Dietary Guild Structure in Fish

Assemblages and Trophic Position

of Constituent Species in Brush
Parks of a Tropical Estuary
MENAKE GAMMANPILA1, U.S. AMARASINGHE2,*, M.J.S. WIJEYARATNE2
1
Regional Research Centre, National Aquatic Resources Research & Development Agency, Kadolkele, Negombo 11500, Sri Lanka
2
Department of Zoology and Environmental Management, University of Kelaniya, Kelaniya 11600, Sri Lanka

E-mail: zoousa@[Link] | Received: 30/12/2017; Accepted: 26/02/2019

*

©Asian Fisheries Society

ISSN: 0116-6514
E-ISSN: 2073-3720
[Link]

Abstract
Trophic guild structure and dietary niche breadth in tropical fish communities are important to identify functional
groups and to understand how trophic positions of constituent species help coexistence. Objective of the present
study was to investigate whether the constituent species of fish assemblages in brush parks could be grouped into
trophic guilds and how these species contribute to structure the fish community along trophic dimensions. Diets of
46 fish species caught in brush parks were analysed and the food items were categorised into 11 broad groups. Based
on the composition of diets, fish were grouped into 8 trophic guilds. Levin’s index of niche breadth indicated that the
constituent species in the trophic guilds for which food was abundant, were generalists, whereas the trophic guilds of
higher trophic levels were specialists. Within each trophic guild, constituent species showed different trophic indices
indicating low inter-specific competition resulting in optimum food resource utilisation.

Keywords: artificial fish shelters; dietary habits; fish aggregation devices; fish community structure; trophic index

studying ecology of fish communities (Garrison and

Introduction Link.2000; Coll et al. 2006).
Following the original definition of environmental guild
Fish aggregating devices (FADs) are mostly artificial
(Root 1967), a trophic guild of biological communities
fish shelters to which many fish species are attracted
can be defined as a group of species that exploit the
and thereby supporting productive fisheries in many
same category of food resources in a similar way.
parts of the world (Dempster and Taquet 2004). Well
Hence grouping of constituent species in a fish
documented FADs established as traditional fishing
community is possible based on their functional role
devices are brush parks in Negombo estuary, Sri
rather than taxonomy (Garrison and Link 2000;
Lanka (Amarasinghe et al. 2002), Acadjas in Benin,
Ramirez-Luna et al. 2008). This is of particular
West Africa (Lalèyè 2000; Niyonkuru and Lalèyè 2010),
importance because the members of any particular
Katha in Bangladesh (Uddin et al. 2015) and Samrah in
guild play more or less similar functional roles providing
Great Lake in Cambodia (Ho 1999; Lamberts 2001;
vital information on the flow of energy and biomass
Baran 2005; Mekong River Commission 2015). FADs
within food webs (Hawkins and MacMahon 1989) as well
produce higher fish yields per unit area compared to
as on species interactions. Many tropical fish
open water fisheries (Welcomme 2002). Fish
communities such as those inhabiting coral reefs
assemblages in brush parks consist of species that
(Longo et al. 2014; Palacios and Zapata 2014; Boaden
are attracted to shelter or to feed on periphytic food
and Kingsford 2015), seagrass beds (Livingston 1982)
materials (Van Dam et al. 2002). In addition,
and floodplains (Jepsen et al. 1997; Whitley and Bollens
opportunistic predators are also attracted to brush
2014), and those that are attracted to artificial fish
parks (Malone et al. 2011). These fish assemblages are
aggregation devices (FADs) (Deudero 2001) exhibit
structured in trophic dimensions for efficient
complex species interactions. Hence, simplification of
utilization of food resources. Gammanpila et al. (2017)
these complex interactions based on the guild
have shown that ecomorphology of the fish species in
structure is recognized as an important approach for
the brush parks of a tropical estuary correlates with

Asian Fisheries Science 32 (2019):8–18 8 8 Asian Fisheries Science 32 (2019):8–18

diet. Knowledge on feeding ecology of the fish species one-third of the intestine, which were considered as
in an assemblage provides insight into trophic the recently ingested food, were used. From each
interactions, which are of immense importance to suspension, three subsamples of one mL each were
fisheries managers opt for ecosystem-based examined under a microscope (10 × 10 magnification)
management (Pikitch et al. 2004). using a Sedgwick-rafter counting chamber for
qualitative and quantitative analysis of stomach/gut
In the present study, the trophic guilds of the fish contents (Wijeyaratne and Costa 1986; Weliange and
assemblages in brush parks in Negombo estuary, Sri Amarasinghe 2003). Empty stomachs or stomach with
Lanka were identified. In addition, the contribution of the almost fully digested food were excluded. Bio-volume
individual fish species to the structure of the fish of each individual food categories was determined
communities along trophic dimensions was using a cell of Pinnularia, a predominant alga
investigated. belonging to Class Bacillariophyceae, as the standard
and expressed as volumetric percentage of the total
food items in the stomach/gut content (Hynes 1950;
Materials and Methods Hyslop 1980; Wijeyaratne and Costa 1986). Relative
Study area bio-volume of each food category in stomach/gut
contents of individual fish was summed across all
Negombo estuary in Sri Lanka, which is located samples and the percentage volumetric proportion of
between latitudes 7°6'–7°12'N and longitudes 79°40'– diet in each species was determined.
79°53' E is a shallow estuary with a mean depth of 0.65
m. It covers an area of 3,164 ha (Goonethilake et al.
2005) with a maximum length of 12.5 km and
maximum width of 3.6 km. It has a shoreline of 24 km
and about 10 % of the estuary has a depth < 0.5 m
(Jayakody 1996).

