Int. J. Aquat. Biol. (2016) 4(2): 69-79; DOI: ISSN: 2322-5270; P-ISSN: 2383-0956 Journal homepage: www.ij-aquaticbiology.com © 2016 Iranian Society of Ichthyology Original Article Loktak Lake, Manipur, northeast India: a Ramsar site with rich rotifer (Rotifera: Eurotatoria) diversity and its meta-analysis Bhushan Kumar Sharma*,1Telsing Paongam Haokip, Sumita Sharma Freshwater Biology Laboratory, Department of Zoology, North-Eastern Hill University, Shillong-793 022, Meghalaya, India. Article history: Received 9 November 2015 Accepted 4 January 2016 Available online 2 5 April 2016 Keywords: Composition Distribution Interesting species New records Richness Wetland Abstract: A total of 162 species (S) of Rotifera belonging to 40 genera and 20 families examined from Loktak Lake, an important floodplain lake of northeast India (NEI) that is one of the richest assemblages of the taxon known from the Indian sub-region. It merits biodiversity value as ~40.0% and ~62.0% of species recorded from India and NEI, respectively. One species is new to India, 23 species are new to Manipur and 14 species are new to Loktak basin. Biogeographically interesting elements included three Australasian, five Oriental, ten palaeotropical and one cosmo-subtropical species. Lecanidae > Lepadellidae > Brachionidae > Trichocercidae collectively comprised 65.4% of S; Lecane > Lepadella > Trichocerca are diverse genera; and paucity of Brachionus spp. is distinct. Loktak Rotifera indicated importance of cosmopolitan, the littoral-periphytonic and small-sized species, and ‘tropical character’. ANOVA recorded significant variations of the rotifer richness amongst three sampling sites of Loktak during June 2010–May 2012 survey. The richness followed osscillaring monthly variations and indicated lack of significant influence of any individual abiotic parameter at all three stations. Introduction Segers et al. (1993) hypothesized tropical and subtropical floodplain lakes to be globally rich habitats for the rotifer diversity. Sharma and Sharma (2014a, 2014b) extended this hypothesis to the floodplains lakes (beels) of the Brahmaputra river basin of northeast India (NEI) with Deepor beel, a Ramsar site and an important wetland of this basin, as one of the globally interesting Rotifera habitat (Sharma and Sharma, 2015). Realizing biodiversity value of the floodplain lakes of NEI vis-a-vis the role of extensive sampling, we undertook meta-analysis of Rotifera diversity of Loktak Lake, a Ramsar site of India and another important floodplain lake of NEI, based on recent collections and our earlier reports (Sharma, 2007, 2009a). An inventory of 162 species recorded till date from this floodplain lake (pat) of Manipur is presented and interesting species are illustrated. The * Corresponding author: Bhushan Kumar Sharma DOI: http://dx.doi.org/10.7508/ijab.2016.02.001 E-mail address: profbksharma@gmail.com nature and composition of the rotifer diversity are discussed with remarks on richness, new records and species of global and regional distribution value. This study merits importance for ecosystem diversity, biogeography and for following meta- analysis of the rotifer diversity of this second well sampled freshwater ecosystem of India. Materials and Methods The present study is a part of limnological survey undertaken (June 2010 - May 2012) and collections on several occasions during 2013-2015 from Loktak Lake (93°46'–93º55'E, 24º25'–24º42'N; area: 286 km2; max. depth: 4.58 m, mean depth: 2.07 m) located in Bishnupur / Imphal districts of Manipur state (NEI). This wetland is characterized by floating mats of vegetation called “Phumdi” which are inhabited by an endangered brow-antlered deer (Rucervus eldi eldi). The common aquatic plants of 70 Sharma et al./ Rotifer diversity of Loktak Lake, Manipur, northeast India this Ramsar site included Ceratophyllum demersum, Eichhornia crassipes, Pistia stratiotes, Salvinia cucullata, Savinia natans, Euryale ferox, Hydrilla verticillata, Nymphoides cristatum, Trapa natans, Potamogeton crispus, Alternanthera philoxeroides, Rumex nepalensis, Ipomoea arguta, Polygonum pulchrum, Utricularia sp., Valllissnaria sp., Oryza rufipegon, Spiranthus sinesis and Nelumbo nucifera. Water samples and qualitative plankton were collected, at regular monthly intervals during June 2010 - May 2012 limnological survey, at three sampling sites namely Loktak A (93°45'56.3''E; 24°32'13.5''N; alt. 726 m ASL), Loktak B (93°47'58.1''E; 24°30'39.1''N; alt. 714 m ASL) and Loktak