Fish
diversity and assemblage structure in Ken River of Panna landscape, central
India
J.A. Johnson 1, Ravi Parmar 2,
K. Ramesh 3, Subharanjan Sen 4 & R. Sreenivasa Murthy5
1,2,3,4 Wildlife Institute of India, Post Box # 18, Chandrabani,
Dehradun, Uttarkhand 248001, India
5 Panna National Park, Madhya Pradesh 488001, India
Email: 1jaj@wii.gov.in (corresponding author), 2 envoravi@gmail.com, 3ramesh@wii.gov.in, 4 ssen@wii.gov.in, 5 rseenu60@gmail.com
Date
of publication (online): 26 October 2012
Date
of publication (print): 26 October 2012
ISSN
0974-7907 (online) | 0974-7893 (print)
Editor:Neelesh
Dahanukar
Manuscript
details:
Ms
# o3024
Received
29 November 2011
Final
received 28 September 2012
Finally
accepted 05 October 2012
Citation: Johnson, J.A., R.
Parmar, K. Ramesh, S. Sen & R.S. Murthy (2012). Fish
diversity and assemblage structure in Ken River of Panna landscape, central
India. Journal of Threatened Taxa 4(13): 3161–3172.
Copyright: © J.A. Johnson, Ravi
Parmar, K. Ramesh, Subharanjan Sen & R. Sreenivasa
Murthy 2012. 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 and Author
Contribution: See end of this article.
Acknowledgements:We are grateful to the Director and Dean, Wildlife Institute of India for their
support and encouragement. We are also thankful to the Madhya Pradesh Forest
Department for permitting us to carry out this study. Constructive comments
from anonymous reviewers in improving the quality of the manuscript are
sincerely appreciated.
Abstract: Fish diversity
and assemblage structure in relation to habitat variables were studied in 15
sites in Panna landscape, central India. The sampling was performed between February–April
2009. Fifty species of fishes belonging
to 32 genera, 15 families and four orders were recorded from the study
area. Cyprinids were the dominant
assemblage members in all study streams (abundance ranges from 56.6–94.5
%). The cyprinid Devario aequipinnatus and the snakehead Channa gachua had highest local dominance
(80% each) in Panna landscape. High
Shannon and Margalef’s diversity was recorded in Madla region of Ken
River. Similarity cluster analysis
explained the study sites along Ken River (Gahrighat, Magradabri and Madla) had
similar faunal assemblage. Canonical Correspondence Analysis (CCA) was performed to study the
species association with a set of environmental variables. The CCA revealed that cyprinid abundance
was associated with stream order, deeper habitat, flow and water temperature.
Keywords: Assemblage
structure, central India, environmental variables, fish diversity, Ken River, Panna landscape.
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Introduction
Freshwater
ecosystem and their resources are an indispensable part of human life and
activity, and health of those freshwater ecosystems is visible in the wellbeing
of the fish assemblage they support. In lotic environment, the diversity, community structure and species
assemblages are influenced by various biotic and abiotic variables (Minns
1989). Water in these habitats may
look homogeneous but they are separated by various environmental factors such
as temperature, depth, current and substrates into a great variety of habitats
(Kottelat & Whitten 1997). Each
habitat has its unique fauna which is adapted to the
various abiotic features of that habitat. Identification of such abiotic gradients that result in structuring of
fish assemblages is one of the main challenges for a fish ecologist. The species richness of a stream within
a river basin may be influenced by local conditions and stream order along a
watershed (Grenouillet et al. 2004). Williams et al. (2003) reported that the historical formation of
a river basin determines the structure of the fish community. The Panna landscape in north-central
Madhya Pradesh is one of the historical landscapes, located on the Vindhyan
Range within the Biogeographic Province 6A Deccan Peninsula - Central Highlands
(Rodgers et al. 2002). A
number of perennial and seasonal streams originate from this landscape and
drain into the Ken River. According
to Satpura hypothesis (Hora 1950), the catchment areas of the Ken River basin
(Vindhyan Hills) in central India play a key role in the migratory route of
several Western Ghats fishes. The
catchment area of this river basin is characterized by extensive seasonal and
annual fluctuations of environmental variables.
