Protistology 1 (4), 179–184 (2000) Protistology August, 2000

Data on the occurrence of species of the aurelia complex world-wide

Ewa Przyboœ and Sergei Fokin1 Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland, 1 Biological Research Institute of St. Petersburg State University, St. Petersburg, Russia

Summary

At present 15 species of the complex are known world-wide (Sonneborn, 1975; Aufderheide et al., 1983). Data on their distribution in the Americas, Africa, and Australia are mainly in the papers cited above, the following ones are cosmopolitan: P. primaurelia, P. biaurelia, P. tetraurelia, and P. sexaurelia. Data on the distribution of species of the P. aurelia complex in Asia are scattered in the literature and rather rare, P. primaurelia, P. biaurelia, P. tetraurelia, P. sexaurelia, and P. novaurelia were there recorded. The greatest data on distribution and frequency of occurrence of the particular species of the P. aurelia complex concerns Europe. As far as Europe is concerned, the following conclusions were drawn: P. novaurelia is a dominant species (found in 178 habitats among 459 studied), P. biaurelia is a frequent one (in 151 habitats), while P. primaurelia (in 103 habitats) is also characteristic but less frequent. Other species known from Europe are rare.

Key words: frequency of species occurrence, geographical distribution, Paramecium aurelia spp. complex

At present 15 species of the P. aurelia complex are occurrence of the particular species of the P. aurelia com- known world-wide (Sonneborn,1975; Aufderheide et al., plex concerns Europe (Table 3). The following species 1983). have been recorded there: P. primaurelia, P. biaurelia, P. Data on their distribution in the Americas, Africa, and triaurelia, P. tetraurelia, P. pentaurelia, P. sexaurelia, P. Australia are mainly in the papers cited above (Table 1). novaurelia, and P. tredecaurelia (Przyboœ, 1991). Recently, Among known species of the complex the following ones P. octaurelia was found in Western Germany (Stoeck and are cosmopolitan: P. primaurelia, P. biaurelia, P. Schmidt, 1998). The number of investigated clones and tetraurelia, and P. sexaurelia. Some of the species were habitats (Table 3) as well as the ratio value (r.v.), i.e. the found in two continents, i.e. P. octaurelia and P. number of habitats for a defined species to the total num- tredecaurelia, while others are known from the USA alone ber of habitats in the area (country or zone) were taken (P. septaurelia, P. decaurelia, P. undecaurelia, P. into consideration (Table 4) when the frequency of spe- dodecaurelia, and P. sonneborni) or only from Australia cies occurrence was estimated. (P. quadecaurelia). As far as Europe is concerned, the following conclu- Data on the distribution of species of the P. aurelia sions were drawn: complex in Asia (Table 2) are scattered in the literature 1. P. novaurelia is a dominant species (r. v. 0.39, found and rather rare. The species were recorded in 31 habitats, in 178 habitats among 459 studied in Europe), P. P. biaurelia appearing in 14, P. primaurelia in 9, while biaurelia is a frequent one (r. v. 0.33, in 151 habitats), both species are rather widely distributed. P. tetraurelia while P. primaurelia (r. v. 0.22, in 103 habitats) is was found in 4 habitats, P. sexaurelia in 3, and P. novaurelia also characteristic but less frequent. in 1 only. The last finding seems very interesting as that 2. P. tetraurelia (r. v. 0.05, in 25 habitats) and P. triaurelia species was regarded as restricted to Europe alone r.v. 0.04, in 17 habitats) are rather rare, while other (Sonneborn, 1975) but more recently Przyboœ (1998b) species very rare, i.e. P. sexaurelia (r. v. 0.01, in 5 recorded it in Turkey (Anatolian Upland). habitats), P. pentaurelia (r. v. 0. 007, in 3 habitats and The greatest data on distribution and frequency of only in southern Europe), P. octaurelia (r.v. 0.004, in

© 2000 by Russia, Protistology. 180 · Ewa Przyboœ and Sergei Fokin

Table 1. Occurrence of species of the Paramecium aurelia complex in the Americas, Africa, Australia (data from: Sonneborn 1975; Aufderheifde et al. 1983)