Barrage (93°45'43.5''E; 24°32'46.9''N; alt. 718 m ASL). In addition, qualitative plankton samples were collected from different parts of Loktak basin during the study period. Water temperature, specific conductivity and pH were recorded with field probes; dissolved oxygen was estimated by Winkler’s method; and total alkalinity, total hardness, calcium and chloride were analyzed following APHA (1992). The plankton samples were collected by towing a nylobolt plankton net (#50 μm) and preserved in 5% formalin; aquatic vegetation was disturbed before each sampling to facilitate collection of planktonic and semi-planktonic rotifers. Individual collections were screened with a Wild stereoscopic binocular microscope; the rotifers were isolated and mounted in Polyvinyl alcohol-lactophenol, and observed with Leica (DM 1000) stereoscopic phase contrast microscope fitted with an image analyzer. The measurements were given in micrometers (μm). The rotifer taxa were identified following Koste (1978), Segers (1995), Sharma (1983, 1998a), Sharma and Sharma (1997, 1999, 2000, 2008, 2013). The rotifer community similarities were calculated vide Sørensen’s index (Sørensen, 1948) and Two-way ANOVA was used to analyze the significance of temporal variations of richness. Ecological relationships between abiotic factors and the rotifer richness were determined by Pearson’s correlation coefficients (r); P values were calculated vide http://faculty.vassar.edu/ lowry/tabs.html and signi- ficance was ascertained after use of Bonferroni corrections. Results The variations (ranges, mean ±SD) of the recorded abiotic factors are presented in Table 1 in comparison with the report of Sharma (2009a). Water temperature at three sampling stations of Loktak Lake varied between 22.6±3.2 – 23.4±3.4°C, pH ranged between 6.8±0.2 – 7.0±0.3 and specific conductivity varied between 80.9±21.7 – 113.1±24.1 µS cm-1 while dissolved oxygen and free carbon dioxide fluctuated between 4.6±2.0 – 6.0±2.7 mg l-1 and 8.8±3.1 - 13.5±5.6 mg l-1, respectively. Total alkalinity, total hardness, calcium and chloride ranged between 65.2±16.5–72.8±24.7 mg l-1; 51.0± 12.8 - 73.1±17.9 mg l-1; 33.7±9.8 - 38.1±9.6 mg l-1 and 19.3±5.5 - 23.7±6.2 mg l-1 between three sampling stations of Loktak, respectively. We report a total of 162 species spread over 40 genera and 20 families from Loktak Lake (Appendix June 2010 – May 2012 (Present study) Sharma (2009a) Parameters↓ Stations→ Loktak A Loktak B Loktak Barrage Nov. 2002-Oct. 03 Mean ± SD Mean ± SD Mean ± SD Mean ± SD Water temperature (°C) 23.1±3.0 22.6±3.2 23.4±3.4 21.4 ± 4.0 pH 6.9±0.3 6.8±0.2 7.0±0.3 6.38 ± 0.23 Specific Conductivity (µS cm-1) 80.9±21.7 86.1±21.7 113.1±24.1 98.9 ± 19.7 Free Carbon dioxide (mg l -1) 10.6±4.3 13.5±5.6 8.8±3.1 9.5 ± 2.1 Dissolved oxygen (mg l-1) 5.2±1.6 4.6±2.0 6.0±2.7 6.2 ± 1.1 Total Alkalinity (mg l-1) 65.2±16.5 72.8±24.7 72.6±12.3 16.0 ± 4.4 Total Hardness (mg l-1) 51.0±12.8 59.2±18.4 73.1±17.9 38.1 ± 8.2 Calcium (mg l-1) 33.7±9.8 37.4±18.9 38.1±9.6 8.9 ± 3.0 Chloride (mg l-1) 23.7±6.2 23.4±7.8 19.3±5.5 14.9 ± 3.1 Table 1. Variations in certain abiotic factors of Loktak Lake. 71 Int. J. Aquat. Biol. (2016) 4(2): 69-79 1) and interesting taxa are illustrated; of these, 152 species are observed in 2010-2015 collections. Further, 135 species are observed at three sampling stations namely Loktak A, Loktak B and Loktak Barrage during June 2010 - May 2012 survey. The rotifer richness (Figs. 1, 2) ranged between 44-79 (60±8), 46-72 (58±7) and 39-67 (50±8) species at three stations, respectively and recorded 73.2-78.5% community similarities vide Sørensen’s index. Peak richness is observed during December 2011, November 2011 and January 2012 at Loktak A, Loktak B and Loktak Barrage, respectively. Mytilina lobata (Fig. 3a) is a new record to the Indian Rotifera. Brachionus caudatus, B. durgae, B. kostei (Fig. 3b), E. meneta, Keratella tecta, Lecane aspasia (Fig. 3c), L. bifurca, L. rhenana (Fig. 3d), L. rhytida (Fig. 3e), L. undulata, Lepadella minuta, L. quadricarinata (Fig. 3f), L. quinquecostata (Fig. 3g), Mytilina acanthophora (Fig. 3h), M. michelangellii (Fig. 3i), Trichocerca edmondsoni (Fig. 3j), T. hollaerti (Fig. 3k), T. maior (Fig. 3l), T. scipio, T. weberi (Fig. 3m), Sinantherina Figure 1. Species richness of Loktak Rotifera (2010-2011). Figure 2. Species richness of Loktak Rotifera (2011-2012). 