Testate Amoebae (Protozoa: Rhizopoda) from Northwest Yunnan, China

Testate Amoebae (Protozoa: Rhizopoda) from Northwest Yunnan, China Jun Yangasbsc,Wenjing Zhangbrc,Weisong Fengb,and Yunfen Shenb-d ABSTRACT

A survey of the testate amoebae of northwest Yunnan in the Hengduan Mountains region of southwest China revealed 151 species belonging to 29 genera. Of these, three genera and 27 species were new to the testate amoebae fauna of China. The widely distributed and dominant species were Centropyxis aculeata aacleata, C, aerophila aerophila, C. sylvatica, Dzflugia elegans, D. globulosa, D. pristis, D. pulex, Phryganella hemisphaerica, Trinema enchelys, and T lineare. The most diverse genera were Dzflugia (41 species), Centropyxis (3 1 species), Arcella (1 8 species), and Euglypha (1 1 species). The number of species in each sample varied greatly. It appears that species diversity of testate amoebae is related not only to macro-environments (e.g., geographic zonation) but also to microenvironments (e.g., microhabitats). INTRODUCTION Northwest Yunnan (25'30'-29' 15'N, 98'05'- 10 l o15'E), covering an area of 68,908 krn2, is located in the southern Hengduan Mountains region of the Eastern Himalayas and comprises the transitional region from the Qinghai-Tibet Plateau to the Yunnan-Guizhou Plateau in southwest China. It is one of only a few places on the planet recognized as both a Global Biodiversity Hotspot (Mittermeier and Mittermeier 1997) and a Global 200 Priority Ecoregion (Olsen and Dinerstein 1998). The main climate of northwest Yunnan is semitropically humid with an annual mean temperature about 15 "Cand an annual mean precipitation about 900 mm. It has unique physical conditions and variety of ecosystems coupled with its ecological function as a north-south corridor for the movement of plants and animals (especially during the ice ages), which gave rise to a rich flora and fauna, including many relict taxa and endemic species (Xiwen and Walker 1986, Zhang et al. 1997, Chen 1998, Zhang 1999). The testate amoebae comprise those amoeboid organisms which are enveloped by a shell or test, within which the body can be completely withdrawn. They are present in a wide range of moist and freshwater habitats, play an important role in their ecology, and are used as an important new class of paleoenvironmental indicator (Ogden and Hedley 1980, Meisterfeld 2000, Patterson and Kumar 2002). Some species have cosmopolitan distributions, but others have restricted geographical distributions. For instance, several species from the Nebelidae occur only in the Southern Hemisphere (Smith and Wilkinson 1986, Foissner 1987, Beyens and Meisterfeld 2001). Unfortunately, few investigations on testaceans from the vicinity of Tibet have been carried out (Laminger 1972a, Wang 1974, Suxena 1979, Shen 1983). Only 16 species of the testate amoebae belonging to two common genera (Arcella and Dzflugia) were mentioned in previous publications on testate amoebae from the Hengduan Mountains region (Hsiao 1949, Zhang and Guo 1986, Yang et al. 2004). The purpose of this work was to provide the faunistic data on testate amoebae in northwest Yunnan and to compare the common species between northwest Yunnan and the regions of Tibet and the Qomolangma Mountain. aDonghu Experimental Station of Lake Ecosystems, Chinese Academy of Sciences, Wuhan 430072, P. R. China. ~nstituteof Hydrobiology, Chinese Academy of Sciences, Wuhan 430072, P. R. China. Graduate School of the Chinese Academy of Sciences, Beijing 100039, P. R. China. Corresponding author; E-mail: [email protected]

