FIRST FINDING OF EPIBIONT PERITRICH AND SUCTORIAN CILIATES (CILIOPHORA) ON OLIGOCHAETES AND HARPACTICOID COPEPODS FROM THE DEEP-WATER HYPOXIC/ANOXIC CONDITIONS OF THE BLACK SEA

Three species of commensal ciliates ( Cothurnia maritima Ehrenberg, 1838 on oligochaete Tubificoides sp.; Paracineta livadiana (Mereschkowsky, 1881) and Corynophrya lyngbyi (Ehrenberg, 1834) on harpacticoid copepods Amphiascella subdebilis (Willey, 1935), Haloschizopera pontarchis Por, 1959, Cletodes tenuipes Scott, 1896 and Enhydrosoma longifurcatum Sars, 1909) were found in the Black Sea deep-water under hypoxic/anoxic conditions for the first time. Corynophrya lyngbyi is reported for the first time in the Black Sea.


Introduction
The Black Sea is known to be permanently anoxic below a water depth of about 120 m in the open sea, and of about 180 m at its shelves, down to its deepest parts at 2200 m depth.
Some authors (Zaitsev et al. 2007, Zaitsev andPolikarvov 2008) assume an absence of eukaryotic life in anoxic Black Sea waters and in hydrogen-sulfide sediments considered then as "toxic" environment.
Ciliates are present in aerobic Black Sea habitats, both in pelagic and benthic environments. The total number of species (29) found in pelagic aerobic waters (Zaika et al. 1976) is low compared to the benthic species diversity (476) Mazei 2003a, b, 2005;Polikarpov et al. 2003).
As a result of investigations of free-living psammophilic ciliate species composition on the Northeastern (Caucasian) coast of the Black Sea at a depth from 0 to 432 m, the highest diversity of ciliates was found in heterogeneous sands at a depth of 3-10 m, the lowest one -in unstable sands of the tidal zone and in silt deeper than 25 m. At the same time no ciliates were found in the zone enriched with H 2 S (deeper than 100 m) (Azovsky and Mazei 2005).
Earlier studies of the oxic/anoxic chemocline of the Black Sea water column revealed only two ciliate species: one of them was being a representative of the Mesodiniidae, and the other one was a small unidentified ciliate with symbiotic bacteria on its surface (Sazhin et al. 1991). However, it is possible that previous deep water sampling methods were inadequate for microorganisms, thus abundance and diversity of ciliates below 100 m water depth were underestimated.
Later additional data concerning taxonomic richness and abundance of deep-water Protozoa and Metazoa in hypoxic and anoxic zones were obtained (Sergeeva et al. , 2011a(Sergeeva et al. , b, 2013. Several ciliate

Research Article
and foraminifera taxa have been known to occur in anoxic sediments of the Black Sea and could live in the Black Sea (e.g. Sergeeva et al. 2011a). The bathymetric distribution of the benthic ciliates at depths from 120 to 2075 m near the Dnieper Canyon and the Sorokin Trough (eastern part of the Black Sea) was described from samples of near bottom water, sediment surface detritus, and the upper layer (0-1 cm) of sediment (Zaika & Sergeeva 2009). The ciliates found in mentioned samples were the representatives of genera Chilodonella Strand, 1928, Trachelocerca Ehrenberg, 1834, Tracheloraphis Dragesco, 1960and Loxophyllum Dujardin, 1841. At the same time more than 30 morphological species were recognized among mentioned materials . The peaks of ciliate abundances were registered at depths of 120, 160-190, and 240 m in the same region .
It should be noted that all ciliate species identified in the Black Sea under hypoxic/anoxic conditions were free-swimming forms.
Thus the present article represents the first investigation of commensal ciliates on hosts from the mentioned peculiar biotope.

Material and methods
The sediment sampling was performed in the Bosporus Strait's outlet area of the Black Sea during the RV 'Maria S. Merian' (Germany) cruise.
For benthos studies we got virtually undisturbed bottom sediments at the stations in the range of depths 80-300 m by TV-multi-corer Ø=9.5cm or gravity corer Ø=7cm. Benthic stations were chosen along this transect, ranging from the oxic to the anoxic zone. Sediment cores, obtained by the multiple-corer, were sectioned at 0-1, 1-2, 2-3, 3-4 and 4-5 cm intervals and all samples immediately preserved in 75% ethanol. The sediments were washed by distilled water through sieves with a mesh size of 1 mm and 63 µm. The organisms recovered in the sieves fractions were stained with vital Rose Bengal and investigated in the Bogorov's chamber using a stereomicroscope.
The isolated organisms were first identified up to higher taxa and then were examined and identified to the lowest possible taxa using the Olympus CX41 microscope. Photomicrographs were obtained by using an Olympus E-410 digital camera. Measurements of ciliates were made using the computer program Scope Photo v. 2.0 for processing of digital images.
For slide preparation the material was mounted in Canada balsam. Permanent slides of oligochaetes and harpacticoids were deposited in the collections of the Department of Ecology of Benthos of the Institute of Biology of the Southern Seas, Sevastopol.
The systematic positions of ciliate species were determined according to review works on peritrichs (Warren and Paynter 1991) and suctorians (Curds 1987, Dovgal 2002, 2013.

Results
The macrobenthic oligochaetes were found in bottom sediments almost at all studied depths (80-253 m). We found three ciliate species from the body surface of oligochaetes and harpacticoids recovered from two stations (Table 1) at the depths of 200 and 248 m. The oligochaetes were identified belonging to the genus Tubificoides Lastčkin, 1937. The 58 oligochaete individuals were analyzed and 13 (22.5%) of them were infected with perictrich ciliates Cothurnia maritima Ehrenberg, 1838.
The systematic positions and descriptions of found ciliate species are given below.
It should be mentioned that in our materials the individuals with furrows of lorica neither were observed.

Measurements
It should be mentioned that length of the stalk in observed individual of the species was not exceeding the height of the body.
Marine species, attached to various substrates such as hydroids and algae. This is the first record of the species in the Black Sea.
The distribution of ciliates on the body surface of both oligochaete and harpacticoid hosts had no a specific distributional pattern at certain areas, and were distributed from the head to the end of the body. In the case of the harpacticoid hosts the suctorian ciliates were attached on the antennae, furcas, carapaces and egg sacs.