Journal of Global Biosciences

ISSN 2320-1355
Volume 4, Number 7, 2015, pp. 2842-2858
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Research Paper
TAXONOMIC AND FUNCTIONAL DIVERSITY OF FISH ASSEMBLAGE IN
THREE INTERCONNECTED TROPICAL RIVERS IN INDIA IN
ACCORDANCE WITH LIMITING SIMILARITY HYPOTHESIS
Subhasree SenGupta1,2 and Sumit Homechaudhuri1
1
Aquatic Bioresource Research Laboratory
Department of Zoology, University of Calcutta
35 Ballygunge Circular Road, Kolkata - 700019, India
2 Department of Zoology, Taki Government College

Taki, North 24 Parganas, West Bengal, India.

Abstract
Freshwater fishes are declining rapidly due to their high sensitivity to the
quantitative and qualitative alteration of aquatic habitats and India is no
exception. The small rivers of this country provide refuge to innumerable fish
species with hardly any documentation. This study was directed to assess fish
biodiversity with reference to species diversity, phylogenetic diversity and
functional diversity of the fish assemblage in three interconnected shallow
rivers in the northern part of West Bengal. Sampling was conducted in between
September 2007 to August 2011and diversity estimators depict higher species
richness and evenness and less dominance during monsoon and autumn with
progressive declining trend through winter, spring and summer. The taxonomic
and functional diversity also depicted similar trend in relation with total fish
catch. However, the average taxonomic distinctness, phylogenetic diversity and
functional diversity estimator indicate the inherent taxonomic diversity and
functional complementarity of the fish assemblage throughout the year
irrespective of total fish catch. This fact signifies the importance of these three
interconnected rivers as a precious habitat for indigenous fish species ensuring
effective niche segregation and coexistence in a stable way for the resident fish
fauna. In order to ensure, long term sustainability of fish stock in this habitat
efforts need to be given to minimize disruption of ecosystems to protect the
precious habitat.
Key words: fish biodiversity, taxonomic diversity, functional diversity, limiting
similarity, Aquatic Ecological System.

INTRODUCTION
Freshwater fish are one of the most threatened taxonomic groups (Darwall and Vie,
2005) because of their high sensitivity to the quantitative and qualitative alteration of aquatic
habitats (Laffaille et al., 2005; Kang et al., 2009; Sarkar et al., 2008). Maintenance of
Hydrological connectivity is essential for the viability and recruitment of native biota (Bunn and
Arthington, 2002) and any alteration may lead to local extinction. Longitudinal and lateral
connectivity between river channels and floodplains leads to characteristic high biodiversity
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pattern (Ward and Stanford, 1995) whereas river regulation leads to reduced native species
abundance and recruitment (Cadwallder and Lawrence, 1990).
The term “biodiversity” should never be equated with species diversity, but can be
considered as 'the sum of the taxonomic or numerical diversity, and the ecological,
genetical, historical and phylogenetic diversity' (Van der Spoel, 1994). The phylogenetic
structure of the assemblage, the functional role of the organism play significant role in shaping
biodiversity of a particular habitat. However, all these different elements are impossible to
encapsulate objectively in the same units to provide a single 'biodiversity' index. Hence, in this
study, data was collected on species richness, phylogenetic and functional diversity of the fish
assemblage of three interconnected rivers, namely Mahananda – Tangon – Punarbhaba, in West
Bengal, India. The importance of this site lies with the connectance between the rivers providing
a corridor for migration of fishes over a large area. Side by side, these small rivers provide
refuge to large number of indigenous fish population (Kundu et al., 2014) and are also
undergoing major habitat alterations due to anthropogenic stresses. Before this rich species
diversity of the region is lost forever, the documentation of the extant fish biodiversity is crucial.

MATERIAL AND METHODS

Study site
The fishes were collected from the Mahananda – Tangon – Punarbhaba river system (MTPRS) in
Malda district of West Bengal, India near Indo-Bangladesh border. The Mahananda originates in
the Himalayas, flows through India and parts of Bangladesh and drains into the Ganges after
flowing through and Malda district in West Bengal (India). The Tangon and Punarbhaba
originate separately in the Thakurgaon district in Bangladesh and enter India through South
Dinajpur and ultimately drain into the Mahananda. The importance of this site lies with the
connectance between the rivers and provides a corridor for fishes over a large area. The depth
of river varies from a maximum 4-4.5 meter during the monsoon, to a minimum of less than 1
meter during summer with a silty-clayey substratum. The basin consists of recent alluvium soil
type flanked by arable land, especially graminoid and scrub vegetation, with sporadic rural
settlements. Throughout the catchment, the stream waters are heavily used for cultivation of
crops. (Fig. 1)

Sampling
Sampling was conducted at 20 sites along different sites of the MTPRS (24° 57' N, 88° 20' E) on
the Indian side in Malda district of West Bengal with the help of local fisher folk in between
September 2007 to August 2013. The annual cycle is divided into five seasons as follows:
monsoon (June-August), autumn (September-November), winter (December-January), spring
(February-March) and summer (April-May). Experimental nettings were undertaken the 5
respective seasons using a gill net of 20 m length with 3 cm spacing between adjacent knots for
sample collection. Each site was sampled for at least 10 times to minimize sampling error. The
nets were placed alongside the Indian bank of the rivers for 12 h from evening (6 pm) to early
morning (6 am) in order to ensure maximum fish catch per unit effort. The specimens were
retrieved from the net, identified morphologically to the lowest taxonomic level following Shaw
and Shebbeare, 1937; Day, 1958 and Talwar and Jhingran, 1991 and then released in wild after
preserving representative specimens in 4% formaldehyde. All species names adhere to Fishbase
(Froese and Pauly, 2015).The fish abundance pattern did not vary significantly among the
various sites, so instead of comparing fish diversity and hydrological parameters among the
study sites, we pooled the data to minimize sampling errors for seasonal comparison.

