix, the data for each species were randomly rearranged between regional lists and similarity was calculated for each lists pair. Five hundred permutations were performed repeatedly, and 1% and 5% probability levels for the expected «random species’ occurrence» similarity were deter-mined and used as criteria for nonrandomness of the values observed (two-tailed test).

To represent the relations between regions, cluster-analysis was performed using the average linkage method.

Results

The total list (Table 1) includes 476 species belonging to 147 genera, 65 families and 15 orders. Among the sea regions, the Romanian and Bulgarian coasts are the richest ones (299 and 253 species, Table 2), whereas the northwestern part is noticeably poorer (83 species found). More than half of the species (292) were found

Table 1. Total list of benthic ciliates of the Black Sea,

Species RC1 BC NW NE References2

Class Kinetofragminophora de Puytorac et al., 1974

Subclass Gymnostomata Butschli, 1889

Order Karyorelictida Corliss, 1974

Family Trachelocercidae Kent, 1881

Kovalevia sulcata (Kovaleva, 1966) Foissner, 1997 X X X X 10; 14; 15; 23-25

Kovalevia teissieri (Dragesco, 1960) Foissner, 1997 X X 10; 15

Proto trachelocerca fasciolata (Sauerbrey, 1928) Foissner, 1996 X X X X 10; 11; 15; 22; 24; 25

Trachelocerca coluber Kahl, 1933 X X X 7; 10; 14; 15; 23; 25

Trachelocerca curvirostre Valkanov, 1934 X 4-6

Trachelocerca geopetiti Dragesco, 1960 X X 10; 15

Trachelocerca incaudata (Kahl, 1933) Foissner, 1997 X X X X 10; 11; 15; 22-25

Trachelocerca minuta Dragesco, 1960 X 25

Trachelocerca multinucleata Dragesco, 1960 X X X 10; 11; 15; 24

Trachelocerca nigricans Kahl, 1933 X 25

Trachelocerca nigrocephala Tucolesco, 1962 X 27

Trachelocerca polyhalina Tucolesco, 1962 X 27

Trachelocerca sagitta (Muller, 1786) Foissner et Dragesco, 1996 X X 15; 25

Trachelocerca tenuicollis Quennerstedt, 1867 X X X 11; 15; 24; 25

Trachelocerca trepida Kahl, 1928 X X 13; 26

Trachelocerca variabilis Kovaleva, 1966 X X 10; 14; 15; 23

Trachelonema minimum Dragesco, 1960 X X X 15; 24; 25

Tracheloraphis aragoi (Dragesco, 1953) Dragesco, 1960 X 10

Tracheloraphis bimicronucleata Raikov, 1969 X X 15

Tracheloraphis crassa Raikov, 1963 X X 10; 15

Tracheloraphis discolor Raikov, 1962 X 15

Tracheloraphis drachi (Dragesco, 1953) Dragesco, 1960 X X X X 10; 15; 24; 25

Tracheloraphis grisea (Kahl, 1933) Dragesco, 1960 X 23

Tracheloraphis kahli Raikov, 1962 X X X 15; 25

Tracheloraphis longicollis (Dragesco, 1960) Foissner et Dragesco, 1996 X X X X 10; 14; 15; 23-25

Tracheloraphis margaritata Kahl, 1933 X X 11; 15; 24;

Tracheloraphis phoenicopterus (Cohn, 1866) Dragesco, 1960 X X X X 7; 8; 11; 14-18; 22-26

Tracheloraphis prenanti Dragesco, 1960 X X X X 10; 15; 24; 25

Tracheloraphis sarmatica Agamaliev et Kovaleva, 1966 X 10

Tracheloraphis swedmarki Dragesco, 1960 X X X 10; 15; 24

Tracheloraphis totevi Kovaleva et Golemansky, 1979 X 15

Tracheloraphis vermiformis Raikov, 1962 X X 10; 15

Tracheloraphis oligostriata (Raikov, 1962) Foissner et Dragesco, 1996 X 25

Family Loxodidae Butschli, 1889

Kentrophoros fasciolatus Sauerbrey, 1928 X X X 11; 15; 24; 25

Kentrophoros fistulosus (Faure-Fremiet, 1950) Foissner, 1995 X X X 10; 14; 15; 23; 25

Kentrophoros flavum Raikov et Kovaleva, 1968 X 15

Kentrophoros gracilis Raikov, 1963 X X X 11; 15; 24; 25

Kentrophoros ponticus Kovaleva, 1966 X X X 10; 15; 25

Loxodes striatus (Engelmann, 1862) Penard, 1917 X X 17; 18; 25

Remanella caudata (Dragesco, 1953) Dragesco, 1960 X 23

Remanella faurei Dragesco, 1954 X X 11; 15; 24;

