Tropical Ecology 52(3): 293-302, 2011 © International Society for Tropical Ecology www.tropecol.com ISSN 0564-3295 Zooplankton communities of Deepor Beel (a Ramsar site), Assam (N. E. India): ecology, richness, and abundance * B. K. SHARMA Department of Zoology, North Eastern Hill University, Permanent Campus, Umshing, Shillong 793022, Meghalaya, India Abstract: Limnological studies undertaken at two sampling stations of Deepor Beel (91º 35′ - 91º 43′ E, 26º 05′ - 260 11′ N) revealed the presence of 171 and 160 species of zooplankton at stations I and II, respectively. Species richness in different months ranged between 96 ± 11 and 97 ± 13 (mean ± SD) and community similarities across months between 48.9 - 88.1 % and 53.1 89.7 % at stations I and II, respectively. Richness, which was mainly influenced by Rotifera (110 species) and Cladocera (45 species), showed significant variations between months but not between stations. Zooplankton (475 ± 114, 459 ± 128 n l-1 at stations I and II, respectively) formed an important quantitative component of the net plankton. Rotifera and Cladocera dominated quantitatively, Copepoda and Rhizopoda formed sub-dominant groups, and Ostracoda and Conchostraca had very low densities. Zooplankton abundance varied significantly between both months and stations. At both stations, richness and abundance inversely correlated with water temperature and rainfall, and positively with specific conductivity and dissolved oxygen. While at station II both richness and abundance also positively correlated with transparency, alkalinity, and hardness, at station I, abundance positively correlated with free CO2. Finally, zooplankton richness and abundance oscillated with annual frequency but showed winter peaks, was not quantitatively dominated by any individual species, and was characterized by higher species diversity with equitable abundance of various species. In view of the paucity of works from the floodplain lakes of India, this study provides important information on zooplankton diversity and ecology of the sampled Ramsar site. Resumen: Estudios limnológicos realizados en dos estaciones de muestreo de Deepor Beel (91º 35′ - 91º 43′ E, 26º 05′ - 26º 11′ N) revelaron la presencia de 171 y 160 especies de zooplancton en las estaciones I y II, respectivamente. La riqueza de especies en diferentes meses varió entre 96 ± 11 y 97 ± 13 (media ± SD), y las similitudes a nivel de comunidad entre meses fluctuaron entre 48.9 - 88.1 % y 53.1 - 89.7 % en las Estaciones I y II, respectivamente. La riqueza, influenciada principalmente por Rotifera (110 especies) y Cladocera (45 especies), mostró variaciones significativas entre meses pero no entre estaciones. El zooplancton (475 ± 114, 459 ± 128 n l -1 en las Estaciones I y II, respectivamente) conformó un componente cuantitativamente importante del plancton. Rotifera y Cladocera dominaron cuantita-tivamente, Copepoda y Rhizopoda formaron grupos subdominantes, y Ostracoda y Conchostraca tuvieron densidades muy bajas. La abundancia del zooplancton varió significativamente tanto entre meses como entre estaciones. En ambas estaciones, la riqueza y la abundancia se correlacionaron inversamente con la temperatura del agua y la precipitación, y positivamente con la conductividad específica y el oxígeno disuelto. Mientras que en la Estación Ii tanto la riqueza como la abundancia también estuvieron correlacionados positivamente con la trans-parencia, la alcalinidad y la dureza, en la Estación I la abundancia se correlacionó positivamente con el CO2 libre. Finalmente, la riqueza y la abundancia del zooplancton oscilaron con una frecuencia anual pero mostraron picos de invierno, no estuvieron dominadas cuantitativamente por ninguna especie particular, y N * Corresponding Author; e-mail: bksharma@nehu.ac.in, profbksharma@gmail.com 294 ZOOPLANKTON COMMUNITIES OF DEEPOR BEEL estuvieron caracterizadas por una diversidad de especies más alta con una abundancia equitativa de varias especies. En virtud de la pobreza de trabajos de los lagos de planicie de inundación en la India, este estudio brinda información importante sobre la diversidad de zooplancton y la ecología del sitio Ramsar muestreado. Resumo: Os estudos limnológicos levados a efeito em duas estações amostra em Deepor Beel (91º 35’ - 91º 43’ E, 26º 05’ - 26º 11’ N) revelou a presença de 171 e 160 espécies de zooplâncton nas Estações I e II, respectivamente. A riqueza específica nos diferentes