Continent or country P. primaurelia P. biaurelia P. P. triaurelis P. tetraurelia pentaurelia P. P. sexaurelia septaurelia P. P. octaurelia P. ** novaurelia P. decaurelia P. undecaurelia dodecaurelia P. P. tredecaurelia quadecaurelia P. P. sonneborni

Northern ++++++++–+++ + America Central ++++ + America Southern ++ + America Hawai + New + Zealand Australia ++ + Kenya + Uganda + Madagascar +

** Known from Europe only (Sonneborn, 1975), found in Asia (Przyboœ, 1998b).

2 habitats), and P. tredecaurelia (r. v. 0.002, in 1 habi- Paramecium aurelia species-complex. J. Protozool. tat). 30, 128–131. 3. Nevertheless, a different number of habitats were stud- Daggett P.-M. (Ed.). 1978. Protozoa and . In: Cata- ied in the particular zones, 21 in the northern, 363 in logue of Strains. I. Thirteenth edition, the American the central, and 75 in the southern zone, the prevail- Type Culture Collection. Rockville, Maryland. pp. 1– ing number of habitats originating in the central zone, 18. and mainly in Poland. Of the total number of 363 stud- Komala Z. 1961. Syngens (varieties) of Paramecium ied habitats in that zone, 286 were situated in Poland. aurelia in Bulgaria. Folia biol. (Krakow). 9, 229–232. We can assume that not all Paramecium species have Komala Z. 1994. Contemporary investigations on the world-wide distribution, as a result of our sampling done Paramecium aurelia in the Tatras. in different parts of the world and extract from the litera- Preliminary Report. Folia biol. (Krakow). 42, 35–39. ture (Przyboœ and Fokin, 2000b). Because some species Komala Z. 1997. A new habitat of of P. aurelia complex as well as a part of other paramecia in Poland. Folia biol. (Krakow). 45, 55–58. cannot be considered as a cosmopolitan, we have to ex- Komala Z.1998a. Report on the Paramecium aurelia spe- pect an interesting sampling in some places of the world cies complex in a water body in the borderland of the which were not yet checked for fauna. Not so much Wyspowy Beskid and Gorce Mts. Folia biol. sampling was done in the Southern Hemisphere so far. (Krakow). 46, 69–71. Thus, it is possible to expect a discovery somewhere even Komala Z. 1998b. Current investigations on the occur- a new Paramecium species. rence of the Paramecium aurelia species complex in Poland. Folia biol.(Krakow).46, 73–76. Komala Z. 1998c. Investigations on the occurrence of the References Paramecium aurelia species complex in the Carpathians. Folia biol. (Krakow). 46, 77–81. Aufderheide K. J., Daggett P.-M. and Nerad T. A. 1983. Komala Z. and Dubis K. 1965. Contribution to the obser- Paramecium sonneborni n. sp., a new member of the vations on the occurrence of Paramecium aurelia Protistology · 181

Table 2. Occurrence of species of the Paramecium aurelia complex in Asia

Number of known strains and habitas (in brackets)

Country References or region P. primaurelia P. P. biaurelia P. tetraurelia P. sexaurelia P. novaurelia Total

Japan 7(4) 7(7) 4(3) 18(4) Sonneborn, 1950; Koœciuszko and Koizumi, 1984 Russia 1(1) 3(3) 4(4) Daggett, 1978; Preer et al. 1974; Koœciuszko, (Far East) 1985; Przyboœ and okin, 1996 Vietnam 1(1) 1(1) Przyboœ and okin, 1996

