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Changing foodwebs of Indian aquatic ecosystems under the
threats of invasive species: An overview
Article in Aquatic Ecosystem Health and Management · September 2021
DOI: 10.14321/aehm.024.02.06
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Changing foodwebs of Indian aquatic ecosystems under
the threats of invasive species: An overview
Preetha Panikkar1, M. Feroz Khan1, U.K. Sarkar2,* B.K. Das
Regional Centre of ICAR-CIFRI, Hessarghatta, Bangalore
2
ICAR-CIFRI, Barrackpore, Kolkata-700120
*Corresponding author: usarkar1@rediffmail.com
Biological invasions are a worldwide threat to the aquatic systems and have the potential to homogenize
entire foodwebs and shift species abundance distributions to more skewed ones. Invasion impacts include
effects on the foodweb structure and ecosystem functioning leading to a loss in native fish biodiversity
and commercially important fisheries in many open water systems. The impacts of invasives are generally
devastating as they spread in the foodweb with each species potentially interacting with multiple species.
The foodweb modeling studies conducted in different inland aquatic ecosystems show that not all exotics
cause a detrimental effect on the resilience of an ecosystem. Information on the foodweb structure and
ecosystem properties is a prerequisite for formulating management protocols for conserving biodiversity,
enhancement programmes and sustaining fisheries. The present paper reviews the impacts of invasive
fishes in Indian aquatic ecosystems in the context of a changing foodweb scenario due to exotic fish species
invasions. The information generated here could be applied for future research on similar ecosystems for
deducing management actions.
Keywords: trophic impact, ecosystem properties, energy flows, resilience
Introduction
India has different climatic zones that support
diverse aquatic habitats rich in aquatic biodiversity
(Sarkar et al., 2017). In recent decades, the
introduction of invasive fish is considered to be a
major cause of species extinction posing a major
ecological threat to the inland aquatic systems
through the displacement of native fauna and
alteration in the dynamics (Singh et al., 2014;
David et al., 2017). The invasions will profoundly
influence the function of the whole foodweb,
and as a consequence the trophic relationships
are impaired. In India, several studies have
highlighted the occurrence of many invasive
flora and fauna in inland ecosystems and also
reported the consequences of the invasion at a
regional scale (Paliwal and Bhandarkar, 2014;
24
Singh et al., 2014; Sandilyan et al., 2018; Sarkar
et al., 2017). Many studies have also shown that
invasive species are a threat to endemic and
threatened species in ecosystems throughout the
world including important biological hotspots
(Denslow, 2007; Raghavan et al., 2008). Biological
invasion generally causes a decrease in the native
biodiversity and population abundances (Kenis
et al., 2009; Vilà etal., 2011; Singh et al., 2014),
and affects the ecosystem functioning (Wardle et
al., 2011; Simberloff et al., 2013). Romanuk et
al., (2009) explored several foodwebs and invader
properties that predispose a species to successfully
invade and establish among the local communities.
The alterations in an ecosystem arising from
invasions usually lead to native species extinctions
and a decline in local species richness.
Exotic fish have invaded inland aquatic systems
Aquatic Ecosystem Health & Management, 24(2): 24–32, 2021. Copyright © 2021 Aquatic Ecosystem Health & Management Society.
ISSN: 1463-4988 print / 1539-4077 online. DOI: 10.14321/aehm.024.02.06
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Panikkar et al. / Aquatic Ecosystem Health and Management 24 (2021) 24–32
25
Fig. 1. Map of India depicting the geographical locations of the reservoirs.
in India from the aquarium traders and aquaculture
farms (Krishnakumar et al., 2009; Singh and Lakra,
2011; Panikkar et al., 2015b). Many of the exotic
fishes have established a good breeding population
in inland systems and caused damage to the aquatic
diversity (Singh and Lakra, 2011; Sandilyan,
2016). The impacts on ecosystems and biodiversity
by invasives result in huge economic (Pimentel et
al., 2000) and socio-cultural losses (Lockwood et
al., 2007). The success of an invasion has a trade
off with increased trophic interactions (Baiser
et al., 2010). The number of connections the
invader possesses influences its performance in the
foodweb invasion. A few studies on this scale have
been undertaken in the reservoir ecosystems over
the last decade describing trophic structure and
foodweb interactions for studying the impacts of
invasions (Khan and Panikkar, 2009; Bijukumar et
al., 2015; Gallardo et al., 2016). These studies have
shown a linkage between foodweb interactions and
invasion.