Variation
in habitat variables such as flow, depth, substrate and water quality may have
a significant impact on both assemblage structure and resource
availability. The influence of
habitat structure and complexity on fish assemblage structure has been tested
mostly in North American streams (Gorman & Karr 1978; Schlosser 1982, 1985;
Capone & Kushlan 1991) and Australian streams (Bishop & Forber 1991;
Pusey et al. 1993). In
India, few studies have been initiated to document the fish diversity and
assemblage structure along the environmental gradients (Johnson 1999;
Arunachalam 2000; Bhat 2003; 2004; Sreekantha et al. 2007; Shahnawaz et al.
2010). Basic information is also
available on fishes of Ramsagar reservoir and fish assemblages of Betwa River,
Khan and Khashipra rivers of Madya Pradesh (Ganasan & Hughes 1998; Garg et
al. 2007; Lakra et al. 2010). However, studies on fish diversity and assemblage structure are still
rudimentary in central and northern India. Therefore, the present study is an attempt to document the fish
assemblage structure in relation to environmental variables of the Ken River of
Panna landscape in central Indian highland.
Material and Methods
Study area
Ken is one the major rivers
of Bundelkhand region of central India, and flows through the Panna National
Park in Madhya Pradesh (Image 1). It is one of the sub-basins of the Yamuna, and the important tributaries
of this river are Sonar, Bearma, Bewas, Kopra, Urmil and others. Among these, the Sonar is the largest
tributary that rises in the Vindhyan Hills in the southwest of Sagar District
and flows through its valley in Damoh District and it travels a distance of
427km and joins the Yamuna at Chilla Village, near Fatehpur in Uttar Pradesh
(Jain et al. 2007). It cuts through
the landscape from south to north and forms the famous Ranneh Falls, which is
located in the Crocodile Park (Ken Ghariyal Sanctuary) of Panna National Park. This river is known as an angler’s
paradise, because the king of freshwater fish Mahseer Tor tor is found
in abundance. The vegetation type
in Panna landscape is characteristic of tropical dry deciduous element. In terms of biome characteristics, it is
classified as ‘high-rainfall dry deciduous forest’ and is largely dependent on
monsoon rainfall during July–September, which usually fluctuates within
the range of 600–1100 mm (Jayapal et al. 2007). Following the monsoon, there is a cool
season until February, followed by dry summer when the temperature often
exceeds 450C (Karanth et al. 2004). Water is a limiting factor during this
season, even though, some stretches of streams inside the park hold some water
as isolated pools. But the main
channel of Ken River retains natural deep pools, rapids and cascades with
heterogeneous riparian cover during summer, which probably provides shelter for
fish when most of the other streams dry up. A total of 15 sites on streams/ rivers
of the Ken River basin within Panna National Park were selected for the present
study (Fig. 1) and the summary of site description isgiven in Table 1. The sampling was
undertaken between February–April 2009, since
during this period most of the streams inside the park retain minimum surface
flow.
Data collection
Fish sampling was performed
in different habitats such as pools, riffles, runs and cascades in 100m reach
of all study sites, using monofilament gill nets of different mesh sizes
(10–34 mm), drag, scoop and cast net. Fish sampling protocol followed the method
of Johnson & Arunachalam (2009). After collection, fish were examined, counted and released back into the
system. A few specimens of
unidentified species were preserved in buffered formalin (10%) and transported
to the laboratory for species confirmation. Species identification and confirmation
were carried out using available literature (Talwar & Jhingran 1991;
Jayaram 1999) and the species valid nomenclatural names were adopted as per the
Catalogue of Fishes of the California Academy of Sciences (Eschmeyer &
Fricke 2011). At each sampling
site, a set of the following environmental variables wererecorded: stream order, altitude, stream width (m), water depth (cm), velocity
(m/Sec), water temperature (0C), conductivity (µS/cm) and riparian
cover (%). Riparian cover was
estimated using spherical densiometer. Sampling protocol for habitat variables followed Pusey et al. (1993).