Brush parks are traditional forms of fish aggregation

devices installed in shallow areas of the estuaries,
rivers and reservoirs using dense masses of
brushwood (Welcomme 2002). About 2,200 brush
parks with a mean surface area of 51.8 m2, covering
11.3 ha and yielding 12.46 t ha-1 yr-1 were reported in
this estuary in 1998 (Amarasinghe et al. 2002).

Sampling of fish
Fish sampling was carried out once a month from
three major brush park fishing areas (Munnakkaraya,
Katunayaka and Talahena; Fig. 1) in the estuary from
April 2014 to April 2016. For ethical clearance,
permission for conducting research was obtained
from the University of Kelaniya, Sri Lanka. Fish that
were harvested by surrounding brush parks with an
encircling net were chosen for dietary analysis.
Accordingly, 64 brush parks were sampled during the
study period, encompassing the wet (May to Fig. 1. Map of Negombo estuary showing locations of brush
September), intermediate (October-November and parks (crosses). The inflowing river (Dandugam Oya) into
March-April) and dry (December to February) seasons the estuary is also shown here. Maps of Sri Lanka and
of the year. Fish specimens were fixed by injecting 10 Gampaha district indicate geographical location of
% buffered formalin into body cavity, packed in 10 % Negombo estuary.
formalin solution and were transported to the
laboratory for dietary analysis. Fish were identified
based on the fish identification guides for Sri Lankan
Trophic classification
waters (Munro 1955; De Bruin et al. 1995). For the analysis of trophic pattern, the stomach/gut
contents were assigned to eleven broader categories:
Dietary analyses phytoplankton, macrophytes, detritus, molluscs, soft-
bodied macrobenthos, small zooplankton,
Stomach contents of each specimen were extracted cladocerans, copepods, ostracods, insects and fish.
and a suspension of 10 mL was prepared by adding 10 The food items consumed were classified into
% buffered formalin. For fish species that did not different trophic levels (Table 1; Winemiller 1990).
have a well-developed stomach, contents in the first

Asian Fisheries Science 32 (2019):8–18 9 9 Asian Fisheries Science 32 (2019):8–18

Table 1. Eleven food categories, their trophic classes and main components of each food category (Modified from Winemiller
1990).

Food category Abbreviation Trophic class Food items

Phytoplankton PP 0 Diatom, blue green algae, green algae, filamentous algae
Macrophytes MP 0 Aquatic plants
Detritus DE 0 Fine organic material
Mollusks ML 1 Bivalves and gastropods
Soft-bodied Macrobenthos MB 1.5 Annelids, Nematodes, Caridina
Small zooplankton SZ 1.5 Brachionus, Trichocerca, Keratella, dinoflagellates
Cladocerans CD 1.5 Daphnia, Moina, Diaphanasoma, Chydoris, Macrothrix
Copepods CP 1.5 Calanoids, Cyclopoids, Harpectocoids
Ostracods OC 1.5 Cypridopsis
Insects IN 1.5 Predominantly aquatic insects
Fish FH 2 Fish scales, fish eggs, small/juvenile fish, fish skeletons