Journal of Freshwater Ecology, Volume 20, Number 3 - September 2005

MATERIALS AND METHODS The 74 samples for this study were collected from 47 localities in northwest Yunnan between 7 August and 8 September 1981. Sampled habitats included a variety of types ponds, streams, lakes, rivers, springs, ditches and others. They were taken with a 64 pm mesh plankton net and scraped from substrates (e.g., stones and macrophytes), fixed with 5% formaldehyde, and preserved in the Herbarium of the Institute of Hydrobiology in Wuhan, China. Identification of testate amoebae was based on BartoS (1954), Beyens and Chardez (1997), Cash and Hopkinson (1905, 1909), Cash, Wailes and Hopkinson (1915, 1918), Decloitre (1962, 1969, 1976a, 1976b, 1977a, 1977b, 1978, 1979a, 1979b, 1981), Deflandre (1928, 1929, 1931, 1936), Foissner and Korganova (2000), Gauthier-Likvre and Thomas (1958), Grospietsch (1964), Liiftenegger and Foissner (1991), Ogden and Hedley (1980), Ogden (1983), Page (1966), Shen (1983), S t ~ ~ i n (1952), ek van Oye (1933, 1958, 1959), Vucetich (1973), Wang and Nie (1933), and Yang et al. (2004). To quantify community differences and similarities between northwest Yunnan and the regions of Tibet (Wang 1977, Shen 1983) and the Qomolangma Mountain (Wang 1974, Shen 1983), we used Snrrenson's coefficient of similarity. The coefficient is described as 2c/(a+b), where a is the number of species in region A, b is the number of species in region B, and c is the number of species in both regions A and B. RESULTS AND DISCUSSION A highly diverse testate amoebae community was found in the samples from northwest Yunnan. A total of 151 species, belonging to 29 genera, was identified (Table l), and the following species were widely distributed and dominant: Centropyxis aculeata aculeata, C. aerophila aerophila, C. sylvatica, Drflugia elegans, D. globulosa, D. pristis, D. pulex, Phryganella hemisphaerica, Trinema enchelys, and T. lineare. In addition, three genera and 27 species of testate amoebae were recorded for the first time in China (Table 1). Thus, the total number of the established testate amoebae in China now amounts to 30 1 species (subspecies) of 44 genera. Most species were found in the genera Drfflugia (4 l), Centropyxis (3 l), Arcella (1 8), Euglypha (1 l), Cyclopyxis (9) and Trinema ( 5 ) . To some extent, this is in accordance with the results reported from Tibet where the most diverse genera were Drfflugia (28), Centropyxis (28), Arcella (22), Nebela (10) and Euglypha (9) (Shen 1983). In the Arctic testate amoebae fauna, the situation seems to be a bit different. Beyens and Chardez (1995) found that Dzflugia ( 4 9 , Centropyxis (45), Euglypha (25), Nebela (22) and Arcella (20) were the most diverse genera in the Arctic between longitudes 27OE and 168"W. Clearly, all regions have the highest number of species in the genera Dzflugia and Centropyxis, which was most similar to that of temperate and boreal zones in the Northern Hemisphere (Beyens et al. 1995). However, the northwest Yunnan testate amoebae community was remarkably different from those ofAustria and New Zealand (van Oye 1956, Laminger 1972b), where the most diverse genus was not Dzflugia. In the high mountains of Austria (Laminger 1972b), the most diverse genera were Centropyxis (17), Nebela (17), Euglypha (13), Dtflugia (1 I), Arcella (5) and Trinema ( 5 ) , and in New Zealand (van Oye 1956), the most diverse genera were Nebela (9) and Euglypha (5). Further, van Oye (1956) believed that the genus Nebela occupied a predominant position and was the most diverse genus among the fauna of New Zealand, which was most similar to that of South America. In our study, the most predominant species, in decreasing order, were DzfJlugia pristis, Centropyxis aculeata aculeata, and D. globulosa. According to Shen (1983), D. globulosa was the first predominant species in northern Tibet and western Tibet, while C. aculeata aculeata was the first predominant species in southern Tibet, eastern Tibet, and