Remanella granulosa (Kahl, 1933) Dragesco, 1960 X X X X 10; 15; 24; 25

Remanella margaritifera (Kahl, 1933) Dragesco, 1960 X X X X 11; 15; 22; 24-26

Remanella minuta (Dragesco, 1954) Dragesco, 1960 X X X 11; 15; 24; 25

Remanella multinucleata (Kahl, 1933) Dragesco, 1960 X X X X 10; 11; 15-17; 24-26

Remanella rotundata Tucolesco, 1962 X 27

Remanella rugosa Kahl, 1933 X 7; 8; 10; 11; 15; 22-26

Remanella rugosa var. unicorpusculata Kahl, 1933 X 7; 8; 15

Species RC1 BC NW NE References2

Family Geleiidae Kahl, 1933

Geleia decolor Kahl, 1933 X X X X 10; 11; 15; 24; 25; 26

Geleia fossata Kahl, 1933 X X X X 7; 8; 10; 11; 15; 24; 25

Geleia nigriceps Kahl, 1933 X X X X 10; 15; 24; 25

Geleia orbis Faure-Fremiet, 1950 X X X X 10; 11; 15; 24; 25

Geleia tenuis Dragesco, 1960 X 15

Geleia swedmarkii Dragesco, 1954 X 25

Incertae sedis in Order Karyorelictida

Cryptopharynx setigerus Kahl, 1926 X X 15; 24; 26

Order Prostomatida Schewiakoff, 1896

Suborder Prostomatina Schewiakoff, 1896

Family Holophryidae Perty, 1852

Holophrya alviolata (Kahl, 1926) Kahl, 1930 X 25

Holophrya atra Svec, 1897 X 17; 18

Holophrya coronata de Morgan, 1925 X 7; 8; 15

Holophrya nigricans Laueterbray, 1908 X 26

Holophrya oblonga Maupas, 1883 X X 7; 8; 15; 24

Holophrya simplex Schewiakoff, 1893 X X 25; 26

Family Metacystidae Kahl, 1926

Metacystis tesselata Kahl, 1926 X X 25; 26

Metacystis truncata Cohn, 1866 X X 16; 17; 26

Metacystis agigeana Tucolesco, 1962 X 27

Metacystis nana Tucolesco, 1962 X 27

Metacystis tekirghiola Tucolesco, 1962 X 27

Vasicola brevis Tucolesco, 1962 X 27

Vasicola lutea Kahl, 1930 X 26

Vasicola parvula Kahl, 1928 X 26

Vasicola procera Tucolesco, 1962 X 27

Vasicola vesiculata Tucolesco, 1962 X 27

Suborder Prorodontina Corliss, 1974

Family Prorodontidae Kent, 1881

Helicoprorodon gigas Kahl, 1933 X X X 7; 8; 10; 11; 15; 24

Helicoprorodon minutus Bock, 1952 X X X 10; 15; 25

Helicoprorodon multinucleatum Dragesco, 1960 X 15

Helicoprorodon orientalis Raikov, 1963 X X 10; 15

Plagiocampa acuminata Kahl, 1933 X 26

Plagiocampa incisa Kahl, 1933 X 26

Plagiocampa margaritata Kahl, 1930 X 26

Plagiocampa marina Kahl, 1930 X X X 13; 16; 17; 18; 25; 26

Plagiocampa minima Kahl, 1927 X 26

Plagiocampa multiseta Kahl, 1930 X X 25; 26

Plagiocampa posticeconica Kahl, 1930 X 26

Plagiocampa rouxi Kahl, 1926 X X 17; 18; 26

Prorodon arenarius Dragesco, 1953 X 23

Prorodon discolor (Ehrenberg, 1831) Kahl, 1930 X X X X 14; 18; 23; 25; 26

Prorodon dubius Kahl, 1930 X 25

Prorodon elegans Kahl, 1928 X 25

Prorodon lemani (Dragesco, 1960) Dragesco, 1965 X 25

Prorodon marinus Claparede te Lachmann, 1858 X X X X 11; 15; 16-18; 23-25

Prorodon mimeticus Kahl, 1930 X 26

Prorodon aff. moebiusi Kahl, 1930 X 25

Prorodon multinucleatus Dragesco, 1954 X X 11; 15; 24;

Prorodon navicularis Tucolesco, 1962 X 27

Prorodon ovum (Ehrenberg, 1831) Kahl, 1930 X X X X 14; 18; 23; 25; 26;

Species

Pseudoprorodon arenicola Kahl, 1933 Pseudoprorodon eforianus Tucolesco, 1962 Rhagdostoma roscoffensis Dragesco, 1960 Rhagdostoma sp.

Spathidiopsis eforianus Tucolesco, 1962 Spathidiopsis luciae Kahl, 1926 Spathidiopsis ovum Kahl, 1926 Spathidiopsis socialis Fabre-Domergue, 1889 Spathidiopsis striatus Cohn, 1866 Trachelotractus entzi (Kahl, 1927) Foissner, 1997 Urotricha sp.

Family Colepidae Ehrenberg, 1838 Coleps hirtus Nitzsch, 1817 Coleps pulcher Spiegel, 1926 Coleps similis Kahl, 1933 Coleps spinosus Vacelet, 1961 Coleps tesselatus Kahl, 1930 Tiarina fusus Claparede te Lachmann, 1857 Order Haptorida Corliss, 1974

Family Enchelyidae Ehrenberg, 1838 Chaenea psammophila Dragesco, 1960 Chaenea robusta Kahl, 1930 Chaenea sapropelica Kahl, 1830 Chaenea simulans Kahl, 1930 Chaenea tesselata (Kahl, 1933) Dragesco, 1965 Chaenea vorax Quennerstedt, 1867 Enchelyodon fascinucleatus Kahl, 1933 Enchelyodon vacuolatus Dragesco, 1960 Enchelys simplex Kahl, 1926 Lacrymaria acuta Kahl, 1933 Lacrymaria caudata (Kahl, 1933) Dragesco, 1960 Lacrymaria cohnii Kent, 1880-1882 Lacrymaria coronata Claparede et Lachmann, 1858 Lacrymaria cucumis Penard, 1922 Lacrymaria kahli (Dragesco, 1954) Dragesco 1960 Lacrymaria lagenula Claparede et Lachmann, 1858 Lacrymaria minima Kahl, 1927

Lacrymaria multinucleata (Dragesco, 1954) Dragesco, 1960 Lacrymaria olor (O.F.M., 1776) Kahl, 1930 Lacrymaria salinarum Kahl, 1928 Lacrymaria versatilis Quennerstedt, 1867 Lacrymaria vertens Stokes, 1885 Lagynophrya halophila Kahl, 1930 Lagynophrya perlata Tucolesco, 1962 Lagynophrya rigida Tucolesco, 1962 Microregma ponticum (Lepsi, 1926) Kahl, 1930 Nannophrya costata Tucolesco, 1962 Plagiopogon loricatus Stein, 1859 Trachelophyllum apiculatum Perty, 1852 Trachelophyllum biacuminatum Tucolesco, 1962 Trachelophyllum triangulatum Tucolesco, 1962 Family Spathidiidae Kahl, 1929 Perispira ovum Stein, 1859 Spathidioides execata Kahl, 1930

RC1 BC NW NE References2

X 15

x 27

X 23

X 17; 18

X 27

X X 25; 26

X X 7; 8; 15; 23

X 26

X X X 7; 8; 15; 25; 26

X X X 7; 8; 11; 15-18; 24; 25

X 16-18

X X X 7; 25; 26

X X X 15-17; 24-26

X X X 15; 24; 25

X X 11; 25

X X X X 15; 17; 18; 22; 24; 25

X 26

X 15

X X 16-18; 25

X 26

X 26

X X 25; 26

X 7; 8; 15

X 26

X 23

X 25

X 25

X X X 11; 15; 24-26

X 25

X X X 11; 15-18; 24-26

X 26

X 23

X X X 1; 7; 15; 24; 25

X X 25; 26

X 25

X X 1; 7; 26

X X 13; 16-18; 26

X 25

X 26

X X 13; 16-18; 25

X 27

X 27

X i;7

X 27

X 26

X 26

X 27

X 27

X 26

X X 14; 26

Species

Spathidium antennatum Kahl, 1926 Spathidium curvatum Kahl, 1928 Spathidium marinum Lepsi, 1926 Spathidium spathula (O.F.M., 1773) Moody, 1912 Spathidium swarczewskyi Tucolesco, 1962 Family Tracheliidae Ehrenberg, 1838 Dileptus binucleatus Kahl, 1930 Dileptus sp.

