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57
Fauna of testate amoebae of western Azerbaijan rivers
Nataly Snegovaya and Ilham Alekperov
Laboratory of Protistology, Institute of Zoology, Azerbaijan National Academy of Sciences, Baku
Summary
Testate amoebae fauna of 5 rivers in Western Azerbaijan was studied. Altogether, 70 species were found. The morphology of 33 species is described. Two families, 6 genera and 23 species are new for science (Centropyxis kurakchayensis sp. n., Difflugia ogdenii sp. n., D. armatostoma sp. n., D. rotiferoformissp. n., D. caucasica sp. n., D. vermiformis sp. n., D. bifurcatasp. n., D. azerbaijanicasp. n., Pontigulasia breviottissp. n., Schwabia sphaerica sp. n., Pelecyamoeba stenostoma sp. n., Armatodifflugia ceratophora sp. n., A. cuneata sp. n., Fabalesquereusia graniformis sp. n., F. compressa sp. n., F. linearis sp. n., Shamkiriella convoluta sp. n., Sh. reticulata sp. n., Sh. phimatophora sp. n., Bipseudostomatella bifurcata sp. n., B. gracilis sp. n., B. cornuta sp. n., Gomocollariella ranaformis sp. n.). Most of the other species are new for the Caucasus and Azerbaijan fauna.
Key words: freshwater testate amoebae, fauna, Caucasus
Introduction
The fauna of testate amoebae of Azerbaijan is studied incompletely. There was a single special research on soil testate amoebae (Zaidov, 1988, 1990, 1995) and several researches of general nature, where soil testate amoebae were studied together with other soil Protozoa (Ibadov, 1983; Mirza-zadeh, 1987). Prior our investigations (Alekperov and Snegovaya, 1999, 2000, 2001, 2002), aquatic testate amoebae were researched in
1930-40 by Veisig and Alizade within the framework of general hydrobiological investigations (Veisig, 1940; Alizade 1934).
Material and Methods
We have studied the fauna of testate amoebae in five rivers of Western Azerbaijan (Fig. 1) in 2002-2004. More than 100 samples of plankton, periphytone and benthos were collected and processed. Testate amoebae
© 2005 by Russia, Protistology
were caught by microcapillars and studied both in vivo and with the use of an original technique based on Chatton-Lwoff "wet" technique (Alekperov et al., 1994, 1996).
In short, the technique used was as follows:
1. If possible, concentrate testate amoebae from natural samples into a watch-glass;
2. Wash concentrate specimens with distilled water;
3. Place a small drop of concentrated specimens on a slide;
4. Quickly add an equally sized drop of molten 5% gelatin (60° C) and mix.
5. Immediately transfer the slide with a thin gelatin layer into cold and leave up to 5 minutes until gelatin has hardened;
6. Then transfer the slide to cold 3-5% cold silver nitrate solution for 25-30 minutes;
7. Wash the slide thoroughly with cold distilled water for 1-3 minutes;
8. Submerge the slide in 1-4 cm cold distilled water in a white-bottomed dish. Irradiate for 20 minutes using sunlight or an ultraviolet source, until gelatin turns golden brown;
9. Transfer slide to 70°-96° and 100° alcohol for 5 minutes each;
10. Clean by xylene for 10 minutes and mount in synthetic neutral mounting medium or Canadian balsam.
In the present report we give descriptions of 33 species, including all new ones. All descriptions are based on the observations of living specimens and the analysis of the material impregnated by silver nitrate. All measurements were made on not less than 5-15 specimens of each species. The type material of all the new taxa has been deposited in the collection of the Protistology laboratory in the Institute of Zoology of NAS of Azerbaijan, Baku.
A SHORT DESCRIPTION OF THE WATER RESERVOIRS INVESTIGATED
The Kurakchay river is a right tributary of the Kura river. It flows into the Varvarian reservoir. It runs on the territory of Evlakh and Goranboy districts. Its length is 186 km, and the catchment area is 2080 km2. Average mineralization of water is 150-350 mg/l. Water is hydrocarbonate-calcium class (Khasanov et al., 1973). The river bottom is stony, silt-covered sandy areas can be also seen. The depth is 0.35-0.75 m. Active reaction (pH) 7.0-7.6, content of O2, 8.18-10.56 mg/l. Aquatic plants include reed, reed mace, rush (Aliev, 2000).
The Korchay river is a left tributary of the Kurakchay river. It flows on the territory of Evlakh and Goranboy districts. The length of the river is 38 km, the catchment area is 51 km2 (Khasanov et al., 1973). The bottom is
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Fig. 1. Map of Azerbaijan.
stony or but pebbly-sandy. The depth of the river is from
0.4 to 0.7 m. Active reaction (pH) is 7.0-7.5, content of O2, 9.2-11.15 mg/l. Aquatic plants are absent (Aliev, 2000).
The Tovuzchay river is a right tributary of the Kura river. The length is 42 km, total catchment area is 278 km2 (Khasanov et al., 1973). The bottom is stony and pebbly-sandy, in some places silty and silty-sandy. The depth is 0.35-0.75 m. Active reaction (pH) is 6.9-7.4, content of O2, 9.55-11.26 mg/l. Reeds, rushes, reed maces are found in the coastal zone (Aliev, 2000).
The Shamkirchay river is a right tributary of the Kura river, it flows on the territory of Gedabek and Shamkir districts of Azerbaijan. The river length is 95 km, catchment area is 1170 km2 (Khasanov et al., 1973). The bottom is stony, sandy, pebbly, pebbly-sandy and in some places silty. The depth is 0.25-0.65 m. Active reaction (pH) is 7.0-7.5, content of O2 is 8.7512.65 mg/l. Aquatic plants are represented by reed mace and rush (Aliev, 2000).
The Kura river is the largest river in Azerbaijan. Its length is 1515 km, the catchment area is 188000 km2 (Kasimov, 1972). Active reaction (pH) is 6.5-7.9. Content of O2 is 6.55-13.86 mg/l (Aliev, 2000). The bottom is silty, silty-sandy, sandy and pebbly. Aquatic plants include rush, reed mace and reed.
Results
Altogether, we have found 70 species, 23 of which are new for science, and 15, for Caucasus fauna (Table 1).
Arcella hemisphaerica Perty 1852 (Fig. 2 a-b; Table 2).
The shell is circular, 40-50 ^m in diameter and 2025 ^m in height. The aperture (10-12 ^m) is invaginated, circular. The cell is attached to the inner shell wall with
Table 1. Specific composition and distribution of testate amoebae in Western Azerbaijan rivers.
Species
1 2 3 4 5
Fam. Arcellidae Ehrenberg 1830
Arcella hemispherica Perty 1852 +
A. megastoma Penard 1926* +
A. vulgaris Ehrenberg 1832 +
A. atava Collin 1914 ** +
A. polypora Penard 1902 + +
A. artocrea Leidy 1876 + +
A. gibbosa Penard 1890 +
Fam. Сyclopixidae Schonborn 1989
Cyclopyxis euristoma Deflandre 1929 + +
C. kahli Deflandre 1929 + +
Trigonopyxis arcula (Leidy 1879) + +
Fam. Сеntropyxidae Deflandre 1953
Centropyxis aculeata (Ehrenberg 1838) + + +
C. aerophila Deflandre 1929 + + + +
C. discoides (Penard 1902) + +
C. ecornis (Ehrenberg 1838) + + +
C. hirsuta Deflandre 1929 + + + +
C. cassis (Wallich 1864) + +
C. mirabilis Barto?1940 ** +
C. compressa van Oye 1948 ** +
C. percolabiensis Dekhtyar 1994 ** + +
C. spinosa Cash 1905 + + + + +
C. elongata (Penard 1890) + + + +
C. kurakchayensis sp. n. +
Fam. Difflugiidae Awerintzev1906
Difflugia acuminata v.inflata Penard 1899 + + +
D. corona Wallich 1864 + + + + +
D. corona v.ecornis Gauthier Lievre et Thomas 1958 ** + +
D. difficilis Thomas 1954 +
D. elegans Penard 1890 + +
D. elegans v. teres Penard 1899 ** + +
D. echinulata Penard 1911 ** +
D. capreolata Penard 1902 ** + +
D. acutissimella Chardez 1985 ** +
D. bicornis Penard 1902 ** + +
D. lobostoma Leidy 1874 +
D. lobostoma var. multilobata Gauthier-Lievre 1958 ** +
D. globularis Wallich 1864 + +
D. gramen Penard 1902 +
D. guttula S.Godeanu 1972 +
D. muriformis G.-L. et T. 1958 ** + + +
D. ogdenii sp. n. + +
D. armatostoma sp. n. +
D. rotiferoformis sp. n. +
D. caucasica sp. n. +
D. vermiformis sp. n. +
D. bifurcata sp. n. +
D. azerbaijanica sp. n. +
Pontigulasia bryophila Penard 1902 + +
P. bigibbosa Penard 1902 + + +
P. compressa (Carter 1864) +
P. breviottis sp. n. +
Cucurbitella mespiliformis v. africana G.-L. et Thomas 1960 ** + +
Schwabia sphaerica sp. n. + +
Pelecyamoeba gen. n.