72 Sharma et al./ Rotifer diversity of Loktak Lake, Manipur, northeast India semibullata, Testudinella amphora (Fig. 3n) and Wolga spinifera (Fig. 3o) are new records from Manipur. Brachionus bidentatus, Epiphanes brachionus, E. incisa, K. lenzi, Lecane aeganea, L. arcula, L. bulla diabolica, L. haliclysta, L. pusilla, L. thienemanni, Lepadella desmeti, L. triba, Platyias leloupi and Trochosphaera aequatorialis are new records from Loktak Lake. Euchlanis semicarinata, Figure 3. (a) Mytilina lobata Pourriot (lateral view), (b) Brachionus bennini Leissling (dorsal view), (c) Lecane aspasia Myers (dorsal view), (d) Lecane rhenana Hauer (dorsal view), (e) Lecene rhytida Harring & Myers (dorsal view), (f) Lepadella quadricarinata (Stenroos) (ventral view), (g) Lepadella quinquecostata (Lucks) (dorsal view), (h) Mytilina acanthophora Hauer (lateral view), (i) Mytilina michelangellii Reid & Turner (lateral view), (j) Trichocerca edmondsoni (Myers) (ventral view), (k) Trichocerca hollaerti De Smet (lateral view), (l) Trichocerca maior Hauer (lateral view), (m) Trichocerca weberi (Jennings) (lateral view); (n) Testudinella amphora Hauer (dorsal view), and (o) Wolga spinifera (Western) (dorsal view) . 73 Int. J. Aquat. Biol. (2016) 4(2): 69-79 Filinia brachiata, Floscularia ringens, Habrotrocha angusticollis, Lecane acanthinula and L. solfatara, Macrochaetus danneelae, Rotaria macroceros and R. tardigrada are not observed in 2010-2015 collections. Discussion Water temperature affirmed sub-tropical nature of Loktak Lake concurrent with its location. The slightly acidic-slightly alkaline, marginally hard and calcium-poor waters are characterized by low ionic concentration as indicated by specific conductivity at all three sampling stations; the latter warranted their inclusion under ‘Class I’ category of trophic classification vide Talling and Talling (1965). This study indicated well-oxygenated waters, low free CO2 and low chloride content. Our report differed in pH, alkalinity, hardness and marginal increase in calcium and chloride values than an earlier report (Sharma, 2009a) based on sampling at one station (Sendra). The present collections revealed a total of 152 rotifer species, including one species new to India, 23 species new to Manipur and 14 species new to Loktak basin. This is in contrast to an earlier report of 120 species from this wetland (Sharma, 2009a) with ~82% community similarity between two lists. Our report raised total richness (S) known to date from this Ramsar site to 162 species which, in turn, is one of the richest Rotifera assemblage known from any freshwater ecosystem of the Indian sub-region following the report of 171 species from Deepor beel (Sharma and Sharma, 2015) - another Ramsar site of India. The rotifer fauna of Loktak reflected 78.7% community similarity with the latter because of common occurrence of several cosmopolitan, cosmotropical and pantropical species. Total richness (S) merits biodiversity value as ~ 40.0% and ~ 62.0% of species of Rotifera known from India and NEI, respectively. The report of 40 genera and 20 families from Loktak Lake reflected rich higher diversity as compared with 50 genera and 23 families as well as 65 genera and 25 families of this phylum known from NEI (Sharma and Sharma, 2014a) and India (BKS, unpublished), respectively while it broadly concurred with 38 genera and 20 families known from Deepor Beel (Sharma and Sharma, 2015). The rich and diverse assemblage of Loktak Rotifera, affirming biodiversity value of this ‘hot- spot’, is hypothesized to habitat diversity and environmental heterogeneity of this Ramsar site. This generalization supported hypothesis of Segers et al. (1993) indicating (sub) tropical floodplains to be the world’s rotifer rich habitats and also concurred with reports from the floodplains of Argentina (Jose De Paggi, 1993, 2001), Brazil (Bonecker et al., 1998), Australia (Shiel et al., 1998), and the Brahmaputra river basin of India (Sharma and Sharma, 2014b). Mytilina lobata is a new addition to the Indian Rotifera. Known from Neotropical region (Segers, 2007), the present report extended its distributional limit to the Oriental region. Twenty-three species are new records from Manipur. Besides, this study extended distribution of fourteen species to Loktak Lake basin (Appendix 1). A total of nineteen globally interesting elements (~12.0% of S) now known from Loktak Lake is of biogeography value than twelve species listed by Sharma (2009a). These included the Australasian Brachionus kostei, Macrochaetus danneelae and Notommata spinata; five Oriental endemics, namely Filinia camasecla, Lecane