The plant materials (primary producers) were included have also adopted a similar criterion for naming
in trophic class 0 (T = 0.00), invertebrates in the trophic guilds.
trophic class 1.5 (T = 1.5) as they were omnivores,
predatory fish in trophic class 2 (T = 2.0). Trophic Dietary niche breadth
index for each species (Ti) was computed using the
following equation (Adams et al. 1983; Winemiller Dietary niche breadth was calculated to determine
1990). the relative level of diet specialisation of fish in the
brush park assemblages. Levin’s niche breadth index
𝑇𝑖 = 1.0 + ∑𝑛𝑗 = 1 𝑇𝑗 (𝑝𝑖𝑗 ) (1) was calculated using the following equation (Levins
1968).
where Tj was the trophic class of prey item j (as
defined in Table 1 above), and pij was the fraction of 𝐵 = 1 / ∑𝑛 𝑃𝑖2 (2)
food (proportion by volume) consumed by species i
consisting of prey item j in the diet (Winemiller 1990). where B is the dietary niche breadth, Pi is the
This index represents the trophic position of each volumetric proportion of food item i in the diet and n
species in the fish assemblage in a trophic is the total number of food items in the diet. B ranges
continuum, which represents the ecological pyramids from 1, when only single resource is used (specialist),
more accurately than by discrete levels (Adams et al. to n, when all different type of resources are used in
1983: Cousins 1987). equal proportion (generalist). In the present study,
species were categorised as highly specialists (range
Trophic guilds of B: 1.0–1.25), specialists (range of B: 1.25–1.50), and
generalists (B > 1.5).
Mean proportions of food item consumed by each of
the 46 species were double square root transformed Relative abundance
to reduce heteroscedasticity as this approach is
effective for datasets with many very low values and a Relative abundance of a particular fish species was
few large values (Quinn and Keough 2002). A similarity calculated as the number of individuals of that
matrix, constructed based on Euclidean distances of species divided by the total number of fishes
the degree of dietary similarities between species collected from brush parks during the study period.
(McQuitty 1966), was used to ordinate the 46 fish
species studied from non-metric multi-dimensional Relationship between trophic index
scaling (nMDS) and the trophic guilds of fish species and dietary niche breadth
were identified. Similarity analysis and nMDS were
performed by PRIMER 5 statistical package (Clarke For each trophic guild, the relationships between the
and Warwick 2001). These trophic guilds were named trophic index and dietary niche breadth of constituent
based on the food item that contributed most to the species were determined using linear and second
diet. In most trophic guilds, the prominent food items order regression analyses to illustrate whether there
contributed over 50 % of the diets. Here, consistency were divergences of dietary habits among constituent
of labelling a trophic guild was not based on species in each guild. As these possible relationships
proportion of taxonomic group of any particular food would indicate that each species occupy different
items. For example, molluscivore and invertivore were trophic levels irrespective of dietary niche breadth,
two trophic guilds in the present analysis although they were used to show that dietary overlaps were
members of both guilds fed on invertebrates. Adite unlikely among constituent species within a trophic
and Winemiller (1997) and Garrison and Link (2000) guild. Regression analyses were performed using
Minitab (Version 16) software package.

Asian Fisheries Science 32 (2019):8–18 10 10 Asian Fisheries Science 32 (2019):8–18