We conducted an investigation on the empirical
studies of a few tropical reservoir ecosystems in
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India on how fish -invasions alter foodwebs in
terms of ecosystem attributes and functions. The
status of the impact of invasive fish on the aquatic
ecosystems in India is synthesized and reviewed
from the perspective of foodwebs. These studies
provide information on the intensity of the impact
of invasives and also address the knowledge gaps
for management through an ecosystem approach.
Trophic interactions between
invasive fishes and other fish
groups in the foodweb
In India, few studies have been conducted to
model the freshwater ecosystems using foodwebbased approach. These studies have reported the
presence and impact of exotic fishes on the other
fishes in the ecosystem. The geographical locations
of these freshwater ecosystems in different states of
India are illustrated in Fig. 1. Invasions have been
found to profoundly influence the function of the
whole foodweb and as a consequence the trophic
26
Panikkar et al. / Aquatic Ecosystem Health and Management 24 (2021) 24–32
interactions are impaired. Invasive fishes are
considered to alter the foodweb of an ecosystem by
reducing the food availability for the native fishes
through competition for food thereby reducing the
native populations and establishing themselves in
the ecosystem. Impacts of invasives are generally
devastating as they spread across the foodweb, as
each species interacts with a number of different
species. We expect negative consequences for the
same reason. Khan and Panikkar (2009) developed
a foodweb model (Fig. 2) of foodweb interactions
in Kelavarapalli, a small reservoir in Tamil Nadu,
India to quantify the energy flows between different
trophic levels in the foodweb and to assess the
impact of invasive fishes on other fishes in the
foodweb.
The introduced Nile Tilapia, Oreochromis
niloticus and the native Etroplus suratensis in the
reservoir were found to have the highest preypredator overlap (Fig. 3) that indicated a high
competition for similar food resources. Indirect
trophic effects through exploitation and competition
also were found to affect the native groups but
the invasives often have been considered to have
positive bottom up effects on secondary consumers
and apex predators in the foodweb. Predation is
considered to cause the most remarkable damage in
invaded aquatic systems, at times cascading down
to the primary producers. Due to the complex preypredator interactions, invasions have been found to
greatly modify the structure of the entire foodweb.
The mixed trophic impact (MTI) studies
which is a form of a sensitivity analysis in
foodweb interactions, explain the influence of one
ecological group on another (Majkowski, 1982).
The MTI study in Kelavarapalli Reservoir, India
indicated that an increase in the biomass of the
highly invasive fish, the African walking catfish
(Clarias gariepinus) would adversely affect
almost all the fish groups in the reservoir. The
Mozambique Tilapia (Oreochromis mossambicus)
negatively impacted the indigenous catfishes
whereas the Nile Tilapia (O. niloticus) was found
not to adversely affect any one of the fish groups
(Khan and Panikkar, 2009). A mass balance model
for the Wyra reservoir in Telangana, India which
has O. mossambicus in the fishery was developed
(Panikkar and Khan, 2008). It is reported that the
tilapia has a negative impact on a few fish groups
such as the murrels, indigenous catfishes and barbs.
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Exotic carps, Cyprinus carpio and Grass Carp,
Ctenopharyngodon idella were reported to have
a negative impact on murrels and minor carps in
Hemavathy reservoir in Karnataka (Khan et al.,
2015). Oreochromis mossambicus in Vellayani
Lake modeled by Bijukumar et al (2015) was
found to have a strong adverse impact on barbs and
carpels, and a slight impact on the other native fishes
like the needle fishes, gobiids, and nandids. An
increase in the biomass of Oreochromis sp. would
have negative impact on many of the indigenous
fishes, Puntius sp., Trichogaster fasciatus, Channa
sp., Chanda ranga, Chanda nama, Batasio batasio
in Bakreswar reservoir in West Bengal, India
(Banerjee et al., 2016).
Fig. 2. Diagrammatic representation of foodweb of Kelavarapalli
reservoir ecosystem in India dominated by exotic fishes (Khan
and Panikkar, 2009) P-Production; Q- consumption are the
trophic flows in t km-1 year-1.