Pelecyamoeba stenostoma gen. et sp. n. +
Armatodifflugia gen. n.
Armatodifflugia ceratophora sp. n. +
A. cuneata sp.n. +
Fam. Lesquereusiidae Jung 1942
Lesquereusia modesta Rhumbler 1895 + + +
L. gibbosa Thomas and Gauthier-Lievre 1959** +
Fabalesquereusia gen. n.
Fabalesquereusia graniformis sp. n. +
F. compressa sp.n. +
F. linearis sp.n. +
Fam. Euglyphidae Wallich 1864
Euglypha filifera Penard 1890 + + +
Fam.Trinematidae Hoogenraad and Groot 1940
Trinema enchelys (Ehrb. 1838) +
Rivers
Species
1 2 3 4 5
Fam.Cyphoderiidae Deflandre 1953
Cyphoderia ampula (Ehrb. 1840) + + +
C. trochus v. amphoralis Penard 1899 ** + + +
Fam.Shamkiriidae fam. n.
Shamkiriella gen. n.
Shamkiriella convoluta sp. n. +
Sh. reticulata sp. n. +
Sh. phimatophora sp. n. +
Fam. Bipseudostomatidae fam. n.
Bipseudostomatella gen. n.
Bipseudostomatella bifurcata sp. n. +
B. gracilis sp. n. +
B. cornuta sp. n. +
Gomocollariella gen. n.
Gomocollariella ranaformis sp. n. +
Rivers
Notes: 1 - Kurakchay river, 2 - Shamkirchay river, 3 - Korchay river, 4 - Kura river, 5 - Tauzchay river. The species marked with two asterisks are first records for the Caucasus fauna; those marked with one asterisk, for the Azerbaijan fauna.
numerous epipodia. Pseudopodia are broad. There are
2 nuclei and several contractile vacuoles.
This species was found only in the Kura river and was always registered in low densities.
Contrary to description by Ogden and Hedley (1980), the aperture lips are absent. There is a spherical cavity occupying the middle part of the shell (seen in lateral view). Before our investigations, this fact was not mentioned (Van Oye, 1948; Godeanu, 1970; Ogden and Hedley, 1980).
Arcella megastoma Penard 1926 (Fig. 2 c-e; Table 2).
Shell diameter is 84-113 ^m, aperture diameter is 38-55 ^m. This species, contrary to other members of Arcella, has a large, slightly invaginated, circular aperture without a collar, about 1/2 of shell diameter. The shell surface is composed of small hexagonal hyaline plates and has numerous pores. The shell is dark-brown.
Endoplasm is gray and contains from 25 up 50 nuclei.
This species was recorded in freshwater basins of Northern Osetia (Tarnogradsky, 1928, 1948, 1959).
Our specimens of A. megastoma differ from those described from the Dnestr and Dunay river basins (Godeanu, 1970; Vikol, 1992) in being almost two times smaller in shell size.
Though this species is common in different regions, this is the first record for the Azerbaijan fauna. Found in the Shamkirchay river.
Arcella atava Collin 1914 (Fig. 2 f-g; Table 2).
Shell diameter is 63 ! m, shell height, 50 ! m, diameter of aperture, 28-30 ^m. The shell is hemispherical and has a shallow invagination with a circular aperture. It has a small collar. The dark-brown shell
surface is composed of hexagonal hyaline plates and has numerous pores.
Endoplasm occupied about 2/3 of the inner shell cavity. Pseudopodia are broad and flat. There are several contractile vacuoles.
This is a relatively rare species. After the first description, it was described only by Bartos (1954). Our specimens show morphological characteristics corresponding to those described by Bartos (1954) and differ only in shell colour and a more rounded shell.
It is the first record of this species for the Caucasus fauna. Found in the Kurakchay river.
Centropyxis hirsuta Deflandre 1929 (Fig. 3 a-b; Table 2).
The shell is light brown or yellow, circular and usually with 3-4 lateral spines. Shell length is 84-90 ^m; breadth, 81.9-84 ^m. The aperture (33-37.9 !m) is invaginated, subterminal, oval. Oral zone is flattened. The cell occupies a small part of the shell and attaches to its inner surface with several epipodia.
Our specimens of C. hirsuta differ from those described in literature (Deflandre, 1929; Bartos, 1954; Ogden and Hedley, 1980) in having much greater shell length and width.
Before our investigation, the species was recorded only from freshwater basins of Georgia (Tarnogradsky, 1957).
Centropyxis mirabilis Bartos 1940 (Fig. 3 c-d; Table 2).
The shell is elliptic, elongated, with a characteristic swelling on the anterior end. Subterminal invaginated aperture (55 ^m) is more or less triangular. The shell has up to 3 lateral spines. Shell length is 125-130 ^m; breadth, 100-110 ^m. The whole shell surface is covered with inorganic particles of variable size and form. Shell
Fig 2. a - Arcella hemisphaerica, lateral view; b - A. hemisphaerica, lateral view; c - A. megastoma, apertural view; d - A. megastoma, lateral view; e - A. megastoma, surface of the shell; f - A. atava, lateral view; g - A. atava, apertural view.
colour is light brown. Long blade-shaped pseudopodia are rarely observed; if present, usually 1-2 in number.
After the first description, this species has not been recorded prior our investigation. Our specimens of C. mirabilis differ from those in the first description (Bartos, 1954) in smaller shell size.
It is a rare species in the study area, being registered only in the Kurakchay river. It is the first record of this species for the fauna of the Caucasus.
Centropyxis percolabiensis Dekhtyar 1994 (Fig. 3 e; Table 2).
Fig. 3. a - Centropyxis hirsuta, lateral view; b - C. hirsuta, apertural view; c - C. mirabilis, apertural view; d - C. mirabilis, lateral view; e - C. percolabiensis, apertural view; f - C. kurakchayensis sp. n., apertural view; g - C. kurakchayensis sp. n., lateral view.
This species was first described from freshwater basins and wet sphagnum in the Ukraine (Dekhtyar, 1994) and has not been recorded in other regions of the world prior to our research. Below we describe C. perco-labiensis from aquatic macrophytes samples in the Shamkirchay and the Korchay rivers.
Shell length is 130-150 ^m; breadth, 100-110 ^m. The shell is elongated, wider at the base. Circular aperture (50-75 ^m) is surrounded with a well-defined collar. Aboral end has 4-7 spines (15-25 ^m) and a fold, separating this area into two asymmetric parts: a larger part with 3 spines, and a smaller one with 2 spines. The shell has a prominence on the dorsal side.
Endoplasm is colourless, granular. Pseudopodia are lobose, usually 1-3. Nucleus about 20 ^m in diameter. There are 2-5 contractile vacuoles.
It is the first record of this species for the fauna of the Caucasus. Our specimens has smaller size than those in the first description by Dekhtyar (1994).
Centropyxis kurakchayensis sp. n. (Fig. 3 f-g; Plate 1 a; Table 2).
Holotype. Slide K-ch N2.
T ype location. On water plants in the Kurakchay river.
Description. The shell is oval, flattened. Length of shell, 190-210 ^m, breadth, 100-120 ^m, height, 4252 ^m. In the middle of the ventral side there is a large invaginated oval aperture, 55-60 + 70-80 ц m. The aperture is bordered by a thin lip well visible from the ventral side. We have observed specimens with spines (3-7), as well as without spines. The whole shell surface is covered with sand grains of different shape. There are 2-3 lobose pseudopodia. Endoplasm light yellow, without inclusions. One nucleus. There are 3-7 contractile vacuoles.
Discussion. There is only one species, C. compressa (Van Oye, 1948), morphologically close to our C. kura-kchayensis. The latter differs from all the other species of the genus in having a strongly flattened shell shape and a central large aperture. From the C. compressa Van Oye 1948 our species differs in having a larger aperture and a different shell shape.
Etymology. The species name refers to the river the species was found in.
Difflugia acuminata v. inflata Penard 1899 (Fig. 4 a; Table 2).
