Zooplankton
diversity of Loktak Lake, Manipur, India
B.K. Sharma 1 &
Sumita Sharma 2
1,2 Freshwater Biology Laboratory,
Department of Zoology, North-Eastern Hill University, Permanent Campus,
Umshing, Shillong, Meghalaya 793022, India
Email: 1 profbksharma@gmail.com(corresponding author), 2 sumitasharma.nehu@gmail.com
Date
of publication (online): 26 May 2011
Date
of publication (print): 26 May 2011
ISSN
0974-7907 (online) | 0974-7893 (print)
Editor: M.M. Saxena
Manuscript details:
Ms # o2457
Received 11 May 2010
Final received 13 January 2011
Finally accepted 27 April 2011
Citation: Sharma, B.K. & S. Sharma (2011). Zooplankton
diversity of Loktak Lake, Manipur, India. Journal of Threatened Taxa 3(5): 1745–1755.
Copyright: © B.K. Sharma & Sumita Sharma 2011. Creative
Commons Attribution 3.0 Unported License. JoTT allows unrestricted use
of this article in any medium for non-profit purposes, reproduction and
distribution by providing adequate credit to the authors and the source of
publication.
Author Details: Drs. BK Sharma and Sumita Sharma specialize in aquatic biodiversity
and limnology and have made significant contributions to faunal diversity,
biogeography and ecology of freshwater zooplankton of India. The corresponding
author is a Professor in Department of Zoology and Dean, School
of Life Sciences, NEHU, Shillong.
Author Contribution: The
present study is the result of work undertaken by the authors at the Freshwater
Biology Laboratory, Department of Zoology, North-Eastern Hill University, Shillong.
Acknowledgements: This study is undertaken partly under the “Potential for
Excellence Program (Focused Area: Biosciences)” of North-Eastern Hill
University, Shillong. The senior
author is thankful to the G.B. Pant Institute of Himalayan Environmental
Development, Almora for a research grant during which this study was
initiated. Thanks are due to the
Head, Department of Zoology, North-Eastern Hill University, Shillongfor necessary laboratory facilities.
Abstract: Zooplankton communities of Loktak Lake showed rich and speciose
biocoenosis (162 and 142 species), high monthly richness (91 ± 13 and 80 ± 10
species) and by higher similarities (51.1–82.0 and 51.8–78.3 %) and
peak richness during winter and autumn over two years of study. Zooplankton
(267 ± 41 n/l) formed a significant
quantitative component (56.0 ± 6.3 %) of
net plankton and showed annual peak abundance during winter. Rotifera
> Cladocera are dominant quantitative groups while Copepoda > Rhizopoda
are sub-dominant groups. We observed significant annual and monthly variations
of zooplankton richness and abundance. This study showed limited influence of individual abiotic factors on zooplankton,
with richness showing a significant
inverse correlation with water hardness and chloride, and abundance inversely
correlated with nitrate. Multiple regressions indicated higher cumulative
effects of 15 abiotic factors on richness and abundance. Our results exhibited
no definite periodicity of richness and abundance of zooplankton and their
constituent groups during two annual cycles. Zooplankton is characterized by
highest species diversity (4.172 ± 0.237), higher evenness and lower dominance.
Keywords: Abundance, community similarities, diversity indices, ecology,
Ramsar site, richness.
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INTRODUCTION
Zooplankton are integral components of
aquatic food webs and contribute significantly to aquatic productivity in
freshwater ecosystems. They have
been studied from various inland aquatic environs of India, but a review of the
limnological literature indicates limited information on their composition,
ecology and role in aquatic productivity in the floodplain lakes in particular
(Sharma & Sharma 2008). The
related contributions from the floodplain lakes of northeastern India (Sharma
& Hussain 2001; Sharma 2005; Sharma & Sharma 2008) are as yet
restricted to the beels of the Brahmaputra river basin of Assam.
The
present study on synecology of zooplankton of Loktak Lake assumes special
limnological significance in view of a lack of investigations in the floodplain
lakes (pats) of Manipur. The
observations are made on temporal variations in richness, community
similarities, abundance, species diversity, dominance and evenness of
zooplankton of this important floodplain lake, a Ramsar site of India. In addition, the influence of abiotic
parameters on richness and abundance of zooplankton are analyzed.