 The nMDS plot (Fig. 2) resulted six major trophic guilds
Results and two minor guilds. The major trophic guilds were
labelled as piscivore (guild i), having eight species with
Trophic guilds more than 50 % fish in their diet. The fish species that
consumed both invertebrates and fish (guild ii) were
Total number of 817 specimens belonging to 46 fish
represented by ten species. The invertivore guild
species of 24 families were used for the dietary
(guild iii) was represented by eight species and they
analysis (Table 2). Number of specimens in each
had more than 60 % invertebrates in their diet. Fishes
species used for this analysis varied from 1 to 107.
of omnivore guild (guild v) consumed food items from
Relative proportions by volume of each of the food
different origins (plant and animal) and were
categories in 46 species are given in Table 3.

Table 2. Fish species examined, their abbreviations (Abv), number of analysed stomachs (N) and size (total length) ranges of fish
species in the assemblages of brush parks in Negombo estuary.

Family Scientific name Abv N Size range (cm)

Acanthuridae Acanthurus gahhm (Forsskål, 1775) Ac 21 6.2–17.4
Ambassidae Ambassis gymnocephalus (Lacepède, 1802) Ag 13 8.1–17.4
Anguillidae Anguilla bicolor bicolor McClelland, 1844 An 1 60.3
Apogonidae Apogon hyalosoma Bleeker, 1852 Ah 3 8.9–13.2
Ariidae Arius jella Day, 1877 Aj 24 14.6–30.5
Arius maculatus (Thunberg, 1792) Am 9 10.6–28.7
Carangidae Caranx sexfasciatus Quoy & Gaimard, 1825 Cs 23 9.2–17.3
Carangoides talamparoides Bleeker, 1852 Ct 2 12.4–17.7
Centropomidae Lates calcarifer (Bloch, 1790) Lc 34 18.1–48.3
Cichlidae Etroplus suratensis (Bloch, 1790) Es 96 5.5–23.5
Oreochromis mossambicus (Peters, 1852) Om 2 25.3–29.5
Oreochromis niloticus (Linnaeus, 1758) On 3 21.4–22.3
Eleotridae Ophiocara porocephala (Valenciennes, 1837) Op 5 14.2–20.7
Gerreidae Gerres abbreviates Bleeker, 1850 Ga 16 7.5–14.4
Gerres filamentosus Cuvier, 1829 Gf 1 8.6
Gerres oyena (Forsskål, 1775) Gy 9 9.9–13.5
Gerres oblongus Cuvier, 1830 Go 6 9.1–15.2
Haemulidae Plectorhinshus gibbosus (Lacepède, 1802) Pg 12 7.1–22.1
Leiognathidae Leiognathus equulus (Forsskål, 1775) Le 1 7.7
Leiognathus fasciatus (Lacepède, 1803) Lg 1 9.1
Lethrinidae Lethrinus rubrioperculatus Sato, 1978 Lh 1 9.0
Lutjanidae Lutjanus argentimaculatus (Forsskål, 1775) La 66 14.3–33.0
Lutjanus fulviflamma (Forsskål, 1775) Lf 19 10.4–15.7
Lutjanus kasmira (Forsskål, 1775) Lk 4 10.6–15.9
Lutjanus russelli (Bleeker, 1849) Lr 41 8.0–26.9
Monodactylidae Monodactylus argenteus (Linnaeus, 1758) Ma 12 6.9–9.2
Mugilidae Liza macrolepis (Smith, 1846) Lm 24 17.1–50.0
Liza melinoptera (Valenciennes, 1836) Lt 32 11.3–45.2
Liza parsia (Hamilton, 1822) Lp 7 13.6–27.3
Cicamugil cascasia (Hamilton, 1822) Lz 2 21.8–23.4
Liza subviridis (Valenciennes, 1836) Ls 25 15.2–49.2
Mugil cephalus Linnaeus, 1758 Mc 12 18.2–54.4
Valamugil buchanani (Bleeker, 1853) Vb 5 12.2–50.3
Valamugil seheli (Forsskål, 1775) Vs 24 12.6–28.5
Plotosidae Plotosus canius Hamilton, 1822 Pc 7 34.4–59.0
Scatophagidae Scatophagus argus (Linnaeus, 1766) Sa 37 7.2–22.7
Serranidae Epinephelus malabaricus (Bloch & Schneider, 1801) Em 41 13.8–40.1
Epinephelus coioides (Hamilton, 1822) Ec 9 10.1–31.5
Epinephelus tauvina (Forsskål, 1775) Et 1 35.7
Sparidae Acanthopagrus berda (Forsskål, 1775) Ab 31 9.2–33.5
Sphyraenidae Sphyraena jello Cuvier, 1829 Sp 1 30.8
Sillaginidae Sillago sihama (Forsskål, 1775) Ss 10 12.5–32.7
Siganidae Siganus javus (Linnaeus, 1766) Sj 107 8.8–22.9
Siganus lineatus (Valenciennes, 1835) Sl 10 10.9–23.2
Siganus vermiculatus (Valenciennes, 1835) Sv 6 11.7–20.3
Terapontidae Terapon puta Cuvier, 1829 Tp 1 10.9