Species diversity, abundance and distribution

In order to assess icthyofaunal diversity in the MTPRS in association with year wise seasonal
variation some of the following diversity indices were used. These were: Shannon-Weaver index
(H') (Shannon and Weaver, 1949), Species evenness or equitability (J) (Pielou, 1969),
Dominance index (D) (Berger and Parker, 1970), Species richness (Margalef, 1958), Fish Species
Richness (Lakra et al., 2010). To normalize variation, the number of species, number of fish and

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species diversity were log10(x+1) transformed (Underwood, 1997) prior to analysis whenever
required. One way analysis of variance (ANOVA) and Duncan’s Post Hoc tests were employed to
check for differences in the abundance of species between all pairs of seasons. The seasonal
variation in fish assemblage structure was graphically represented by the application of cluster
analysis based on Bray-Curtis similarity index and non metric Multi dimensional scaling (MDS).
In order to overcome sampling errors, non-parametric methods like Chao 1, Chao 2, Jackknife,
Bootstrap estimators and Rarefaction curve were used to ascertain actual species richness. All
the computation was performed using EstimateS (version 8) and SPSS (Statistical Package for
Social Sciences, Version 13.0; Norušis 2000) and PRIMER 5 software. IUCN and NBFGR (Lakra et
al., 2010; Viswanath et al., 2010) proposed threat criteria of species was considered.

Assessment of Taxonomic diversity of the fish assemblage

In order to assess phylogenetic structure of the assemblage as a measure of biodiversity, five
indices were employed, viz. Phylogenetic diversity (Faith, 1992), average phylogenetic diversity
(AvPD), Taxonomic distinctness (Δ) (Warwick and Clarke, 1995) Average taxonomic
distinctness (Δ+), Variation in taxonomic distinctness (VarTD, Λ+) (Clarke and Warwick, 2001).

Calculation of Functional Diversity of the fish assemblage

Representative specimens (n = 10) of all fish species were fixed in 4% formaldehyde and
transferred to the laboratory for further morphometric analysis taking care of following
parameters as depicted by Mouillot et al., 2007 for calculation of functional diversity following
Mason et al., 2003. The parameters were biomass, ratio of standard length to body depth, caudal
fin aspect ratio, eye diameter and the mean value of each trait were taken for each species. We
considered that intraspecific differences are much lower than interspecific differences for
functional traits (Dumay et al., 2004).

RESULTS
A total of 52 species distributed amongst 7 orders, 23 families, 45 genera and 52 species
(Table 1) were obtained from MTPRS in this study. Cyprinidae (22.05%) was the most abundant
fish family followed by Bagridae (12.31%) and Schilbeidae (11.28%) respectively (Figure 2). A
comparison of species richness across seasons indicated seasonal variation with species
richness being greatest during the monsoon and autumn showing a declining trend through
winter, spring and summer (P<0.05, df=4) following Kruscal Wallis ANOVA according to Zar
(1999). Out of 52 fish species found in the river, Siluriform catfishes M. cavasius, M. tengara, M.
vittatus, C. batrachus, O. pabo, O. pabda, W. attu; members of order Cypriniformes like C. reba, E.
danricus, P. chola, S. bacaila and Perciformes like C. nama, C. fasciatus, C. punctatus, C. striatus, N.
nandus were found throughout the year (Table 1). Out of these 16 species 5 belonged to
vulnerable, 2 belonged to endangered category (Table1) as per NBFGR report (Lakra et al.,
2010) and Goswami et al., 2012 signifying it as a preferred home for threatened fishes also.
The fish diversity was also analyzed from diversity estimators. The Shannon-Weaver
index, species richness and evenness (Table 3) was found to be higher in monsoon (3.7, 66, 0.94
respectively) and the least in summer (2.42, 18, 0.84 respectively) signifying existence of more
diverse fish assemblage in monsoon. Consequently, autumn (0.046) and monsoon (0.117)
showed least value of dominance index signifying higher evenness in fish assemblage structure.
Shannon–Weaver diversity index showed positive correlation (r= 0.97) with the Evenness, Fish
species richness (FSR) (r= 0.99) and Species richness (r= 0.69) index (Table 4). However, the
Dominance index was found to be negatively correlated (r= -0.55, -0.52) with Shannon–Weaver
diversity index and FSR (Table 4). This pattern of species richness was confirmed by the
rarefaction curves, which suggest that summer had the lowest diversity, with an asymptote in
the region of 7 species, whilst the asymptotes during monsoon, autumn, winter and spring were
in the region 52, 50, 38, 21 species respectively. Similar trends were shown by Jackknife1, 2 and
Chao 1 estimators, sample based rarefaction curve, bootstrap values, MM runs, ACE and ICE
mean values when plotted in a curve (Figure 3). The Bray Curtis (Figure 4) similarity index and

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MDS plot (Figure 5) demonstrated a close resemblance of species composition between

monsoon and autumn, but dissimilarity between monsoon-summer and winter-summer.
Estimation of taxonomic diversity of a fish assemblage also depicted similar trend. The
phylogenetic diversity (PD) and total taxonomic distinctness of the fish assemblage was found
to be at higher side in monsoon (3150 and 4753.9) and autumn (3100 and 4648) with
progressive declining trend in winter (2650 and 3990.7) and spring (2000 and 2880.6) and
lowest value in summer (1050 and 1570.6) (Fig. 6). On the other hand, average PD and average
taxonomic distinctness (AvTD) was found to be fairly constant with highest value in winter
(62.5 and 91.42) and lowest value in summer (58.33 and 87.25). The variation in taxonomic
distinctness (VarTD), (Λ) was found to be highest in summer (425.8) with least number of
species and lowest in monsoon (259.6) with highest species richness. The AvTD and VarTD
value for all the seasons (except summer) were found to lie well within the 95% probability
funnel with 1000 simulations with summer lying just beneath the funnel (Figure 7). Thin line
denotes the theoretical mean for such random selections. Mean FDvar (Figure 8) were calculated
for the selected characters and found to be relatively high in all the five seasons with highest
value in Monsoon (0.846) and least in Summer (0.826).