Fig. 3. Prey-predator niche overlap plot in exotic fishes
dominated in Kelavarapalli reservoir, India (Khan and Panikkar,
2009)[The ecological groups in box II have no overlap and are
independent for both prey and predators. Groups in box III have
a high overlap for both preys and predators. The numbers 2 to
10 represents different ecological groups as follows (2) Exotic
African catfish (3) Indigenous catfishes (4) Snake head fishes
(5) Exotic Mozambique Tilapia (6) Exotic Nile Tilapia (7) Pearl
spots (8) Major carps (9) Dipterans (10) Zoo-benthos].
Panikkar et al. / Aquatic Ecosystem Health and Management 24 (2021) 24–32
Effect of invasive fishes on the
foodweb structure
Foodweb structure is an important constituent
in understanding any ecosystem and ecologists
around the world are investigating on this aspect
(Christensen et al., 2005; Khan and Panikkar, 2009;
Figueroa et al., 2009; Panikkar et al., 2015a,b;
Murphy et al., 2019). The ecological summary
indices for different foodwebs of the aquatic
ecosystems studied in India are presented in the
Table 1.
The introduction of fish in a system through
stock supplementation can cause substantial
effects on the foodweb (Mbabazi et al., 2004). A
mass-balanced model study in Hemavathy, a large
reservoir in Karnataka, southern state of India
was conducted for two periods (before and after
the introduction of fish seeds) using ECOPATH
to assess the impact of non-native fish seed
stocking on the foodweb of the ecosystem (Khan
et al., 2015). In this particular reservoir, nonnative carps Cyprinus carpio and the grass carp,
Ctenopharyngodon idella were stocked along with
the Indian major carps (Catla catla, Labeo rohita
and Cirrhinus mrigala). The biomass of the non –
native carp group was estimated as 0.447 and 2.56 t
km-1 year-1 in the fishery during 1982-83 (pre-stock
phase) and 2002-03 (post-stock phase) respectively
(Khan et al., 2015). The mixed trophic impact
routine showed that an increase in biomass of the
non-native carps showed a slight negative impact
on the eels but did not impact any other fish groups
in the ecosystem.
In the Hemavathy reservoir of Karnataka, India
(Khan et al., 2015), the trophic flows basically
happened in the first four trophic levels and the food
-web was characterized by the predominance of low
trophic level organisms during the two different
phases which were modelled. The comparative
trophic models showed a variation in the foodweb
structure and system summary statistics in the
reservoir during the two phases despite the changes
in biomass of the natives and non-native fishes
(Khan et al 2015). Connectance index (CI) and
omnivory index (OI) are ecological indicators that
are correlated with the maturity of the reservoir
ecosystem as described by Odum (1971). The OI
and CI values showed a decline after the non-native
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27
fish seed stocking in Hemavathy reservoir and the
difference in these values indicate that the poststock phase of the ecosystem displayed less weblike features which is a positive impact of fingerling
stocking. The Vellayani lake in Kerala which has
an invasive population of O. mossambicus modeled
by Bijukumar et al (2015) had the highest OI for
the newly introduced Indian major carps (0.38)
indicating that they exploit the niches available in
the ecosystem winning the competition with other
fishes.
Impact of invasive fish on the
‘health’ of an ecosystem
The health of an ecosystem has been measured
using Odum’s attributes of system maturity
(Christensen, 1995). The Ecopath model provides
various tools which help in developing various
attributes that can be used in assessing the state
of health of an ecosystem. A healthy ecosystem
is not easily vulnerable to disturbances, both
anthropogenic and natural. A comparative analysis
of different time periods for the same system
can help managers to assess the health status of
the system (Christian et al., 1996). The network
analysis which indicate the health status consisting
of algorithms that describe the system attributes
through foodweb dynamics (Christensen, 1998).
These are effective tools in assessing the impacts of
exotic species induced perturbation on ecosystems
(Miehls, 2009). A comparative statement of the
ecosystem health indicators for the different inland
ecosystems modeled in India are presented in
Table.1. This section gives an account of various
ecological indicators that helps in assessing the
health of an ecosystem.
As described in the previous section CI and OI
are also indicators of ecosystem maturity and these
values tend to increase as the ecosystem matures
(Odum, 1971). The system’s OI of Vellayani
lake was 0.205 suggesting a low omnivory in
the system. The CI was 0.395 indicating a high
diversity of trophic compartments that can be
expected in tropical lakes with rich biodiversity.
These indicators point to the immaturity of the lake.