acanthinula, Lecane bulla diabolica and L. niwati and L. solfatara; the palaeotropical Euchlanis semicarinata, Lecane lateralis, L. simonneae, L. unguitata, Lepadella bicornis, L. discoidea, L. vandenbrandei, Testudinella brevicaudata, Trichocerca abilioi and T. hollaerti are palaeotropical species; and Brachionus durgae is a cosmo (sub) tropical species. Of these, Brachionus kostei, Euchlanis semicarinata, Notommata spinata, Lecane niwati, L. solfatara Lepadella vandenbrandei, Notommata spinata, Testudinella brevicaudata, Trichocerca abilioi and T. hollaerti are interestingly restricted in their distribution in India exclusively to NEI. Ascomorpha ecaudis, Brachionus mirabilis, Filinia brachiata, F. saltator, Keratella lenzi, Lecane 74 Sharma et al./ Rotifer diversity of Loktak Lake, Manipur, northeast India aeganea, L. aspasia, L. bifurca, L. doryssa, L. elegans, L. haliclysta, L. rhenana, L. rhytida, L. simonneae, L. pusilla, L. tenuiseta, L. thienemanni, Lepadella benjamini, L. bicornis, L. costatoides, L. dactyliseta, L. desmeti, L. elongata, L. quadricarinata, L. quinquecostata, Lophocharis salpina, Mytilina acanthophora, M. bisulcata, M. michelangellii, Macrochaetus longipes, Platyias leloupi, Testudinella amphora, T. parva, T. emarginula, T. tridentata, Trichocerca bicristata, T. edmondsoni, T. insignis, T. flagellata, T. maior, T. scipio, T. tenuior, T. weberi, Tripleuchlanis plicata and Trochosphaera aequatorialis are examples of regional distributional importance in the Indian sub-continent. Of these, Testudinella amphora is a recently reported from India from Assam (Sharma et al. 2015); Lecane aeganea, Trichocerca hollaerti and T. maior are new additions to the Indian Rotifera from Mizoram state of NEI (Sharma and Sharma 2015). Our present tally of 152 species of Loktak Rotifera reflected its diverse nature than the reports of 111 species from the floodplains of Argentina (Jose De Paggi, 1993); 124 species (lake) and 136 species from Oguta and Iyi-Efi lakes, respectively of the Niger delta (Segers et al., 1993); 130 species from Lake Guarana, Brazil (Bonecker et al., 1994); 106 taxa from Thale-Noi Lake, a Ramsar site in Thailand (Segers and Pholpunthin, 1997); 104 species from Laguana Bufeos, Bolivia (Segers et al., 1998); and 114 taxa examined from the Rio Pilcomayo National park (a Ramsar site), Argentina (Jose De Paggi, 2001). The richness is higher than 67-103 species (Sharma, 2005), 69-93 species (Sharma and Sharma, 2008) and 60-100 species (Sharma et al., 2015) examined from various beels of the Brahmaputra river basin, Assam; and 62-73 species reported from 14 floodplain lakes (pats) of Manipur (Sharma, 2009b). We caution on over- emphasis on comparisons of richness as it was likely to be influenced by sampling intensity (Dumont and Segers, 1996). We attribute high richness of Rotifera of Loktak to the rotiferologist effect following Fontaneto et al. (2012) though our biodiversity update is also the result of the sampling intensity. Loktak Rotifera is characterized by diverse Lecanidae (47 species) > Lepadellidae (27 species) > Brachionidae (17 species) > Trichocercidae (15 species) which collectively formed 65.4% of S. It differed from the importance of Lecanidae > Lepadellidae > Trichocercidae > Brachionidae listed earlier from this Ramsar site (Sharma, 2009a) with distinct increase in the richness of the first two families in our recent collections. Nevertheless, marginal increase in the brachionid diversity deserved caution due to their yet restricted occurrence and of Brachionus spp. in particular. This feature differed from high Brachionidae richness (28 species) reported from Deepor beel (Sharma and Sharma, 2015). Notommatidae = Euchlanidae > Testudinellidae = Trochosphaeridae = Mytilinidae, other diverse families (~20.0% of S), deserved attention in Loktak Lake. The littoral-periphytonic Lecane (47 species) > Lepadella (22 species) > Trichocerca (17 species), together, included ~52.0% of S of Loktak Rotifera and thus supported hypothesis of Green (2003) on the possibility of assemblage rules for the periphytic community. Interestingly, their significance broadly concurred with the reports from the floodplains of Niger delta (Segers et al., 1993), Broa reservoir, Brazil (Segers and Dumont, 1995), River Nan, Thailand (Sanoamuang, 1998), Bolivia (Segers et al., 1998) and Okavango Delta of South Africa (Green, 2003); it also concurred with the report from Deepor beel (Sharma and Sharma, 2015). The rich diversity of ‘tropic-centered’ Lecane, more cosmopolitan species (~60% of S), and collective importance (~24% of S) of cosmotropical and pantropical species assigned ‘tropical character’ to Loktak Rotifera. This conclusion is concurrent with several tropical faunas (Green, 1972; Pejler, 1977; Fernando, 1980; Dussart et al., 1984; Segers, 1996, 2001) globally as well as from India (Sharma, 1996, 1998b, 2005; Sharma and Sharma, 2008; 2014a, 2014b, 2015). These remarks are supported by relative paucity of ‘temperate-centered’ Keratella in our collections. 