DISCUSSION
The MTPRS is an aggregation of small river systems ultimately draining to the Ganges
providing corridor for migration of fishes over large area. Very few studies were made in the
rivers of sub-Himalayan Indo-Gangetic plains of Bengal, which revealed huge freshwater fish
faunal diversity in the shallow rain fed rivers (Menon, 1974, 1999; Chakraborty and
Bhattacharjee, 2008). The present study reported capture of 52 fish species in the river
distributed amongst 7 orders, 23 families, 45 genera (Table 1). This data was comparable with
distribution pattern of different finfishes in river of various climatological regions as depicted in
Table 2. This comparison established MTPRS as a precious habitat for fish species conservation.
Cyprinidae (22.05%) was the most abundant fish family and similar results were noted in other
river systems in Southeast Asia (Bhat, 2003).
Several statistical estimators have been used for calculating and extrapolating species
richness taking into account the possible proportion of rare species and make conservative
estimates of the true species richness of an area (Colwell and Coddington, 1994). The fish
diversity estimators across seasons indicated higher diversity and lower dominance of fish
fauna during the monsoon and autumn with progressive declining trend through winter, spring
and summer respectively. This observation was also confirmed by the rarefaction curves, which
suggest that summer had the lowest diversity, with an asymptote in the region of 7 species,
whilst the asymptotes during monsoon, autumn, winter and spring were in the region 52, 50,
38, 21 species respectively (Figure 3). More diversity and evenness in the fish assemblage
elaborated less dominance of resident fish species and vice versa. Nair et al. (1989) and
Chowdhury et al. (2010) have shown similar results in their study. The higher diversity in
monsoon was due to increased niche availability supported by huge nutrient flux from
allochthonous sources along with surface run-off. Flanked by crop fields and numerous
wetlands, the river banks are well vegetated which helps retain precipitation in the drainage
basin for longer time so as to enhance the nutrient quality of the rivers as observed earlier
(Shaji and Easa, 1995, 1998; Arunachalam, 2000; Bhat, 2003; Kar et al., 2006; Shahnawaz et al.,
2010). This was followed by autumn as the post-monsoonal elevated nutrient pool supported
survival of a diversified fish assemblage. However, the constantly declining diversity pattern
over winter, spring and summer could be attributed to overfishing and progressive depletion in
nutrient pool as well as water availability. The shallowness of the riverine banks also provides
suitable sites for fish breeding during monsoon. (Kundu et al., 2014). The Bray Curtis (Figure 4)
similarity index and MDS plot (Figure 5) demonstrated a close resemblance of species
composition between monsoon and autumn due to similarity in habitat quality, environmental
conditions and species composition, but dissimilarity between monsoon-summer and winter-
summer.

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However, it should be noted that richness is not the only measurable component of
community level biodiversity. The phylogenetic structure of the assemblage is also clearly
important, and an assemblage comprising a group of closely related species must be regarded as
less ‘biodiverse’ than an assemblage of the same number of more distantly related species. The
importance of measuring taxonomic diversity of the assemblage gained impetus in last decade
for setting conservation priorities (Vane-Wright et al., 1991; Williams et al., 1991) and
environmental monitoring (Warwick and Clarke, 1995; Clarke and Warwick, 1998; 1999).
Taxonomic distinctness (∆+) was used to summarize the pattern of relatedness in the sample.
The fish assemblage under study, elaborate highest taxonomic distinctness and phylogenetic
diversity in monsoon and autumn with progressive declining trend through winter, spring and
summer (Figure 6) in relation with total fish catch. However, the mean statistic, average
phylogenetic diversity (AvPD) and average taxonomic distinctness (AvTD) value was found to
be fairly constant for most of the seasons signifying inherent taxonomic diversity in the fish
assemblage throughout the year despite fluctuation in species richness value. The variation in
taxonomic distinctness (VarTD, Λ+) can be defined as the variance in the pairwise path lengths
and reflects the unevenness of the taxonomic tree. VarTD showed highest value in summer with
progressive declining trend through spring, winter, autumn and least in monsoon. This trend
found support in Clarke and Warwick (2001) with negative correlation coefficient between
taxonomic distinctness and variance close to 1. This indicated greater variance in taxonomic
composition of the fish assemblage with summer containing representation of eight families and
eighteen species. On the other hand, monsoon showed less variance with representation of 23
families and 52 species. Increased variability might also be a symbol of stress as found in marine
system by Clarke and Warwick (2001). The term "stress" in this study might be on account of
overfishing and progressive depletion in nutrient pool as well as least water availability in
summer. The null hypothesis that the taxonomic distinctness of the fish assemblage is not
significantly different from the global list was tested by repeated subsampling (1000
simulations) of species list of appropriate size. In this study, 95% confidence funnel (Figure 7)
was constructed across the full range of species found throughout the year. Similarly, VarTD
(Λ+) values for all seasons with 1000 simulations fall within the funnel with summer falling
above the upper limit of funnel, indicating higher than expected variation in taxonomic
composition.
The phenomenon of increased phylogenetic diversity with increasing environmental
stability could be substantiated over an evolutionary and ecological time scale from the study of
marine realm vis-à-vis terrestrial biota (Briggs, 1994). In the marine realm there are 34 phyla.
Although these vary in their total numbers of species, none of them is overwhelmingly
predominant. However, the terrestrial biota, which has been subjected to a much higher
degree of environmental variability in comparison with sea over the last 450 million years,
comprises only 15 phyla, with more than 90% of the species belonging to the phylum
Arthropoda (Briggs,1994). The study testified the aforementioned hypothesis in having
increased taxonomic distinctness in monsoon and autumn with lowest value in summer
signifying more stability in the riverine system in monsoon and autumn. A stable system could
accommodate higher fish diversity, i.e. higher taxonomic richness ensuring higher taxonomic
distinctness and less variance as found in the case of monsoon and autumn. On the other hand,
water level drops down in summer with minimum influx of nutrient leading to lowering of fish
diversity in the river. This results in further loss of nutrients in species poor season (Thorp et
al., 1994). Thus, taxonomic distinctness of the fish assemblage is lowered with corresponding
increment in VarTD. The self- conservative nature of the lotic system is well established.
The fundamental idea behind the study of biodiversity patterns is the presumed
connection between the shape of species assemblages and the functional ways in which they are
organized, as competitors or members of a web of interactions and to how species are facing
similar environmental constraints. Taxonomic and ecological characterisations of community
composition are complementary and are useful in conservation context (Angermeier and
Winston, 1998; King et al., 2009). Functional diversity is certainly a key for understanding
ecosystem processes. According to the limiting similarity hypothesis (MacArthur and Levins,