Family Didiniidae Poche, 1913 Askenasia stellaris (Leegaad, 1920) Kahl, 1930 Didinium balbiani (Fabre-Domergue, 1888) Kahl, 1930 Didinium balbiani var. rostratum Kahl., 1926 Didinium nasutum (O.F.M., 1786) Kahl, 1930 Mesodinium cinctum Calkins, 1902 Mesodinium pulex Claparede et Lachmann, 1858 Mesodinium pulex f. pupula (Kahl, 1933) Dragesco, 1963 Mesodinium rubrum Claparede et Lachmann, 1858 Order Pleurostomatida Schewiakoff, 1896 Family Amphileptidae Butschli, 1889 Acineria incurvata Dujardin, 1841 Acineria uncinata Tucolesco, 1962 Centrophorella fasciolata Sauerbrey, 1928 Heminotus caudatus (Kahl, 1933) Dragesco, 1960 Amphileptus marina (Kahl, 1928) Carey, 1991 Litonotus anguilla (Kahl, 1930) Carey, 1991 Litonotus cygnus (Wrezesnioski, 1870) Kahl, 1931

Litonotus elongatus Dragesco, 1954 Litonotus fasciola (Ehrenberg, 1838) Wrzesnioski, 1870 Litonotus fusidens (Kahl, 1926) Carey, 1991 Litonotus lamella (Ehrenberg) Schewiakoff, 1896 Litonotus loxophylliforme (Dragesco, 1960) Carey, 1991 Litonotus naviculiformis Tucolesco, 1962 Litonotus pictus Gruber, 1884

Loxophyllum dragescoi (Faure-Fremiet, 1908) Carey, 1991

Loxophyllum elongatum Tucolesco, 1962

Loxophyllum fasciolatum Kahl, 1933

Loxophyllum grande Entz., 1879

Loxophyllum helus Stokes, 1884

Loxophyllum helus var. rotundatum Kahl, 1933

Loxophyllum kahli Dragesco, 1960

Loxophyllum levigatum (Sauerbrey, 1928) Dragesco, 1960 Loxophyllum lanceolatum (Dragesco, 1954) Dragesco, 1960 Loxophyllum multinucleatum Kahl, 1928 Loxophyllum multiplicatum Kahl, 1931 Loxophyllum setigerum Quennerstedt, 1867

Loxophyllum trinucleatum Mansfeld, 1923 Loxophyllum uninucleatum Kahl, 1928 Loxophyllum vermiforme Sauerbrey, 1928 Subclass Vestibulifera de Puytorac et a., 1974 Order Trichostomatida Butschli, 1889

Family Plagiopylidae Schewiakoff, 1896 Plagiopyla ovata Kahl, 1931

RC1 BC NW NE References2

X 26

X 26

X i;7

X 12

X 27

X 16; 17

X X 25; 26

X 26

X X 25; 26

X 26

X 25

X 25

X X X 11; 15; 16-18; 24-26

X X X 15; 24-26

X 11; 24

X X 1; 3; 7; 26

X 27

X 26

X X 11; 15; 24

X 26

X X 23; 25

X X X X 10; 14; 15; 17; 18; 23;

25; 26;

X 15

X 1; 16; 17

X X 7; 8; 17; 18; 25

X X 11; 15-17; 24; 26

X 13

X 27

X X 23; 26

X 18

X 27

X X 25; 26

X 26

X X X X 11; 15; 23-26

X 26

X X X 10; 15; 25

X X X X 10; 13-16; 23-26

X 25

X X X 18; 25; 26

X 7; 8; 15

X X X X 10; 11; 14; 15; 18; 23-

26

X 18

X X X 18; 25; 26

X 15

X X 15; 18; 26

Species

RC1

BC

NW

NE

References2

Plagiopyla sp.

Sonderia Alfredkahli (Kahl, 1933) Tucolesco, 1961

Sonderia cyclostoma Kahl, 1930

Sonderia kahli (Kahl, 1933) Tucolesco, 1961

Sonderia macrochilus Kahl, 1930

Sonderia mira Kahl, 1928

Sonderia schizostoma Kahl, 1930

Sonderia sinuata Kahl, 1930

Sonderia tubigula Kahl, 1930

Sonderia vestita Kahl, 1928

Sonderia vorax Kahl, 1928

Sonderiella scandens Kahl, 1926

Family Coelosomididae Corliss, 1961 Coelosomides marina (Anigstein, 1912) Strand, 1928 Paraspathidium fuscum (Kahl, 1928) Dragesco, 1960

Family Trimyemidae Kahl, 1933

Trimyema alfredkahli Tucolesco, 1962 Trimyema compressum Lackey, 1925 Trimyema kahli Tucolesco, 1962 Trimyema marina Kahl, 1931 Trimyema minuta Kahl, 1931 Sertumia tekirghiolica Tucolesco, 1962

Order Colpodida de Puytorac et al., 1974 Family Colpodidae Ehrenberg, 1838 Colpoda cucullus O.F.M., 1786 Colpoda inf lata Stokes, 1885 Colpoda maupasi Enriques, 1908 Colpoda simultans Kahl, 1931 Colpoda steini Maupas, 1883

Family Cyrtolophosididae Stokes, 1888 Cyrtolophosis bursaria Schewiakoff, 1893 Cyrtolophosis elongata Schewiakoff, 1892 Cyrtolophosis mucicola Stokes, 1885 Subclass Hypostomata Schewiakoff, 1896 Order Synhymeniida de Puytorac et al., 1974 Family Scaphidiodontidae Deroux, 1978 Chilodontopsis numerosus Ozaki and Yagiu, 1941 Order Nassulida Jankowski, 1967 Suborder Nassulina Jankowski, 1967 Family Nassulidae de Fromentel, 1874 Nassula citrea Kahl, 1931 Nassula labiata Kahl, 1933 Nassula rosea Tucolesco, 1962 Nassula sp.