MATERIALS AND METHODS
This study is a part of a limnological survey
undertaken (November 2002–October 2004) in Loktak Lake (93046’–93055’E
& 24025’–24042’N) located in Bishnupur/Imphal
districts of Manipur of India. Various common aquatic plants of this Ramsar site included Eichhornia crassipes, Hydrilla
verticellata, Euryale ferox, Vallisnaria spiralis, Utricularia flexuosa, Trapa
natans, Lemna trisula, Pistia striates, Salviniasp.Nymphaea spp., Nymphoides spp., Nelumbo mucifera, Potamagetonspp.andAzolla pinnata.
The observations were undertaken at one sampling
site at Sendra (93047’45.61’’E & 24030’56.75’’N). Water samples were collected at
regular monthly intervals and were analyzed for various abiotic factors
following APHA (1992); water temperature, specific
conductivity, pH and dissolved oxygen were recorded by the field probes. Qualitative (by towing) and
quantitative plankton samples (by filtering 25l water each) were collected
monthly by nylobolt plankton net (mesh size 50µm) and were preserved in 5%
formalin. The former were screened
for various species and quantitative samples were analyzed for their
abundance. Zooplankton species were
identified following the works of Koste (1978), Michael & Sharma (1988),
Sharma (1998) and Sharma & Sharma (1999a, 1990b, 2000, 2008). Community similarities (Sorensen’s
index), species diversity (Shannon’s index), dominance (Berger-Parker’s index)
and evenness (Pileou’s index) were calculated following Ludwig & Reynolds (1988) and Magurran (1988). Significance of temporal
variations of biotic parameters was ascertained by ANOVA (two-way). The hierarchical cluster analysis,
based on zooplankton community similarities, was done using SPSS (version
11.0). Ecological correlations
between abiotic and biotic parameters were determined by simple correlation
coefficients (r); their P values were calculated vide http://faculty.vassar.edu/lowry/tabs.html. and significance was ascertained after use of Bonferroni
correction (p < 0.0033). Multiple regression (R2) was used to ascertain cumulative
effect of 15 abiotic factors i.e., water temperature (X1), rainfall
(X2), pH (X3), specific conductivity (X4),
dissolved oxygen (X5), free carbon dioxide (X6),
alkalinity (X7), hardness (X8), phosphate (X9),
nitrate (X10), sulphate (X11), silicate (X12),
chloride (X13), dissolved organic matter (X14) and total dissolved solids (X15) on biotic factors.
RESULTS AND DISCUSSION
Abiotic parameters
Mean water temperature affirms sub-tropical
range of Loktak Lake. Slightly acidic and soft waters of this Ramsar site are
characterized by low specific conductivity (Table 1) depicting low ionic
concentrations. Our results also
indicate moderate dissolved oxygen, low free CO2, low concentrations
of micro-nutrients and other abiotic factors.
Zooplankton richness and community
similarities
Among 189 species of Zooplankton documented from
Loktak Lake (Sharma unpublished), 169 species observed at the sampled site
reflect speciose and diverse nature of their biocoenosis. Zooplankton comprise dominant
qualitative component of net plankton (237 species) and significantly influence
temporal variations of the latter (r = 0.960, p < 0.0001). Overall zooplankton richness is the
highest known till date from any floodplain lake or any individual aquatic
ecosystem of India and, hence, reflects greater environmental heterogeneity of
this Ramsar site. The richness is,
however, notably higher than an unpublished report of 55 species (including
undetermined and doubtful species) from this lake (Singh 1991). Besides, it is distinctly higher than the
records from other Indian floodplain lakes i.e., 51 species (Khan 1987) and 26
species (Yousuf et al. 1986) from Kashmir; 19 species (Baruah et al. 1993) and
31 species (Sanjer & Sharma 1995) from Bihar; 49 species (Sharma &
Hussain 2001) from Assam, and 71 species (Khan 2003) from West Bengal. Qualitative dominance of zooplankton in
net plankton communities of Loktak Lake concurs with the findings of Sharma
& Sharma (2008) but differs from higher phytoplankton richness observed by
Baruah et al. (1993), Sinha et al. (1994) and Sharma & Hussain (2001). In
general, zooplankton biocoenosis exhibits typical ‘tropical character’ and
greater richness of cosmopolitan species while cosmotropical and pantropical
species are well represented.