Thailand 2(2) 2(2) Daggett, 1978; Przyboœ and okin, 2000 a,b India 9(1) 9(1) Sonneborn, 1974 Turkmenia 1(1) 1(1) Daggett, 1978 Georgia 3(3) 3(3) Daggett, 1978; Koœciuszko, 1985 Turkey 1(1) 1(1) Przyboœ, 1998b Lebanon 2(1) 2(1) Sonneborn, 1974 Israel 2(2) 1(1) 3(3) Przyboœ, 1995; Przyboœ and okin, 1999 Total 12(9) 15(14) 5(4) 11(3) 1(1) 44(31)

syngens in Italy. Folia biol. (Krakow). 13, 265–267. cal Garden of the Jagiellonian University in Krakow. Komala Z. and Dubis K. 1966. Syngens of Paramecium Folia biol. (Krakow). 48 (in press). aurelia in some regions of Moscow and Leningrad. Koœciuszko H. 1964. Syngens of Paramecium aurelia in Folia biol. (Krakow). 14, 227–228. Hungary. Folia biol. (Krakow). 12, 17–22. Komala Z. and Koœciuszko H. 1963. Investigations on the Koœciuszko H. 1985. Species of the Paramecium aurelia occurrence of different varieties of Paramecium complex in some regions of the USSR. Folia biol. aurelia in Poland. Proceedings of the First Interna- (Krakow). 33, 117–122. tional Conference on Protozoology. Prague, 1961. pp. Koœciuszko H. and Koizumi S. 1984. Habitats of the Para- 96–98. mecium aurelia complex in Japan. Folia biol. Komala Z. and Przyboœ E. 1992. Species of the Parame- (Krakow), 32, 57–62. cium aurelia complex in the Middle Sudeten of Koœciuszko H. and Prajer M. 1988. Habitats of species of Czecho-Slovakia. Folia biol. (Krakow). 40, 129–135. the Paramecium aurelia complex and some other Komala Z. and Przyboœ E. 1994. Paramecium triaurelia Paramecium species in Finland. Folia biol. (Krakow). and other species of the aurelia complex in the 36, 65–72. Kaczawskie Mts and Plateau (Western Sudeten). Fo- Koœciuszko H. and Prajer M. 1991. New habitats of spe- lia biol. (Krakow). 42, 27–33. cies of the Paramecium aurelia complex in Komala Z. and Przyboœ E. 1999. Paramecium biaurelia Scandinavia (southern Sweden). Folia biol. (Krakow). and other accompanying zooplankton in the ponds of 39, 25–27. the Botanical Garden in Krakow. Folia biol. (Krakow). Preer J. R. Jr., Preer L. B. and Jurand A. 1974. Kappa and 47, 51–52. other in Paramecium aurelia. Bact. Komala Z. and Przyboœ E. 2000. Further investigations Rev. 38, 113–163. on the zooplankton of the water bodies of the Botani- 182 · Ewa Przyboœ and Sergei Fokin

Table 3. Occurrence of species of the Paramecium aurelia complex in Europe

Number of studied clones and habitats (in brackets)

Region References (temperature zone) P. primaurelia P. biaurelia P. triaurelia P. tetraurelia P. pentaurelia P. sexaurelia P. octaurelia P. novaurelia P. tredecaurelia Total

Northern Sweden 41 62 101 Koœciuszko and Prajer, 1991 (2) (2) (3)* Norway 1 1 Sonneborn, 1956** (1) (1) Finland 11 6 54 71 Koœciuszko and Prajer, 1988 (2) (1) (2) (5) Russia 112 13 1 58 184 Komala and Dubis, 1966; (7) (4) (1) (7) (12)* Koœciuszko, 1985; Przyboœ and Fokin, 1996 Central Great 1 3 1 2 7 Sonneborn, 1956; Tait et al. Britain (1) (3) (1) (2) (7) 1971** Poland 341 557 15 77 763 1753 Przyboœ and Komala, 1993 (56) (56) (3) (9) (122) (286)* (ref. since 1959), 1996, 1997; Komala and Przyboœ, 1994, 1999, 2000; Komala 1997, 1998 a, 1998b, 1998c Ukraine 2 8 16 32 58 Przyboœ and Chornobai, 1994, (2) (2) (5) (8) (17)* 1998; Przyboœ, 1997 Czech 86 18 1 95 200 Komala and Koœciuszko, 1963; Republic (13) (2) (1) (9) (21)* Komala and Przyboœ, 1992; Przyboœ and Komala, 1992 Slovakia 7 51 58 Dubis and Komala, 1964 (data (3) (6) (7)* from: Komala, 1994) Germany 12 1 14 11 17 56 Przyboœ and okin, 1997; (7) (1) (2) (2) (7) (19)* Sonneborn, 1956, 1974**; Stoeck and Schmidt, 1998 France 1 1 1 1 4 Sonneborn, 1956; Rafalko and (1) (1) (1) (1) (4) Sonneborn, 1959 Holland 1 1 Sonneborn et al. 1959** (1) (1) Switzerland 1 1 Sonneborn, 1956 (1) (1)