meses oscilou entre 96 ± 11 e 97 ± 13 (média ± DP) e as semelhanças de comunidade ao longo dos meses situou-se entre os 48,9 - 88,1 % e os 53,1 - 89,7 % nas Estações I e II, respectivamente. A riqueza, que era principalmente influenciada pela Rotifera (110 espécies) e a Cladocera (45 espécies), mostrou variações significativas entre meses mas não entre Estações. O zooplâncton (475 ± 114,459, 459 ± 128 n ℓ-1 nas estações I e II, respectivamente) forma uma componente quantitativa importante do plâncton líquido. Os Rotifera e Cladocera dominavam quantitativamente, os Copepoda e Rhizopoda formam os subgrupos dominantes, apresentando os Ostracoda e os Conchostraca densidades muito baixas. A abundância do zooplâncton variou significativamente quer entre meses e quer entre estações. Em ambas as Estações, a riqueza e abundância estavam inversamente correlacionadas com a temperatura e a queda pluviométrica, e positivamente com a condutividade específica e o oxigénio dissolvido. Enquanto na Estação II, quer a riqueza, quer a abundância estavam também positivamente correlacionadas com a transparência, alcalinidade e dureza, já na Estação I, a abundância estava positivamente correlacionada com o CO2 livre. Finalmente, a riqueza e abundância do zooplâncton oscilaram com uma frequência anual mas mostraram picos no inverno que não estavam quantitativamente dominados por nenhuma espécie individual, e estava caracterizado por elevada diversidade de espécies com abundância equivalente das várias espécies. Com vista à insuficiência de trabalhos nos lagos de encharcamento da Índia, este estudo proporciona informação importante sobre a diversidade do zooplâncton e ecologia do sítio Ramsar amostrado. Key words: Abundance, Ramsar site, richness, synecology, zooplankton. Introduction Freshwater ecosystems are colonized by a diverse array of aquatic organisms. Amongst these, zooplankton, which function as primary consumers, comprise an integral component of aquatic food-webs and contribute significantly to biological productivity. Inspite of several studies from fresh-water environs of India, there is still little information on the ecology and role of zooplankton in the aquatic productivity of the Indian floodplain lakes (Sharma & Sharma 2008). Studies of zooplankton ecology from the floodplains of North-eastern India have so far been restricted to the reports of Sharma & Hussain (2001) and Sharma & Sharma (2008); in addition, Sharma & Sharma (2001, 2005) and Sharma (2005, 2009a, 2009b) dealt with Rotifera diversity. Thus, in view of the paucity of works from India, the present study of zooplankton communities of Deepor Beel, a Ramsar Site and an important floodplain lake of the Brahmaputra river basin of Assam, assumes special importance. This report presents information on zooplankton richness, community similarities and abundance, and the constituent zooplankton groups at two sampling stations. Analysis includes the temporal variation of species diversity, dominant groups, and the influence of seventeen abiotic parameters on zooplankton richness and abundance. Materials and methods Limnological investigations were undertaken during November 2002 - October 2003 in Deepor Beel (longitude: 91º 35′ - 91º 43′ E, latitude: 26º 05′ SHARMA 295 Fig. 1. A-Map of India indicating location of Assam state and map of Assam indicating location of Deepor beel; B- Map of Deepor beel indicating the sampling stations I and II. Table 1. Abiotic factors studied at Deepor Beel (Nov 02 - Oct 03) (Mean ± SD). Factors Rainfall (mm) Water temperature (0C) pH Transparency (cm) Specific Conductivity (µS cm-1) Dissolved oxygen (mg l-1) Free CO2 (mg l-1) Alkalinity (mg l-1) Hardness (mg l-1) Chloride (mg l-1) Phosphate (mg l-1) Sulphate (mg l-1) Nitrate (mg l-1) Silicate (mg l-1) B.O.D5 (mg l-1) Dissolved organic matter (mg l-1) Station I 204.5 ± 160.4 27.2 ± 4.6 6.89 ± 0.18 51.9 ± 26.2 99.2 ± 13.2 Station II 204.5 ± 160.4 27.4 ± 5.1 6.93 ± 0.21 52.7 ± 25.3 96.8 ± 15.5 6.7 ± 1.6 7.2 ± 2.1 66.3 ± 12.1 62.1 ± 9.9 34.6 ± 5.2 0.18 ± 0.07 10.2 ± 3.2 0.72 ± 0.12 3.02± 1.02 3.11 ± 0.59 3.84 ± 0.80 7.0 ± 1.1 6.8 ± 1.9 68.9 ± 10.3 61.2 ± 12.3 35.1 ± 5.0 0.19 ± 0.10 9.9 ± 3.4 0.74 ± 0.14 3.10 ± 1.27 3.21 ± 0.46 3.90 ± 0.64 Total dissolved solids (mg l-1) 2.37 ± 0.29 2.57 ± 0.30 - 26º 11′ N; area: 40 km2; altitude: 42 m above sea level) in the Kamrup district of lower Assam, N. E. India (Fig. 1 A & B). Various aquatic macrophytes, namely Hydrilla