Analysis
Information on structure of
fish assemblages was extracted by adopting different univariate indices, namely
local distribution index, Margalef’s species richness index and Shannon
diversity index. The calculation of
these indices followed the methods of Johnson & Arunachalam (2009), and
Muchilisin & Azizah (2009). Local distribution index was calculated by D= (Ni.st/N.st) x 100, where
Ni.st is total number of sites where the fishes are found; N.st is total number
of sites. Margalef’s species
richness was calculated using the equation R = (S-1)/ln N, where S is the
number of species, N is the total number of individuals. The Shannon index of diversity was
obtained by the following equation H’ = -∑pi ln (pi), where pi = ni/N; ni is the number of individuals of ‘i’th species and N = ∑ni.
The indices were used to compare the species distribution, richness and
diversity across the study sites. Quantitative data of species along with the number of individuals
belonging to each species were used to calculate percent similarity index using
Bray-Curtis similarity index based on Padhye et al. (2006). Dendrograms were constructed to
understand the similarity of fish assemblage structure between the sampling
sites. This was done using
Bray-Curtis similarity index using non-transformed species abundance data
(Anderson 2001) using PAST programme. Further, the species abundance and environmental variables with
separated sites were submitted to Canonical Correspondence Analysis (CCA),
which is a direct gradient ordination technique that extracts the best
synthetic gradients from field data on biological communities and environmental
features: it forms a linear combination of environmental variables that
maximally separate the niche of the species (ter
Braak & Verdonschot 1995). It
is also a powerful exploratory tool for simplifying complex data sets and has
the advantage over integrated analysis of both species and environmental data
at each site (Taylor et al. 1993). In order to reduce the complexity of ordinance biplot, only cyprinid
species were included in CCA. The
resulting species abundance-environmental variables biplot is an ordination
diagram in which species and sites are represented by points with respect to
the supplied explanatory variables, represented by arrows. The arrows point in the direction of
maximum variation in value of the corresponding variable. The arrow of a variable runs from the
centre of the diagram to an arrow head, the coordinates of which are the
correlation of the variable with axes (ter Braak 1986; ter Braak &
Verdonschot 1995). The CCA was obtained with XLSTAT® 2012 version programme.
Results
A total of 50 species of
primary freshwater fishes belonging to 32 genera, 15 families and four orders
were recorded from the study area (Table 2). Among the species, Devario
aequipinnatus and Channa gachua had highest local dominance (80% for
each) followed by Esomus danricus (66.6%) and Garra mullya(60%). They were represented in
most of the study sites. The
Mahseer Tor tor (Image 2) was recorded from Gahrighat, Magradabri
and Madla in Ken River and also in Pandav Fall Stream. Number of species, species abundance,
cyprinid abundance, Shannon diversity and Margalef’s richness index for study
sites are given in Table 3. The
total number of species as well as abundance was highest in the Madla area of
Ken River, whereas the lowest was recorded in Mahuapani Stream. Similarly, the Madla area of Ken had a
high Shannon diversity index (3.48), whereas the Mahuapani stream registered a
low Shannon diversity index (0.99). Cyprinids were the dominant members of the assemblage structure in the
study area and comprised 56.6–94.5 % in the assemblage structure. The maximum cyprinid population was
recorded from Jamunahi stream, while low cyprinid population was observed in
Nararan stream (Table 3). Among the species, the distribution
of 14 species (Acanthacobitis botia, Clupisoma montana, C. garua,
Crossocheilus latius, Devario devario, Garra gotyla, Glyptothorax telchitta,
Labeo angra, L. rohita, Lepidocephalichthys guntae, Ompok pabda, Osteobrama
cotio, Pseudambassis baculis and Salmosphasia balookee) werereported in the Himalayan river system and they were not reported from
peninsular India. Similarly, three
species commonly found in rivers of peninsular India (Nemacheilus denisoni,
Rita gogra and Salmosphasia boopis) were recorded for the first time
in the Yamuna River basin. Thus,
short notes on meristic and morphometric features of new records are given at
this section.
Notes on new records (Image 3
a–c)
Salmosphasia boopis: Body elongate and laterally
compressed. Body scales large,
40–41 scales present across the lateral line. Dorsal fin inserted well anterior to the
origin of anal fin. Fin rays counts: dorsal - iii/7; pectoral - i/14; pelvic -
i/8; anal - iii/12.