Asian Fisheries Science 32 (2019):8–18 11 11 Asian Fisheries Science 32 (2019):8–18

Table 3. Volumetric proportions of different food categories and trophic index of each fish species assemblage in brush parks in
Negombo estuary (PP = Phytoplankton, MP = Macrophytes, DE = Detritus, ML = Mollusks, MB = Macrobenthos, SZ = Small
zooplankton, CD = Cladocerans, CP = Copepods, OS = Ostracods, IN = Predominantly aquatic insects and FH = Fish).
Abbreviations (Abv) of the names of fish species are as given in Table 2. In each row, the major food items are indicated in bold.

Food items (proportions) Trophic

Abv
PP MP DE ML MB SZ CD CP OS IN FH index
Ac 0.53 0.18 0.17 0.00 0.06 0.00 0.02 0.01 0.00 0.00 0.04 1.23
Ag 0.06 0.02 0.01 0.00 0.72 0.00 0.00 0.03 0.00 0.00 0.16 2.45
An 0.00 0.00 0.00 0.05 0.10 0.00 0.00 0.00 0.00 0.00 0.85 2.90
Ah 0.00 0.00 0.00 0.00 0.85 0.00 0.00 0.00 0.00 0.00 0.15 2.58
Aj 0.03 0.01 0.02 0.04 0.36 0.00 0.00 0.03 0.00 0.01 0.51 2.66
Am 0.00 0.00 0.00 0.00 0.13 0.00 0.00 0.00 0.00 0.00 0.87 2.93
Cs 0.02 0.02 0.01 0.00 0.45 0.00 0.00 0.00 0.00 0.00 0.50 2.68
Ct 0.00 0.00 0.00 0.00 0.70 0.00 0.00 0.00 0.00 0.00 0.30 2.65
Lc 0.00 0.00 0.00 0.00 0.14 0.00 0.00 0.00 0.00 0.00 0.86 2.92
Es 0.30 0.24 0.39 0.02 0.02 0.00 0.00 0.01 0.00 0.01 0.02 1.11
Om 0.11 0.13 0.58 0.08 0.00 0.00 0.00 0.10 0.00 0.00 0.00 1.22
On 0.12 0.08 0.68 0.05 0.00 0.00 0.00 0.00 0.00 0.00 0.07 1.18
Op 0.00 0.00 0.00 0.53 0.21 0.00 0.00 0.00 0.00 0.00 0.26 2.36
Ga 0.21 0.01 0.01 0.03 0.26 0.00 0.00 0.05 0.00 0.01 0.42 2.35
Gf 0.27 0.00 0.00 0.00 0.36 0.00 0.00 0.05 0.00 0.00 0.32 2.25
Gy 0.62 0.00 0.00 0.00 0.05 0.00 0.00 0.33 0.00 0.00 0.00 1.57
Go 0.12 0.00 0.25 0.21 0.12 0.00 0.00 0.07 0.00 0.00 0.22 1.94
Pg 0.04 0.00 0.01 0.00 0.93 0.00 0.00 0.00 0.00 0.00 0.02 2.43
Le 0.12 0.00 0.00 0.00 0.01 0.01 0.00 0.08 0.16 0.00 0.61 2.62
Lg 0.18 0.00 0.03 0.08 0.00 0.00 0.00 0.14 0.00 0.00 0.57 2.43
Lh 0.01 0.00 0.00 0.00 0.81 0.00 0.00 0.00 0.00 0.00 0.18 2.57
La 0.04 0.03 0.00 0.00 0.25 0.00 0.00 0.00 0.00 0.00 0.68 2.74
Lf 0.00 0.00 0.00 0.00 0.47 0.00 0.00 0.00 0.00 0.00 0.53 2.76
Lk 0.00 0.00 0.00 0.02 0.37 0.00 0.00 0.00 0.00 0.00 0.62 2.80
Lr 0.02 0.00 0.00 0.00 0.61 0.00 0.00 0.00 0.00 0.00 0.36 2.65
Ma 0.18 0.25 0.15 0.02 0.24 0.00 0.00 0.00 0.00 0.00 0.16 1.70
Lm 0.21 0.23 0.46 0.01 0.05 0.00 0.00 0.04 0.00 0.00 0.01 1.16
Lt 0.16 0.10 0.64 0.02 0.01 0.00 0.00 0.07 0.00 0.00 0.01 1.15
Lp 0.06 0.21 0.62 0.02 0.00 0.00 0.00 0.06 0.00 0.00 0.04 1.18
Lz 0.15 0.01 0.83 0.00 0.00 0.00 0.00 0.01 0.00 0.00 0.00 1.02
Ls 0.21 0.25 0.45 0.03 0.00 0.00 0.00 0.07 0.00 0.00 0.00 1.13
Mc 0.31 0.08 0.55 0.00 0.04 0.00 0.00 0.03 0.00 0.00 0.00 1.10
Vb 0.04 0.22 0.75 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00
Vs 0.21 0.15 0.58 0.00 0.02 0.00 0.00 0.01 0.00 0.03 0.01 1.11
Pc 0.00 0.00 0.00 0.02 0.93 0.00 0.00 0.00 0.00 0.00 0.04 2.50
Sa 0.43 0.37 0.13 0.00 0.05 0.00 0.00 0.01 0.00 0.00 0.01 1.10
Em 0.00 0.00 0.00 0.00 0.50 0.00 0.00 0.00 0.00 0.00 0.50 2.75
Ec 0.00 0.00 0.00 0.00 0.49 0.00 0.00 0.00 0.00 0.00 0.51 2.80
Et 0.00 0.00 0.00 0.00 0.44 0.00 0.00 0.00 0.00 0.00 0.56 2.78
Ab 0.03 0.12 0.07 0.04 0.01 0.00 0.09 0.00 0.00 0.00 0.62 2.45
Sp 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 3.00
Ss 0.07 0.17 0.00 0.00 0.61 0.00 0.00 0.01 0.00 0.00 0.14 2.20
Sj 0.40 0.21 0.28 0.01 0.07 0.00 0.00 0.02 0.00 0.00 0.01 1.17
Sl 0.47 0.44 0.08 0.00 0.01 0.00 0.00 0.00 0.00 0.00 0.00 1.01
Sv 0.58 0.17 0.16 0.02 0.01 0.00 0.00 0.00 0.01 0.00 0.04 1.13
Tp 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 3.00

represented by two species, whereas herbivore guild other two minor guilds were represented by one
(guild vi) was represented by five species having more species each. The diet of Ophiocara porocephala
than 60 % plant material (phytoplankton and (Valenciennes, 1837) which contained 53 % molluscs
macrophytes) in their diet. The detritivore guild (guild was labelled as molluscivore (guild iv) and that of
vii) consisting of fish that had more than 40 % detritus Gerres oyena (Forsskål, 1775) which contained 62 %
in their diet was composed of eleven species. The

Asian Fisheries Science 32 (2019):8–18 12 12 Asian Fisheries Science 32 (2019):8–18

Fig. 2. Non-metric multi-dimensional scaling (nMDS) ordination based on dietary habits of 46 fish species in the brush parks of
the Negombo estuary. Stress value of 0.07 indicates that two dimensional nMDS plot is sufficient to ordinate the fish species.
The eight clusters represent trophic guilds. The abbreviations of species names belonging to individual trophic guilds are as
given in Table 2.