verticellata, Eichhornia crassipes, Vallisnaria spiralis, Utricularia flexuosa, Trapa bispinosa, Euryale ferox, Najas indica, Monochoria hastaefolia, Ipomoea fistulosa, Hygroryza aristata, Polygonum hydropiper, and Limnophila sp., cover this floodplain lake. Due to local constraints, the observations for this study were limited (November 2002 - October 2003) to two sampling stations (I and II). The sampled sites were characterized by common occurrence of H. verticellata, E. crassipes, U. flexuosa, T. bispinosa, E. ferox, N. indica, and P. hydropiper. Station II, however, differed from the former in occasional (temporal) lack of the rest of the observed macrophyte species, namely V. spiralis, M. hastaefolia, I. fistulosa, H. aristata, and Limnophila sp.. Though there is no official monitoring of water table of this floodplain lake, general minimum and maximum water levels were observed in Deepor beel during April 2003 and August 296 ZOOPLANKTON COMMUNITIES OF DEEPOR BEEL Table 2. Temporal variation of zooplankton at Deepor Beel (Nov 02 - Oct 03) (Range, Mean ± SD). Qualitative Zooplankton Total richness Monthly richness Community similarity (%) Station I Station II 171 species 68 - 112 96 ± 11 48.9 - 88.1 160 species 68 - 113 97 ± 13 53.1 - 89.7 Rotifera Total richness Monthly richness 110 species 43 - 65 56 ± 6 100 species 38 - 60 52 ± 7 Cladocera Total richness Monthly richness 45 species 17 - 41 29 ± 6 43 species 20 - 41 32 ± 6 Quantitative Net Plankton (n l-1) 708 - 961 Zooplankton (n l-1) Percentage Species Diversity 239 - 657 33.2 - 68.4 3.548 - 4.229 812 ± 80 696 - 1058 475 ± 114 57.9 ± 9.0 3.991 ± 0.181 801 ± 123 255 - 687 35.4 - 66.9 3.529 - 4.219 459 ± 128 56.5 ± 8.8 3.973 ± 0.193 Different Groups Rotifera (n l-1) Percentage 105 - 318 231 ± 60 42.8 - 65.2 48.7 ± 6.1 106 - 325 198 ± 70 37.9 - 49.6 42.5 ± 4.1 Cladocera (n l-1 ) Percentage 43 - 252 142 ± 59 14.6 - 38.3 28.7 ± 7.0 56 - 233 142 ± 48 22.0 - 37.6 30.6 ± 4.9 Copepoda (n l-1) Percentage 49 - 95 66 ± 17 7.6 - 31.4 15.1 ± 6.5 66 - 101 81 ± 13 11.9 - 29.-0 18.7 ± 4.9 Rhizopoda (n l-1) 7 - 41 29 ± 14 5 - 66 35 ± 18 Ostracoda (n l-1) Conchostraca (n l-1) 2 - 10 6±3 0-4 2003, respectively; the water levels were identical at both the stations. Further, the two sampling stations experienced similar inputs of rainwater and floods. Water samples were collected monthly from the selected sampling sites and were analyzed for the following abiotic factors: water temperature, rainfall, pH, transparency, specific conductivity, dissolved oxygen, free CO2, alkalinity, hardness, chloride, phosphate, sulphate, nitrate, silicate, BOD5, dissolved organic matter, and total dissolved solids. Water temperature, specific conductivity, and pH were recorded through field probes, transparency was noted with a Secchi disc, dissolved oxygen was estimated by Winkler’s method, and the other parameters were analyzed following APHA (1992). Three qualitative (by towing) and quantitative plankton (by filtering 25 l water each) samples were collected monthly at each station using a nylobolt plankton net (No. 25). Zooplankton samples were preserved in 5 % formalin, screened, and then identified following Koste (1978), Michael & Sharma (1988), Sharma (1998) and Sharma & Sharma (1999a, 1999b, 2000, 2008). 2 - 10 5±2 0-2 Quantitative samples were analyzed for abundance of net plankton (phyto- and zooplankton), zooplankton, and zooplankton constituent groups. Community similarity (Sorensen’s index) and species diversity (Shannon’s index) were calculated following Ludwig & Reynolds (1988) and Magurran (1988). ANOVA was used to analyse the significance of temporal variation of the biotic communities. Simple correlation coefficients (r1 and r2, respectively for stations I and II) were calculated between all abiotic and biotic parameters. Results and discussion Water samples analyzed from Deepor Beel are characterized by low ionic concentrations (Table 1) and, thus, warrant the inclusion of this Ramsar site under the ‘Class I’ category of trophic classification following Talling & Talling (1965). Mean water temperature confirms the tropical range concurrent with the lake’s geographical location. The nearly neutral and marginally hard waters of this floodplain lake show moderate values for dissolved oxygen, low free CO2, and low concen- SHARMA Fig. 2. Monthly variations in zooplankton richness of Deepor Beel (Nov 02 - Oct 03). tration of micronutrients. In general, the ranges