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1967), species were always in competition and the magnitude of competition would be higher
between those, who are more similar to each other, i.e. functionally redundant. Hence, species
eventually co-existing in a stable way would be those that are dissimilar to each other and
therefore allowed niche segregation minimizing competition. The results of this study suggested
high functional diversity for the fish assemblage with highest value in Monsoon (0.846) and
least in Summer (0.826) signifying preponderance of limiting similarity hypothesis and high
functional complementarity of the resident fish fauna and indicative of ecosystem processes in
the river network.

CONCLUSION
At the end of this length discussion on different facets of biodiversity, the
stretches of Mahananda, Tangon and Punarbhaba rivers seems to be preferred habitats for
feeding and reproduction of a large number of indigenous fishes some of which belong to the
threatened categories amongst the Indian fishes. This could be attributed to the fact that i) an
aggregation of small river systems ultimately draining to the Ganges providing corridor for
migration of fishes over large area, ii) shallowness of the rivers providing good spawning site,
iii) input of allochthonous nutrients along with surface run off, iv) stratifications imparted by
submerged and riparian vegetation. In order to ensure, long term sustainability of fish stock in
this habitat efforts need to be given either to ensure minimum biological and economical over-
fishing as well as minimize disruption of ecosystems to protect the precious habitat. Hence, this
study identify MTPRS as an Aquatic Ecological System (AES) having i) stream networks less
than 1000 Km2, ii) identical hydrologic, nutrient and temperature regime should be treated as
an actual conservation target according to Nature Conservancy Classification (Higgins et al.,
2005). The conserved fish stock in rivers could also serve as ‘banks’ of organisms for the
replenishment of unprotected or degraded areas (Nevill and Philips, 2001) with the aim to i)
provide refuge, ii) allow population to increase for further introduction to newer habitats as a
part of rehabilitation programme. This type of investigation at local community level indicated
patterns of diversity which would help conservation biologists to develop management
strategies for effective long-term conservation of biological diversity (Magurran, 2004).

ACKNOWLEDGEMENT
The authors are thankful to the Head of the Department, Department of Zoology, University of
Calcutta, Principal, Bethune College, Kolkata and and Mr. Dipak Roy of Roy’s Fish Farm, Malda
for the facilities provided. S.S.G. is thankful to University Grants Commission - Minor Research
Project.

REFERENCES
(1) Angermeier, P L. and Winston, M R. (1998). Local versus regional influences on local
diversity in stream fish communities of Virginia. Ecology, 79: 911-927.
(2) Arthington, A H., Balcombe, S R., Wilson, G A., Thomas, M C. and Marshall, J.
(2005). Spatial and temporal variation in fish-assemblages structure in isolated
waterholes during the 2001 dry season of an arid—zone floodplain river, Cooper
Creek. Australia. Marine and Freshwater Research, 56: 25–35.
(3) Arunachalam, M. (2000). Assemblage structure of stream fishes in the Western
Ghats (India). Hydrobiologia, 430: 1-31.
(4) Arunachalam, M., Johnson, J A. and Sankaranaryanan, A. (1997). Fish diversity in
rivers of Northern Karnataka, Western Ghats. International Journal of Ecology and
Environmental Sciences, 23: 327–333.
(5) Berger, W H. and Parker, F L. (1970). Diversity of planktonic Foramnifera in deep
sea sediments. Science, 168: 1345-1347.
(6) Bhat, A. (2003). Diversity and composition of freshwater fishes in river systems of
Central Western Ghats, India. Environmental Biology of Fishes, 68: 25-38.
(7) Briggs, J C. (1994). Species diversity: land and sea compared. Systematic Biology, 43:
130-135.

http://mutagens.co.in 2847
Journal of Global Biosciences Vol. 4(7), 2015 pp. 2842-2858
ISSN 2320-1355