Table 1. Species list of testate amoebae found in northwest Yunnan, China. Arcella arenaria Greeff, 1866 A. arenaria sphagnicola Deflandre, 1928 *A. artocrea aplanata Grospietsch, 1954 A. catinus Penard, 1890 A. discoides Ehrenberg, 1871 *A. discoides pseudovulgaris Deflandre, 1928 A. discoides scutellIJbrmis Playfair, 1918 A. hemisphaerica Perty, 1852 A. hemisphaerica gibba Deflandre, 1928 *A. hemisphaerica intermedia Deflandre, 1928 A. hemisphaerica undulata Deflandre, 1928 A. megastoma Penard, 1902 A. polypora Penard, 1890 A. rotundata Playfair, 1918 A. rotundata aplanata Deflandre, 1928 *A. tuberosus Decloitre, 1954 A. vulgaris Ehrenberg, 1832 A. vulgaris undulata Deflandre, 1928 Assulina muscorum Greeff, 1888 A. seminulunt (Ehrenberg) Leidy, 1872 Bullinularia indica Penard, 1907 B. minor (Penard) Hoogenraad et De Groot, 1948 Canzpascus minutus Penard, 1899 Centropyxis aculeata aculeata (Ehrenberg) Stein, 1857 C. aculeata grandis Deflandre, 1929 *C. aculeata minima van Oye, 1953 C. aculeata oblonga Deflandre, 1929 C. aculeata oblonga f. minor Kufferath, 1929 *C. aculeata tropica Deflandre, 1929 C. aerophila aerophila Deflandre, 1929 *C. aerophila grandis StCp~nek,1963 C. aerophila sphagnicola Deflandre, 1929 C. cassis (Wallich) Deflandre, 1929 C. cassis compressa Shen, 1981 *C. cassis gigantea Decloitre, 1966 C. cassis spinifera Playfair, 1918 *C. circularis Decloitre, 1968 C. constricta (Ehrenberg) Deflandre, 1929 *C. constricta minima Decloitre, 1953 C. cordformis cyctostoma Hu et Shen, 1997 C. discoides (Penard) Deflandre, 1929 C. ecornis (Ehrenberg) Leidy, 1879 C. ecornis Ieidyi Thomas, 1957 *C. ecornis minuta Golemansky, 1962 C. elongata (Penard) Thomas, 1959 *C. gauthieri Thomas, 1959 C. hemisphaerica (Barnard) Wailes, 1 9 13 C. hirsuta Deflandre, 1929 C. marsupformis ( ~ a l l i c h Deflandre, ) 1929 * New species to the testacean fauna of China ** New genera to the testacean fauna of China

C. minuta Deflandre, 1929 C. platystonza (Penard) Deflandre, 1929 C. spinosa (Cash) Deflandre, 1929 C. sylvatica (Deflandre) Thomas, 1955 C. triangularis Shen, 1981 Corythion dubium Tarhek, 1881 ** Cryptodlflugia ovijormis Penard, 1890 Cucurbitella mespiliformis Penard, 1902 Cyclopyxis arcelloides (Penard) Deflandre, 1929 C. deflandrei van Oye, 1956 C. eurysroma Deflandre, 1929 *C. eurystoma gauthieriana Bonnet et Thomas, 1960 C. eurystoma parvula Bonnet et Thomas, 1960 C. kahli Deflandre, 1929 *C. kahli crucigenia StCpanek, 1963 C. kahli cyclostoma Bonnet et Thomas, 1960 *C. plana BartoS, 1963 Cyphoderia ampulla vitraea Wailes et Penard, 1911 Difflugia acuminata Ehrenberg, 1838 D. amphoralis globosa Gauthier-Litvre et Thomas, 1958 D. ampuliula Playfair, 1918 D. avellana Penard, 1890 D. baciliariarum Perty, 1849 D. brevicolla Cash, 1909 D. curvicaulis Penard, 1899 D. dlficilis Thomas, 1954 D. elegans Penard, 1890 D. elegans angustata Deflandre, 1926 D. elegans bicornis Jung, 1936 D. fallax Penard, 1890 D. gassowkii (Gassowsky) Ogden, 1983 D. glans Penard, 1902 D. globulosa Dujardin, 1837 D. gramen Penard, 1902 D. hiraethogii Ogden, 1983 D. kabylica Gauthier-Litvre et Thomas, 1958 D. lacustris (Penard) Ogden, 1983 D. lanceoiata Penard, 1890 D. lemani Blanc, 1892 *D. levanderi Playfair, 1918 D. limnetica (Levander) Penard, 1902 D. lithophila Penard, 1902 D. lobostoma Leidy, 1879 D. lucida Penard, 1890 D. mammillaris Penard, 1893 D. manicata Penard. 1902 D. minuta Rampi, 1950 (continued)