Suborder Microthoracina Jankowski, 1967 Family Leptopharyngidae Kahl, 1926 Leptopharynx eurystomus Kahl, 1931

Family Microthoracidae Wrzesnioski, 1870 Drepanomonas revoluta Penard, 1922

Order Cyrptophorida Faure-Fremiet, 1956 Suborder Chlamydodntina Deroux, 1976 Family Chiliodonellidae Deroux, 1970

25

26 26 26 26

25

26

25; 26;

26

26

26

26

15; 24

7; 8; 10; 11; 15-18; 24-26

27 18; 26 27 18; 26 26 27

18

17; 18 12 12 16-18

12; 26

12

18; 26

25

26

26

27

25

12

Species RC1 BC NW NE References2

Chilodonella aplanata Kahl, 1931 X 26

Chilodonella calkinsi Kahl, 1928 x 26

Chilodonella cucullulus (O.F.M., 1786) Kahl, 1931 x X X 17; 18; 25; 26

Chilodonella nana Kahl, 1928 X X 16-18; 26

Chilodonella osrima Tucolesco, 1962 X 27

Odontochlamys gouraudi Certes, 1891 X 12

Family Chlamydodontidae Stein, 1859

Chlamydodon cyclops Entz., 1884 X X 7; 8; 15; 26

Chlamydodon erythromaculatus Tucolesco, 1962 X 27

Chlamydodon mnemosyne Ehrenberg, 1837 X X 17; 18; 26

Chlamydodon obliquus Kahl, 1931 X 26

Chlamydodon triquertus (O.F.M., 1786) Dragesco, 1960 X X 7; 9; 26

Chlamydodon sp. X 25

Family Lynchellidae Jankowski, 1968

Lynchella eforiana Tucolesco, 1962 X 27

Suborder Dysteriina Deroux, 1976

Family Hartmannulidae Poche, 1913

Hartmannula acrobates Entz., 1884 X 26

Hartmannula ocellata Tucolesco, 1962 X 27

Paratrochilia chilodontoides Kahl, 1933 X 26

Trochilioides artemiae Tucolesco, 1962 X 27

Trochilioides dubia Wallengern, 1903 X 1; 2; 7

Trochilioides striata (Buddenbrock, 1920) Kahl, 1928 X 26

Family Dysteriidae Claparede et Lachmann, 1858

Dysteria kahli Tucolesco, 1962 X 27

Dysteria monostyla (Ehrenberg) Stein, 1859 X X 1; 7; 26

Dysteria peneckei (Lepsi, 1926) Kahl, 1931 X 2; 7

Dysteria procera Kahl, 1931 X 26

Dysteria proraefrons James-Clark, 1866 X 26

Dysteria pusilla Claparede et Lachmann, 1858 X 26

Trochilia sigmoides Dujardin, 1841 X 26

Trochilia sp. X 1

Class Oligohymenophora de Puytorac et al., 1974

Subclass Hymenostomata Delage et Herouard, 1896

Order Hymenostomatida Delage et Herouard, 1896

Suborder Tetrahymenina Faure-Fremiet et Corliss, 1956

Family Tetrahymenidae Corliss, 1952

Colpidium campylum (Stokes, 1886) Bresslau, 1922 X 26

Colpidium colpoda (Ehrenberg, 1838) Stein, 1860 X 26

Stegochilum fusiforme Schewiakoff, 1892 X 17; 18

Tetrahymena patullum (Claparede et Lachmann, 1859) Corliss, 1971 X 7; 9; 16; 17

Tetrahymena pyriformis (Ehrenberg, 1830) Furgason, 1940 X 12

Family Glaucomidae Corliss, 1971

Glaucoma scintillans Ehrenberg, 1838 X 2; 12; 17; 18

Suborder Peniculina Faure-Fremiet et Corliss, 1956

Family Parameciidae Dijardin, 1840

Paramecium aurelia O.F.M., 1773 X 4-7

Paramecium bursaria (Ehrenberg, 1831) Focke, 1836 X 26

Paramecium calkinsi Woodruff, 1921 X X 16-18; 26

Paramecium caudatum Ehrenberg, 1838 X X 17; 18; 26

Paramecium putrinum Claparede et Lachmann, 1858 X 26

Family Frontoniidae Kahl, 1926

Frontonia algivora Kahl, 1931 X 26

Frontonia arenaria Kahl, 1933 X X X 10; 14; 15; 23; 24; 26

Species RC1 BC NW NE References2

Frontonia depressa (Stokes, 1886) Kahl, 1931 X 17; 18

Frontonia elliptica Beardsley, 1902 X 26

Frontonia fusca Quennerstedt, 1869 X X X 7; 9; 25; 26;