Nemacheilus denisoni: Body loach-like, marked with
12 broad vertical bands and a black band at the base of caudal fin. A black spot present at base of origin
of dorsal fin and rows of small spots also present on dorsal and caudal
fins. Fin rays counts: dorsal -
iii/8; pectoral - i/8; pelvic - i/7; anal - iii/5.
Rita gogra: Head depressed, occipital
process subcutaneous, extends to predorsal plate. Dorsal and pectoral fins bear strong
osseous spine. Barbelsthree pairs, maxillary barbel extends to operculum. Mandible reaches base of pectoral fin
and nasal barbel short. Fin rays
counts: dorsal - i/6; pectoral - i/10; pelvic - i/7; anal - iii/9.
Cluster analysis of species
composition in Ken River basin revealed that fish assemblages of Ken River had
two distinct clusters based on the Bray-Curtis similarity (Fig. 2). The sites along Ken River (Gahrighat,
Magradabri and Madla) had more similar faunal assemblage and they were grouped
together. The rest of the streams
that drained into Ken River had a similar assemblage and they formed a major
group in the cluster. Further, the
streams associated with Ken River in the northeastern region of Panna landscape
such as Khaiya, Ghatera, Bargadi, Silatanala, Judinala and Imalia had a similar
assemblage and they formed a separate cluster within the major group. At the same time Nararan, Gehranala and
Pandav Fall streams had distinct fish assemblages whichwere grouped separately.
Environmental variables vs. fish abundance
Most environmental characters
measured exhibited a high level of differences across the fifteen study sites
(Table 4). River sites (Gahrighat,
Magradabri and Madla) had more water depth and channel width than other
sites. High velocity was recorded
in Gahrighat (0.68m/sec) followed by Magrdabri (0.52m/sec). In contrast, high percentage
of canopy cover (94%) and low water temperature (19.80C) wereobserved in Imalia stream. The
species and site scores biplot based on CCA of the cyprinid fish composition
displayed 12.1% of weighted variance in the abundance and 49% in weighted
averages and class total of species with respect to the environmental
variables. The eigenvalues of axis
1 and 2 accounted 0.62 and 0.20 respectively. Indeed, the biplot generated by CCA
suggested that stream order, flow, depth and width
were the most important variables for the first axis. Flow was a very important variable for
the second axis, although riparian cover and altitude were still influenced
(Fig. 3). The biplot of the species and site score produced from CCA show the
distribution of species and sites in ordination space (Fig. 3). In this plot 27 cyprinid species have
been depicted to provide insight into their composition and distribution. The fish abundance sites such as
Gahrighat, Magradabri and Madla along Ken River (site label 1, 12 & 14 in
Fig. 3) were associated with more deeper habitat, flow, stream order and
temperature, whereas the abundance were not influenced by altitude,
conductivity and riparian cover (Fig. 3). Further, it also explained that the species such as Tor tor, Salmophasia
boopis, Salmophasia balookee, Salmophasia bacaila, Labeo angra andGibelion catla were displaced high weighted average for flow, whereas Bangana
dero, Labeo calbasu and Puntius sarana scored a high weighted
average for width, depth and temperature. The distribution of Danio rerio, Devario
aequipinnatus, Esomus danricus and Rasbora daniconius were
influenced by altitude, riparian cover and conductivity.