Table 1, continued D. murlformis crucilobata Gauthier-Lievreet Thomas, 1958 D. oblonga Ehrenberg, 1838 D. oblonga nodosa Leidy, 1879 D. pristis Penard, 1902 D. pulex Penard, 1902 D. rubescens Penard, 1902 D. sarissa Tai, 1931 D. stoutii Ogden, 1983 D. tenuis (Penard) Ogden, 1983 D. urceolata Carter, 1864 D. varians Penard, 1902 D. viscidula Penard, 1902 Euglypha acanthophora (Ehrenberg) Perty, 1849 E. acanthophora brevispina Penard, 1902 E. ciliata (Ehrenberg) Leidy, 1878 E. compressa Carter, 1869 * E. cristata acicularis Wailes, 1912 E. filifera Penard, 1890 E. laevis (Ehrenberg) Perty, 1849 E. rotunda Wailes et Penard, 191 1 *E. rotunda obliqua Decloitre, 1956 E. strigosa (Ehrenberg) Leidy, 1879 E. tuberculata Dujardin, 1841 Heleopera petricola Leidy, 1879 *Hoogenraadia afvicana Gauthier-Litvre et Thomas, 1958 Hyalosphenia elegans Leidy, 1879 H. minuta Cash, 1891 H. subjiava Cash, 1909 Lesquereusia epiforniumPenard, 1893 * New species to the testacean fauna of China ** New genera to the testacean fauna of China

L. spiralis (Ehrenberg) Biischli, 1880 *Nebela galeafa Penard, 1890 N. militaris Penard, 1890 N. fubulosa Penard, 1890 'Nefzelia labeosa Beyens ef Chardez, 1997 *N. ovformis (Cash) Ogden, 1979 N. tuberculata (Wallich) Netzel, 1983 Phryganella hemisphaerica Penard, 1902 I? paradoxa Penard, 1902 Plagiopyxis callida Penard, 1910 l? declivis Thomas, 1955 Pontigulasia bigibbosa Penard, 1902 l? spectabilis Penard, 1902 Pseudodzjlugia fascicularis Penard, 1902 I? gracilis Schlumberger, 1845 Quadrulella syrnmefrica (Wallich) Schulze, 1875 **Schoenborina humicola (Schonborn) Decloitre, 1964 **Schwabiaterricola Bonnet et Thomas, 1955 Tracheleuglypha dentafa(Vedjowsky) Deflandre, 1928 Trigonopyxis arcula (Leidy) Penard, 1912 Trinema complanatum Penard, 1890 *T. con~planatunzinaequalis Decloitre, 1969 1: enchelys (Ehrenberg) Leidy, 1878 * I : intermedia Decloitre, 1965 7: lineare Penard, 1890 Wailesellaeboracensis (Wailes) Deflandre, 1928

southeastern Tibet. There has obviously bear a shift in the most dominant species from northwest to southeast among these regions. In most regions of Tibet the predominant genera were Centropyxis, Dzflugia and Arcella, which were considered as faunal elements of the Palaearctic realm; but in southern Tibet, eastern Tibet and southeastern Tibet the genus Trinema has second and third positions with respect to abundance, which reflected the faunal feature of the Oriental realm (Shen 1983). Northwest Yunnan is located in the transitional region between the Palaearctic and Oriental realms; therefore, its diversified testate amoebae species are probably shared by both realms. The number of species in each sample varied greatly. There was one sample with maximum of 50 species, accounting for 33.1% of the total species from northwest Yunnan. Another sample had 32 species, and one had 30 species. However, fewer than 10 species were found in 56.9% of the samples, except for the samples where no testate amoebae were observed (23 out of 74). Furthermore, more than one third of all species (57 species) were observed only in one sample, showing a very patchy distribution of most species in northwest Yunnan. Furthermore, no species occurred in all samples, and only three species (Dzflugiapristis, Centropyxis aculeata aculeata and Trinema lineare) had frequencies of > 20% (Fig. 1). The testacean fauna of northwest Yunnan was rich and diverse, possibly due to the richness in microhabitats diversity.