(8) Bunn, S E. and Arthington, A H. (2002). Basic principles and ecological

consequences of altered flow regimes for aquatic biodiversity. Environmental
Management, 30(4): 492–507.
(9) Cadwallader, P L., Lawrence, B. (1990). Fish. In: Mackay N, Eastburn D Eds. The
Murray Murray-Darling Basin Commission, Canberra. pp 316–335.
(10) Chakraborty, T. and Bhattacharjee, S. (2008). The ichthyofaunal diversity of
freshwater rivers of South Dinajpur District of West Bengal, India. Journal of
Bombay Natural History Society, 105: 292-298.
(11) Chowdhury, M S N., Hossain, M S., Das, N G. and Barua, P. (2010). Environmental
variables and fisheries diversity of the Naaf River Estuary, Bangladesh. Journal of
Coastal Conservation, 15: 163-180.
(12) Clarke, K R. and Warwick, R M. (2001). A further biodiversity index applicable to
species lists: variation in taxonomic distinctness. Marine Ecology Progress Series,
216: 265-278.
(13) Clarke, K R. and Warwick, R M. (1999). The taxonomic distinctness measure of
biodiversity: weighting of step lengths between hierarchical levels. Marine Ecology
Progress Series, 184: 21–29.
(14) Clarke, K R. and Warwick, R M. (1998). A Taxonomic Distinctness Index and its
Statistical Properties. Journal of Applied Ecology, 35(4): 523-531.
(15) Colwell, R K. and Coddington, J A. (1994). Estimating the extent of terrestrial
biodiversity through extrapolation. Biological Sciences, Royal Society of London, 345:
101-118.
(16) Darwall, W R T. and Vie, J C. (2005). Identifying important sites for conservation of
freshwater biodiversity: extending the species based approach. Fisheries
Management Ecology, 12: 287–293.
(17) Day F. (1958). The Fishes of India, vol. 1 and 2. William Dawson, London.
(18) Dumay, O., Tari, P S., Tomasini, J A. and Mouillot, D. (2004). Functional groups of
lagoon fish species in Languedoc Roussillon, southern France. Journal of Fish
Biology, 64: 970-983.
(19) Faith, D P. (1992). Conservation evaluation and phylogenetic diversity. Biological
Conservation, 61: 1–10.
(20) Froese, R. and Pauly, D. (2015). FishBase. World Wide Web electronic publication.
www.fishbase.org, version (05/2015)
(21) Goswami, U C., Basistha, S K., Bora, D., Shyamkumar, K., Saikia, B. and Changsan, K.
(2012). Fish diversity of North East India, inclusive of the Himalayan and Indo
Burma biodiversity hotspots zones: A checklist on their taxonomic status,
economimportance, geographical distribution, present status and prevailing
threats. International Journal of Biodiversity and Conservation, 4(15): 592-613.
(22) Higgins, J V., Bryer, M T., Khoury, M L. and Fitzhugh, T W. (2005). A freshwater
classification approach for biodiversity conservation planning. Conservation Biology,
19: 432-445.
(23) Kang, B., He, D., Perrett, L., Wang, H., Hu, W., Deng, W. and Wu, Y. (2009). Fish and
fisheries in the Upper Mekong: current assessment of the fish community, threats
and conservation. Reviews in Fish Biology and Fisheries, 19: 465–480.
(24) Kar, D., Nagarathna, A V., Ramachandra, T V. and Dey, S C. (2006). Fish diversity and
conservation aspect in an aquatic ecosystem in north-eastern India. Zoos’ Print
Journal, 21: 2308-2315.
(25) King, A J., Tonkin, Z. and Mahoney, J. (2009). Environmental flow enhances native
fish spawning and recruitment in the Murray River, Australia. River Research and
Application, 25: 1205-1218.
(26) Kundu, N., Majumdar, D. and Homechaudhuri, S. (2014). Seasonal changes in fish
assemblages and trophic guild in some interconnected rivers in eastern
India. Indian Journal of Fisheries, 61(3): 35-44.

http://mutagens.co.in 2848
Journal of Global Biosciences Vol. 4(7), 2015 pp. 2842-2858
ISSN 2320-1355