75 Int. J. Aquat. Biol. (2016) 4(2): 69-79 High richness of the littoral-periphytonic and fewer euplanktonic species in Loktak are hypothesized to paucity of definite limnetic habitats (De Manuel, 1994). The occurrence of both non- planktonic species and planktonic taxa in the littoral weedy margin of this Ramsar site affirmed occupation of different niches as hypothesized by Bonecker et al. (1998). These features concurred with the reports from Deepor beel (Sharma and Sharma, 2013, 2015) and other floodplain lakes of the Brahmaputra river basin (Sharma and Sharma, 2014b). Loktak Rotifera is notable for large number of small-sized species particularly of Lecane, Lepadella, Trichocerca and Colurella. This feature is hypothesized to predation by juvenile fish and invertebrates (Baumgartner et al., 1997) but specific observations are desired to confirm predation effect. Our June 2010 - May 2012 limnological survey registered significant variations of the rotifer richness amongst three Loktak stations during the study period (F2, 46=11.473, P=9.07E-05) as well as during two successive years (F 2, 22=4.327, P=0.026; F2, 22=7.051, P=0.004) as against insignificant variat- ions reported earlier (Sharma, 2009a). Further, it recorded significant monthly variations at two sampling sites: Loktak B (F1, 11=3.536, P=0.023) and Loktak Barrage (F1, 11=6.618, P=0.002) and follow- ed osscillaring variations with maxima during December 2011, November 2011 and January 2012 at Loktak A, Loktak B and Loktak Barrage, respectively. This study indicated lack of significant influence of any individual abiotic factor on the richness in contrast to inverse correlation with rainfall, pH, hardness, nitrate, chloride and total dissolved solids and positive correlation with dissolved oxygen recorded vide Sharma (2009a). The differences suggested that the rotifers are generalists in terms of abiotic factors vs. their occurrence with factors associated with microhabitat being more important. To sum up, the specious and diverse Rotifer assemblage of Loktak Lake, with various new records and species of global and regional interest, imparted biodiversity and biogeography merit to our meta-analysis. With our bias towards inclusion of monogonont plankton and semi-plankton taxa, this study suggested scope for further up-date of the rotifer inventory of this ‘hot-spot’ based on specific sampling of periphytic, sessile, colonial and benthic communities. We estimate the report of 220+ species from this Ramsar site while analysis of the ‘rotifer- macrophytes associations’ merits biodiversity interest due to lack of such studies in India. Acknowledgements We thank the Head, Department of Zoology, North- Eastern Hill University, Shillong for laboratory facilities. The samples for this study were collected by TPH. The authors have no conflict of interests. References A.P.H.A. (1992). Standard methods for the examination of water and wastewater. 18th Ed., American Public Health Association, Washington D.C. 1198 p. Baumgartner G., Nakataki K., Cavicchiolo M., Baugartner M.S. (1997). Some aspects of the ecology of fish larvae in the floodplain of the high Parana river, Brazil. Review Brazilian Zoology, 14: 551-563. Bonecker C.C., Lansac-Tôha F.A., Staub A. (1994). Qualitative study of Rotifers in different environments of the high Parana River floodplain (Ms), Brazil. Revista UNIMAR, 16(1): 1-16. Bonecker C.C., Lansac-Tôha, F.A., Rossa D.C. (1998). Planktonic and non-planktonic rotifers in two environments of the upper Parana River floodplain, state of Mato Grosso do Sul, Brazil. Brazilian Archives of Biology and Technology, 41: 447-456. De Manuel J. (1994). Taxonomic and zoogeographic considerations on Lecanidae (Rotifera: Monogononta) of the Balearic archipelago, with description of a new species, Lecane margalefi n. sp. Hydrobiologia, 288: 97-105. Dumont H.J., Segers H. (1996). Estimating lacustrine zooplankton species richness and complementarity. Hydrobiologia, 