Frontonia leucas (Ehrenberg, 1833) Ehrenberg, 1838 X X 17; 18; 25

Frontonia marina Fabre-Domergue, 1891 X X X X 7; 9; 10; 14-17; 23-26

Frontonia microstoma Kahl, 1931 X 26

Frontonia vacuolata Dragesco 1960 X 23

Family Urocentridae Claparede et Lachmann, 1858

Urocentrum turbo (O.F.M., 1773) Kahl, 1931 X 17; 18

Family Lembadionidae Jankowski et Corliss, 1979

Lembadion sp. X 17; 18

Order Scuticociliatida Small, 1967

Suborder Philasterina Small, 1967

Family Philasteridae Kahl, 1931

Philaster resedaceus Tucolesco, 1962 X 27

Philaster salinus Tucolesco, 1962 X 27

Paralembus digitiformis Kahl, 1933 X 13

Parauronema virginianum Thompson, 1967 X 13; 16; 17; 18

Family Uronematidae Thompson, 1964

Homalogastra setosa Kahl, 1926 X 12

Uronema acuminata Kahl, 1926 X X 14; 23; 25

Uronema acutum Budd., 1920 X 26

Uronema constanziana Tucolesco, 1962 X 27

Uronema elegans Maupas, 1883 X X 1; 3; 7; 17; 18; 26

Uronema filificum Kahl, 1931 X 26

Uronema halophila (Kahl, 1931) Jankowski, 1964 X 17; 18

Uronema marina Dujardin, 1841 X X X X 14-18; 23-26

Uronema nigricans Thompson et Evans, 1968 X 1; 2

Uronema tortum (Maupas, 1883) Kahl, 1931 X 26

Uropedalium opisthostomum (Lepsi, 1926) Kahl, 1931 X 1; 2; 7

Uropedalium pyriforme Kahl, 1928 X 26

Family Cohnilembidae Kahl, 1933

Cohnilembus longivelatus Kahl, 1933 X 13

Cohnilembus pusillus Quennerstedt, 1869 X X 18; 26

Cohnilembus reesi Kahl, 1933 X X 1; 26

Cohnilembus stichotricha Kahl, 1928 X 26

Cohnilembus subulatus Kent, 1882 X 26

Cohnilembus versinus (O.F.M., 1786) Kahl, 1933 X X 1; 3; 7; 16-18; 26

Cohnilembus vexillifer Entz, 1879 X 26

Family Loxocephalidae Jankowski, 1964

Loxocephalus ellipticus Kahl, 1933 X 18

Cardiostomatella vermiforme Kahl, 1928 X X X 16-18; 25; 26

Family Cinetochilidae Perty, 1852

Cinetochilum margaritaceum (Ehrenberg, 1831) Perty, 1852 X X 12; 17; 18; 26

Cinetochilum marinum Kahl, 1931 X X 13; 16-18; 26

Platynematum marinum Kahl, 1933 X 26

Sathrophilus marinum Groliere et Detcheva, 1979 X 20

Family Pseudocohnilembidae Evans et Thompson, 1964

Pseudocohnilembus pusillus (Quennerstedt, 1869) Evans et

Thompson, 1964 X X 1; 2; 7; 16-18; 26

Insertae sedis in Suborder Philasterina

Anophryoides salmacida (Mugard) de Puytorac et Groliere X 18

Aristerostoma marina Kahl, 1931 X 13; 16-18

Protocruzia adherens (Mansfeld, 1923) Kahl, 1930 X X 13; 16-18; 26

Species RC1 BC NW NE References2

Protocruzia contrax (Mansfeld, 1923) Kahl, 1930 X 26

Protocruzia labiata Kahl, 1932 X 26

Protocruzia pigerrima (Cohn, 1866) da Cunha, 1915 X X 13; 16-18; 26

Protocruzia tuzeri Villeneuve-Brachon, 1940 X 21

Suborder Pleuronematina Faure-Fremiet in Corliss, 1956

Family Pleuronematidae Kent, 1880-1882

Pleuronema coronatum Kent, 1881 X X X X 11; 17; 15; 18; 22-26

Pleuronema crassum Dujardin, 1836 X X X 1; 7; 11; 15; 17; 18; 24; 25

Pleuronema marinum Dujardin, 1841 X X X 4-6; 11; 15; 24-26

Pleuronema nana Tucolesco, 1962 X 27

Pleuronema smalli Dragesco, 1968 X 16; 17; 19

Family Cyclidiidae Ehrenberg, 1838

Cristigera cirrifera Kahl, 1928 X X 25; 26

Cristigera media Kahl, 1928 X X 25; 26

Cristigera setosa Kahl, 1928 X X 25; 26

Cristigera setosa f. tricincta Kahl, 1928 X 26

Cristigera sulcata Kahl, 1933 X X 25; 26

Cyclidium candens Kahl, 1928 X X 25; 26

Cyclidium citrullus (Cohn, 1866) Kahl, 1931 X X 16-18; 26

Cyclidium curvatum Mansfeld, 1922 X 26

Cyclidium elongatum (Schewiakoff, 1889) Kahl, 1931 X X 3; 7; 12; 16-18; 26

Cyclidium glaucoma O.F.M., 1786 X X X 1; 3; 7; 9; 12; 17; 18; 25; 26

Cyclidium lanugenosum Penard, 1922 X 26

Cyclidium longisetosum Penard, 1922 X 25

Cyclidium oligotrichum Kahl, 1928 X 26

Cyclidium simulans Kahl, 1928 X 26

Cyclidium veliferum Kahl, 1933 X X 25; 26

Family Histiobalantiidae de Puytorac et Corliss, 1979

Histiobalanthium majus Kahl, 1931 X 25

Histiobalanthium marinum Kahl, 1933 X 22

Subclass Peritricha Stein, 1859

Order Peritrichida Stein, 1859

Family Vorticellidae Ehrenberg, 1838

Pseudovorticella plicata Gourret et Roeser, 1886 X 13

Vorticella campanula Ehrenberg, 1831 X X 3; 7; 14; 17; 18; 23

Zoothamnium affine Stein, 1854 X 1; 7

Family Scyphidiidae Kahl, 1933

Scyphidia rugosa Dujardin, 1841 X 12

Family Vaginicolidae de Fromentel, 1874

Cothurnia compressa Claparede et Lachmann, 1858 X 7; 9

Cothurnia innata (O.F.M., 1786) Hofker, 1930 X 6

Platycola decumbens Ehrenberg, 1830 X 17; 18

Pyxicola socialis (Gruber, 1879) Kent, 1881 X 4-7

Vaginicola ingenita O.F.M., 1786 X 6

Vaginicola ovata Dons, 1922 X 7; 9

Class Polyhymenophora Jankowski, 1967

Subclass Spirotricha Butschli, 1889

Order Heterotrichida Stein, 1859

Suborder Heterotrichina Stein, 1859

Family Spirostomidae Stein, 1867

Angisteinia clarissima (Anigstein, 1912) Isquith, 1968 X X X X 7; 8; 10; 11; 14; 15; 23-25

Species RC1 BC NW NE References2

Angisteinia clarissima f. arenicola (Kahl, 1932) Isquith, 1968 X X X 10; 15; 24

Angisteinia salinarum (Florentin, 1899) Isquith, 1968 X X 13; 26

Blepharisma hyalinum Perty, 1849 X 12

Blepharisma musculus Penard, 1922 X 13

Blepharisma steini Kahl, 1932 X 17; 18

Blepharisma vestitum Kahl, 1932 X 16; 17

Blepharisma violacea Tucolesco, 1962 X 27

Gruberia calkinsi Beltran, 1933 X X 10; 14; 15; 23; 26

Gruberia uninucleata Kahl, 1932 X X X X 10; 11; 14-18; 23; 24

Parablepharisma chlamydophorum Kahl, 1932 X 26

Parablepharisma collare Kahl, 1932 X 26

Parablepharisma kahli Tucolesco, 1962 X 27

Parablepharisma pellitum Kahl, 1932 X 26

Spirostomum ambiguum (O.F.M., 1786) Ehrenberg, 1838 X 17; 18

Spirostomum teres claparede et Lachmann, 1858 X 4; 6; 7

Family Metopidae Kahl, 1927

Metopus contortus (Quennerstedt, 1867) Kahl, 1932 X X X 15; 25-27

Metopus halophila (Kahl, 1925) Corliss, 1960 X 26

Metopus hasei Sondheim, 1929 X 27

Metopus striatus McMurrich, 1884 X 26

Metopus setosus Kahl, 1927 X 25

Metopus vestitus Kahl, 1935 X 26

Family Condylostomidae Kahl in Doflein et Reichenow, 1929

Condylostoma acuta Dragesco, 1960 X 25

Condylostoma arenaria Spiegel, 1926 X X X X 4; 6-8; 10; 11; 14; 15;