of most abiotic factors show insignificant differences at the two stations and broadly agree with earlier reports from other floodplain lakes of Assam (Sharma & Sharma 2001, 2008; Sharma 2005). Plankton samples from Deepor Beel show (Table 2) the existence of a speciose and diverse zooplankton biocoenosis (171 species) and, thus, reflect the overall environmental heterogeneity and habitat diversity of this Ramsar site. Total zooplankton richness (171 species), the second highest known from any floodplain lake or individual aquatic ecosystem in India, follows that of 212 species for Loktak Lake (Sharma 2009a & b) another important Ramsar site and floodplain lake located in N. E. India. Richness in Deepor Beel is, however, distinctly more than that reported from several other floodplain lakes of India: 51 species from Trigamasar and Naranbagh lakes (Khan 1987), and 26 species from Mirgund Wetland (Yousuf et al. 1986) of Kashmir; 19 (Baruah et al. 1993) and 31 species (Sanjer & Sharma 1995) from Kawar Lake, Bihar; 49 species from Samuajan Beel, Upper Assam (Sharma & Hussain 2001); and 71 species from Beri Gopalpur and Sosadanga, West Bengal (Khan 2003). Though Deepor Beel is in fact more species rich, the differences between this study and other studies may result also from incomplete species inventories, inadequate sampling, and overlooking identification of smaller species. Nevertheless, a comparison with the zooplankton species richness reported by the author in Sharma & Sharma (2008) (which used similar 297 methodology) from 15 other floodplain lakes of the Brahmaputra river basin of Assam (102 - 156 species) underlines the relatively high species diversity of Deepor Beel. As in the findings of Sharma & Sharma (2008) and Sharma (2009a), zooplankton forms the dominant qualitative component (phytoplankton + zooplankton = 230 species) of the net plankton in Deepor Beel. These results, in turn, differ from the higher phytoplankton richness reported in certain other floodplain lakes from Bihar (Baruah et al. 1993; Sinha et al. 1994) and Assam (Sharma & Hussain 2001). Rotifera (110 species) and Cladocera (45 species) contribute the most to zooplankton richness. The micro-faunal diversity of these two groups in Deepor Beel, as well as their nature and composition, were discussed in Sharma & Sharma (2005, 2008). Zooplankton richness in Deepor Beel shows little annual variation between station I (171 species) and station II (160 species). Comparisons between sampling stations indicate broadly similar monthly ranges, mean values, and standard deviations of zooplankton richness (68 - 112, 96 ± 11 species; 68 - 115, 97 ± 13 species). On the other hand, significant monthly variation (F11, 23 = 23.966, P < 0.005) occurred in zooplankton composition. This statement holds valid for the present stations and may not reflect the general environmental heterogeneity of Deepor Beel unless ascertained by analysis of collections from other parts of this Ramsar site. Richness (Fig. 2) oscillates with annual frequency with winter peaks in February (station I) or December (station II) and minima in summer (April). While the peaks concur with luxuriant winter growth of aquatic macrophytes, the latter coincides with the lowest water level. Lack of information on the seasonal variation in zooplankton richness in other Indian floodplain lakes prohibits comparison of this study with other areas, but winter peaks concur with the author’s observations in Loktak Lake (Sharma unpublished). Rotifera species (56 ± 6, 52 ± 7 species) form the main qualitative component of zooplankton at both stations and exert the dominant influence on zooplankton temporal variation (monthly zooplankton richness and Rotifera richness, r1 = 0.969, r2 = 0.918). Additionally, Cladocera (34 ± 6, 38 ± 6 species) contributes significantly to the zooplankton richness (r1 = 0.923, r2 = 0.966). The qualitative importance of Rotifera in Deepor Beel agrees with that reported for several other floodplain lakes (Khan 2002, 2003; Sharma 2000, 2005, 2009a, b; Sharma & Sharma 2001, 2008). 