Shell length is 180-210 !m; breadth, 125-140 !m. The shell is cylindrical, with a small curved tail on the aboral end. Apical aperture is 60-70 !m in diameter. Shell surface is coated with sand particles of various size and shape, smalleron in the anterior region.
Endoplasm colourless, light green with symbiotic zoochlorellae. Pseudopodia are thin and long, usually
2-3 in number.
Our specimens differ from those described from Moldova freshwater basins (Vikol, 1992) in having smaller shell size and a different shell shape.
This variety was found in small shallow bays of Touzchay, Kurakchay and Shamkirchay rivers.
Difflugia corona Wallich 1864 (Fig. 4 b-d; Table 2).
Shell length of our specimens of D. corona is 130145 ^m, the breadth of equatorial part is 125-130 ^m. Apical aperture (40-45 ^m) is circular and has a small indented collar with 11-12 lobes. There are up to 6 (usually 3) spines on the aboral part of the shell. The shell is covered with sand particles of variable shape and size. The cytoplasm occupies about 2/3 of the inner shell cavity and contains a single nucleus and several contractile vacuoles. There are numerous thin lobopo-dia.
We found D. corona, as well as D. corona v. ecornis Gauthier-Lievre et Thomas (Fig. 3 d). This variety differs from the typical form in the absence of spines, smaller size of shell (110-115 ^m) and aperture (25-30 ^m).
Our specimens differ from the first description of D. corona v. ecornis by Gauthier-Lievre and Thomas
(1958) found in the freshwater of Gabon (Africa) in having a smaller shell and aperture size. This is the first record of this variety for the fauna of Caucasus.
Difflugia difficilis Thomas 1955 (Fig. 5 a; Table 2).
The shell is colourless or light brown, oval, with a small outgrowth on the aboral part. Shell length is 120125 ^m; breadth, 85-90 ^m. Apical aperture is 30-35 !m in diameter and is surrounded by a small collar. Shell surface is covered with siliceous elements of various size. Endoplasm light brown, granular. Thin pseudopodia are usually 2-5 in number. Nucleus with a small central nucleolus. There are 6-9 contractile vacuoles.
Our specimens differ from D. difficilis described by Gauthier-Lievre and Thomas (1958), Chardez (1987) and Green (1996) in having a larger shell and aperture size and a different shell shape.
We found this species in the sediments in the Kurakchay river.
Difflugia elegans Penard 1830 (Fig. 5 b-c; Table 2).
This species is common in the study area. Shell length is 100-110 ^m, breadth, 50-60 ^m. The shell has a well-defined tail-like protuberance on the posterior part, 20-25 ^m in length. There is a small neck surrounding the apical aperture (35-40 ^m). The shell surface is covered with sand particles and diatom frustules.
Endoplasm colourless. There are 2-3 short lobopodia. Several contractile vacuoles.
In the same samples D. elegans v. teres Penard, 1899 was observed (Fig. 4 c), differing from the typical form in being wider (85 ^m) in the equatorial shell part.
Fig 4. a - Difflugia acuminata v. inflata, lateral view; b - D. corona, lateral view; c - D. corona, apertural view; d - D. corona v. ecornis, lateral view.
Our specimens of D.elegans are similar to those described from freshwater basins of Moravia (Bartos,
1954).
Difflugia echinulata Penard 1911 (Fig. 5 d; Table 2).
The shell is drop-shaped, its length is 140-170 ^m,
breadth in equatorial part, 80-100 ^m. Apical aperture (30-40 ^m) is circular and usually surrounded by small particles. Posterior part of the shell bears 8-12 spines of different length. They are tubular extensions of the organic shell cover. The whole shell surface is covered by dark sand particles.
Fig. 5. a - Difflugia difficilis, lateral view; b - D. elegans, lateral view; c - D. elegans v. teres, lateral view; d - D. echinulata, lateral view; e - D. capreolata, lateral view.
Our specimens show morphological characteristics D. echinulata described by Velho and Lansac-Toha
which correspond to the species described from Africa (1996) from the Parana river (Brasil) in shell shape and
(Gauthier-Lievre and Thomas, 1958) and differ from more numerous spines.
This species was found in peryphiton of shallow bays of the Shamkirchay river. It is the first record for the fauna of the Caucasus.
Difflugia capreolata Penard 1902 (Fig. 5 e; Table 2).
The shell is pear-shaped. Shell length is 190-200 !m; breadth, 125-135 ^m. Apical aperture (63-68 ^m) invaginated. There is a neck. The dark-brown shell is covered with variously shaped plates.
The cytoplasm is colourless, granular. Two or five short lobopodia. There are 1-3 contractile vacuoles.
Our specimens of D. capreolata differ from those described in the literature (Gauthier-Lievre and Thomas, 1958; Vikol, 1992) in having smaller shell size.
This species was found in the small lagoons of the Shamkirchay river. It is the first record for the fauna of the Caucasus.
Diflugia acutisimella Chardez 1985 (Fig. 6 a; Table 2)
The shell is elongated, cigar-shaped, with a narrowed aboral end. Shell length is 110-135 ^m; breadth, 60-75 !m. Subapical aperture (25-35 ^m) is surrounded with a collar, consisting of small mainly globular particles. The shell wall is made of transparent siliceous plates with a mesh of organic cement.
The cytoplasm is colourless, granular. Pseudopodia long, usually 2 or 3. There are several contractile vacuoles.
This species was described for the first time by Chardez (1985) and, as far as we know, has not been registered in the other regions before our investigations. Our specimens differ from those of Chardez (1985) in having smaller size and constant.
This species was found on the sediment of the Kurakchay river. It is the first record for the fauna of the Caucasus.
Difflugia bicornis Penard 1902 (Fig. 6 b; Table 2).
The shell is almost cylindrical, with two spines on the aboral end. Shell length (without spines) is 125-140 !m; breadth, 90-100 ^m; length of spines, 15-23 ^m. Apical aperture is circular, 65-80 ^m in diameter. The shell is covered with sand particles of variable size and diatom frustules. The cytoplasm is colourless and occupies a small part of the inner shell cavity. There are 1-3 long lobopodia and 6-9 contractile vacuoles.
This species was noted in the freshwaters ofNorthern-West Hungary by Torok (1997).
It is a rare species, occurring only in some samples from the Shamkirchay and Tauzchay rivers. This is its first record for the fauna of the Caucasus.
Difflugia lobostoma var. multilobata Gauthier-Lievre 1958 (Fig. 6 c; Table 2).
The shell has roughly globular shape and is 90-120 !m in diameter. Apical aperture (20-27 ^m) with 5 up to 7 (usually 6) lobes. The aperture surrounded by a thin collar made up of very small particles. Shell wall with sand particles of various size and diatom frustules.
The cytoplasm is light yellow. There are 2 or 3 short lobose pseudopodia.
Our specimens show morphological characteristics which correspond to the species described earlier in the literature (Gauthier-Lievre and Thomas, 1958; Vucetich, 1975; Velho and Lansac-Toha, 1996).
It is the first record of this species for the fauna of the Caucasus.
Difflugia muriformis Gauthier-Lievre et Thomas
1958 (Fig. 6 d; Table 2).
The shell is approximately ellipsoid, colourless, with small regular rounded structures, distributed in the 12-16 rows, from aperture to aboral part, along the whole surface. Each row consists of 11-12 structures. Shell length is 100-115 ^m, breadth in the equatorial part 85-95 ^m. Aperture apical, 25 ^m in diameter, with 12-16 rounded, well defined lobes. There is a thin collar (5-10 ^m).
The cytoplasm is colourless, without inclusions. There are 1 or 2 long lobose pseudopodia.
Our specimens differ from those described by Gauthier-Lievre and Thomas (1958), Vucetich (1973) and Velho and Lansac-Toha (1996) in having an ellipsoid shell shape and a smaller diameter of aperture.
This species was found in sediment and aquatic macrophytes samples of the Shamkirchay, Korchay and Kurakchay rivers, being a common taxon in the study area. It is the first record for the Caucasus fauna.
Difflugia ogdenii sp. n. (Fig. 6 e-f; Plate 1 b-c; Table 2).
Holotype. Slide K-ch N 8.
Type location. In the sediments of the Kurakchay and Korchay rivers.
Description. The shell is bag-shaped, with a wide, flat and oblique anterior part. Length of shell is 100140 ^m; breadth, 90-100 ^m. A wide invaginated apical aperture has a diameter of 50-60 ^m and is surrounded by a broad (20-30 ^m) collar. Shell wall formed by sand particles of variable size; smaller particles in the aperture region. Lobose pseudopodia usually no more than two.