17; 18; 23-26

Condylostoma magnum Spiegel, 1926 X X X 7; 9; 14; 26

Condylostoma oxyoura Dragesco, 1960 X 15

Condylostoma patens (O.F.M., 1786) Dujardin, 1841 X X 7; 9; 16; 17; 26

Condylostoma patulum Claparede te Lachmann, 1858 X 7; 9

Condylostoma psammophila Bock, 1952 X 25

Condylostoma remanei Spiegel, 1928 X X X 7; 8; 10; 11; 15-18; 23; 24

Condylostoma rugosa, Kahl, 1932 X 26

Family Stentoridae Cams, 1863

Stentor coeruleus (Pallas, 1766) Ehrenberg, 1830 X 4-7; 17; 18

Stentor polymorphus (O.F.M., 1773) Ehrenberg, 1830 X 5; 6

Stentor roeselii Ehrenberg, 1835 X 4; 6; 7

Family Bursariidae Dujardin, 1840

Thylakidium truncatum Schewiakoff, 1892 X 17; 18

Family Peritromidae Stein, 1867

Peritromus faurei Kahl, 1932 X 15; 24; 26

Peritromus kahli Tucolesco, 1962 X 27

Peritromus sandinae Tucolesco, 1962 X 27

Suborder Armophorina Jankowski, 1964

Family Caenomorphidae Poche, 1913

Caenomorpha levanderi Kahl, 1927 X 25

Suborder Coliphorina Jankowski, 1967

Family Folliculinidae, Dons, 1914

Folliculina producta Wridgt, 1859 X 4; 6; 7

Suborder Licnophorina Corliss, 1957

Family Licnophoridae Butschli, 1887

Licnophora sp. X 1

Species RC1 BC NW NE References2

Order Oligotrichida Butschli, 1887

Suborder Oligotrichina Butschli, 1887

Family Halteriidae Claparede et Lachmann, 1858

Halteria grandinella (O.F.M., 1773) Dujardin, 1841 X X 12; 17; 18; 25

Family Strombidiidae Kahl in Doflein et Reichenow, 1929

Strobilidium minimum Gruber, 1884 X 26

Strobilidium polyhalinum Tucolesco, 1962 X 27

Strombidium arenicola Dragesco, 1960 X 11; 15; 24

Strombidium calkinsi Faure-Fremiet, 1932 X 25

Strombidium cinctum Kahl, 1932 X 25

Strombidium conicum (Lochmann, 1908) Wulff, 1919 X X 25; 26

Strombidium costa turn Tucolesco, 1962 X 27

Strombidium elegans Florentin, 1899 X X 25; 26

Strombidium lagenula Faure-Fremiet, 1924 X 26

Strombidium obliquum Kahl, 1932 X 26

Strombidium opisthostomum Tucolesco, 1962 X 27

Strombidium sauerbreyae Kahl, 1932 X X X 10; 11; 15; 24

Strombidium sulcatum Claparede et Lachmann, 1858 X X X 22; 25-27

Strombidium turbo Claparede te lachmann, 1859 X 4; 6; 7

Strombidium viridae Stein, 1867 X 25

Order Hypotrichida Stein, 1859

Suborder Stichotrichina Faure-Fremiet, 1961

Family Spirofilidae von Gelei, 1929

Kahliela leptocirra Tucolesco, 1962 X 27

Stichotricha opisthotonoides Smith, 1877 X 26

Stichotricha simplex Kahl, 1932 X 16; 17

Family Strongylidiidae Faure-Fremiet, 1961

Urostrongylum caudatum Kahl, 1932 X 25

Urostrongylum contortum Kahl, 1928 X 26

Family Urostylidae Butschli, 1889

Urostyla halseyi Calkins, 1929 X 12

Family Holostichidae Faure-Fremiet, 1961

Amphisiella annulata (Kahl, 1928) Kahl, 1932 X 17; 18

Amphisiella capitata (Perejaslawzeva, 1886) Kahl, 1932 X X 10; 13; 14; 16-18; 23

Amphisiella marioni Gourret et Roeser, 1888 X 17; 18

Gonostomum affine (Stein, 1859) Sterki, 1878 X 12

Holosticha diademata (Rees, 1883) Kahl, 1935 X X X 16-18; 25; 26

Holosticha fasciola Kahl, 1935 X 25

Holosticha kessleri (Wrzesnioski, 1877) Kahl, 1935 X X 14; 23; 25

Holosticha manca Kahl, 1932 X 26

Holosticha teredorum Tucolesco, 1962 X 27

Keronopsis arenivorus Dragesco, 1954 X X 11; 15; 24

Keronopsis flavicans Kahl, 1932 X 26

Keronopsis monilata Kahl, 1928 X 26

Keronopsis ovalis Kahl, 1932 X 16; 17

Keronopsis pernix (Wrzesniowski, 1877) Kahl, 1932 X X 7; 8; 14; 23

Paruroleptus lacteus Kahl, 1832 X 12

Psammomitra retractilis (Claparede et Lachmann, 1835) Borror, 1972 X 17; 18

Pseudokeronopsis rubra (Ehrenberg, 1835) Borror et Wicklow, 1983 X X X 10; 14-18; 23; 26

Trachelostyla caudata (Kahl,1935) Maeda et Carey, 1984 X X X X 10; 11; 14; 15; 22-26

Trachelostyla pediculiformis (Kahl, 1935) Maeda et Carey, 1984 X X X 7; 8; 15-18; 25-26

Uroleptopsis citrina Kahl, 1932 X 13; 17; 18

Uroleptus sp. X 16; 17; 18

Species RC1 BC NW NE References2

Family Keronidae Dujardin, 1840

Epiclintes ambiguus (O.F.M., 1786) Butschli, 1889 X X 10; 15; 24

Suborder Sporadotrichina Faure-Fremiet, 1961

Family Oxytrichidae Ehrenberg, 1838

Gastrostyla dubia (Dragesco, 1954) Maeda and Carey, 1984 X X X 15; 23; 24

Gastrostyla pulchra (Perejaslawzewa, 1886) Kahl, 1935 X X 25; 28

Histriculus similis (Quennerstedt, 1867) Kahl, 1930-1935 X 26

Histriculus sp. X 17; 18

Opisthotricha halophila Kahl, 1932 X 26

Oxytricha aerugenosa Wrzesniowski, 1870 X 25

Oxytricha discifera Kahl, 1935 X X X 10; 15; 25

Oxytricha gibba (O.F.M., 1773) Stein, 1859 X X 11; 15; 24

Oxytricha marina Kahl, 1932 X 26

Steinia marina Kahl, 1932 X X 16-18; 25

Stylonychia mytilus (O.F.M., 1773) Ehrenberg, 1830 X X X 4-7; 12; 17; 18; 25; 26

Stylonychia purtina Stokes, 1885 X 14; 23

Tahysoma rigescens Kahl, 1932 X 13; 16-18

Urosoma caudata Stokes, 1887 X X 25; 26

Family Aspidiscidae Ehrenberg, 1838

Aspidisca aculeata Ehrenberg, 1838 X 26

Aspidisca binucleata Kahl, 1932 X 26

Aspidisca cicada (O.F.M., 1786) Claparede et Lachmann, 1858 X X 14; 23; 26; 27

Aspidisca dentata Kahl, 1928 X X 17; 18; 25

Aspidisca fusca Kahl, 1928 X X 17; 18; 26

Aspidisca lyncaster (O.F.M., 1779) Stein, 1895 X X 11; 15; 24

Aspidisca nana Tucolesco, 1962 X 27

Aspidisca pulcherrima Kahl, 1932 X X 13; 26

Aspidisca sedigita Quennerstedt, 1867 X X X 13; 25; 26

Aspidisca steini Buddenbrock, 1920 X X 7; 8; 17; 18; 25

Aspidisca tuberosa Kahl, 1932 X 26

Family Euplotidae Ehrenberg, 1838

Diophrys appendiculata (Ehrenberg, 1838) Kahl, 1935 X X X 1; 7-9; 15-18; 25; 26

Diophrys scutoides Agamaliev, 1967 X 25

Diophrys scutum Dujardin, 1841 X X X X 7; 10; 11; 14; 15-18;