298 ZOOPLANKTON COMMUNITIES OF DEEPOR BEEL Table 3. Zooplankton community similarities (%) at station I (Nov 02 - Oct 03). Nov Dec Jan Feb March April May June July Aug Sept Oct Nov - Dec 66.7 - Jan 73.0 76.2 - Feb 88.1 74.6 77.4 - March 80.7 66.7 76.7 85.7 - Apr 51.0 66.7 59.3 56.0 62.5 - May 85.7 60.7 71.2 80.0 64.1 55.3 - June 65.5 69.0 72.1 63.2 61.8 65.3 66.7 - July 78.0 67.8 77.4 82.8 75.0 56.0 72.7 63.2 - Aug 73.4 77.2 76.7 82.1 66.7 58.3 71.7 69.1 85.7 - Sep 76.4 69.1 75.9 81.5 76.9 65.2 70.6 60.4 74.1 73.1 - Oct 74.1 66.7 66.7 79.2 70.6 48.9 68.0 61.5 75.5 74.5 57.1 - July 71.2 69.1 71.7 80.8 74.5 66.7 66.7 76.0 - Aug 76.2 78.0 77.2 71.4 58.2 53.1 65.4 70.4 66.7 - Sep 76.7 82.1 77.9 79.2 73.1 65.2 69.2 72,7 70.6 69.1 - Oct 74.6 72.7 75.5 84.6 74.5 66.7 66.7 60.0 72.0 70.4 78.4 - Table 4. Zooplankton community similarities (%) at station II (Nov 02 - Oct 03). Nov Dec Jan Feb March April May June July Aug Sept Oct Nov - Dec 71.9 - Jan 71.0 89.7 - Feb 75.4 73.7 80.0 - March 76.7 60.7 70.4 75.5 - At both stations, zooplankton richness is negatively correlated with water temperature (r1 = - 0.705, r2 = -0.776) and rainfall (r1 = -0.523, r2 = - 0.654) but positively correlated with specific conductivity (r1 = 0.497, r2 = 0.647) and dissolved oxygen (r1 = 0.501, r2 = 0.764). Richness is also positively correlated with transparency (r2 = 0.650), alkalinity (r2 = 0.608), and hardness (r2 = 0.632) at station II. At stations I and II respectively, this study found 48.9 - 88.1 % and 53.1 - 89.7 % similarity in zooplankton community across monthly samples (Tables 3 & 4). Further, 34.8 and 33.3 % instances in the matrices indicate 60 - 70 % similarity and an additional 40.9 and 54.5 % of instances at the two stations respectively indicate 70 - 80 % similarity. In summary, the majority (75 - 88 %) of instances show 60 - 80 % similarity. This generally high level of similarity suggests limited monthly variations in species composition contrasted by lower simila- Apr 63.0 56.0 62.5 68.1 73.9 - May 70.0 64.3 63.0 71.7 76.9 69.6 - June 67.8 65.4 71.7 65.4 70.6 66.7 70.6 - rities, due to decreased species richness, during the four month summer period (March - June). At stations I and II, respectively, peak similarity values occurred between November - February and December - February and minima were recorded in AprilOctober and April - August. Zooplankton composition shows closeness between November - February and July - August at station I and between DecemberJanuary and February - October at station II. At both stations, and apparently due to the fewer number of species present in April, the samples collected in April exhibit the greatest divergence in their composition. Zooplankton (239 - 657, 475 ± 114 n l-1 and 255 - 687, 459 ± 128 n l-1) forms the main quantitative component of net plankton of Deepor Beel (33.2 68.4, 57.9 ± 9.0 % 35.4 - 66.9, 56.5 ± 8.8 %, at the two stations respectively), thus, significantly contributing to net planktons’ temporal variation (monthly net plankton abundance and zooplankton SHARMA Fig. 3. Monthly variations of zooplankton abundance of Deepor Beel (Nov 02 - Oct 03). Fig. 4. Monthly variations of zooplankton species diversity of Deepor Beel (Nov 02 - Oct 03). abundance, r1 = 0.906, r2 = 0.919). The lowest percentage composition was recorded in April. At both stations, in the six months of November through February and July through August, zooplankton contributed more than 60.0 % of the net plankton abundance. In general, quantitative dominance of zooplankton agrees with earlier reports from Assam (Sharma & Hussain 2001). However, such patterns 299 contrast with the higher phytoplankton abundance noticed in some floodplain lakes and wetlands of Kashmir (Kaul & Pandit 1982), Bihar (Baruah et al. 1993; Rai & Dutta-Munshi 1982; Sanjer & Sharma 1995), and West Bengal (Sugunan 1989; Vass 1989). Generally, zooplankton abundance of Deepor Beel is higher than that reported from Kawar wetland of Bihar (Baruah et al. 1993) and certain beels of Assam (Sharma & Hussain 2001; Sharma & Sharma 2008). On the other hand, abundance is lower than that reported from various floodplain lakes of Bihar (Rai & Dutta Munshi 1982; Sanjer & Sharma 1995), Kashmir (Khan 1987) and West Bengal (Khan 2002; Sugunan 1989; Vaas 1989). At both stations, zooplankton abundance (Fig. 3) oscillates with annual frequency with a rather halting general increase to winter peaks in December at station I and January at station II. Density registers significant monthly variation (F11, 23 = 24.386, P < 0.005) but insignificant variation between stations. This study shows no definite seasonal periodicity other than higher winter densities. The high winter densities at both stations reflect a significant negative correlation of zooplankton abundance with water temperature (r1 = - 