341: 125-132. Dussart B.H., Fernando C.H., Matsumura-Tundisi J., Shiel R.J. (1984). A review of systematics, distribution and ecology of tropical freshwater zooplankton. Hydrobiologia, 113: 77-91. Fernando C.H. (1980). The freshwater zooplankton of Sri 76 Sharma et al./ Rotifer diversity of Loktak Lake, Manipur, northeast India Lanka, with a discussion of tropical freshwater zooplankton composition. Internationale Revue Hydrobiologie, 65: 411-426. Fontaneto D., Márcia Barbosa, A., Segers H., Pautasso M. (2012). The ‘rotiferologist’ effect and the other global correlates of species richness in rotifers. Ecography, 35: 174-182. Green J. (1972). Latitudinal variation in associations of planktonic Rotifera. Journal of Zoology, London, 167: 31-39. Green J. (2003). Associations of planktonic and periphytic rotifers in a tropical swamp, the Okavango Delta, Southern Africa. Hydrobiologia, 490: 197-209. José De Paggi S. (1993). Composition and seasonality of planktonic rotifers in limnetic and littoral region of a floodplain lake (Parana River System). Revue Hydrobiologie Tropicale, 26: 53-64. José De Paggi S. (2001). Diversity of Rotifera (Monogononta) in wetlands of Rio Pilcomayo national park, Ramsar site (Formosa, Argentina). Hydrobiologia, 462: 25-34. Koste W. (1978). Rotatoria. Die Rädertiere Mitteleuropas, begründet von Max Voigt. Überordnung Monogononta. Gebrüder Borntaeger, Berlin, Stuttgart. I. 673 p. Pejler B. (1977). On the global distribution of the family Brachionidae (Rotatoria). Archiv für Hydrobiology, supplement, 53: 255-306. Sanoamuang L. (1998). Rotifera of some freshwater habitats in the floodplains of the River Nan, northern Thailand. Hydrobiologia, 387/388: 27-33. Segers H. (1995). Rotifera 2: Lecanidae. 6. In: H.J. Dumont, T. Nogrady (Eds.). Guides to identification of the Microinvertebrates .of the Continental waters of the world. SPB Academic Publishing bv. Amsterdam, the Netherlands. pp: 1-226. Segers H. (1996). The biogeography of littoral Lecane Rotifera. Hydrobiologia, 323: 169-197. Segers H. (2001). Zoogeography of the Southeast Asian Rotifera. Hydrobiologia, 446/447: 233-246. Segers H. (2007). Annotated checklist of the rotifers (Phylum Rotifera), with notes on nomenclature, taxonomy and distribution. Zootaxa, 1564: 1-104. Segers H., Dumont H.J. (1995). 102+ rotifer species (Rotifera: Monogononta) in Broa reservoir (SP., Brazil) on 26 August 1994, with the description of three new species. Hydrobiologia, 316: 183-197. Segers H., Ferrufino N.L., De Meester L. (1998). Diversity and Zoogeography of Rotifera (Monogononta) in a flood plain lake of the Ichilo River, Bolivia, with notes on little known species. Internationale Revue Hydrobiologie, 83(5-6): 439- 448. Segers H., Nwadiaro C.S., Dumont H.J. (1993). Rotifera of some lakes in the floodplain of the river Niger (Imo State, Nigeria). II. Faunal composition and diversity. Hydrobiologia, 250: 63-71. Segers H., Pholpunthin P. (1997). New and rare Rotifera from Thale Noi Lake, Pattalang Province, Thailand, with a note on the taxonomy of Cephalodella (Notommatidae). Annals Limnologie, 33(1): 13-21. Sharma B.K. (1983). The Indian species of the genus Brachionus (Eurotatoria: Monogononta: Brachionid- ae). Hydrobiologia, 104: 31-39. Sharma B.K. (1996). Biodiversity of freshwater Rotifera in India - a status report. Proceedings of the Zoological Society, Calcutta 49: 73-85. Sharma B.K. (1998a). Freshwater Rotifers (Rotifera: Eurotatoria). In: Fauna of West Bengal. State Fauna Series, 3(11): 341-461. Sharma B.K. (1998b). Faunal Diversity in India: Rotifera. In: J.R.B. Alfred, A.K. Das, A.K. Sanyal (eds.). Faunal diversity of India. ENVIS Centre, Zoological Survey of India, Calcutta, pp. 57-70. Sharma B.K. (2005). Rotifer communities of floodplain lakes of the Brahmaputra basin of lower Assam (N.E. India): biodiversity, distribution and ecology. Hydrobiologia, 533: 209-221. Sharma B.K. (2007). Notes on rare and interesting Rotifers (Rotifera: Eurotatoria) from Loktak lake, Manipur – A Ramsar site. Zoo’s Print, 22(9): 2816– 2820. Sharma B.K. (2009a). Diversity of Rotifers (Rotifera: Eurotatoria) of Loktak lake, North- Eastern India. Tropical Ecology, 50(2): 277-285. Sharma B.K. (2009b). Rotifer communities of floodplain lakes of Manipur (North-East India): biodiversity, distribution and ecology. Journal of Bombay Natural History Society, 106(1): 45-56. Sharma B.K., Sharma S. (1997). Lecanid rotifers (Rotifera: Monogononta: Lecanidae) from North Eastern India. Hydrobiologia, 