23-26

Discocephalus rotatorius Ehrenberg, 1831 X X 7; 25; 29

Euplotes affinis Dujardin, 1842 X 26

Euplotes alatus Kahl, 1932 X 26

Euplotes balteatus Dujardin, 1841 X X 26; 13

Euplotes charon (O.F.M., 1786) Ehrenberg, 1830 X X X X 1; 7; 8; 11; 14; 15; 2326;

Euplotes crassus (Dujardin, 1841) Kahl, 1932 X 13

Euplotes cristatus Kahl, 1932 X 16; 17; 18

Euplotes elegans Kahl, 1932 X 26

Euplotes elegans f. littoralis Kahl, 1932 X 26

Euplotes extensus Fresenius, 1865 X 26

Euplotes gracilis Kahl, 1932 X 26

Euplotes harpa Stein, 1859 X X X 1; 7; 8; 11; 14; 15-18;

23; 24; 26

Euplotes moebiusi Kahl, 1932 X X 7; 9; 26

Euplotes moebiusi f. quadricirratus Kahl, 1932 X 26

Euplotes patella (O.F.M., 1773) Ehrenberg, 1838 X 17; 18

Euplotes trisulcatus Kahl, 1932 X X X 13; 16; 17; 18; 25; 26

Species

RC1

BC

NW

NE

References2

Euplotes vannus O.F.M., 1786

Swedmarkia arenicola Dragesco, 1954 Ur onychia magna Pierantoni, 1909 Uronychia transfuga (O.F.M., 1786) Stein, 1859

1; 7; 11; 14; 15; 23; 24; 26 13 18; 26

10; 11; 14-18; 23-26

1 RC - Romanian coasts, BC - Bulgarian coasts, NW - northwestern part (the Gulf of Odessa and the Crimea), NE -northeastern part (Russian shore sector from Adler to Anapa).

2 1 - Lepsi, 1926a; 2 - Lepsi, 1926b; 3 - Jeliaskova-Paspalewa, 1933; 4 - Valkanov, 1934; 5 - Valkanov, 1935; 6 - Valkanov, 1936; 7 - Valkanov, 1957; 8 - Czapik, 1952a; 9 - Czapik, 1952b; 10 - Kovaleva, 1966; 11 - Bacescu et al, 1967; 12 -Detcheva, 1973; 13 - Detcheva, 1977; 14 - Djurtubaev, 1977; 15 - Kovaleva and Golemansky, 1979; 16 - Detcheva, 1980; 17 - Detcheva, 1981; 18 - Detcheva, 1983; 19 - Groliere and Detcheva, 1974; 20 - Groliere and Detcheva, 1979; 21 - Groliere et al., 1974; 22 - Gulin et al., 1986; 23 - Djurtubaeva and Mokhammed, 1995; 24 - Petran, 1997; 25 - Azovsky and Mazei, 2003; 26 - Tucolesco, 1961; 27 - Tucolesco, 1962; 28 - Perejaslawzewa, 1886; 29 - Valkanov, 1954.

in one region, whereas only 34 species occurred in all the four regions. Most of them are widespread or cosmopolitan, specifically psammophilous forms.

Taxonomic diversity. Analysis of taxonomical diversity statistics showed that NW and NE regions were distinguished obviously and highly significantly. Their average taxonomic diversity was much lower than it could be expected from the null-hypothesis (the actual points lay far from the 5% funnel, Fig. 2A).

Their diversity variances were also significantly above the expected level (Fig. 2B), suggesting a very uneven representation of different higher taxa. Indeed, in NW region, the orders Colpodida, Nassulida and Cryptophorida are absent, Scuticociliatidae is represented by only 4 species (out of 59 noted in the total list); Haptorida - by 3 species out of48; Trichostomatida

- by the one species out of 20. On the other hand, more than 50% of the species fall into the orders Karyo-relictida and Hypotrichida (Table 1). A similar picture could be seen for NE region: Colpodida and Peritricida orders are absent, there is only one Nassulida species (out of 6), two Cryptophorida species (out of 27), and four

Trichostomatida species (out of 21). At the same time, almost half of the fauna is constituted by Karyorelictida, Prostomatida and Haptorida species; Synhymeniida is represented in this region only.

Faunas of Bulgarian and Romanian coasts are richer and more taxonomically diverse. Among the genera reported for these regions, 38 genera (33%) were found only at BC and 21 (22%), only at RC.

S/G ratio for the sea fauna on the whole was 3.24, being rather close to the values obtained for other relatively well-studied seas: Caspian - 3.36; White -3.18; Baltic - 2.89; North Atlantic - 3.33 (Agamaliev, 1983; Azovsky, 1996). However, for the separate sea parts except Bulgarian coast, the S/G ratios were significantly higher than expected (Fig. 2C), indicating that more congenerics were found at each of these regions.

Comparison of species composition between regions.

Comparison of the regions using S0rensen’s similarity index showed that the two richest regions, Romanian and Bulgarian costs, were also the most similar by their species composition, whereas the poorest north-western

Table 2. Characteristics of taxonomic diversity of the Black Sea benthic ciliates.

Whole sea Regions

RC BC NW NE

Number of species 476 299 253 83 144

Species found in one region only 292 148 98 11 35

Number of genera 147 98 114 39 60

Genera found in one region only 62 21 38 0 3

Species/genera ratio 3.24 3.05 2.22 2.13 2.40

Average taxonomic diversity 4.465 4.443 4.469 4.227 4.348

Variance of taxonomic diversity 0.644 0.664 0.682 1.102 0.791

part significantly differed from the others (Table 3, Fig. 3A). As can be seen from the dendrogram, all regions were grouped according to their total richness. Indeed, when the Simpson’s index was used, which is insensitive to the disparity in total number of species, this difference disappeared (Fig. 3B). All pair-wise similarities were significantly higher than it could be expected from «random species’ occurrence» null-hypothesis.