Monthly zooplankton richness varies between 69–121
(85±13) species during the study period (Table 2); it registers significant
annual (F1, 23. = 26.712, p< 0.005) and monthly variations (F11,
23. = 10.752, p < 0.005). Richness ranges between 91±13 and 85±13 species during two years and
shows annual maxima during winter (December 2002) and autumn (November 2003) respectively. Qualitative diversity is higher
throughout first year except during July. In general, this study records (Figs.
1 & 2) relatively lower number of species from February–July during
first year while this trend is noticed from January–August in the
succeeding year. Individual abiotic factors exert limited influence on
zooplankton richness; it registers only significant negative correlations with
hardness (r = -0.650, p = 0.0011) and chloride (r = -0.723, p = 0.0002). On the other hand, multiple regression (Table 3) indicates higher cumulative effect of
15 abiotic factors on richness (R2 = 0.863).
The collections examined from the selected study
site indicate Rotifera (104 species) > Cladocera (41 species) > Rhizopoda
(17 species). The
faunal diversity of Rotifera and Cladocera of Loktak Lake is explained
separately by Sharma (2009a), and Sharma & Sharma (2009a)
respectively. Monthly richness
(Figs. 1 & 2) of Rotifera ranges between 41–79 (53±9) species during
the study period with annual ranges of 57±10 and 48±6 species respectively
(Table 2). Cladocera monthly
richness ranges between 17–31 (22±4) species and annual mean richness
between 23±5 and 22±4 species respectively while Rhizopoda record
lower mean annual richness (7±1 and 6±1 species). The
qualitative dominance of the rotifers concurs with the reports of Sharma
(2000a, 2000b, 2005), Sharma & Sharma (2001, 2005, 2008) and Khan (2002,
2003). Further, zooplankton or
their constituent groups follow no definite pattern of periodicity of richness
in this Ramsar site.
Zooplankton communities indicate (Tables 4 &
5) only marginally different i.e., 51.1–82.0 % and 51.8–78.2 %
annual similarity ranges (vide Sorenson’s index). Peak similarities are observed between December–October
and February–September while minima are noticed between April–August
and May–July communities during two years respectively. Similarity
matrices exhibit broadly concurrent maximum instances (50.0% and 48.3%) of 60–70
% similarity in both years but show different patterns of cluster analysis
(Figs. 3 & 4). The
hierarchical cluster analysis shows (Fig. 3) closeness in zooplankton
composition between December–October, May–June, March–April
and August–September; the last group, however, shows distinct divergence
from rest of the monthly collections during 2002–03. In the following year (Fig. 4), greater
zooplankton similarity is noticed between November–September, December–August,
March–April and May–October while June–July communities
indicate diverse composition.
Zooplankton abundance
Abundance of zooplankton (Table 2) ranges
between 204–319 (267±41) n/l and indicates significant annual (F1,
23 = 32.096, p < 0.005) and monthly variations (F11, 23 =
7.132, p < 0.005). The density
is apparently higher than the reports of Yadava et al. (1987), Baruah et al.
(1993) and Sharma & Hussain (2001) and it is, however, lower than the
results of Khan (1987), Sanjer & Sharma (1995) and Khan (2002). The
abundance shows (Fig. 5) broadly trimodal annual patterns with peaks during
winter (December 2002 and December 2003). The stated patterns differ from
bimodal periodicity noticed by Yadava et al. (1987) and Sanjer & Sharma
(1995) while these concur with the findings of Sharma & Hussain (2001). Abundance is noticed to be relatively
higher (287±34 n/l) throughout the second year except only during March.