Przyboœ E. 1968. The occurrence of syngens of Parame- Przyboœ E. 1995. Species of the Paramecium aurelia com- cium aurelia in Rumania. Folia biol. (Krakow). 16, plex in Israel (Ciliophora, Protista). Israel J. Zool. 41, 131–136. 205–206. Przyboœ E. 1980. Distribution of species of the Parame- Przyboœ E. 1996. The Paramecium aurelia habitat in cium aurelia complex in Spain. Folia biol. (Krakow). Greece. Folia biol. (Krakow). 44, 105–106. 28, 405–412. Przyboœ E. 1997. The Paramecium aurelia species com- Przyboœ E. 1990. Intraspecies differentiation of Parame- plex on the Volyn-Podolian Upland. Folia biol. cium sexaurelia (Ciliophora). Arch. Protistenkd. 138, (Krakow). 45, 59–63. 123–125. Przyboœ E. 1998a. New habitats of Paramecium biaurelia Przyboœ E. 1991. Studies on the Paramecium aurelia spe- in Italy. Folia biol. (Krakow). 46, 87–89. cies complex in Spain (Ciliophora). Arch. Protistenk. Przyboœ E. 1998b. The first habitat of Paramecium 140, 151–156. novaurelia of the P. aurelia spp. complex in Asia Tur- Przyboœ E. 1993. Species of the Paramecium aurelia com- key). Folia biol. (Krakow). 46, 91–95. plex in Spain. Microbiologia SEM. 9, 113–117. Protistology · 183

Table 3. Contination

Southern Italy 19 10 1 28 Sonneborn, 1956; Sonneborn (4) (3) (1) (8) et al., 1959; Komala and Dubis, 1965; Przyboœ, 1998a Hungary 42 15 1 16 74 Koœciuszko, 1964 (13) (5) (1) (9) (27)* Bulgaria 2 3 1 1 7 Komala, 1961**; Komala and (2) (3) (1) (1) (5)* Koœciuszko, 1963 Romania 6 18 9 3 26 62 Przyboœ, 1968 (2) (4) (2) (1) (4) (12)* Greece 21 21 Przyboœ, 1996; Przyboœ, and (1) (1) Lekka, 2000 Spain 96 91 9 48 1 2 39 286 Przyboœ, 1980, 1990, 1991, (11) (7) (3) (4) (1) (2) (4) (22)* 1993 Total 628 870 69 187 5 37 11 1167 1 2975 (103) (151) (17) (25) (3) (5) (2) (178) (1) (459)* * Some species appear together in one habitat. ** Without detailed data concerning the number of the studied clones.

Table 4. Frequency of occurrence of species of the Paramecium aurelia complex in the temperature zones in Europe

Ratio value of particular species Total number of studied Zone clones and habitats (in brackets) P. primaurelia P. biaurelia P. P. triaurelia P. tetraurelia P. pentaurelia P. sexaurelia P. octaurelia P. novaurelia P. tredecaurelia

Northern 0.33 0.43 0.05 0.05 –––0.52 – 369 (21)

Central 0.18 0.34 0.03 0.05 – 0.005 0.005 0.41 0.003 2138 (363)

Southern 0.43 0.27 0.07 0.08 0.04 0.04 – 0.24 – 478 (75)

Total ratio 0.22 0.33 0.04 0.05 0.007 0.01 0.004 0.39 0.002 2975 (459) value

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Address for correspondence: Ewa Przyboœ. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow 31–016, Slawkowska 17, Poland. E-mail: [email protected]