So, the ciliofaunas of all four regions could be treated as derivatives from the common species pool, the differences between them mainly resulting from the inequality in total number of species. More detailed investigations are necessary to reveal any possible differences in faunistic composition.

Discussion

Analysis ofbetween-region similarity indices shows that the regional faunas could be treated as derivative subsets from the total species pool, and the differences observed in their species composition may be partially explained by differences in the summary sampling effort and hence in total number of species found. Nevertheless, the regions appear to be much less similar by their species composition than it was reported, for example, for different parts of the Caspian Sea (Agamaliev, 1983). Taxonomic diversity indices also indicate a significant disproportion in the taxonomic composition. High S/G ratios also confirm the fact that some genera were overrepresented in regional lists while some others were under-represented. It may be caused by discordance in the approaches used by the researches who worked at different times and in different regions. The specificity of Romanian and Bulgarian fauna mentioned above may be explained by the fact that some authors had considered as «benthic» many species found in rather specific habitats (brackish-water littoral sites, macroalgae, attached Peritricha, etc.) (Tucolesco, 1961, 1962; Petran, 1963, 1968, 1976, 1997; Detcheva, 1981, 1983). This taxonomic dispro-portion may be a result of different attention paid to some groups of ciliates by one or another author due to the personal differences in experience and techniques used. This is supported by the fact that most groups under-represented in NE and NW regions are small-sized, not specifically psammophilious forms. Besides, some studies carried out in these regions were extended to the upper sublittiral zone where the specific psammophilous forms, such as Kariorelictids, are rather numerous (Djurtubaeva and Mokhammed, 1995; Azovsky and Mazei 2003), whereas investigations on RC and BC regions were only restricted to the littoral zone.

More than half of all the species were found in one part of the sea only, and only 7% of species were

common for all four regions investigated. This situation is rather typical for insufficiently explored regions. For example, out of178 species found in Britain, only about 50% have been found at more than one site (Patterson et al., 1989). Analysis of the literature shows that the proportion of species, reported as new for the sea, does not change noticeably during the last half of the century.

Fig. 2. Taxonomic diversity characteristics of the whole sea and separate regions ordered by total species number: A - average taxonomic diversity, AvTD; B - variance of diversity, VarTD; C - species/ genera ratio. The 95 % probability funnels (solid lines) and the mean expected levels (dotted lines) are shown.

Table 3. Faunistic similarities between the regions. Above the diagonal - Czekanowski-Serensen index, below the diagonal -Simpson's index.

Regions RC BC NW NE

RC - 0.44 + 0.26 - - 0.39

BC 0.48 + - 0.38 0.42

NW 0.60 ++ 0.77 ++ - 0.40

NE 0.61 ++ 0.59 ++ 0.54 ++ -

Notes: Signs after the values mark the significant deviations from the expectations of the random occurrence model (one symbol for 95% and two - for 99% confidence funnel).

These values, expressed as the percent of the total number of species found in the study, make up: 24.8% (Tucolesco, 1961, 1962); 18.6% (Petran, 1968); 19.2% (Kovaleva and Golemansky, 1979); 26.3 (Detcheva, 1977, 1980, 1983); 16.3 (Djurtubaeva and Mokhammed, 1995); 22.9 (Azovsky and Mazei, 2003). Such constant increment of faunistic richness indicates that the Black Sea ciliofauna is still explored rather imperfectly. Among the species we found recently for the first time, there were such widespread forms as Didinium nasutum, Tracheloraphis oligostriata, Histiobalanthium majus; and also some ciliates common for other seas but earlier registered in the Black Sea only once (Gastrostyla pulchra, Discocephalus rotatorius) (Azovsky and Mazei, 2003). This fact is another evidence of the imperfection of our knowledge about the fauna.

These circumstances can apparently explain the noticeable discrepancy in the estimates, reported earlier, of the known richness of the fauna: 226 species -Tucolesco, 1961; 300 - Burkovsky, 1984; 109 - Kovaleva and Golemansky, 1979; 116 - Agamaliev, 1983; over 160

- Djurtubaeva and Mokhammed, 1995). These estimates were obviously based on rather incomplete data and are greatly understated. Our total list includes now 476 species. It is obvious that this list must be viewed with some caution and needs a critical review. In particular, this concerns some early inves-tigations (Lepsi, 1926a, 1926b; Tucolesco, 1961, 1962; and some others). Many «new» ciliates reported by these authors are poorly described and can hardly be distinguished with confidence at the present-day level oftaxonomic require-ments. For example, among the ciliates found at Romanian coast, a half(149) were only reported by Tucolesco (1961, 1962). Among them, 47 species were first described by him (Tucolesco, 1962) and have never been mentioned later by anyone else, possibly because of poorly described diagnostic features. Thus these species were not included in the species list of marine benthic ciliates compiled by Carey (1991).

The statements about the distribution of species or any detailed between- or within-sea comparison must

be done with some caution because of undersampling and uncertainty over taxonomic practice. Inconsistent standards of taxonomic judgement may lead to biased estimates of species diversity (Patterson et al., 1989; Finlay, 1998; Foissner, 1999). Nevertheless, it is reasonably safe to suggest that the number of the Black Sea benthic ciliates recorded at present could be roughly considered as at least 400-450 bona fide species, even if some dubious species discussed above are not taken into account. To all appearance, a considerable increase of this figure can be expected. However, even now this conservative value is appreciably higher than the species richness of such relatively well-studied seas as the Caspian (351 species, Agamaliev, 1983; Alekperov and Asadullaeva, 1996; Agamaliev and Suleymanova, in press), the Baltic (318 species, Agamaliev, 1983) and the White Sea (about 260 species, Burkovsky, 1984; Mazei, 2002). Thus, the Black Sea benthic ciliofauna may be considered as the richest one known to date.

Acknowledgements

This work was supported by the Russian Fund for Basic Research (grant № 03-04-48018).

BC

NE----------

NW------------------------

I—I—I—I—I—I—I—I—I—I—f

0.44 0.39 0.34

B

BC

NW

RC

NE---------------------------------

i---------1---------1---------1--------1--------1

0.75 0.70 0.65 0.60 0.55 0.50

Fig. 3. Results of cluster-analysis of the regions based on species composition similarity. A - Czekanowski-S0rensen index; B - Simpson’s index.

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Address for correspondence: Andrey I. Azovsky. Department of Hydrobiology, Biology Faculty, Moscow State University, Moscow, 119899, Russia. E-mail: [email protected]

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