0.782, r2 = -0.876). Abundance at both stations is also negatively correlated with rainfall (r1 = -0.573, r2 = -0.647) and positively correlated with dissolved oxygen (r1 = 0.662, r2 = 0.712), transparency (r1 = 0.609, r2 = 0.605), and specific conductivity (r1 = 0.474, r2 = 0.640). In addition at station II, abundance is negatively correlated with free CO2 (r2 = -0.470) and positively correlated with alkalinity (r2 = 0.545) and hardness (r2 = 0.598). At both stations, zooplankton abundance is also positively correlated with its richness (r1 = 0.863, r2 = 0.889). Zooplankton communities of Deepor Beel are characterized by higher species diversity (3.548 4.238, 3.991 ± 0.181 and 3.529 - 4.219, 3.973 ± 0.192) than other beels of Assam (Sharma & Sharma 2008) and, at both stations, show broadly identical values. The zooplankton community shows significant monthly variation (F11, 23 = 15.185, P < 0.005) but insignificant variation between stations. In general, at station I, relatively high diversities (more than 4.0) are observed in January through February and July through October and, at station II, from January through March and July through December. Zooplankton diversity follows oscillating annual patterns at both stations, peaking in July at station I and October at station II. Both have April minima (Fig. 4). 300 ZOOPLANKTON COMMUNITIES OF DEEPOR BEEL Fig. 5. Monthly variations of zooplankton groups’ abundance of Deepor Beel (Nov 02 - Oct 03) at station I. Fig. 6. Monthly variations of zooplankton groups’ abundance of Deepor Beel (Nov 02 - Oct 03) at station II. The high species diversity of zooplankton in Deepor beel, characterized by low densities of majority of species of different groups, may be ascribed to fine niche partitioning amongst species in combination with high micro- and macro-scale habitat heterogeneity in this Ramsar site with a well developed littoral profile and occurrence of various macrophytes as hypothesized by Segers (2008). This interesting feature (mainly influenced by higher Rotifera diversity) also concurs with the results of Dumont & Segers (1996) in a tropical lake with developed weedy littoral. At both stations, zooplankton species diversity is positively correlated with richness of zooplankton (r1 = 0.656, r2 = 0.610), Rotifera (r1 = 0.571, r2 = 0.623) and Cladocera (r1 = 0.590, r2 = 0.575) and, at station I, is significantly correlated with zooplankton abundance (r1 = 0.467). At both stations, Rotifera (231 ± 60 and 198 ± 90 n l-1) constitute the dominant quantitative group (48.7 ± 6.1 % and 42.5 ± 4.1 %) of zooplankton, thus, significantly contributing to the overall temporal variation (r1 = 0.896, r2 = 0.970). At both stations and in accordance with the results of Khan (1987), Sanjer & Sharma (1995), Sharma & Sharma (2001, 2008) and Sharma (2005, 2009a), rotifers reveal (Figs. 5 & 6) an oscillating annual periodicity with January (winter) peaks. Throughout the study period and at both sites, species of Brachionidae and Lecanidae, in the stated order, constitute the major components of rotifer diversity. In contrast to Sharma (1992), no definite seasonal periodicity of abundance of loricate or illoricate rotifers was found. Furthermore, only a few rotifer species seem to be of relative quantitative importance but no individual species shows any distinct dominant role. Cladocera forms a second important group (142 ± 59 and 142 ± 48 n l-1), comprising 28.7 ± 7.0 % and 30.6 ± 4.9 % of general abundance, and considerably influencing zooplankton temporal variation (monthly zooplankton abundance and Cladocera abundance). At both stations respectively, Cladoceran density oscillates with annual frequency and shows (Figs. 5 & 6) peaks in winter (January and December) and minima during April (summer). Their abundance is higher than that reported in the floodplain lakes of Kashmir (Khan 1987), Bihar (Baruah et al. 1993; Sinha et al. 1994), and Assam (Sharma & Hussain 2001; Sharma & Sharma 2008). Cladocera abundance is largely influenced by species of Chydoridae and Daphniidae. As with Rotifera, only a few cladoceran species show relatively high individual densities but no individual species shows any distinct dominant role. At the two stations respectively, Copepoda abundance ranges between 66 ± 17 n l-1 and 81 ± 13 n l-1 comprising 15.1 ± 6.5 % and 18.7 ± 4.9 % of zooplankton. Copepoda abundance oscillates with annual frequency peaking in May (station I) and February and