Table 2. The number of common species of testate amoebae and Sgrenson's similarity coefficients between northwest Yunnan and different regions of Tibet and the Qomolangma Mountain. Region Northern Tibet Western Tibet Southern Tibet, the upper Brahmaputra Basin Northern Qomolangma Mountain Southern Qomolangma Mountain Southeastern Tibet Southern Tibet, the middle Brahmaputra Basin Eastern Tibet

Species 17 35 23 31 35 56 59 68

Coefficient 0.20 0.37 0.26 0.32 0.35 0.49 0.49 0.56

In northwest Yunnan, many species were identical to those reported from several regions of Tibet (Wang 1977, Shen 1983) and the Qomolangma Mountain (Wang 1974, Shen 1983). Species common to northwest Yunnan and southern Tibet of the middle Brahmaputra basin and eastern Tibet accounted for 37.7% and 44.4% of the total species from the northwest Yunnan, respectively. Our calculated Ssrenson's coefficient of similarity ranged from 0.20 to 0.56 (Table 2), demonstrating that the fauna of northwest Yunnan and eight regions of Tibet and the Qomolangma did not have strong similarities. Local ecological factors play an important role in shaping community structure and in determining species richness of unicellular eukaryotes (Hillebrand and Azovsky 2001). We therefore suggest that the variation we observed for species in these two regions may be reflecting variation in available microhabitats.

23.0 21.6

20.3

Number of species

Figure 1. Frequencies of 15 1 testacean species in 74 samples from northwest Yunnan, China. ACKNOWLEDGEMENTS We thank Wenxuan Cao, Jiayou Cheng, and Yiyu Chen for collecting samples. Special thanks go to Harry Goon who made linguistic corrections. This work was supported by the National Natural Science Foundation of China (30270185), the Knowledge Innovation Program of the Chinese Academy of Sciences (KSCX 2-2-06), and the Foundation of Donghu Experimental Station of Lake Ecosystems, the Chinese Academy of Sciences (200202).