algae, was labelled as algivore (guild viii). It must be Accordingly, Sphyraena jello Cuvier, 1829 and Terapon
noted that this classification of trophic guilds was not puta Cuvier, 1829 in piscivores trophic guild were high
strictly based on taxonomic groups. Adite and specialists, Arius maculatus (Thunberg, 1792), Lates
Winemiller (1997) also categorised fish species calcarifer (Bloch, 1790) and Anguilla bicolor bicolor
feeding on various taxonomic groups of invertebrates McClelland, 1844 were specialists, whereas
into several trophic guilds such as invertebrate Leiognathus fasciatus (Lacepède, 1803),
feeders, worm feeders, insectivores and Acanthopagrus berda (Forsskål, 1775) and Leiognathus
molluscivores. equvlus (Forsskål, 1775) in the same trophic guild were
moderate generalists. In invertivore guild, Plotosus
canius Hamilton, 1822 and Plectorhinshus gibbosus
Trophic index, dietary niche breadth (Lacepède 1802) were high specialists, Apogon
and relative abundance hyalosoma (Bleeker, 1852) and Lethrinus
rubrioperculatus Sato, 1978 were specialists. Only fish
Trophic indices estimated on the basis of dietary species, Cicamugil cascasia in detritivore guild
composition using equation 1 above, that ranges from showed specialist dietary habits. In herbivore (guild
1 to 3 are given in Table 3. The lowest trophic indices vi), omnivore (guild v) and invertebrate and fish (guild
(1.00–1.25) were recorded in the detritivore and ii) trophic guilds, all species were generalists (Fig. 3B).
herbivore guilds while the highest indices (2.5–3.0) As the molluscivore and algivore guilds had only one
were recorded in the piscivore guild. The algivore species each, they were not treated as specialists.
guild had a trophic index of 1.57 while that of omnivore
guild ranged from 1.5 to 2.00. The trophic index of Relative abundance of fish species in piscivore and
molluscivore guild was 2.36 and that of invertivore invertivore guilds was low while that of some species
guild ranged from 2.2 to 2.65. Broad ranges of trophic in herbivore, omnivore and detritivore guilds were
indices (Fig. 3A) indicated that fish species in each high (Fig. 3C). The herbivore guild was dominated by
trophic guild fed on a variety of food items. Siganus javus (20.4 %) and Scatophagus argus
(Linnaeus, 1766) (9.5 %). The omnivore guild was
Levin’s index for dietary niche breadth (B) ranged from dominated by Monodactylus argenteus (Linnaeus,
1 in piscivore guild to 5.16 in omnivore guild (Fig. 3B). 1758) (17.5 %) and in detritivore guild, E. suratensis was
The narrow ranges for dietary niche breadth were the most abundant having 11.5 % of total number of
recorded for the herbivore and invertivore trophic fish in the assemblage. In piscivore guild, only Arius
guilds. B value of 1 indicates that the fish species maculatus (6.4 %) registered over 5 % of abundance.
feeds only on a single prey type (high specialists) and None of the species in invertivore and invertebrate
B value of > 1.5 indicates that the fish species ingests and fish feeding guilds had a relative abundance
a diverse range of food types (generalists). greater than 5 % (Fig. 3C).

Asian Fisheries Science 32 (2019):8–18 13 13 Asian Fisheries Science 32 (2019):8–18

Fig. 3. Trophic indices (A), Levin’s dietary niche breadths (B) and relative abundance (C) of the 46 fish species collected from the
brush parks of the Negombo estuary. Abbreviations of species names are as given in Table 2.

These indicate that the species occupying lower values. As there were only two species in the
trophic levels in the food web (i.e., herbivore, omnivore omnivore trophic guilds (Figs. 2 and 3) species in
and detritivore guilds) were more abundant whereas omnivore and herbivore guilds were combined to
those occupying higher trophic levels were less determine the relationship between Ti and B and a
abundant. significant positive second-order relationship (P <
0.05) was found between Ti and B (Fig. 4D). This
Trophic continuum and dietary niche indicates that the constituent species of this trophic
breadth guild show opposite of those in the piscivore guild. In
this guild, there were no specialists and the species
The significant negative linear and negative second which occupied higher trophic levels in the trophic
order relationships (P < 0.05) were observed continuum had wider dietary niche breadths as they
respectively between Ti and dietary niche breadth (B) fed on both plant matter and animal matter (Table 3).
in the piscivore guild and fish consumed invertebrates In the detritivore trophic guild, a significant parabolic
and fish guild (Figs. 4A and 4B). These indicate that relationship (P < 0.05) was observed between Ti and
fish species in both guilds which fed on several types dietary niche breadth (Fig. 4E). In this trophic guild,
of prey occupied wide ranges of trophic indices (Ti) in there were only one species which had lower (< 1.5)
the upper trophic levels. These also show that the dietary niche breadth, which could be treated as
species in higher trophic levels were specialists specialists while rest of the species were generalists
having narrow dietary niche breadths. On the other having higher B values. Presumably, there was no
hand, species with lower Ti, fed different food items, food limitation for constituent species in this trophic
mainly on soft-bodied macrobenthos and fish (Table guild.
3). In the invertivore guild, a significant negative
second order relationship (P < 0.05) was observed
between Ti and dietary niche breadth (Fig. 4C). This
Discussion
indicated that individual species in this trophic guild
The structure of fish communities aggregated in fish
also occupied different levels in the trophic
shelters is not expected to be similar to those in
continuum. They were also either specialists or
natural habitats because the species that are
generalists. Among the invertivores, the species with
attracted to such structures are mostly “shade-philic”
the highest dietary niche breadth (Sillago sihama
species and perhaps predators in search of prey
(Forsskål, 1775)) had a relatively low Ti while some
organisms. The trophic guild structure of fish
others with high dietary niche breadth also had
communities associated with natural reefs and
relatively high Ti values. Four species which can be
artificial reefs such as shipwrecks was also found to
considered as specialists also had intermediate Ti