Discussion. This species is similar to an unidentified Difflugia sp. shown in a photograph in a publication by
Fig. 6. a - Difflugia acutisimella, lateral view; b - D. bicornis, lateral view; c - D. lobostoma v. multilobata, lateral view; d - D. muriformis, lateral view; e - D. ogdenii sp. n., lateral view; f - D. ogdenii sp. n., apertural view.
Table 2. Biometric characteristics of the species investigated.
Arcella hemisphaerica
Character X M max min S V n
Diameter of shell 47 47 50 40 3.23 1.02 10
Diameter of aperture 11 11 12 10 0.94 1.30 10
Depth 23 24 25 20 2.0 0.62 10
Apertural invagination 11 11 15 8 3.13 0.99 10
Arcella megastoma
Character X M max min S V n
Diameter of shell 101 101 113 84 9.64 2.49 15
Diameter of aperture 48 50 55 38 5.9 1.52 15
Depth 21 21 25 18 2.06 0.53 15
Apertural invagination 12 12 17 11 1.91 0.49 15
Arcella atava
Character X M max min S V n
Diameter of shell 63 63 64 61 0.83 0.26 10
Diameter of aperture 29 29 30.4 28 0.71 0.22 10
Depth 50 50 50.7 48 0.78 0.25 10
Apertural invagination 28 28 28.8 27.6 0.37 0.12 10
Centropyxis hirsuta
Character X M max min S V n
Length of shell 82 81 84 89.6 1.95 0.62 10
Breadth of shell 83 83 84 81.9 0.68 0.22 10
Depth of shell 48 49 51 44 2.82 0.89 10
Length of aperture 35 36 37.8 33 1.51 0.48 10
Breadth of aperture 40 39 42 37 1.75 0.55 10
Length of spines 18 16 25.2 12.6 4.65 1.47 10
Centropyxis mirabilis
Character X M max min S V n
Length of shell 127 127 130 125 2.37 0.75 10
Breadth of shell 106 106 110 100 4.28 1.35 10
Height of peak 59 60 60 57 1.45 0.46 10
Breadth of aperture 53 53 55 50 2.18 0.69 10
Height of aperture 61 61 63 60 1.34 0.42 10
Centropyxis percolabiensis
Character X M max min S V n
Length of shell 143 146 150 130 7.0 2.20 10
Breadth of shell 107 108 110 100 3.53 1.12 10
Diameter of aperture 65 67 75 50 8.76 2.77 10
Length of spine 22 24 25 15 3.68 1.17 10
Centropyxis kurakchayensis sp. n.
Character X M max min S V n
Length of shell 200 196 210 190 8.72 3.90 5
Breadth of shell 113 116 120 100 8.07 3.61 5
Height of shell 50 52 52 42 5.0 2.5 5
Length of aperture 76 76 80 70 3.94 1.76 5
Breadth of aperture 58 58 60 55 2.28 1.02 5
Difflugia acuminata v. inflata
Character X M max min S V n
Length of shell 198 198 210 180 10 2.83 15
Breadth of aperture 135 135 140 125 5.20 1.34 15
Diameter of aperture 66 65 70 60 3.60 0.93 15
Difflugia corona
Character X M max min S V n
Length of shell 141 142 145 130 4.42 1.40 10
Breadth of shell 128 130 130 125 2.33 0.74 10
Diameter of aperture 43 45 45 40 2.33 0.74 10
Difflugia difficilis
Character X M max min S V n
Length of shell 123 123 125 120 2.17 0.69 10
Breadth of shell 88 88 90 85 2.0 0.63 10
Diameter of aperture 33 34 35 30 1.94 0.61 10
Difflugia elegans
Character X M max min S V n
Length of shell 107 109 110 100 3.27 1.04 10
Breadth of shell 56 56 60 50 4.40 1.39 10
Diameter of aperture 38 39 40 35 2.25 0.71 10
Length of spine 23 24 25 20 2.28 0.72 10
Difflugia echinulata
Character X M max min S V n
Length of shell 160 160 170 140 11.6 4.36 7
Breadth of shell 89 90 100 80 9.32 3.52 7
Diameter of aperture 37 40 40 30 4.88 1.84 7
Length of spine 16 15 20 10 4.50 1.70 7
Difflugia capreolata
Character X M max min S V n
Length of shell 19е 19е 200 190 3.а9 1.23 10
Breadth of shell 130 129 135 125 3.95 1.25 10
Diameter of aperture ее ее еа е3 1.95 0.е2 10
Difflugia acutissimella
Character X M max min S V n
Length of shell 12е 130 135 110 9.е2 4.30 5
Breadth of shell еа 70 75 е0 е.52 2.92 5
Diameter of aperture 30 30 35 25 5.77 2.89 5
Difflugia bicornis
Character X M max min S V n
Length of shell 133 135 140 125 5.70 1.80 10
Breadth of aperture 94 94 100 90 3.33 1.05 10
Length of spines 19 20 23 15 3.33 1.05 10
Difflugia lobostoma var. multilobata
Character X M max min S V n
Diameter of shell 10е 108 125 90 13.2 4.17 10
Diameter of aperture 22 22 27 20 2.40 0.7е 10
Difflugia muriformis
Character X M max min S V n
Length of shell 107 105 115 110 5.30 1.37 15
Breadth of shell а9 90 95 а5 3.72 0.9е 15
Diameter of aperture 23 23 25 20 2.2е 0.58 15
Difflugia ogdenii sp. n.
Character X M max min S V n
Length of shell 125 129 140 100 15.2 4.80 10
Breadth of shell 100 100 110 90 а.03 2.54 10
Diameter of aperture 55 55 60 50 4.01 1.27 10
Breadth of collar 24 24 30 20 3.57 1.13 10
Difflugia armatostoma sp. n.
Character X M max min S V n
Length of shell 121 125 130 100 12.5 5.57 5
Breadth of shell (wide part) а7 а5 95 а0 5.94 2.ее 5
Breadth of shell (narrow part) 55 55 е0 45 5.93 2.е5 5
Diameter of aperture 34 35 37 30 2.77 1.24 5
Difflugia rotiferoformis sp. n.
Character X M max min S V n
Length of shell 141 142 150 130 7.29 2.30 10
Breadth of shell а2 а2 90 75 е.05 1.91 10
Diameter of aperture 33 34 35 30 2.10 0.ее 10
Length of “tail” 17 17 1а 1е 0.е3 0.20 10
Difflugia caucasica sp. n.
Character X M max min S V n
Diameter of shell 103 104 110 95 5.а7 1.86 10
Diameter of aperture 33 33 35 30 2.27 0.72 10
Difflugia vermiformis sp. n.
Character X M max min S V n
Length of shell 131 132 135 125 3.е0 1.14 10
Breadth of shell а1 а1 а5 75 3.7а 1.20 10
Thickness of shell 1е 15 17 12 1.65 0.52 10
Diameter of aperture 31 32 35 25 3.е5 1.16 10
Length of "tail" 1е 1е 17 12 1.71 0.54 10
Thickness of collar 11 12 13 а 2.00 0.е3 10
Difflugia bifurcata sp. n.
Character X M max min S V n
Length of shell 121 122 125 115 4.13 1.31 10
Breadth of shell 157 158 160 150 3.е0 1.14 10
Diameter of aperture 31 30 33 29 1.83 0.58 10
Difflugia azerbaijanica sp. n.
Character X M max min S V n
Length of shell 172 173 180 160 8.17 3.33 е
Breadth of shell 132 133 140 120 8.17 3.33 е
Length of spines 23 24 25 20 2.45 1.0 е
Diameter of aperture ае а7 90 а0 4.е4 1.89 е
Pontigulasia bryophila
Character X M max min S V n
Length of shell 9а 9а 110 а0 9.е9 2.5 15
Breadth of aperture е4 е7 70 55 6.11 1.58 15
Length of neck 30 30 35 23 4.1 1.06 15
Pontigulasia compressa
Character X M max min s V n
Length of shell 140 143 145 130 6.85 2.17 10
Breadth of shell 96 95 100 90 4.38 1.38 10
Thickness of shell 68 70 70 65 2.37 0.75 10
Diameter of aperture 38 40 40 35 2.17 0.69 10
Pontigulasia breviottis sp. n.
Character X M max min s V n
Length of shell 86 87 90 80 4.05 1.28 10
Breadth of shell 63 63 65 60 2.17 0.69 10
Length of neck 27 28 29 25 1.83 0.58 10
Diameter of neck 44 45 47 40 2.83 0.89 10
Cucurbitella mespiliformis v.africana
Character X M max min s V n
Diameter of shell 76 77 80 70 4.16 1.32 10
Thickness of collar 13 14 15 10 2.28 0.72 10
Height of collar 23 25 25 20 2.58 0.82 10
Diameter of aperture 23 24 25 20 2.41 0.76 10
Pelecyamoeba stenostoma sp. n.