356: 159-163. Sharma B.K., Sharma S. (1999). Freshwater Rotifers (Rotifera: Eurotatoria). In: Fauna of Meghalaya. State Fauna Series, 4(9): 11-161. Sharma B.K., Sharma S. (2000). Freshwater Rotifers 77 Int. J. Aquat. Biol. (2016) 4(2): 69-79 (Rotifera: Eurotatoria). In: Fauna of Tripura: State Fauna Series. Zoological Survey of India, Calcutta, 7 (4): 163-224. Sharma B.K., Sharma S. (2014a). Northeast India - An important region with a rich biodiversity of Rotifera. In: B.K. Sharma, H.J. Dumont, R.L. Wallace (Ed.). Rotifera XIII: Rotifer Biology- A structural and functional Approach. International Review of Hydrobiology, 99(1-2): 20-37. Sharma B.K., Sharma S. (2014b). Floodplains of the Brahmaputra river basin-globally interesting ecotones with rich Rotifer (Rotifera: Eurotatoria) biodiversity. In: R.K. Sinha, B. Ahmed (Eds.). Rivers for Life - Proceedings of the International Symposium on River Biodiversity: Ganges–Brahmaputra-Meghna River System, Ecosystems for Life, A Bangladesh-India Initiative, IUCN, International Union for Conservation of Nature, pp. 258-270. Sharma B.K., Sharma S. (2015). Biodiversity of freshwater rotifers (Rotifera: Eurotatoria) of Mizoram, Northeast India: composition, new records and interesting features. International Journal of Aquatic Biology, 3(5): 301-313. Sharma B.K., Sharma S., Hatimuria M.K. (2015). Rotifer assemblages (Rotifera: Eurotatoria) of the floodplain lakes of Majuli River Island, the Brahmaputra river basin, northeast India. International Journal of Aquatic Biology, 3(1):1-13. Sharma S., Sharma B.K. (2008). Zooplankton diversity in floodplain lakes of Assam. Records of the Zoological Survey of India, Occasional Paper No. 290: 1-307. Sharma S., Sharma B.K. (2013). Faunal Diversity of Aquatic Invertebrates of Deepor Beel (a Ramsar site), Assam, northeast India. Wetland Ecosystem Series, 17: 1-226. Shiel R.J., Green, J.D., Nielsen, D.L. (1998). Floodplain biodiversity: why are there so many species? Hydrobiologia, 387/388: 39-46. Sørensen T. (1948). A method of establishing group of equal amplitude in plant sociology based on similarity of species content and its application to analyze the vegetation of Danish commons. Biologiske Skrifter, 5: 1-34. Talling J.F., Talling I.B. (1965). The chemical composition of African lake waters. Internationale Revue der Gesamten Hydrobiologie, 50: 421-463. 78 Sharma et al./ Rotifer diversity of Loktak Lake, Manipur, northeast India Class: Monogononta Order: Ploimida Family: Brachionidae 1. Anuraeopsis fissa Gosse, 1851 2. Brachionus angularis Gosse, 1851 3. B. bidentatus Anderson, 1889 *** 4. B. calyciflorus Pallas, 1766 5. B. caudatus Barrois & Daday, 1894 ** 6. B. falcatus Zacharias, 1898 7. B. durgae Dhanapathi, 1974 ** 8. B. kostei Shiel, 1983 ** 9. B. mirabilis Daday, 1897 10. B. quadridentatus Hermann, 1783 11. Keratella cochlearis (Gosse, 1851) 12. K. lenzi Hauer, 1953 *** 13. K. tropica (Apstein, 1907) 14. K. tecta (Gosse, 1851) ** 15. Platyias leloupi (Gillard, 1967) *** 16. P. quadricornis (Ehrenberg, 1832) 17. Plationus patulus (O.F. Müller, 1786) Family: Epiphanidae 18. Epiphanes brachionus (Ehrenberg, 1837) *** Family: Euchlanidae 19. Beauchampiella eudactylota (Gosse, 1886) 20. Dipleuchlanis propatula (Gosse, 1886) 21. Euchlanis dilatata Ehrenberg, 1832 22. E. incisa Carlin, 1939 *** 23. E. meneta Myers, 1930 ** 24. E. semicarinata Segers, 1993 # 25. E. triquetra Ehrenberg, 1838 26. Tripleuchlanis plicata (Levande, 1894) Family: Mytilinidae 27. Lophocharis salpina (Ehrenberg, 1834) 28. Mytilina acanthophora Hauer, 1938 ** 29. M. bisulcata (Lucks, 1912) 30. M. lobata Pourriot, 1996 * 31. M. michelangellii Reid & Turner, 1988 ** 32. M. ventralis (Ehrenberg, 1830) Family: Trichotriidae 33. Macrochaetus danneelae Koste & Shiel, 1983 # 34. M. longipes Myers, 1934) 35. M. sericus (Thorpe, 1893) 36. Trichotria tetractis (Ehrenberg, 1830) 37. Wolga spinifera (Western, 1894) ** Family: Lepadellidae 38. Colurella adriatica (Ehrenberg, 1837) 39. C. obtusa (Gosse, 1886) 40. C. sulcata (Stenroos, 1898) 41. C. uncinata (O. F. Müller, 1773) 42. Lepadella acuminata (Ehrenberg, 1834) 43. L. apsicora Myers, 1934 44. L. apsida Harring, 1916 45. L. bicornis Vasisht & Battish, 1971 46. L. benjamini Harring, 1916 47. L. costatoides Segers, 1992 48. L. dactyliseta (Stenroos, 1898) 49. L. desmeti Segers & Chittapun, 2001 *** 50. L. discoidea Segers, 1993 51. L. eurysterna Myers, 1942 52. L. ehrenbergi (Perty, 1850) 53. L. heterostyla (Murray, 1913) 54. L. lindaui Koste, 