Asian Fisheries Science 32 (2019):8–18 14 14 Asian Fisheries Science 32 (2019):8–18

Fig. 4. The relationships between trophic index and dietary niche breadth in the five trophic guilds. (A): Piscivore, (B):
Invertebrates and fish feeders, (C): Invertivore, (D): Herbivore and omnivore and (E) Detritivore. Here, species in trophic guilds,
herbivore and omnivore were pooled due to less number of species in each guild.

be different from each other (Arena et al. 2007; Simon temporary visitors (Amarasinghe et al. 2002;
et al. 2013), and therefore, the energy flow in the two Gammanpila et al. 2017).
fish shelters may not be the same. Fish communities
in shelters such as brush parks might be structured in Results of the present study revealed that, despite
such a way that the constituent species would find their relatively short stay in the brush parks, the fish
refuge from predators and enhance schooling assemblages associated with the brush parks are
behaviour (Dempster and Taquet 2004). In some composed of eight distinct trophic guilds. Knowledge
regions such as West Africa, where brush parks are on trophic guilds are particularly useful to simplify the
placed for longer periods of > 12 months (Niyonkuru structure and dynamics of complex tropical aquatic
and Lalèyè 2010), community structure of fish ecosystems (Garrison and Link 2000). As the
assemblages can be expected to be somewhat members of each trophic guild play somewhat similar
stabilized. However, brush parks in Negombo estuary roles, understanding the guild structure is useful to
are maintained for much less time period (3–97 days), postulate how fisheries exploitation regimes impact
and as such, fish species that are attracted to these on trophic dynamics. One of the limitations of the
brush parks in search of food and shelter are present study was the small sample size of some
species. However, it may not have a significant

Asian Fisheries Science 32 (2019):8–18 15 15 Asian Fisheries Science 32 (2019):8–18

impact because of the low variability of dietary level in conventional food web studies (Winemiller
composition of individual fish species. Earlier studies 1990).
on food and feeding habits of Sphyraena jello (Bachok
et al. 2004; Mohanraj and Prabhu 2012), Anguilla Results of the present study also showed that the
bicolor (Rupasinghe and Ettygalle 2006), Epinephelus constituent species of trophic guilds exhibit broad
sp., Leiognathus equulus (Forsskål, 1775) and L. dietary niche breadth. Trophic generalists are
fasciatus (Hajisamae and Ibrahim 2008) also indicated characterised by high dietary overlap whereas trophic
that these species belong to the similar trophic guilds specialists often require trophic divergence for their
as found in the present study. co-existence (Weliange and Amarasinghe 2003). It
can therefore be stated that although the fish
Coexistence of constituent species with more or less assemblages aggregated in artificial fish shelters
similar dietary habits in a fish community is generally such as brush parks consist of trophic guilds, they are
possible when there is a niche segregation along structured through trophic divergence. In animal
temporal, spatial and trophic dimensions (Edirisinghe communities, trophic divergence is known to be an
and Wijeyaratne 1986; Piet et al. 1999). A trophic guild important process that generates diversity (Grant and
consists of species that exploit similar food resources Grant 2006), and it also has important implications in
in a similar way (Root 1967; Garrison and Link 2000) ecological dynamics by reducing inter-specific
and as such, one can expect that dietary overlaps competition for food (Post et al. 2008). Divergence in
among species of a trophic guild may lead to inter- the use of food resources that results less
specific competition. However, in fish communities exploitative competition appears to exist in brush
with complex interactions, ecological strategies parks too. This is of particular importance because
appear to exist for relaxing inter- and intra-specific management of tropical fish stocks and the aquatic
competition (Carr et al. 2002; Boaden and Kingsford ecosystems that support them will require a good
2015). Although each trophic guild is represented by knowledge of food web ecology (Winemiller and
species with similar dietary habits (Root 1967), results Jepsen 1998). Ecologically, food and feeding not only
of the present study suggests that even within a affect the energy transfer and subsequent growth
trophic guild, constituent species occupy different and survival of prey and predators, but also the
trophic positions relaxing competition for food distribution, abundance and demographics of fishes
resources when a trophic continuum is considered. (Brodeue et al. 2017).
This is particularly so for fish species occupying
higher trophic levels where competition for food Findings of the present study therefore have
resources might be keen as opposed to those occupy significant implications in fisheries management
in lower trophic levels with abundant food resources. because fisheries associated with natural and
artificial fish shelters contribute significantly to
In many oceans including the Mediterranean sea enhance fisheries production in many parts of the
(Riera et al. 1999; Massuti et al. 1999), Indian Ocean world (Dempster and Taquet 2004). Furthermore,
(Ariyarathna and Amarasinghe 2012) and Central trophic guild structure in fish assemblages have a
America (Hunter and Mitchell 1967), flotsam potential utility value as a rapid appraisal method to
associated fisheries are established. Despite the assess the impact on the ecosystems. For example,
relatively short duration of fish assemblages in these Welcomme et al. (2006) have shown that
artificial shelters, coexistence of constituent species environmental guilds of fish assemblages can be used
might conform to divergence of resource exploitation as a tool to assess the impacts on fluvial ecosystems
especially the food. Trophic guild structure broadly and that they can assist in predicting responses of
indicates dietary niche divergence of coexisting river fish biota to anthropogenic changes such as
species along the trophic dimension. It is also river damming and water diversion. Also, knowledge
suggested that in artificial fish shelters such as FADs, about community-wide interactions that regulate the
flotsam, floating fish cages, petroleum drilling dynamics of exploited fishery resources is recognized
platforms and brush parks, availability of food to be important for fisheries management (Jennings
resource is limited and therefore these shelters can and Kaiser 1998). As such, findings of the present
be considered as ‘ecological traps’ (Marsac et al. 2000; study can be used as a baseline for future assessment
Menard et al. 2000; Hallier and Gaertner 2008). of environmental impacts, which would be useful in
Nevertheless, the concept of ‘ecological trap’ is ecosystem-based management of brush park
shown not to be always true when fish assemblages fisheries.
attracted to fish shelters are considered (Schaefer
and Fuller 2002). Similarly, it can be argued that
similar dietary habits of the constituent species in a
Conclusion
trophic guild may increase inter-specific competition
Trophic guild, by definition is represented by species
for food. Present analysis however, has shown that
with similar dietary habits. In the present study, it was
within a trophic guild, component species occupy
revealed that fish species attracted to brush parks in
slightly different trophic levels, which are defined by
a Sri Lankan estuary were organized in different
trophic indices (Fig. 3A). Here trophic index (Ti) was
trophic guilds and the constituent species of a
defined as a continuum rather than a discrete trophic