Character X M max min s V n
Length of shell 87 90 90 80 4.47 2.0 5
Length of neck 48 50 50 45 2.74 1.23 5
Diameter of aperture 21 20 22 20 0.89 0.40 5
Armatodifflugia ceratophora sp. n.
Character X M max min s V n
Length of shell 78 80 80 75 2.58 1.05 6
Breadth of shell 110 110 115 105 4.47 1.83 6
Length of ledge 26 26 27 22 1.98 0.81 6
Diameter of aperture 24 25 25 20 2.04 0.83 6
Armatodifflugia cuneata sp. n.
Character X M max min s V n
Length of shell 106 108 110 95 5.85 2.39 6
Breadth of shell 112 112 115 105 4.08 1.67 6
Length of ledge 19 20 20 15 2.04 0.83 6
Diameter of aperture 15 15 16 12 1.47 0.60 6
Height of collar 7 6 10 6 2.07 0.84 6
Schwabia sphaerica sp. n.
Character X M max min s V n
Length of shell 101 100 110 90 7.54 2.39 10
Breadth of shell 100 100 100 98 0.70 0.22 10
Diameter of aperture 20 22 22 18 1.90 0.60 10
Lesquereusia gibbosa
Character X M max min s V n
Length of shell 106 105 115 100 4.83 1.53 10
Thickness of shell 86 86 90 80 2.89 0.91 10
Diameter of aperture 22 23 25.2 20 1.5 0.48 10
Fabalesquereusia graniformis sp. n.
Character X M max min s V n
Length of shell 110 110 115 100 6.12 2.74 5
Breadth of shell 83 85 85 80 2.74 1.23 5
Diameter of aperture 23 25 25 20 2.74 1.23 5
Fabalesquereusia compressa sp. n.
Character X M max min s V n
Length of shell 120 120 125 110 6.12 2.74 5
Breadth of shell 105 105 110 100 5.0 2.24 5
Thickness of shell 55 55 60 50 5.0 2.24 5
Diameter of aperture 37 37 40 35 2.45 1.23 5
Fabalesquereusia linearis sp. n.
Character X M max min s V n
Length of shell 79 80 80 75 2.19 0.98 5
Breadth of shell 64 65 65 60 2.19 0.98 5
Diameter of aperture 22 23 23 20 1.64 0.74 5
Cyphoderia trochus v. amphoralis
Character X M max min s V n
Length of shell 75 76 75.8 74.9 0.24 0.08 10
Thickness of shell (wide part) 38 38 37.9 37.1 0.28 0.09 10
Thickness of shell (narrow part) 17 17 16.9 16.3 0.20 0.06 10
Diameter of aperture 13 13 12.9 12.3 0.19 0.06 10
Shamkiriella convoluta sp. n.
Character X M max min s V n
Length of shell 108 110 115 100 7.58 3.39 5
Breadth of shell 86 88 90 80 4.22 1.89 5
Diameter of aperture 39 40 45 35 4.18 1.87 5
Shamkiriella reticulata sp. n.
Character X M max min S V n
Length of shell 135 135 140 125 6.12 2.74 5
Breadth of shell 112 115 115 105 4.47 2.0 5
Diameter of aperture 45 45 50 40 5.0 2.24 5
shamkiriella phimatophora sp. n.
Character X M max min S V n
Length of shell 118 120 125 100 10.4 4.64 5
Breadth of shell 84 85 90 80 4.18 1.87 5
Diameter of aperture 27 25 30 25 2.74 1.23 5
Bipseudostomatella bifurcata sp. n.
Character X M max min S V n
Diameter of shell 88 90 95 80 7.58 3.39 5
Height of shell 68 70 70 65 2.74 1.23 5
Diameter of aperture 27 27 30 25 2.50 1.12 5
Bipseudostomatella gracilis sp. n.
Character X M max min S V n
Diameter of shell 130 130 135 120 6.12 2.74 5
Diameter of aperture 27 27 30 25 2.51 1.12 5
Bipseudostomatella cornuta sp. n.
Character X M max min S V n
Height of shell 84 85 90 75 6.52 2.92 5
Diameter of shell 87 87 90 85 2.51 1.12 5
Diameter of aperture 43 43 45 40 2.51 1.12 5
Height of neck 23 23 25 20 2.51 1.12 5
Gomocollariella ranaformis sp. n.
Character X M max min S V n
Diameter of shell 75 75 80 70 5.0 2.24 5
Length of neck 65 65 70 60 5.0 2.24 5
Height of neck 43 45 45 40 2.74 1.23 5
Diameter of aperture 21 20 22 20 1.10 0.49 5
Notes: x- arithmetic mean, M - median, max - maximum, min - minimum, S - standard deviation, V- coefficient of variance, n - number of specimens.
Ogden (1984). It may be the same species, although our specimens have a larger shell size and a smaller size of covering plates.
This species clearly differs from all other representatives of the Difflugia genus in shell shape.
Etymology. The species is named in honour of Dr. C. Ogden, an English protozoologist.
Difflugia armatostoma sp. n. (Fig. 7 a; Plate 1 d; Table 2).
Holotype. Slide K-ch N 5.
Type location.This species was found in the sediments of the Kurakchay river.
Description. Shell pear-shaped. Shell length 100130 ^m, breadth in the widest part, 80-95 ^m. Apical aperture (30-37 ^m) is surrounded by well-developed teeth-like indentations. The shell surface is covered with quartz particles, smaller in the aperture region.
The cytoplasm is dark-brown, granular. There are 3-5 short lobose pseudopodia. Nucleus with numerous nucleoli.
Discussion. Our species differs from D. manicata Penard, 1902 in the shell shape and better developed apertural "teeth". It differs from all the other species of the genus by the presence of a well-developed teethlike indentations around the aperture and in shell shape.
Etymology. Species name refers to well-developed indentations around the aperture.
Difflugia rotiferoformis sp. n. (Fig. 7 b; Plate 1, e; Table 2).
Holotype. Slide K-ch N 13, 14.
Type location. On the aquatic plants in the small bays of Kurakchay river.
Description. Shell light yellow, elongated, resembling a rotifer in general outlook, with the aboral part narrowed into a small tail (15-20 ^m). Length ofshell 130150 ^m, breadth in the equatorial part, 75-90 ^m. Apical aperture (30-35 ^m) is oval and bordered by small pieces of quartz.
Endoplasm colourless, without inclusions. There are 2 or 3 short lobose pseudopodia. There are several contractile vacuoles.
Discussion. There are several representatives of the Difflugia genus with a "tail". Our species differs from:
1. D. oblonga v acuminata Ehrenberg 1838 in having smaller shell size and a blunt tail end.
2. D. mamillaris Penard 1902 in shell shape and larger shell size.
3. D. difficilis Thomas 1955 in larger shell size and absence of apertural collar.
Etymology. Species name refers to shell shape resembling a rotifer.
Difflugia caucasica sp. n. (Fig. 7 c; Plate 1 f; Table 2).
Holotype. Slide K-ch N 1.
Type location. In the sediments in the shallow lagoons of the Kurakchay river.
Description. The shell is spherical, 95-110 ^m in diameter. Aperture sub-apical, 30-35 ^m in diameter, without a collar. Shell wall formed by transparent polygonal plates.
The cytoplasm is light green, granular. There are 1 or 2 long lobose pseudopodia. There are 5 to 7 contractile vacuoles.
Discussion. This species resembles D. globularis (Wallich 1864), D. globulosa Dujardin 1937 and D. gramen Penard 1902, differing from the latter species in having a simple aperture, and from the former two, in shell size and structure and the subapical aperture.
Etymology. The species name refers to the region where the species was discovered.
Difflugia vermiformis sp. n. (Fig. 7 d-f; Table 2).
Holotype. Slide K-ch N 10.
Type location. On aquatic plants in the shallow bays of the Kurakchay rivers.
Description. The shell is elongated, flattened and curved, with a blunt spine on the aboral end. Shell length is 125-135 ^m; breadth, 75-85 ^m. Apical aperture (2535 ^m) is ovoid and surrounded with a well-defined thin collar. Shell wall formed by transparent plates of variable size.
The cytoplasm is colourless, without inclusions. There are 1-2 short lobopodia.
Discussion. D. vermiformis clearly differs from all known representatives of Difflugia genus in its shell shape.
Etymology. Species name refers to worm-like shell shape.
Difflugia bifurcata sp. n. (Fig. 7 g; Plate 2 a; Table 2).
Holotype. Slides K-ch N 12.