Zooplankton comprise main quantitative component
(39.4 - 65.2, 56.0±6.3) % of net plankton (480±74 n/l) and contribute
significantly to their temporal variations (r = 0.652, p = 0.0012). The quantitative dominance of
zooplankton of Loktak Lake, in turn, concurs with the results of Sharma &
Hussain (2001). This salient
feature is in contrast to higher phytoplankton abundance reported from the
floodplain lakes and wetlands from different parts of India i.e., Kashmir (Kaul
& Pandit 1982), Bihar (Baruah et al. 1993; Sanjer & Sharma 1995), West
Bengal (Sugunan 1989) and Assam (Yadava et al. 1987). Abundance records only significant inverse correlation with
nitrate (r = -0.697, p = 0.0003) showing very limited influence of individual
abiotic while multiple regression indicates (Table 3)
higher cumulative effect of 15 abiotic factors on zooplankton abundance (R2= 0.851).
Amongst different groups (Figs. 6 & 7), Rotifera
form (46.6±4.1%)) an important quantitative component of zooplankton and
contribute significantly (r = 0.974) to their abundance; Cladocera (27.1±5.0 %)
and Copepoda (17.4±7.2 %) also contribute to their abundance while Rhizopoda
(Table 2) comprise between 8.3±2.0 %. Other groups of zooplankton, namely, Ostracoda, Gastrotricha and
Conchostraca indicate very poor abundance (Table 2). The synecology of Rotifera and
micro-crustaceans are dealt with separately by Sharma (2009b) and Sharma &
Sharma (2009b) respectively. In general, zooplankton and their constituent groups follow
no definite pattern of quantitative periodicity in Loktak Lake.
Species diversity, evenness and
dominance
Zooplankton showthe highest species diversity (3.750–4.639, 4.172±0.237) so far known from
any floodplain lake or aquatic ecosystem of northeastern India and elsewhere
from this country (Sharma unpublished). This salient feature affirms higher environmental heterogeneity of
Loktak Lake. It registers significant monthly (F11, 23 = 3.762, p <
0.01), shows (Table 2) relatively higher values (4.186±0.278) during the second
year and exhibits broadly bimodal but different annual patterns (Fig. 8) with
peaks during winter (December 2002) and post-monsoon (October 2004) and minima
during summer (May). In general,
higher species diversity (< 4.0) is observed during November-February, July
and September–October during both years and again during May and June (2ndyear) and August (1st year). The notable feature of higher species diversity with relatively
lower densities of majority of species noticed in this study may be ascribed to
fine niche portioning amongst zooplankton species in combination with high
micro- and macro-scale habitat heterogeneity as hypothesized by Segers (2008).
This study depicts (Table
2) higher evenness (0.813–0.992, 0.939±0.032) of zooplankton; this
feature affirms equitable abundance of various species and concurs with the
results of Sharma & Hussain (2001) and Sharma & Sharma (2008). It follows (Fig. 9) bimodal and multimodal
annual patterns with peaks during August and October during the two years
respectively while minima are noticed during summer (May). Zooplankton show
(Table 2) lower dominance (0.044–0.170, 0.079±0.033) indicating lack of
quantitative importance of individual species. This feature again concurs with the results of Sharma &
Hussain (2001) and Sharma & Sharma (2008). It registers only significant
monthly variations (F11, 23 = 3.896, p < 0.01), and follows (Fig.
10) different annual patterns with maxima during April and May during two years
respectively.
To sum up, zooplankton form important
qualitative and quantitative components of net plankton, exhibit highly
speciose character with richest diversity and quantitative dominance of
Rotifera > Cladocera, and indicate no definite periodicity of richness or
abundance of zooplankton or their constituent groups. zooplankton are characterized by
highest species diversity, higher evenness and lower dominance, and exhibit
lower densities of majority of species. This study registers limited influence
of individual abiotic factors but records higher cumulative influence of 15
abiotic factors on richness and abundance. The present observations limited to one sampling station,
though provide useful information on composition, production and ecology of
zooplankton yet may not reflect full view of heterogeneity of this interesting
Ramsar site. Further studies in
different parts (pats) of Loktak basin are, hence, desired and have been
initiated.
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