(27) Laffaille, P., Acou, A., Guillouet, J. and Legult, A. (2005). Temporal change in
European eel, Anguilla anguilla, stock in a small catchment after installation of fish
passes. Fisheries Management Ecology, 12: 123–129.
(28) Lakra, W S., Sarkar, U K., Gopalakrishnan, A. and Kathivelpandian, A. (2010).
Threatened Freshwater Fishes of India, NBFGR, Lucknow, pp. 1-20.
(29) Lakra, W S., Sarkar, U K., Kumar, R S., Pandey, A., Dubey, V K. and Gusain, O P.
(2010). Fish diversity, habitat ecology and their conservation and management
issues of a tropical River in Ganga basin, India. Environmentalist, DOI
10.1007/s10669-010-9277-6.
(30) MacArthur, R. and Levins, R. (1967). Limiting similarity convergence and
divergence of coexisting species. American Naturalist, 101: 377-387.
(31) Magurran, A E. (2004). Measuring biological diversity in time and space. In:
Magurran AE, McGill BJ (Eds) Biological Diversity: Frontiers in Measurement and
Assessment, Oxford, Oxford University Press, pp 85–104.
(32) Margalef, R. (1958). Information theory in ecology. General Systematics, 3: 36-71.
(33) Mason, N W H., MacGillivray, K., Steel, J B., Wilson, J B. (2003). An index of
functional diversity. Journal of Vegetation Science, 14: 571-578.
(34) Menon, A G K. (1999). Freshwater fishes of India. Zoological Survey of India,
Kolkata, 366 pp.
(35) Menon, A G K. (1974). A checklist of the fishes of the Himalayan and Indo-Gangetic
plains. Inland Fisheries Society, Barrackpore, 136 pp.
(36) Mouillot, D., Dumay, O. and Tomasini, J A. (2007). Limiting similarity, niche filtering
and functional diversity in coastal lagoon fish communities. Estuarine, Coastal and
Shelf Science, 71: 443-456.
(37) Nair, N B., Arunachalam, M., Madhusoodanan, N K C. and Suryanarayanan, H.
(1989). Seasonal variation and species diversity of fishes in the Neyyer River of the
Western Ghats. Tropical Ecology, 30: 69–74.
(38) Nevill, J. and Phillips, N. (2001). Representative freshwater reserves: better late
than never! Paper presented to the International River Symposium, Hilton Hotel
Brisbane, 29-31 August 2001.
(39) Norušis, M J. (2004). SPSS 13.0 Advanced Statistical Procedures
Companion. Englewood Cliffs: Prentice Hall.
(40) Oliveira, E F., Gaulart, E. and Minte-Vera, C V. (2004). Fish diversity along spatial
gradients in the Itaipu Reservior, Parana, Brazil. Brazilian Journal of Biology,
64(3A): 447–458.
(41) Pielou, E C. (1969). An Introduction to Mathematical Ecology. Wiley, New York.
(42) Sarkar, U K., Pathak, A K. and Lakra, W S. (2008). Conservation of freshwater fish
resources of India: new approaches, assessment and challenges. Biodiversity
Conservation, 17: 2495-2511.
(43) Shahnawaz, A., Venkateshwarlu, M., Somashekar, D S. and Santosh, K. (2010). Fish
diversity with relation to water quality of Bhadra River of Western Ghats (INDIA).
Environmental Monitoring Assessment, 161: 83-91.
(44) Shaji, C P. and Easa, P S. (1998). Status and distribution of endemic freshwater
fishes in Kerala, Western Ghats, India. Tigerpaper, 25: 21-24.
(45) Shaji, C P. and Easa, P S. (1995). Freshwater fish diversity in Wayanad, Kerala, South
India. Journal of Zoological Society, Kerala, 5:30-36.
(46) Shannon, C E. and Weaver, W. (1949). A Mathematical Model of Communication.
University of Illinois Press, Urbana
(47) Shaw, G E. and Shebbeare, E O. (1937). The fishes of North Bengal. Journal of Royal
Asiatic Society of Bengal, III, 1-137.
(48) Talwar, P K. and Jhingran, A. (1991). Inland fishes of India and adjacent countries.
Oxford and IBH Publishing Co. Pvt. Ltd., New Delhi, 2 volumes: xix + 1158 pp.

http://mutagens.co.in 2849
Journal of Global Biosciences Vol. 4(7), 2015 pp. 2842-2858
ISSN 2320-1355

(49) Thirumala, S., Kiran, B R. and Kantaraj, G S. (2011). Fish diversity in relation to
physico-chemical characteristics of Bhadra reservoir of Karnataka, India. Advances
in Applied Science Research, 2(5): 34-47.
(50) Thorp, J H., Black, A R., Haag, K H. and Wehr, J D. (1994). Zooplankton assemblages
in the Ohio River: Seasonal, tributary, and navigation dam effects. Canadian Journal
of Fisheries and Aquatic Sciences, 51: 1634–1643.
(51) Underwood, A J. (1997). Experiments in Ecology: Their Logical Design and
Interpretation Using Analysis of Variance. Cambridge University Press, Cambridge
(52) Van der Spoel, S. (1994). A biosystematic basis for pelagic biodiversity. Bijdragen
tot de Dierkunde, 64: 3-31.
(53) Vane-Wright, R I., Humphries, C J. and Williams, P H. (1991). What to protect?
Systematics and the agony of choice. Biological Conservation, 55: 235–254.
(54) Vishwanath, W., Lakra, W S. and Sarkar, U K. (2007). Fishes of Northeast India.
National Bureau of Fish Genetic Resources, Lucknow, India, p. 264.
(55) Ward, J V. and Stanford, J A. (1995). Ecological connectivity in alluvial river
ecosystems and its disruption by flow regulation. Regul River 11: 105–119.
(56) Warwick, R M., Clarke, K R. (1995). New ‘biodiversity’ measures reveal a decrease in
taxonomic distinctness with increasing stress. Marine Ecology Progress Series, 129:
301–305.
(57) Williams, P H., Humphries, C J., Vane-Wright, R I. (1991). Measuring biodiversity:
Taxonomic relatedness for conservation priorities. Australian Systematic Botany,
4(4): 665 – 679.
(58) Zar, J H. (1999). Biostatistical Analysis, 4th edn. Pearson Education, Singapore.