Type location. In the sediments in the small shallow lagoons of the Kurakchay river.
Description. The shell is pear-shaped. Aboral zone is divided into two parts. Length of shell 115-125 ^m, breadth in equatorial part, 150-160 ^m. Apical aperture is circular, its diameter is 29-33 ^m. The whole shell surface is covered with siliceous plates.
The cytoplasm is dark-brown. There are 3 or 5 long lobose pseudopodia and 3-4 contractile vacuoles.
Discussion. This species clearly differs from all representatives of the Difflugia genus in shell shape.
Etymology. Species name refers to bifurcate shell shape.
Difflugia azerbaijanica sp. n. (Fig. 8 a-b; Plate 2 b-c; Table 2).
Holotype. Slide K-ch N 1.
Type location. In a small marginal lake of the Kurakchay river basin on aquatic plants.
Description. The shell asymmetrical, with a well-defined bifurcation from equatorial part into two halves, one of them ending with two spines (20-25 ^m) and the second part, which is wider, with one spine. Shell length with spines is 160-180 ^m; breadth, 120-140 ^m. Apical aperture 80-90 ^m.
The spines are always 3 in number, two of them located closely to each other, and the third one, slightly apart. The shell is covered with sand particles of variable shape and size; smaller particles are near the aperture.
The cytoplasm is transparent. There are 1-3 wide lobopodia and several contractile vacuoles.
Discussion. This species is closely related to D. foissneri Chardez 1987 found in freshwater basins of Belgium (Chardez, 1987). Our species differs from it in smaller shell size, asymmetric location oflarger spines and a well-defined bifurcation of the aboral shell part.
In our opinion, Centropyxis marsupiformis described by Brazilian specialists (Velho and Lansac-Toha, 1996) from the high Parana river could be D. foissneri or D. azerbaijanica sp. n. Unfortunately, the authors did not give the shell shape laterally, so correct identification cannot be done. One of the specimens described from Brazil is clearly not C. marsupiformis.
Etymology. The species name refers to the country where the species was discovered.
Pontigulasia bryophila Penard 1902 (Fig. 8 c; Table 2).
In ecological investigations, this species is comparatively often referred too, but there are no modern morphological descriptions. Below we give a short description of this species found in the small bays on the water plants in the Kurakchay and Shamkirchay rivers.
Shell length is 80-120 ^m; breadth, 55-70 ^m. The shell is elongated, cylindrical with a short curved anterior end. Apical aperture is 25 ^m in diameter. There is an internal perforated diaphragm at the junction
Fig. 7. a - Difflugia armatostoma sp. n., lateral view; b - D. rotiferoformis sp. n., lateral view; c - D. caucasica sp. n., lateral view; d - D. vermiformis sp. n., dorsal view; e - D. vermiformis sp. n., lateral view; f - D. vermiformis sp. n., apertural view; g - D. bifurcata sp. n., lateral view.
Plate 1. a - Centropyxis kurakchayensis sp. n., apertural view; b - Difflugia ogdenii sp. n., apertural view; c - D. ogdenii sp. n., lateral view; d - D. armatostoma sp. n., lateral view; e - D. rotiferoformis sp. n., lateral view; f - D. caucasica sp. n., lateral view.
of the neck. Neck length is 25-30 ^m. Shell wall with sand particles of variable size. Cytoplasm grey.
Our specimens of P. bryophila correspond to the species described by Bartos (1954), differing only in smaller shell size.
Pontigulasia compressa (Carter 1864) (Fig. 8 d; Table 2).
This species was first found in Azerbaijan in a freshwater basin of the Apsheron peninsula (Alekperov and Snegovaya, 2002). Below we give a description of the species found in the Shamkirchay river.
Plate 2. a - Difflugia bifurcata sp. n., lateral view; b - D. azerbaijanica sp. n., general view; c - D. azerbaijanica sp. n., lateral view; d - Pontigulasia breviottis sp. n., general view; e - Pelecyamoeba stenostoma sp. n., dorsal view; f - P. stenostoma sp. n., lateral view; g - P. stenostoma sp. n., apertural view.
The shell is pear-shaped, flattened. Apical aperture (35-40 ^m) is circular or ovoid. The neck (50 ^m) is linked to the rest of the shell in a V-form. There is a perforated diaphragm internally at the junction of the neck. Shell length is 130-145 ^m, breadth of the equatorial part, 90-100 ^m, height, 65-70 ^m. Shell wall with quartz grains.
The cytoplasm is light green, granular. There are 3-5 elongate pseudopodia.
Pontigulasia breviottis sp. n. (Fig. 8 a-b; Plate 2 d; Table 2).
Holotype. Slides K-ch N 10.
Type location. On aquatic plants in the small bays of the Kurakchay river.
Description. Shell ellipsoid, a well-defined neck with two small ear-shaped protuberances. Shell length 80-90 ^m; breadth, 60-65 ^m. The aperture is circular (3 ^m). There is an internal diaphragm at the junction of the shell with the neck. Dark-brown shell surface with sand particles of variable size.
The cytoplasm colourless, granular. There are 1-3 elongate pseudopodia. Nucleus with nucleoli.
Discussion. Our species is morphologically closely related to P. spectabilis Penard 1902 but is twice larger in shell size and the shell shape is different. It differs from all other representatives of the Pontigulasia genus in an uncommon neck shape.
Etymology. Species named refers to two short earlike protuberances of the shell.
Cucurbitella mespiliformis v. africana Gauthier-Lievre et Thomas 1960 (Fig. 9 c; Table 2).
Shell spherical, 70-80 ^m in diameter. A circular aperture (20-25 ^m) is surrounded with a well-defined collar consisting of 4 lobes. The shell surface is composed of sand particles of different size and diatom frustules.
The cytoplasm is dark-brown, granular. There are 1-3 short lobose pseuodopodia.
Our specimens differ from those in the first description (Gauthier-Lievre and Thomas, 1960) and those from freshwater basins of Brasil (Velho and Lansac-Toha, 1996) in having a much smaller shell size and twice as large height of the collar.
This species was found in the shallow lagoons on the sand in the Korchay and Kurakchay rivers. Before our investigations only the typical form was recorded from freshwaters ofthe Northern Caucasus (Tarnograd-sky and Popov, 1932). This is the first record of this variety for the fauna of Caucasus.
Pelecyamoeba gen. n.
Diagnosis. Shell pyriform in frontal view, hatchetshaped in lateral view, with a conical elongated neck.
There is a well-defined expansion in equatorial part of the shell and a longitudinal invagination on the aboral end. Aperture oval or lenticular. Its edge is surrounded with a small collar. Freshwater forms.
Pelecyamoeba gen. n. belongs to the family Difflu-giidae, differing from all its genera in shell morphology.
Type species Pelecyamoeba stenostoma gen. et sp. n.
Etymology. Genus name refers to hatchet-like shell shape.
Pelecyamoeba stenostoma gen. et sp. n. (Fig. 9 d-f; Plate 2 e-g; Table 2).
Holotype. Slide K-ch N 12.
Type location. Periphyton on the underwater stones in the shallow lagoons of the Kurakchay river.
Description. The shell is pyriform, hatchet-shaped in lateral view. Total shell length is 80-90 ^m, the length of neck, 45-50 ^m. Apical aperture (20-22 ^m) lenticular, surrounded with a thin collar. There is a longitudinal invagination on the aboral end. Shell surface covered with transparent particles of variable size.
The cytoplasm is light green, with several contractile vacuoles. There are 1-3 short lobose pseudopodia.
Etymology. Species name refers to a narrow slotshaped aperture.
Armatodifflugia gen. n.
Diagnosis. Shell rounded, with a pronounced lateral conical protuberance on its lower part. Upper and lower parts separated with a well-defined fold similar to that in Pontigulasia. Aperture apical, a collar around it consisting of highly refractive particles. Shell wall usually transparent. Shell surface covered with closely adhering polygonal mineral plates joined with organic cement. Freshwater forms.
The Armatodifflugia genus is included into the Difflugiidae Wallich 1864. Its differs from the remaining genera of the Difflugiidae by a very characteristic shell shape. Type species Armatodifflugia ceratophora.
Etymology. Genus name refers to a feature of shell morphology, the presence of a pronounced protuberance.
Armatodifflugia ceratophora sp. n. (Fig. 10 a-b; Plate
3 b-c; Table 2).
Holotype. Slide K-ch N 3.
Type location. Peryphiton and sediments in the shallow lagoons of the Kurakchay river.