1981 55. L. minuta (Weber & Montet, 1918) ** 56. L. ovalis (O.F. Müller, 1786) 57. L. patella (O.F. Müller, 1773) 58. L. quadricarinata (Stenroos, 1898) ** 59. L. quinquecostata (Lucks, 1912) ** 60. L. rhomboides (Gosse, 1886) 61. L. triba Myers, 1934 *** 62. L. triptera Ehrenberg, 1832 63. L. vandenbrandei Gillard, 1952 64. Squatinella mutica (Ehrenberg, 1832) Family: Lecanidae 65. Lecane acanthinula (Hauer, 1938) # 66. L. aculeata (Jakubski, 1912) 67. L. aeganea Harring, 1914 *** 68. L. arcula Harring, 1914 *** 69. L. aspasia Myers, 1917 ** 70. L. bifurca (Bryce, 1892) ** 71. L blachei Berzins, 1973 72. L. bulla (Gosse, 1851) L. bulla diabolica (Hauer, 1936) *** 73. L. closterocerca (Schmarda, 1859) 74. L. crepida Harring, 1914 75. L. curvicornis (Murray, 1913) 76. L. decipiens (Murray, 1913) 77. L. doryssa Harring, 1914 78. L. elegans Harring, 1914 79. L. flexilis (Gosse, 1886) 80. L. furcata (Murray, 1913) 81. L. haliclysta Harring & Myers, 1926 *** 82. L. hamata (Stokes, 1896) 83. L. hornemanni (Ehrenberg, 1834) 84. L. inermis (Bryce, 1892) 85. L. inopinata Harring & Myers, 1926 86. L. lateralis Sharma, 1978 87. L. leontina (Turner, 1892) 88. L. ludwigii (Eckstein, 1883) 89. L. luna (O.F. Müller, 1776) 90. L. lunaris (Ehrenberg, 1832) 91. L. monostyla (Daday, 1897) Appendix 1: Systematic list of Rotifera recorded from Loktak Phylum: Rotifera Super-class: Eurotatoria 79 Int. J. Aquat. Biol. (2016) 4(2): 69-79 92. L. nitida (Murray, 1913) 93. L. niwati Segers, Kothetip & Sanoamuang, 2004 94. L. obtusa (Murray, 1913) 95. L. ohioensis (Herrick, 1885) 96. L. papuana (Murray, 1913) 97. L. ploenensis (Voigt, 1902) 98. L. pusilla Harring, 1914 *** 99. L. quadridentata (Ehrenberg, 1830) 100. L. rhenana Hauer, 1929 ** 101. L. rhytida Harring & Myers, 1926 ** 102. L. ruttneri Hauer, 1938 103. L. signifera (Jennings, 1896) 104. L. simonneae Segers, 1993 105. L. solfatara (Hauer, 1938) # 106. L. stenroosi (Meissner, 1908) 107. L. tenuiseta Harring, 1914 108. L. thienemanni (Hauer, 1938) *** 109. L. undulata Hauer, 1938 ** 110. L. unguitata (Fadeev, 1925) 111. L. ungulata (Gosse, 1887) Family: Notommatidae 112. Cephalodella forficula (Ehrenberg, 1830) 113. C. gibba (Ehrenberg, 1830) 114. C. mucronata Myers, 1924 115. Monommata longiseta (O.F. Müller, 1786) 116. M. maculata Harring & Myers, 1930 117. Monommata sp. 118. Notommata spinata Koste & Shiel, 1991 Family: Scaridiidae 119. Scaridium longicaudum (O.F. Müller, 1786) Family: Gastropodidae 120. Ascomorpha ecaudis Perty, 1850 Family: Trichocercidae 121. Trichocerca abilioi Segers & Sarma, 1993 # 122. T. bicristata (Gosse, 1887) 123. T. cylindrica (Imhof, 1891) 124. T. edmondsoni (Myers, 1936) ** 125. T. elongata (Gosse, 1886) 126. T. flagellata Hauer, 1938 127. T. hollaerti De Smet, 1990 ** 128. T. insignis (Herrick, 1885) 129. T. longiseta (Schrank, 1802) 130. T. maior Hauer, 1936 ** 131. T. rattus (O.F. Müller, 1776) 132. T. scipio (Gosse, 1886) ** 133. T. similis (Wierzejski, 1893) 134. T. tenuior (Gosse, 1886) 135. T. weberi (Jennings, 1903) ** Family: Asplanchnidae 136. Asplanchna priodonta Gosse, 1850 Family: Synchaetidae 136. Ploesoma lenticulare Herrick, 1855 137. Polyarthra vulgaris Carlin, 1943 138. Synchaeta pectinata Ehrenberg, 1832 Family: Dicranophoridae 139. Dicranophoroides caudatus (Ehrenberg, 1834) 140. Dicranophorus forcipatus (O.F. Müller, 1786) Order: Flosculariaceae Family: Flosculariidae 141. Floscularia ringens (Linnaeus, 1758) # 142. S. semibullata (Thorpe, 1893) ** 143. Sinantherina spinosa (Thorpe, 1893) 144. S. socialis (Linnaeus, 1758) Family: Conochilidae 145. Conochilus unicornis Rousselet, 1892 Family: Trochosphaeridae 146. Filinia brachiata (Rousselet, 1901) # 147. F. camasecla Myers, 1938 148. F. longiseta (Ehrenberg, 1834) 149. F. opoliensis (Zacharias, 1898) 150. F. saltator (Gosse, 1886) 151. Trochosphaera aequatorialis Semper, 1872 *** Family: Testudinellidae 152. Testudinella amphora Hauer, 1938 ** 153. T. brevicaudata Yamamoto, 1951 154. T. emarginula (Stenroos, 1898) 155. T. parva (Ternetz, 1892) 156. T. patina (Hermann, 1783) 157. T. tridentata Smirnov, 1931 Class: Bdelloidea Order: Philodinida Family: Philodinidae 158. Philodina citrina Ehrenberg, 1832 159. Rotaria macroceros (Gosse, 1851) # 160. R. neptunia (Ehrenberg, 1830) 161. R. tardigrada (Ehrenberg, 1832) # Family: Habrotrochidae 162. Habrotrocha angusticollis (Murray, 1905) # * New record from India; **New record from Manipur state; *** New record from Loktak; # not observed in present collections