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Table 1: Seasonal occurrence of fish fauna in the Mahanada – Tangon - Punarbhaba river along with their Family name, Economic use and
threat status as per NBFGR report, 2010.
Sl Mons Autu Winte Sprin Sum Threat
Species Family Economic use
No. oon mn r g mer Status
1 Arius gagora Hamilton-Buchanan, 1822 Ariidae Fisheries: commercial * * * * a NT
2 Mystus cavasius Hamilton-Buchanan, 1822 Bagridae Fisheries: commercial * * * * * VU
3 Mystus tengara Hamilton-Buchanan, 1822 Bagridae Fisheries: commercial * * * * * LC
4 Mystus vittatus Bloch, 1797 Bagridae Fisheries, aquarium: commercial * * * * * LC
5 Rita rita Hamilton-Buchanan, 1822 Bagridae Fisheries: commercial * * * * a VU
6 Sperata aor Hamilton-Buchanan, 1822 Bagridae Fisheries: commercial; gamefish * * * a * VU
7 Chaca chaca Hamilton-Buchanan, 1822 Chacidae Aquarium: commercial * * * * a EN
8 Clarias batrachus Linnaeus, 1758 Clariidae Fisheries, aquaculture, aquarium: commercial * * * * * LC
Heteropneusti Fisheries, aquaculture, aquarium: commercial * * * a a VU
9 Heteropneustes fossilis Bloch, 1794
dae
10 Pangasius pangasius Hamilton-Buchanan, 1822 Pangasiidae Fisheries, aquaculture: commercial; gamefish * * a a a VU
11 Ailia coila Hamilton-Buchanan, 1822 Schilbeidae Fisheries: commercial * * * * a VU
12 Ailiichthys punctata Day, 1871 Schilbeidae Fisheries: commercial * * * * a VU
13 Clupisoma garua Hamilton-Buchanan, 1822 Schilbeidae Fisheries: commercial; gamefish * * a a a VU
14 Eutropiichthys vacha Hamilton-Buchanan, 1822 Schilbeidae Fisheries: commercial; gamefish * * * * a VU
15 Neotropius atherinoides Bloch, 1794 Schilbeidae Fisheries: minor commercial; aquarium: potential * * * * a LC
16 Silonia silondia Hamilton-Buchanan, 1822 Schilbeidae Fisheries: commercial; gamefish * * * * a VU
17 Ompok pabda Hamilton-Buchanan, 1822 Siluridae Fisheries: commercial * * * * * VU
18 Ompok pabo Hamilton-Buchanan, 1822 Siluridae Fisheries: commercial * * * * * EN
19 Ompok bimaculatus Bloch, 1794 Siluridae Fisheries, aquaculture, aquarium: commercial * * * * a EN
20 Wallago attu Schneider, 1801 Siluridae Fisheries: commercial; gamefish * * * * * VU
Lepidocephalichthys guntea Hamilton- Cobitidae Aquarium: commercial * * * * a VU
21
Buchanan, 1822
22 Botia dario Hamilton-Buchanan, 1822 Cobitidae Aquarium: commercial * * * * a VU
Amblypharyngodon mola Hamilton-Buchanan, Cyprinidae Fisheries: commercial * * * a a LC
23
1822
24 Cabdio morar Hamilton-Buchanan, 1822 Cyprinidae Fisheries: commercial * * a a a LC
25 Cirrhinus cirrhosus Bloch, 1795 Cyprinidae Fisheries: commercial * * * a a VU
26 Cirrhinus reba Hamilton-Buchanan, 1822 Cyprinidae Fisheries: commercial * * * * * VU
Cyprinidae Fisheries, aquaculture, aquarium: commercial; * * * a a NE
27 Cyprinus carpio carpio Linnaeus, 1758
gamefish
28 Esomus danricus Hamilton-Buchanan, 1822 Cyprinidae Fisheries: minor commercial; aquarium: commercial * * * * * VU
29 Labeo calbasu Hamilton-Buchanan, 1822 Cyprinidae Fisheries: commercial; aquaculture: commercial * * * * a LC

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30 Bangana dero Hamilton-Buchanan, 1822 Cyprinidae Fisheries: commercial; bait: usually * * a a a VU

31 Labeo rohita Hamilton-Buchanan, 1822 Cyprinidae Fisheries, aquaculture: commercial; gamefish * * * a a LC
32 Puntius chola Hamilton-Buchanan, 1822 Cyprinidae Fisheries, aquarium: commercial * * * * * VU
33 Systomus sarana Hamilton-Buchanan, 1822 Cyprinidae Fisheries, aquarium: commercial; gamefish * * * a a VU
34 Salmophasia bacaila Hamilton-Buchanan, 1822 Cyprinidae Fisheries: commercial * * * * * DD
Ambassidae Fisheries: minor commercial; aquarium: public * * * * * LC
35 Chanda nama Hamilton-Buchanan, 1822
aquariums
36 Parambassis ranga Hamilton-Buchanan, 1822 Ambassidae Fisheries: subsistence fisheries; aquarium: commercial * * * a a LC
37 Anabas testudineus Bloch, 1795 Anabanitidae Fisheries, aquaculture, aquarium: commercial * * * a a VU
Trichogaster fasciata Bloch and Schneider, OsphronemidaFisheries: commercial * * * * * NK
38
1801 e
39 Channa orientalis Bloch and Schneider, 1793 Channidae Fisheries, aquarium: commercial * * * a a NK
40 Channa punctatus Bloch, 1793 Channidae Fisheries, aquaculture, aquarium: commercial; * * * * * NK
41 Channa striatus Bloch, 1793 Channidae Fisheries, aquaculture, aquarium: commercial * * * * * NK
42 Glossogobius giuris Hamilton-Buchanan, 1822 Gobiidae Fisheries, aquaculture, aquarium: commercial * * * a a NT
Mastacembeli Fisheries, aquarium: commercial * * * * a VU
43 Mastacembelus armatus Lacepede, 1800
dae
44 Badis badis Hamilton-Buchanan, 1822 Badidae Fisheries: of no interest; aquarium: commercial * * a a a NK
45 Nandus nandus Hamilton-Buchanan, 1822 Nandidae Fisheries, aquarium: commercial * * * * * NK
46 Gudusia chapra Hamilton-Buchanan, 1822 Clupeidae Fisheries: subsistence fisheries * * a * a VU
Clupeidae Fisheries: highly commercial; aquaculture: * a a a a VU
47 Tenualosa ilisha Hamilton-Buchanan, 1822
experimental
48 Setipinna phasa Hamilton-Buchanan, 1822 Engraulididae Fisheries: commercial * * * a a LC
49 Chitala chitala Hamilton-Buchanan, 1822 Notopteridae Fisheries, aquaculture: commercial; gamefish * * * * a EN
50 Notopterus notopterus Pallas, 1769 Notopteridae Fisheries, aquaculture: commercial; gamefish * * * * a EN
Tetraodontida Fisheries: minor commercial; * * a a a NK
51 Tetraodon cutcutia Hamilton-Buchanan, 1822
e
52 Xenentodon cancila Hamilton-Buchanan, 1822 Belonidae Fisheries: minor commercial; aquarium: potential * * * a a VU
* = present, a= absent, EN= Endangered, VU= Vulnerable, NT= Near threatened, DD= data deficient, LC= Least Concerned, NK = Not known