The informational indices also called the flow
indices, such as, the ascendency and overhead
have also been shown to be related to maturity
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14
13
Kelavarapalli
430
reservoir, Tamil Nadu
9540
9162
30
331.2
6740
Ravishankarsagar
reservoir,
Chattisgarh
Hemavathy reservoir,
Karnataka
Veli Lake, Kerala
Vellayani Lake ,
Kerala
Bakreswar Reservoir,
West Bengal
7.5
3.64
1.72
O. mossambicus
Oreochromis sp.
9.73
21.22
12.26
72.58
5.33
Cyprinus carpio,
Ctenopharyngodon
idella
None
80.33
76.07
16.24
TPP/
TB
10.36
7.78
Oreochromis
niloticus,
O. mossambicus,
Clarias gariepinus
None
1.367
Oreochromis
mossambicus,
TPP/
TR
0.05
0.021
0.012
0.005
0.005
0.006
0.011
TB/
TST
0.176
0.395
NA
0.264
0.299
0.296
0.246
CI
0.109
0.205
NA
0.112
0.162
0.156
0.191
OI
8.44
2.52
NA
NA
1.99
3.04
NA
FCI
3.08
2.24
NA
3.75
2.29
2.31
3.42
MPL
25
39.87
NA
38.2
46.8
35.92
24.75
75
60.13
NA
61.8
53.2
64.08
75.25
Ascen- Overdancy head
Banerjee et
al 2016
Bijukumar et
al., 2015
Aravindan,
1993
Khan et al.,
2015
Panikkar et
al., 2015a
Khan and
Panikkar,
2009
Panikkar and
Khan, 2008
Reference
(Abb: TPP- Total Primary production, TR- Total Respiration, TB- Total Biomass, TST -Total System Throughput, CI-Connectance Index,
OI- Omnivory index, FCI-Finns cycling Index, MPL-Mean Path Length, NA -Not available)
27
17
14
13
17
1074
Wyra reservoir,
Telengana
No. of
Exotic fish in the
Ecological system
groups
Area
(ha)
Inland ecosystem
Table 1. Ecological indicator for ecosystem health assessment of the inland aquatic systems modelled in India.
28
Panikkar et al. / Aquatic Ecosystem Health and Management 24 (2021) 24–32
Panikkar et al. / Aquatic Ecosystem Health and Management 24 (2021) 24–32
of an ecosystem (Christensen, 1995). The highest
overhead values were obtained for Wyra followed
by Kelavarapalli reservoir ecosystems which has a
composition of exotics in the fishery (Table 1).
Finn’s cycling index, as described by Finn
(1976), is the fraction of the recycled throughput
of an ecosystem and is an indicator of the system’s
resilience and stability (Odum, 1969). The high
value of FCI represents a mature and healthy
ecosystem and low values are typical of vulnerable
ecosystems that are sensitive to nutrient input
alterations as reported by Christensen and Pauly
(1993). FCI of Vellayani lake was 2.52 % of the
total system throughput of the system and the MPL
was 2.24 which further indicates the developing
stage of the lake. Ravishankarsagar reservoir
with an FCI of 1.99 is indicative of a developing
ecosystem. FCI of Bakreswar reservoir system
was relatively very high (8.44%) showing least
vulnerability compared to the other ecosystems.
The flow of energy through a foodweb is
expressed by Finn (1976) in terms of Mean Path
Length (MPL) which is the number of trophic
interactions of each ecological group. The addition
of fish into the system did not affect the stability of
the system as in the case of Hemavathy reservoir
which showed an increase in MPL by 14%.
(Khan et al., 2015). The system was found to be
more stable after systematic fish stocking (which
included non- native carps) in the reservoir.
We conducted a study on the fish species
composition and catch structure of Manchanbele
reservoir in Karnataka, India and developed the
foodweb model. This reservoir was dominated by
exotic fish (O. niloticus) in the late 1990. Due to
structured stocking with Indian major carps (IMC)
there was a shift in fishery from the dominance of
exotic fishes to IMCs (ICAR-CIFRI, 2016). The
exotics to IMC ratio in 2010-11 was 4:3 which
gradually changed to 1:3 in 2017-18. The foodweb
model of Manchanbele reservoir showed a CI
value of 0.31 that is comparable with the values of
Ravishankarsagar reservoir (Panikkar et al., 2015a)
as depicted in Table 1. This shows that the presence
of exotics may not adversely affect the resilience of
the ecosystem.