Description. Shell rounded, with a pronounced lateral conical protuberance. Shell length (from aperture to aboral end) is 75-80 ^m. Breadth of shell in the equatorial part (including the conical protuberance) is 105-115 ^m, the length of protuberance is 22-27 ^m. An oval aperture (20-25 ^m) is surrounded with a low collar which consists of irregular highly refractive particles. There is a U-like fold on the border between
Fig. 8. a - Difflugia azerbaijanica sp. n., dorsal view; b - D. azerbaijanica sp. n., lateral view; c -Pontigulasia bryophila, lateral view; d - P. compressa, dorsal view.
the upper and the lower part of shell. Shell wall transparent, with siliceous particles and organic cement.
The cytoplasm is grey, granular. Pseudopodia were not observed.
Etymology. Species name reflects the shape of the protuberance on the shell.
Armatodifflugia cuneata sp. n. (Fig. 10 c-d; Plate 3 d-e; Table 2).
Holotype. Slide K-ch N 4.
Type location. Peryphiton of the shallow lagoons in the Kurakchay river.
Description. Shell length (from aperture to aboral
Fig. 9. a - Pontigulasia breviottis sp. n., dorsal view; b - P. breviottis sp. n., lateral view; c - Cucurbitella mespiliformis v. africana, lateral view; d - Pelecyamoeba stenostoma sp. n., lateral view; e - P. stenostoma sp. n., dorsal view; f - P. stenostoma sp. n., apertural view.
Fig. 10. a - Armatodifflugia ceratophora sp. n., lateral view; b - A. ceratophora sp. n., apertural view; c -A. cuneata sp. n., lateral view; d - A. cuneata sp. n., apertural view.
end) is 95-110 ^m. Shell breadth in the equatorial part (including the conical protuberance) is 105-115 ^m. Length of protuberance is 15-20 ^m. Apical aperture (12-16 ^m) oval, with a well-defined high (6-10 ^m) collar, consisting of small particles. U-like fold on the upper and the lower part is not well-defined, contrary to A. ceratophora. The shell surface is covered with
diatom frustules and siliceous particles joined with organic cement.
There are 5-9 contractile vacuoles. Nucleus located in the middle part of the cell.
Discussion. This species differs from the type species A. ceratophora in shell shape, a better defined collar and a less defined equatorial fold.
Etymology. Species name reflects the wedge-like shape of the shell protuberance.
Schwabia sphaerica sp. n. (Fig. 11 a; Plate 3 a; Table 2).
Holotype. Slides Kh-ch N 18.
Type location. On aquatic plants in the Shamkirchay and Kurakchay rivers.
Description. The shell is ovoid, round in crosssection. Shell length is 90-110 ^m; breadth, 98-100 ^m. Apical aperture (18-22 ^m) with a spherical outline but without a collar. Shell wall chitinous, with small sand plates and diatom frustules. The cytoplasm grey, without inclusion. There are several contractile vacuoles.
Discussion. This species differs from most representatives of the Schwabia genus, except S. robusta Chardez 1964, in larger shell size. S. sphaerica sp. n. differs from other species in shell structure and presence of sand plates in shell wall.
Etymology. Species name refers to spherical shell shape.
Lesquereusia gibbosa Thomas et Gauthier-Lievre
1959 (Fig. 10 b).
Retort-shaped shell with a short curved neck. Shell length is 100-115 ^m, the width 80-90 ^m. A circular aperture (20-25 ^m) is bordered with elongated siliceous particles. The same particles cover the whole shell surface. The cytoplasm is colourless. There are several contractile vacuoles and 1-3 lobose pseudopodia.
Our specimens of L. gibbosa have smaller shell size than those described by Thomas and Gauthier-Lievre
(1959).
This species was found on water plants in the Kurakchay river. This is the first record for the fauna of Caucasus.
Fabalesquereusia gen. n.
Diagnosis. The shell bean-shaped.Terminal aperture rounded or oval, indented. No neck. Shell wall with siliceous particles and diatom frustules.
The genus clearly differs from the closely related Lesquereusia genus in absence ofneck. Freshwater forms.
Type species F. graniformis sp. n.
Etymology. The genus name refers to the bean-shaped shell.
Fabalesquereusia graniformis sp. n. (Fig. 11 c-d; Plate
3 f; Table 2).
Holotype. Slide K-ch N 3.
Type location. Sediments and aquatic plants in the Kurakchay river.
Description. The shell bean-shape, flattened laterally. Shell length 100-115 ^m, breadth in equatorial part 80-85 ^m. Terminal aperture circular, 20-25 ^m in diameter, indented. There is a shell fold below aperture.
No neck. The shell surface is covered with siliceous particles and diatom frustules. Impregnated specimens with black "points" on the different shell zones.
The cytoplasm colourless, granular and located in the wide part of the shell. The cell attaches to the shell with 4-6 epipodia. Pseudopodia lobose, granular. There are 6-8 contractile vacuoles.
Etymology. The species name refers to seed-like shell shape.
Fabalesquereusia compressa sp. n. (Fig. 12 a-b; Plate 4 a; Table 2).
Holotype. Slide K-ch N 14.
Type location. Peryphiton of the shallow lagoons of the Kurakchay river.
Description. Shell bean-shaped and flattened. Shell length is 110-125 ^m, breadth in the equatorial part, 100-110 ^m. Shell height 50-60 ^m. The terminal aperture (35-40 ^m) indented. There is a shell fold below the aperture. No neck. Shell surface covered with siliceous particles and some diatom frustules.
Cytoplasm dark-brown, granular, located in the wide part of the shell. The cell attaches to the shell with
3-5 epipodia. There are 1-3 lobose pseudopodia and 46 contractile vacuoles.
Discussion. F. compressa sp. n. clearly differs from the type species F. graniformis sp. n. described above in having a larger aperture and a stronger lateral shell compression.
Etymology. Species name refers to the flattened shape of shell.
Fabalesquereusia linearis sp. n. (Fig. 12 c-d; Plate
4 b; Table 2).
Holotype. Slide K-ch N 10.
Type location. On sediments and grey sand in the shallow lagoons of the Kurakchay river.
Description. The shell bean-shaped, flattened. Shell length, 75-80 ^m; breadth, 60-65 ^m. Terminal aperture (20-23 ^m) oval, indented. There is no fold characteristic of F. graniformis sp. n. and F. compressa sp. n. Shell surface mainly covered with short straight diatom frustules.
The cytoplasm is usually grey, with 3-5 contractile vacuoles. Pseudopodia were not observed.
Discussion. This species differs from F. graniformis sp. n. and F. compressa sp. n. in absence of the frontal fold, shell shape and a different structure of shell wall, although all three species were found in identical environments.
Etymology. Species name refers to a characteristically "lined" shell surface.
Cyphoderia trochus v. amphoralis Penard 1899 (Fig. 13 a; Table 2).
Fig. 11. a - Schwabia sphaerica sp. n., lateral view; b - Lesquereusia gibbosa, lateral view; c -Fabalesquereusia granifromis sp. n., lateral view; d - F. graniformis sp. n., apertural view.
The shell is retort-shaped with curved apical end. The aperture circular, 12-13 ^m in diameter. Shell wall composed of circular siliceous plates arranged in diagonal rows. There are numerous pores which stain black after silver impregnation.
The cytoplasm is light-yellow, located in the middle of the shell. There are 5-9 !m long, thin branching pseudopodia. One nucleus.
Our specimens have smaller length and aperture diameter than those described by Chardez (1991). In the fauna of the Caucasus the species was found for the first time.
Shamkiriidae fam. n.
Diagnosis. The shell with characteristic longitudinal folds, sometimes flattened. Aperture oval to triangular, with a well-defined collar. The shell surface is covered with transparent plates of variable size. Freshwater forms.
According to morphology of shell and aperture, the new family may be related to the Difflugiidae. It clearly differs from all known families in completely unusual morphology of the shell.
Type genus Shamkiriella gen. n.
Fig. 12. a - Fabalesquereusia compressa sp. n., lateral view; b - F. compressa, sp. n., apertural view; c -F. linearis sp. n., lateral view; d - F. linearis sp. n., apertural view.
Shamkiriella gen. n.
Diagnosis. Same as family.
Type species. Shamkiriella convoluta gen. et sp. n. Etymology. The genus is named after the region found.
Shamkiriella convoluta sp. n. (Fig. 13 b-c; Plate 4 c-d; Table 2).
Holotype. Slides Sh-ch N 1.
Type location. Small bays with aquatic plants of the Shamkirchay river.