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Table 2: Comparison with distribution pattern of finfishes in aquatic bodies of different

climatological region with present study.
Number of
Location References
finfishes
Punarbhaba River, West Bengal, India 52 Present study
Naaf River Estuary, Bangladesh 98 Chowdhry et al. 2010
Bhadra River, Western Ghats, India 56 Shahnawaz et al. 2010
Bhadra Reservoir, Karnataka, India 33 Thirumala et al. 2011
Copper Creek, Australia 14 Arthington et al. 2005
Itaipu reservoir, Brazil 85 Oliveira et al. 2004
Sharavati River, Western Ghats, India 51 Bhat 2003
Aghanashini River, Western Ghats, India 52 Bhat 2003
Bedti River, Western Ghats, India 63 Bhat 2003
Kali River, Western Ghats, India 53 Bhat 2003
Rivers of North Karnataka 20 Arunachalam et al. 1997
Neyyer River, India 33 Nair et al. 1989

Table 3: Seasonal variation in diversity estimators calculated from fish abundance data.
Seasons
Diversity Estimators Monsoon Autumn Winter Spring Summer
Dominance index 0.117 0.046 0.221 0.135 0.199

Shannon-Weaver index
3.7 3.63 3.44 2.99 2.43

Pielous Evenness index 0.94 0.92 0.91 0.86 0.84

Species richness index

28.23 66 32.01 18.69 13.08

Fish Species Richness 52 51 44 32 18

Table 4: Correlation coefficient between diversity indices with respect to fish abundance
in the Punarbhaba river.
Shanon-Weaver Evenness Dominance index Species richness
index index
Fish species richness 0.99* 0.97* -0.55* 0.72*
Shanon-Weaver 0.97* -0.52** 0.69*
index
Evenness index -0.44*** 0.63**
Dominance index -0.72*
* Correlation is significant at the 0.05 level (2 tailed).
** Correlation is significant at the 0.01 level (2 tailed).
*** Not significant

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Figure 1:: Map showing location of Mahananda-Tangon-Punarbhaba

Mahananda river system (MTPRS)
along with study sites.

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Figure 2: Abundance of existing fish family in different seasonal regime in MTPRS.

Belonidae
60
Tetraodontidae
Notopteridae
Engraulididae
50
Clupeidae
Nandidae
Mastacembelidae
40
Gobiidae
Channidae
Belontiidae
30
Anabantidae
Ambassidae
Cyprinidae
20
Cobitidae
Siluridae
Schilbeidae
10
Pangasiidae
Heteropneustidae
Clariidae
0
Chacidae
Monsoon Autumn Winter Spring Summer
Bagridae

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Figure 3: Pattern difference in a) Chao 1, b) Individual based rarefaction curve, c) Jacknife, d)

Bootstrap estimators from samples collected from MTPRS in various seasons.
60
60 a) b)
Species accumulation

Monsoon 40
40
Autumn
Winter
20 20
Spring
Summer

0 0
1 9 17 1 9 17
60 Number of samples 60 Number of samples
c) d)
a)
Species accumulation

40 40

20 20

0 0
1 9 17 1 9 17

Number of samples Number of samples

Figure 4: Dendrgram showing resemblance between seasonal fish assemblage structure based
on Bray_Curtis similarity index.

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Figure 5: Non-metric
metric multidimensional scaling (nMDS) of samples collected at different seasons
based on abundance of fishes using Bray–
Bray Curtis similarity at 20,40, 60 and 80 % of similarity.
M1-M24= monsoon, A1-A24=
A24= Autumn, W1-W24=Winter,
W1 Sp1-Sp24=
Sp24= Spring, Sm1-Sm24=
Sm1
Summer samples.

Figure 6: Seasonal variation on a) Phylogenetic diversity and b) Taxonomic distinctness of the

fish assemblage
ssemblage in the MTPRS.

a) 5000 93
Variation in Taxonomic Distinctness

65
Average Taxonomic Distinctness
Total Taxonomic Distinctness
Average Phylogenetic Diversity
Total Phylogenetic Diversity

4000 b)
62 91
2500

3000
59
and

89
2000
1500 56

87
53 1000

500 50 0 85
Monsoon Autumn Winter Spring Summer Monsoon Autumn Winter Spring Summer
Total phylogenetic diversity Average phylogenetic diversity Total Taxonomic distinctness AverageTaxonomic distinctness
Variation in Taxonomic distinc

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Figure 7:: Funnel plot showing (a) Average Taxonomic Distinctness (∆( +) and (b) Variation in
Taxonomic distinctness of the fish assemblage in the MTPRS. Thin line in middle indicate mean
of 1000 simulations confirming theoretical unbiasedness. Continuous line indicates 95%
probability limits for each.

Variation in Taxonomic Distinctness

Average Taxonomic Distinctness

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