September (station II) SHARMA with minima in October (both stations) (Figs. 5 & 6). The sub-dominant role of Copepoda in this study contradicts their dominant role reported earlier in certain floodplain lakes of Bihar (Baruah et al. 1993), Assam (Sharma & Hussain 2001), and West Bengal (Khan 2003). At both stations, cyclopoids play a dominant role and nauplii occur throughout the whole study period indicating an active reproductive phase of this group. Another sub-dominant group is Rhizopoda; at stations I and II, abundances range respectively, between 29 ± 14 n l-1 and 35 ± 18 n l-1 (Figs. 5 & 6) and comprise 5.9 ± 2.2 % and 7.3 ± 18 % of zooplankton peaking in September at both stations. Other zooplankton groups such as Ostracoda and Conchostraca have very poor abundance. The paucity of detailed analysis of zooplankton in the Indian floodplain lakes highlights the importance of this study, which, in turn, indicates the distinctly rich and diverse nature of zooplankton biocoenosis of Deepor Beel, the speciose character of Rotifera and Cladocera, and the qualitative and quantitative predominance of zooplankton in net plankton communities. Zooplankton richness and abundance show winter peaks that occur simultaneously with lower water temperature. The lack of clear temporal patterns, the different monthly trends, and certain other variations noticed at the two sampling stations need further confirmation. The present results may not reflect holistic environmental heterogeneity of this Ramsar site unless extended to more sampling stations with particular reference to variations in the macrophyte associations. In view of the existing lacunae, investigations have since been initiated by the author. Conclusions Zooplankton comprises an important quantitative component of net plankton, showing a diverse and speciose character, with a rich faunal diversity and the quantitative dominance of Rotifera and Cladocera. In general, richness or abundance of zooplankton or its constituent groups is found to oscillate with annual frequency but more observations may be required to corroborate this conclusively. Results show high species diversity with relatively low densities and equitable abundance for the majority of species. Individual abiotic factors have a limited influence on zooplankton richness and abundance. The present results provide useful information on zooplankton diversity particularly in view of the paucity of a detailed 301 community analysis in the Indian floodplain lakes. In order to acquire better understanding of holistic environmental heterogeneity of this Ramsar site, investigations, however, need to be extended to more sampling stations with particular reference to variations in the macrophyte associations. Acknowledgements This study is undertaken partly under the “Potential for Excellence Program (Focus Area: Biosciences) of North-Eastern Hill University, Shillong. The author is thankful to the G. B. Pant Institute of Himalayan Environmental Development, Almora, for a research grant during which this study was initiated. The author is grateful to Dr. (Mrs.) Sumita Sharma, North Eastern Regional Centre, Zoological Survey of India, Shillong, for her useful comments. Thanks are also due to the Head of the Department of Zoology, North-Eastern Hill University, Shillong, for the necessary laboratory facilities. Finally, the author wishes to thank an anonymous reviewer for critical comments and constructive suggestions. References A. P. H. A. 1992. Standard Methods for the Examination of Water and Wastewater. 18th edn. American Public Health Association, Washington D. C. Baruah, A., A. K. Sinha & U. P. Sharma. 1993. Plankton variability of a tropical wetland, Kawar (Begusarai), Bihar. Journal of Freshwater Biology 5: 2732. Dumont, H. J. & H. Segers. 1996. Estimating lacustrine zooplankton species and complementarity. Hydrobiologia 341: 125-132. Kaul, V. & A. K. Pandit. 1982. Biotic factors and food chain structure in some typical wetlands of Kashmir. Pollution Research 1: 49-54. Khan, M. A. 1987. Observations on zooplankton composition, abundance and periodicity in two floodplain lakes of the Kashmir Himalayan valley. Acta Hydrochemica Hydrobiologica 15: 167-174. Khan, R. A. 2002. The ecology and faunal diversity of two floodplain Ox-bow lakes of South-Eastern West Bengal. Records of the Zoological Survey of India, Occasional Paper No. 195: 1-57. Khan, R. A. 2003. Faunal diversity of zooplankton in freshwater wetlands of Southeastern West Bengal. Records of the Zoological Survey of India, Occasional Paper No. 204: 1-107. Koste, W. 1978. ROTATORIA. Die Rädertiere Mitteleuropas, begründet von Max Voigt. Überordnung 302 ZOOPLANKTON COMMUNITIES OF DEEPOR BEEL Monogononta. Gebrüder Borntraeger, Berlin, Stuttgart. Ludwig, J. A. & J. F. Reynolds. 