LITERATURE CITED BartoS, E. 1954. Korefionoice Radu Testacea. Vydalo Vydavatelstvo Slovenskej AkadCmie Vied Bratislava, 187 pp. Beyens, L. and D. Chardez. 1995. An annotated list of testate amoebae observed in the Arctic between the longitudes 27"E and 168"W. Arch. Protistenkd. 146: 219-233. Beyens, L. and D. Chardez. 1997. New testate amoebae taxa from the polar regions. Acta Protozool. 36: 137-142. Beyens, L. and R. Meisterfeld. 2001. Protozoa: Testate amoebae. In: J.P. Smol, H.J.B. Birks, and W.M. Last. (eds.) Tracking Environmental Change Using Lake Sediments. Kluwer Academic Publishers, Dordrecht, The Netherlands, pp. 121-153. Beyens, L., D. Chardez, D. de Baere, and C. Verbruggen. 1995. The aquatic testate amoebae fauna of the Strsmness Bay area, South Georgia. Antarct. Sci. 7: 3-8. Cash, J. and J. Hopkinson. 1905. The British Freshwater Rhizopoda and Heliozoa. Vol. 1 Rhizipoda, part I. The Ray Society London, 150 pp. Cash, J. and J. Hopkinson. 1909. The British Freshwater Rhizopoda and Heliozoa. Vo1.2 Rhizipoda, part 11. The Ray Society London, 168 pp. Cash, J., G.H. Wailes, and J. Hopkinson. 1915. The British Freshwater Rhizopoda and Heliozoa. Vo1.3 Rhizipoda, part 111. The Ray Society London, 156 pp. Cash, J., G.H. Wailes, and J. Hopkinson. 1918. The British Freshwater Rhizopoda and Heliozoa. Vo1.4 Rhizipoda, part IV. The Ray Society London, 130 pp. Chen, Y.Y. 1998. The Fishes of the Henduan Mountains Region. Science Press, Beijing, 264 pp. Decloitre,.L. 1962. Le genre Euglypha Dujardin. Arch. Protistenkd. 106: 5 1- 100. Decloitre, L. 1969. Contribution a la faune de l'Iran. Rhizopodes thecamoebiens. Bulletin du Museum National &Histoire Naturelle (2e sdrie) (Paris) 4 1: 362-37 1. Decloitre, L. 1976a. Le genre Arcella Ehrenberg. Arch. Protistenkd. 118: 291-309. Decloitre, L. 1976b. Le genre Euglypha. Arch. Protistenkd. 118: 18-33. Decloitre, L. 1977a. Le genre Cyclopyxis. Arch. Protistenkd. 119: 31-53. Decloitre, L. 1977b. Le genre Nebela. Arch. Protistenkd. 119: 325-352. Decloitre, L. 1978. Le genre Centropyxis I. Arch. Protistenkd. 120: 63-85. Decloitre, L. 1979a. Le genre Centropyxis 11. Arch. Protistenkd. 121: 162-192. Decloitre, L. 1979b. Mises ajour au 31.12.1978 des mises i jour au.31.12.1974 concernant les genres Arcella, Centropyxis, Cyclopyxis, Euglypha et Nebela. Arch. Protistenkd. 122: 387-397. Decloitre, L. 1981. Le genre Trinema Dujardin, 1841. Arch. Protistenkd. 124: 193-218. Deflandre, G. 1928. Le genre Arcella Ehrenberg. Arch. Protistenkd. 64: 152-287. Deflandre, G. 1929. Le genre Centropyxis Stein. Arch. Protistenkd. 67: 322-375. Deflandre, G. 1931. Thecamoebiens nouveaux ou peu connus, I. Ann. de Protistol. 3 : 81-109. Deflandre, G. 1936. Etude monographique sue le genre Nebela Leidy (RhizopodaTestacea). Ann. de Protistol. 5: 201-322. Foissner, W. 1987. Soil protozoa: fundamental problems, ecological significance, adaptations in ciliates and testaceans, bioindicators and guide to the literature. Progr. Protistol. 2: 69-2 12. Foissner, W. and G.A. Korganova. 2000 The Centropyxis aerophila complex (Protozoa: Testacea). Acta Protozool. 39: 257-273. Gauthier-Likre, L, and R. Thomas. 1958. Les genres Dzflugia, Pentagonia, Maghrebia et Hoogenraadia (Rhizopodes testacCs) en Afrique. Arch. Protistenkd. 103: 241-370. Grospietsch, T. 1964. Die Gattungen Cryptodlflugia und Drflugiella (Rhizopoda Testacea): Zool. Anz. 172: 243-257.