Asian Fisheries Science 32 (2019):8–18 16 16 Asian Fisheries Science 32 (2019):8–18

particular trophic guild show broad dietary niche Boaden, A.E. and M.J. Kingsford. 2015. Predators drive community
breadth. When a trophic continuum is considered, it structure in coral reef fish assemblages. Ecosphere 6(4):1–33 (Article
was evident that within a trophic guild, constituent 46).
species occupy different trophic positions relaxing Brodeur, R.D., B.E. Smith, R.S. Mcbride, R. Heintz and E. Farley Jr. 2017.
competition for food resources. The species in lower New perspectives on the feeding ecology and trophic dynamics of
trophic levels were mostly generalists while those in fishes. Environmental Biology of Fishes 100:293–297.
higher trophic guilds were specialists. Trophic Carr, M.H., T.W. Anderson and M.A. Hixon. 2002. Biodiversity,
generalists in the lower trophic levels without having population regulation, and the stability of coral-reef fish
food resource limitations were evidently communities. Proceedings of the National Academy of Sciences of
characterized by high dietary overlap whereas trophic the United States of America 99: 11241–11245.
specialists with food limitation at higher trophic levels Clarke, K.R. and R.M. Warwick. 2001. Changes in marine communities:
exhibited greater trophic divergence for relaxing An approach to statistical analysis and interpretation, 2nd edn.
inter-specific competition. These findings can be PRIMER-E Ltd., Plymouth Marine Laboratory, Plymouth, United
used as a baseline for future assessment of Kingdom. 176 pp.
environmental impacts, which would be useful in Coll, M., I. Palomera, S. Tudela and F. Sarda. 2006. Trophic flows,
ecosystem-based management of brush park ecosystem structure and fishing impacts in the South Catalan Sea,
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Acknowledgements Ecology and Evolution 2:312–316.

De Bruin, G.H.P., B.C. Russell and A. Bogusch. 1995. FAO species
identification field guide for fishery purposes. The Marine Fishery
The authors gratefully acknowledge the financial
Resources of Sri Lanka. FAO, Rome, Italy. 400 pp., 32 colour plates.
support provided by the National Aquatic Resources
Dempster, T. and M. Taquet. 2004. Fish aggregation device (FAD)
Research and Development Agency, Sri Lanka to carry
research: gaps in current knowledge and future directions for
out this study. Assistance of Messrs. W.A.
ecological studies. Reviews in Fish Biology and Fisheries 14:21–42.
Sumanadasa and A.M.S. Pushpananda in field studies
Deudero, S. 2001. Interspecific trophic relationships among pelagic fish
is also gratefully acknowledged. We are thankful to
species underneath FADs. Journal of Fish Biology 58:53–67.
brush park fishermen in the Negombo estuary for
Edirisinghe, E.A.D.N.D. and M.J.S. Wijeyaratne. 1986. Food resource
their cooperation during data collection. The authors
partitioning among the fishes co-existing in brush parks, an artificial
express their sincere appreciation to Mr. R.P.K.C.
habitat in a lagoon in Sri Lanka. Journal of Inland Fisheries 3:115–125.
Rajapakse, Department of Zoology and Environmental
Gammanpila, M., U.S. Amarasinghe and M.J.S. Wijeyaratne. 2017.
Management, University of Kelaniya, Sri Lanka for his
Morphological correlates with diet of fish assemblages in brush park
technical support in preparing the manuscript.
fisheries of tropical estuaries. Environmental Biology of Fishes
100:1285–1299.

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