Discussion
Ken River in Panna landscape
has a diverse fish fauna of high conservation importance. When compared to other sub-basins of the
Yamuna River basin, Ken River has lesser species richness than that of Champal
basin and Betwa, where 71 and 60 species of fishes were reported respectively,
but the fish assemblages were similar to that of these neighbouring basins
(Dubey & Verma 1959; Vyas et al. 2012). In this study, cyprinids dominate the
assemblage structure in Panna landscape as they occupy all possible habitats
due to their high adaptive variability (Johnson & Arunachalam 2009). Three of the cyprinids species such as Devario
aequipinnatus, Esomus danricus and Garra mullya were widely
distributed in most of the study sites and they also have widespread
distribution in India (Talwar & Jhingran 1991; Jayaram 1999) and they are a
common and abundant species in Indian waters. Such extensive distribution and their
common high abundance suggest that most of these species are capable of
tolerating a wide range of environmental conditions (Pusey et al. 1993). However, in the present study a comparatively
low percentage of cyprinid population was recorded in Nararan Stream. This is mainly due to the introduction
of Channa gachua and C. punctatus in the temple tank, which in
turn contribute to the assemblages structure of this
stream. A noteworthy observation is
the occurrence of Salmophasia boopis, Nemacheilus denisoni and Rita gogra (known so far from the rivers of peninsular India)
in Vindhyan ranges (Yamuna River basin) and this is interesting in the context
of Ichthyogeography. The Salmophasia
boopis was reported from streams/ rivers originating from
the Western Ghats of Tamil Nadu, Kerala, Karnataka and Maharashtra(Dahanukar 2011a). Similarly, Nemacheilus
denisoni is also widely distributed in the Western Ghats
rivers (Menon 1987). On the other hand, Rita gograwas reported from Krishna, Godavari and Narmada river basin of Deccan regions
and distributed in Maharashtra, Karnataka, Andhra Pradesh, Madhya Pradesh and
Chhattisgarh states (Dahanukar 2011b). These distribution records of peninsular forms in Vindhyan Ranges recollect
the possibility of prehistoric river valley modifications as proposed by Hora
(1950). However, number of species
per family recorded in Ken River basin revealed that more number of species
were recorded belonging to the cyprinid family as compared to other families,
which clarify that speciation occurred in this landscape in recent times.
Another interesting finding
is the occurrence of a viable population of the Mahseer Tor tor in Ken
River of Panna landscape. In the
present study this species was recorded from Gahrighat, Magradabri and Madla
along the Ken River and Pandav Fall stream. It is a mighty game fish, which has
provided a worthwhile source of sport for international anglers. It reaches up to 78kg or more and there
is a record of anglers having captured a 45kg fish (Menon 1999; Rayamajhi et
al. 2010). Twenty years ago, they
figured prominently in commercial catches in certain stretches of the Narmada
and Tapti rivers, but the landings are reported to have declined remarkably in
recent years (Talwar & Jhingran 1991). Presently, size ranges from 1.5–2 kg have been reported from its
native ranges and this species distribution is restricted only in certain
pockets of Protected Areas in Narmada, Tapti, Betwa and Chambal rivers in
Central India. The reduction in
population is mainly due to degradation of feeding and breeding habitat due to
construction of barrages and dams that in turn affect the breeding migration of
this species towards upstream and finally that leads to patchy and fragmented populations
in different rivers. This patchy
distribution and selective harvesting of this big sized barb have also led to
the disappearance of this species from its native ranges. However, in the present study we noted
different life history stages of Tor tor in Gahrighat region of Ken
River indicating that this species breeds upstream in the main Ken River, which
provides some promise for survival of this species. However, further study on habitat use,
spawning site selection and life history traits of this species in Ken River is
warranted.
The present study also
revealed that the main Ken River had high diversity of fish compared to its
associated streams, which shows that the diverse habitat conditions such as
rocky and deep pools, velocity, etc. in Ken River support great variety of
fauna (Kaemingk et al. 2007). Moreover, it is a well established fact that
there is occurrence of higher species richness at the confluence of tributary
streams with the main river than in the tributary streams (Falke & Gido
2006). Further, the low diversity
of fish documented in streams associated with Ken River is mainly due to the
drying of streams during summer.
In addition to that, a
variety of factors like water quality, habitat availability, flow variability
and nutrient supplies from riparian habitats control the abundance and
distribution of stream fishes. Such
environmental variables are easier to predict than other biotic variables like
predation and competition. Earlier,
single water quality parameter was used to correlate with fish abundance
(Echelle et al. 1972; Matthews 1987). Hawkes et al. (1986) hypothesised that the combination of different
water quality parameters is likely to operate “in concert with each other as a
multivariate system and not as isolated unvariate variables”. In this study, the combinations of eight
environmental variables were used to correlate with cyprinid abundance among
the study streams. The CCA analysis
revealed that the environmental variables such as water depth, flow, water
temperature and stream order substantially influence fish assemblage
structuring in the Ken River basin. These variables have previously been considered important factors in
structuring fish assemblage (Matthews 1998; Angermeier & Winston 1999;
Marchetti & Moyle 2001; May & Brown 2002). In harsh and variable environments such
as streams and rivers, abiotic factors are likely to play an important role in
determining fish assemblage structure.
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