Hillebrand, H. and A.I. Azovsky. 200 1. Body size determines the strength of the latitudinal diversity gradient. Ecography 24: 25 1-256 Hsiao, S.C. 1949. A limnological study of Erh Hai, 2 biological features. J. Anim. Ecol. 18: 89-98. Laminger, H. 1972a. Notes on some terrestrial Testacea (Protozoa, Rhizopoda) from Nepal, Himalaya (Lhotse Shar). Arch. Protistenkd. 114: 486-488. Laminger, H. 1972b. Ein beitrag zur Kenntnis der Hochgebirgs-Testaceen~sterreiche. Arch. Protistenkd. 114: 101- 151. Liifienegger, G. and W. Foissner. 1991. Morphology and biometry of twelve soil testate amoebae (Protozoa, Rhizopoda) from Australia, Africa, and Austria. Bull. Br. Mus. (Nat. Hist.) Zool. 57: 1-16. Meisterfeld, R. 2000. Order Arcellinida Kent, 1880. In: J.J. Lee, G.F. Leedale, and P. Bradbury. (eds.) An Illustrated Guide to the Protozoa (2nd), Allen Press, Lawrence Kansas, USA, pp. 827-860. Mittermeier, R.A. and C.G. Mittermeier. 1997. Megadiversity: Earth's Biologically Wealthiest Nations. CEMEX, S.A. Ogden, C.G. 1983. Observations on the systematics of the genus Dzflugia in Britain (Rhizopoda, Protozoa). Bull. Br. Mus. (Nat. Hist.) Zool. 44: 1-73. Ogden, C.G. and R.H. Hedley. 1980. An Atlas of Freshwater Testate Amoebae. British Museum (Natural History) and Oxford University Press, London and Oxford, 222 PP. Olsen, D.M. and E. Dinerstein. 1998. The Global 200: a representation approach to conserving the earth's most biologically valuable ecoregions. Conserv. Biol. 12: 502-515. Page, F.C. 1966. Cryptodzflugia operculata n. sp. (Rhizopodea: Arcellinida, Cryptodifftugiidae) and the status of the genus Cryptodiflugia . Trans. Am. Microsc. SOC.85: 506-515. Patterson, R.T. and A. Kumar. 2002. A review of current testate rhizopod (thecamoebian) research in Canada. Paleogeogr. Paleoclimatol. Paleoecol. 180: 225-25 1. Shen, Y.F. 1983. Protozoa of the Tibetan Plateau. In: X.Z. Jiang, Y.F. Shen, and X.J.Gong. (eds.) Aquatic Invertebrates of the Tibetan Plateau. Science Press, Beijing, pp. 48-100. Smith, H.G. and D.M. Wilkinson. 1986. Biogeography of testate rhizopods in the es southern temperate and Antarctic zone. Colloque les ~ c o s ~ s t e mTerrestres Subantarctiques 58: 83-96. St&pknek,M. 1952. Testacea of the pond of Hradek at Kunratice (Prague). Acta Musei Nationalis Pragae, Series B 8: 1-55 Suxena, M.R. 1979. Algae and Testacea from high altitudes of Himalayas. I (collected by N.C.C. Punch Chulli Expedition, W. Himalaya 1970). Hydrobiologia 65: 107-128. van Oye, P. 1933. Rhizopodes du district sub-alpin de la Belgique. Archiv fir Naturgeschichte,Abtailung B, NF 2: 538-573. van Oye, P. 1956. On the thecamoeban fauna of New Zealand with description of four new species and biogeografical discussion. Hydrobiologia 8: 16-37. van Oye, P. 1958. ~ t u d sur e les Rhizopodes des rnarais du Sud-ouest d'uvira (Congo-belge). Hydrobiologia 10: 85- 127. van Oye, P. 1959. Faune Rhizopodique du plateau de Kundulungu (Congo Belge). Hydrobiologia 13: 239-286. Vucetich, M.C. 1973. Estudio de tecamebianos Argentinos, en especial 10s del Dominion Pampasico. Rev. Mus. La Plata Nueva Ser. Secc. Zool. 11: 287-332. Wang, C.C. 1974. Protozoa from Qomolangma Feng (Mount Jolmolungma) Area. In: The Scientific Reports of the Comprehensive Scientific Expendition of the Qomolangma

Feng Area (1966-1968), Biology and Alpine Physiology, Science Press, Beijing, pp. 145-172. Wang, C.C. 1977. Protozoa from some districts of Tibetan Plateau. Acta Zool. Sin. 23: 131-160. Wang, C.C. and D.S. Nie. 1933. Studies of Sarcodina of Nanking, I. Cont. Biol. Lab. Sci. Soc. China 9: 341-387. Xiwen, L. and D. Walker. 1986. The plant geography of Yunnan Province, Southwest China. J. Biogeogr. 13: 367-397. Yang, J., W.S. Feng, and Y.F. Shen. 2004. On new records of the genus Dzflugia from China (Sarcomastigophora, Rhizopoda, Lobosea, Arcellinida, Difflugiidae). Acta Zootax. Sin. 29: 239-247. Zhang, L. and X.M. Guo. 1986. A survey of zooplankton in Mahu. In: Studies In Qinghai-Xizang (Tibet) Plateau Special Issue of Hengduan Mountains Scientific Expedition (11). Science and Technology Press, Beijing, pp. 543-548. Zhang, R.Z. 1999. Zoogeography of China. Science Press, Beijing, 394 pp. Zhang, R.Z., D. Zheng, Q.Y. Yang, and Y.H. Liu. 1997. Physical Geography of Hengduanshan Mountains. Science Press, Beijing, 151 pp.

Received: 7 December 2004

Accepted: 4 March 2005