1988. Statistical Ecology: a Primer on Methods and Computing. John Wiley & Sons, New York. Magurran, A. E. 1988. Ecological Diversity and its Measurement. Croom Helm Limited, London. Michael, R. G. & B. K. Sharma. 1988. Indian Cladocera (Crustacea: Branchiopoda: Cladocera). Fauna of India and Adjacent Countries Series. Zoological Survey of India, Calcutta. Rai, D. N. & J. M. Dutta-Munshi. 1982. Ecological characteristics of `Chaurs’ of North Bihar. pp. 89-95. In: B. Gopal, R. E. Turner, R. G. Wetzel & D. F. Winghon (eds.) Wetlands - Ecology and Management. Vol. II. International Scientific Publications and National Institute of Ecology, Jaipur, India. Sanjer, L. R. & U. P. Sharma. 1995. Community structure of plankton in Kawar lake wetland, Begusarai, Bihar: II. Zooplankton. Journal of Freshwater Biology 7: 165-167. Segers, H. 2008. Global diversity of rotifers (Rotifera) in freshwater. Hydrobiologia 595: 49-59. Sharma, B. K. 1992. Systematics, distribution and ecology of freshwater rotifera in West Bengal. pp. 231 273. In: S. R. Mishra & D. N. Saksena (eds.) Recent Advances in Aquatic Ecology. Ashish Publishing House, New Delhi. Sharma, B. K. 1998. Freshwater rotifers (Rotifera: Eurotatoria). Fauna of West Bengal State Fauna Series 3: 341-461. Zoological Survey of India, Calcutta. Sharma, B. K. 2000. Synecology of rotifers in a tropical floodplain lake of Upper Assam (N. E. India). The Indian Journal of Animal Sciences 70: 880-885. 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. 2009a. Diversity of rotifers (Rotifera, Eurotatoria) of Loktak Lake, Manipur, North eastern India. Tropical Ecology 50: 277-285. Sharma, B. K. 2009b. Rotifer communities of floodplain lakes of Manipur (N. E. India): biodiversity, distribution and ecology. Journal of the Bombay Natural History Society 106: 45-56. Sharma, B. K. & Md. Hussain. 2001. Abundance and ecology of zooplankton in a tropical floodplain lake, Assam (N. E. India). Ecology, Environment & Conservation 7: 397-403. Sharma, B. K. & S. Sharma. 1999a. Freshwater Rotifers (Rotifera : Eurotatoria). State Fauna Series: Fauna of Meghalaya 4: 11-161. Zoological Survey of India, Calcutta. Sharma, B. K. & S. Sharma. 1999b. Freshwater Cladocerans (Crustacea: Branchiopoda: Cladocera). State Fauna Series: Fauna of Meghalaya 4: 469-550. Zoological Survey of India, Calcutta. Sharma, B. K. & S. Sharma. 2000. Freshwater Rotifers (Rotifera: Eurotatoria). State Fauna Series: Fauna of Tripura 7: 163-224. Zoological Survey of India, Calcutta. Sharma, B. K. & S. Sharma. 2001. Biodiversity of Rotifera in some tropical floodplain lakes of the Brahmaputra river basin, Assam (N. E. India). Hydrobiologia 446 / 447: 305-313. Sharma, B. K. & S. Sharma. 2005. Faunal diversity of Rotifers (Rotifera: Eurotatoria) of Deepor Beel, Assam (N. E. India)- a Ramsar site. Journal of the Bombay Natural History Society 102: 169-175. Sharma, B. K. & S. Sharma. 2008. Faunal diversity of Cladocera (Crustacea: Branchiopoda) of Deepor Beel, Assam (Northeast India)- a Ramsar site. Journal of the Bombay Natural History Society 105: 196-201. Sharma, S. & B. K. Sharma. 2008. Zooplankton diversity in floodplain lakes of Assam. Records of the Zoological Survey of India, Occasional Paper No. 290: 1307. Sinha, A. K., A. Baruah, D. K. Singh & U. P. Sharma. 1994. Biodiversity and pollution status in relation to physico-chemical factors of Kawar lake (Begusarai), North Bihar. Journal of Freshwater Biology 6: 309331. Sugunan, V. V. 1989. Limnological features in beels: Biotic factors. Bulletin of Central Inland Capture Fisheries Research Institute, Barrackpore 63: 128135. Talling, J. F. & I. B. Talling. 1965. The chemical composition of African lake waters. Internationale Revue Gesammten Hydrobiologia 50: 421-463. Vass, K. K. 1989. Beel fisheries resources in West Bengal. Bulletin of Central Inland Capture Fisheries Research Institute, Barrackpore 63: 29-35. Yousuf, A. R., G. Mustafa Shah & M. Y. Qadri. 1986. Some limnological aspects of Mirgund wetland. Geobios New Reports 5: 27-30. (Received on 28.05.2009 and accepted after revisions, on 16.09.2010)