Description. Shell elongated (100-115 ^m), flattened laterally, with a short (15 ^m) tail-like protuberance on the aboral end and a triangular apical aperture (35-45 !m). There is a well-defined collar. In frontal view, two folds along longitudinal axis can be seen (Fig. 12 b). One of them begins at the aperture margin and ends near the tail-like protuberance on the aboral end. The second fold is smaller, begins approximately in the equatorial part and ends on the aboral tail-like protuberance. In lateral view, two folds can also be seen, beginning from
the aperture margin and ending at a distance of about 1/3 of shell length from the aboral end.
The shell surface is covered with transparent particles ofvariable size. The cytoplasm with numerous contractile vacuoles.
Etymology. Species name refers to "folds" on shell surface.
Shamkiriella reticulata sp. n. (Fig. 13 d-e; Plate 5 a-b; Table 2).
Holotype. Slide K-ch N 1.
Type location. Sediments of shallow bays in the Kurakchay river.
Description. Shell elongated, flattened laterally. Shell length, 125-140 ^m, breadth in the equatorial part, 105-115 ^m. On the aboral end there is a short tail-like protuberance. Apical aperture (40-50 ^m) oval, with a well-defined collar, consisting, contrary to the other two species of Shamkiriella, of separate transparent refractive particles. In frontal view, one fold is visible, beginning in the anterior quarter of the shell and ending near the aboral part. In lateral view, the shell with two well-defined long folds, beginning below the aperture, and with three short folds. Shell surface covered with transparent small particles.
The cytoplasm colourless, with 5-7 contractile vacuoles.
Etymology. Species name refers to the reticulate shell surface outlook.
Shamkiriella phimatophora sp. n. (Fig. 14 a-b; Plate
4 e; Table 2).
Holotype. Slide K-ch N 14.
Type location. On aquatic plants and sediments in the Kurakchay river.
Description. The shell elongated, with length 100125 ^m and breadth 80-90 ^m. Oral end flattened, with triangular aperture (25-30 ^m) surrounded by a well defined collar. In frontal view (Fig. 13 a), two folds can be seen, one beginning at about 1/3 of shell length from the aperture and ending near the aboral end with a taillike caudal protuberance. The second fold begins below the aperture, runs along the shell and units with the other fold near aboral end. In lateral view (Fig. 13 b), two folds can be seen, beginning from aperture margin, running along longitudinal axis and uniting some distance from the aboral end. One more fold can be seen on the right, in the posterior third of the shell. It ends in the tail-like caudal protuberance. Shell wall with transparent, sometimes light green particles.
Cytoplasm light, with numerous contractile vacuoles.
Discussion. This species clearly differs from S. convoluta sp. n. in shell morphology, especially arrangement of the frontal folds.
Etymology. Species name refers to the presence of folds on the shell surface.
Fam. Bipseudostomatidae fam. n.
Diagnosis. Shell with two apertures, connected to the shell with well-defined neck(s). The neck may be solid with two apertures (Gomocollariella gen. n.) or each of apertures has a separate neck. No clear collar around apertures. Shell wall composed of sand particles joined with organic cement. Freshwater forms.
Bipseudostomatidae fam. n. includes two genera: type genus Bipseudostomatella gen. n. and Gomocolla-riella gen. n.
Bipseudostomatella gen. n.
Diagnosis. Shell with two separate apertures located at opposite parts of the shell. Each aperture connected to the rest of the shell with a neck.
Type species B. bifurcata gen. et sp. n.
Remarks. In a paper by Cash and Hopkinson (1909) a form with two apertures is illustrated, described as a Difflugia petricola. In our opinion, it is a representative of the new Bipseudostomatella genus.
Etymology. Genus named refers to a feature of shell morphology: the presence of two apertures.
Bipseudostomatella bifurcata gen. et sp. n. (Fig. 14 c-d; Plate 5 c; Table 2).
Holotype. Slide K-ch N 12.
Type location. Sand of shallow lagoons in the Kurakchay river.
Description. Shell rounded and flattened, with a diameter of 80-95 ^m and a height of 65-70 ^m in equatorial part. There are two separate apertures (2530 ^m in diameter) on the opposite sides. Each aperture has a neck (30 ^m) connecting it with the rest of the shell. All shell surface covered with siliceous particles that are smaller on the necks.
Cytoplasm colourless. Pseudopodia were not observed. There are 5-8 contractile vacuoles.
Etymology. Species name refers to bifurcate shell shape.
Bipseudostomatellagracilis sp. n. (Fig. 15 a, b; Plate
5 d; Table 2).
Holotype. Slide K-ch N 5.
Type location. Aquatic plants in shallow lagoons of the Kurakchay river.
Description. Shell more or less spherical, 120-135 !m in diameter, with two separate apertures (25-30 ^m in diameter). Each aperture is connected with the rest of the shell by a neck (50 ^m). Aperture margins with small particles. The fold between the neck and the shell proper is wavy, contrary to B. bifurcata, which has a flat fold. Shell wall with siliceous plates and diatom frustules.
Plate 3. a - Shwabia sphaerica sp. n., lateral view; b - Armatodifflugia ceratophora sp. n., lateral view; c - A. ceratophora sp. n., apertural view; d - A. cuneata sp. n., lateral view; e - A. cuneata sp. n., apertural view; f - Fabalesquereusia graniformis sp. n., lateral view.
The cytoplasm colourless, without inclusion. There Discussion. B. gracilis sp. n. differs from the type
are 1-3 contractile vacuoles. Pseudopodia were not species B. bifurcata sp. n. in having a spherical, not observed. Nucleus with nucleoli. flattened shell and larger shell size.
Plate 4. a - Fabalesquereusia compressa sp. n., lateral view; b - F. linearis sp. n., lateral view; c -Shamkiriella convoluta sp. n., general view; d - Sh. convoluta sp. n., lateral view; e - Sh. phimatophora sp. n., lateral view.
Etymology. Species name refers to the graceful outlook of the shell.
Bipseudostomatella cornuta sp. n. (Fig. 15 c; Plate
5 e; Table 2).
Holotype. Slide K-ch N 12.
Type location. Between aquatic plants in shallow lagoons of the Kurakchay and Korchay rivers.
Description. General shell outlook resembles that of B. bifurcata. Shell diameter in the equatorial part is
85-90 ^m, shell height, 75-90 ^m. There are two apertures (40-45 ^m in diameter) connected to the shell by separate necks (20-25 ^m). The necks are rather close to each other and not at the opposite parts of the shell, as in the other two species of the genus. The shell surface is covered with closely adhering siliceous particles.
The cytoplasm is colourless, granular. There are 78 contractile vacuoles.
Discussion. B. cornuta clearly differs from the other two species ofthis genus in shell shape and aperture location.
Fig. 13. a - Cyphoderia trochus v. amphoralis, lateral view; b - Shamkiriella convoluta sp. n., lateral view; c - Sh. convoluta sp. n., dorsal view; d - Sh. reticulata sp. n., lateral view; e - Sh. reticulata sp. n., dorsal view.
Etymology. Species name refers to the presence of Gomocollariella gen. n.
two necks resembling horns. Diagnosis. Shell almost spherical, with two apertures
Fig. 14. a - Shamkiriellaphimatophora sp. n., lateral view; b - Sh. phimatophora sp. n., dorsal view; c ■ Bipseudostomatella bifurcata sp. n., lateral view; d - B. bifurcata sp. n., apertural view.
located on a single strong neck. Shell wall chitinous, Gomocollariella ranaformis gen. et sp. n. (Fig. 15 d;
with some mineral particles. Freshwater forms. Plate 5 f; Table 2).
Type species. Gomocollariella ranaformis sp. n. Holotype. Slide K-ch N 5.
Etymology. Genus name refers to a feature of shell Type location. On aquatic plants in shallow lagoons
morphology: the presence of a single neck with two of the Kurakchay river.
apertures. Description. The shell is almost spherical, 70-80 ^m
Fig. 15. a - Bipseudostomatella gracilis sp. n., aboral view; b - B. gracilis sp. n., apertural view; c - B. cornuta sp. n., lateral view ; d - Gomocollariella ranaformis sp. n., lateral view.
in diameter. There is a strong broad neck (60-70 ^m in length and 40-45 ^m in height) with two apertures (2022 ^m). Aperture shape oval to triangular. The shell is chitinous, light-brown, with a few quartz particles.
The cytoplasm is grey, granular. There are 2-6 lobose pseudopodia and 4-6 contractile vacuoles.
Etymology. Species name refers to the shell shape resembling a frog.
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Address for correspondence: Nataly Snegovaya, Ilham Alekperov. Laboratory of Protistology, Institute of Zoology, Azerbaijan National Academy of Sciences, Baku, 370073. E-mail: snegovaya@yahoo.com, l_alekperov@yahoo.com
Editorial responsibility: Andrew Goodkov