In Wyra Reservoir, Panikkar and Khan (2008)
observed that the system as a whole presents mean
transfer efficiencies very analogous in the two
phases (a very small difference of 0.1% the total
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29
system’s carrying capacity) which shows that the
general foodweb structure remains similar despite
the changes of abundance of the various indigenous
fish groups such as murrels, cichlids, catfishes,
gobids, major carps, barbs, minnows and prawns)
including the invasives (tilapia) in the system.
Foodweb approach for managing
exotic fish invasions
Management of exotic species invasion in
inland waters is a challenge worldwide and hence
needs considerable attention. The impacts of an
invasive fish could refer to positive or negative
environmental and socio-political alterations
that these species can cause in the systems
(Shackleton, 2007;Binimelis et al., 2008; Kull
et al., 2011, Gallardo et al., 2016). Based on the
situation, exotic species can cause alterations in the
ecosystem properties of the invaded systems or on
human livelihood (Blackburn et al., 2014; Bacher
et al., 2018; Shackleton et al., 2019). The range and
intensity of these intrusions make management of
inland aquatic ecosystems a challenging task and
these challenges can be tackled through ecosystem
management approach (Christensen et al., 1996;
Khan and Panikkar, 2009).
Globally, there are several guidelines and
management protocols formulated to deal with
the invasive species (Piria et al., 2017). In India,
presently, there are no proper management policy to
address the issues associated with the introduction
and establishment of exotic species in inland
aquatic systems. The United Nations’ Sustainable
Development Goals for 2030 highlights the need
for protecting aquatic life and invasive species
management (Ives et al., 2019). To meet the
objective of ecosystem approach of management
and to assess the impact of exotic species invasion
in inland systems on the foodweb, it is imperative
to understand the trophic function of the ecological
change that is happening within the system (Khan
and Panikkar, 2009). Ecosystem models play a vital
role in the ecosystem approach to management
of aquatic systems and they provide inputs in
identifying the properties and potential changes
in complex ecosystems that cannot be identified
with single-species models (Christensen and Pauly,
1998).
30
Panikkar et al. / Aquatic Ecosystem Health and Management 24 (2021) 24–32
Ecosystem modeling using the Ecopath with
Ecosim (EwE) approach has been largely used as
a tool to assess the ecosystem properties, explore
the trophic interactions, fisheries monitoring and
management (Christensen et al., 2005; Heymans
et al., 2016). The present review on ecopath
modeling studies conducted in the freshwater
ecosystems of India to illustrate the intensity of
the impacts also address the knowledge gaps and
suggest recommendations for better management
and prevention of exotic invasion in the country.
The information derived from this method aids
in developing conservation and management
guidelines to mitigate the effects of the invasion.
The EwE is being attempted to provide information
on how ecosystems are likely to respond to changes
in fishery management practices and therefore,
suggested as a tool for designing ecosystem
approach to fisheries (Plaganyi, 2007). The need
to understand and quantify ecosystem behaviour
and conditions has come to the forefront of
environmental planning and policy formulations,
particularly in view of the contemporary paradigm
of
ecosystem-based
fishery
management
(Bijukumar et al., 2015).
Adequate information on foodweb structure and
ecosystem function provide vital information on the
effects of invasives on the ecosystem and provide
a baseline for management of aquatic resources
(Winemiller and Polis, 1996; Ives et al.,2019).
Such information has relevance for developing
management strategies aimed at managing exotic
invasive fishes and conserving the biodiversity
(Khan and Panikkar, 2009; Ives et al., 2019).
Conclusions
The inland open waters in India are vulnerable to
exotic fish invasion and there is a need to develop a
national level database on invasive fish in different
aquatic systems. To effectively manage the invasive
in a system, there has to be some general rules on
regional basis as to which management responses
are efficient to deal with invasion events. This
involves steps taken to (i) prevent invasive species
from entering a new habitat (ii) if introduced,
efforts to remove if they become well established
(iii) actions to limit negative impacts and to retain
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any benefits of the species if they become widely
established (Shackleton et al., 2017; Novoa et al.,
2019).
We found that many empirical studies on bioinvasions have focused on the species specific
aspects while a few have adopted the foodweb
perspective. Species-specific approaches run a risk
of bias and tend to ignore the interaction between
different causes of invasion at the ecosystem
level. The foodweb approach is one of the most
intuitive conceptualizations of an ecosystem. The
information presented and discussed here forms
a basis for future research planning on changing
foodweb interactions with the introduction of
exotics in the aquatic system.
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