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Cogent Food & Agriculture

ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/oafa20

Diet composition and feeding habits of the African


sharptooth catfish Clarias gariepinus (Burchell,
1822) from Ribb Reservoir, South Gondar, Ethiopia

Agumassie Tesfahun & Sale Alebachew

To cite this article: Agumassie Tesfahun & Sale Alebachew (2023) Diet composition and
feeding habits of the African sharptooth catfish Clarias gariepinus (Burchell, 1822) from
Ribb Reservoir, South Gondar, Ethiopia, Cogent Food & Agriculture, 9:2, 2284228, DOI:
10.1080/23311932.2023.2284228

To link to this article: https://doi.org/10.1080/23311932.2023.2284228

© 2023 The Author(s). Published by Informa


UK Limited, trading as Taylor & Francis
Group.

Published online: 27 Nov 2023.

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Tesfahun & Alebachew, Cogent Food & Agriculture (2023), 9: 2284228
https://doi.org/10.1080/23311932.2023.2284228

FOOD SCIENCE & TECHNOLOGY | RESEARCH ARTICLE


Diet composition and feeding habits of the
African sharptooth catfish Clarias gariepinus
(Burchell, 1822) from Ribb Reservoir, South
Received: 13 June 2023 Gondar, Ethiopia
Accepted: 13 November 2023
Agumassie Tesfahun1* and Sale Alebachew2
*Corresponding author: Agumassie
Tesfahun, Department of Biology,
College of Natural and
Abstract: A total of 525 catfish samples were collected with various fishing
Computational Sciences, equipment, including hooks, longlines, and different mesh sizes of gillnets (4–12
P. O. Box 272, Debre Tabor, Ethiopia
E-mail: agumas2012@yahoo.com cm). Out of these specimens, 379 (72.2%) stomachs contained prey items, whereas
146 (27.8%) were empty stomachs. Overall, fish prey, zooplankton, detritus and
Reviewing editor:
María Luisa Escudero Gilete, phytoplankton were the four most important food items, accounting for 48.1%,
Nutrition and Bromatology,
Universidad de Sevilla, Spain 21.8%, 17.1% and 5.9% of C. gariepinus diets by volume respectively. Diet composi­
tion varied across different size classes of the fish. The smallest fish (<37 cm
Additional information is available at
the end of the article standard length) mainly consumed detritus, mud (sediment), and zooplankton
which comprise 41.2%, 29.3%, and 19.8% of the total volume, respectively. The
larger fish (>37 cm SL) primarily fed on fish prey (14.0–74.1%) followed by zooplank­
ton (11.2–21.3%) by volume. The relative importance of detritus, mud, zooplankton,
phytoplankton, and insects decreased with increasing fish size from the Ribb
Reservoir. The food and feeding habits of C. gariepinus significantly differed between

ABOUT THE AUTHORS PUBLIC INTEREST STATEMENT


Agumassie Tesfahun received his MSc Degree in Ethiopia’s agricultural sector is still insufficient to
Fisheries, Limnology, and Aquatic-eco-toxicology supply food to this fast growing population.
from Hawassa University, Ethiopia. Currently, he Clarias gariepinus African sharptooth mudcatfish
is the Assistant Professor at the Department of is one of the most commercially important fish
Biology, Debre Tabor University, Ethiopia. Besides, species in Ethiopia. Fisheries and aquaculture are
he has been working in community service and alternatives to meet the high protein demand.
research on fisheries biology of the most com­ The African catfish Clarias gariepinus (family
mercially important fish species from Ribb Clariidae) is one of Ethiopia’s most important
Reservoir Tana basin, Ethiopia, and aquaculture commercial fisheries. Understanding fish feeding
establishment (earthen pond system) from habits and their trophic interactions within food
Fogera District, South Gondar, Ethiopia. webs is important for fisheries management and
Sale Alebachew was awarded his MSc Degree in establishing sound aquaculture practices. The
Animal Production from Debre Markos University, food and feeding habits of this fish species is
Agumassie Tesfahun Ethiopia. Now, he is a lecturer and researcher at impacted by the expansion of irrigation practices
the Department of Animal Sciences, Debre Tabor such as high sedimentation load, poor land use
University, Ethiopia. Moreover, he has been practice, lack of vegetation cover, and the con­
working on fisheries biology of the most com­ struction of dams and weirs within the rivers.
mercially important fish species from Ribb This fish resource is not well utilized due to the
Reservoir Tana basin, Ethiopia, and aquaculture scientific knowledge gap. Therefore, water buffer
establishment (earthen pond system) from zone management is needed to improve food
Fogera District, South Gondar, Ethiopia. and feeding habits of this fish species for better
sustainable utilization.

© 2023 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution
License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribu­
tion, and reproduction in any medium, provided the original work is properly cited. The terms on
which this article has been published allow the posting of the Accepted Manuscript in
a repository by the author(s) or with their consent.

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dry and wet seasons. Fish prey and detritus were mainly consumed in the dry
season and contributed to 63.6% and 21.9% of the total volume respectively.
Zooplankton and phytoplankton were the most preferred food items during the
wet season, contributing 71.8% and 22.2% of the total volume. Generally,
C. gariepinus appears to be an omnivore, the species exhibits ontogenetic dietary
shifts with larger specimens being more carnivorous, and the species exhibits diet­
ary plasticity across wet and dry seasons, which may be linked to food availability
from the Ribb Reservoir.

Subjects: Zoology; Marine & Aquatic Science; Ecology - Environment Studies

Keywords: Catfish; Detritus; Diet preferences; Diet composition; Ethiopia; Ribb Reservoir

1. Introduction
The African catfish Clarias gariepinus (1822) is extremely widespread throughout most of Africa
(Spataru et al., 1987; Willoughby & Tweddle, 1978). In Ethiopia, it is widespread in almost all water
bodies (lotic and lentic systems) such as in the rift valley, Abay, Awash, Baro-Akobo, Omo-Gibe,
Tekeze, and Wabishebele-Genale basins (Awoke, 2015; Golubtsov & Mina, 2003). In many regions
of Africa, it is one of the most important commercial freshwater fish species (Dadebo et al., 2014;
Yesuf et al., 2023). Consequently, it is also the second commercially important fish species from
the newly established Ribb reservoir, the Lake Tana Basin (Alebachew et al., 2022; Tesfahun &
Alebachew, 2023). Understanding fish feeding habits and their trophic interactions within food
webs is important for fisheries management and establishing sound aquaculture practices
(Tesfahun & Temesgen, 2018). According to Adeyemi et al. (2009), the food and feeding habits
of fish provides vital evidence for management in a controlled environment and for the formula­
tion of suitable nutrition provided for the fishin aquaculture. C. gariepinus feeds on a wide spec­
trum of prey items such as phytoplankton, macrophytes, zooplankton, insects, fish, detritus,
amphibians, molluscs, birds, and sediment (Admassu et al., 2015; Dadebo et al., 2014; Eyayu,
(2019); Teka & Admassu, 2016; Tekle-Giorgis et al., 2016). The food and feeding habits of this
catfish species depend on the season of the year and the spatial differences (Dereba, 2019;
Houlihan et al., 2001; Kamal et al., 2010). Several studies were conducted with respect to the
food and feeding habits of the C. gariepinus in Ethiopian water bodies (Abera, 2007; Admassu et al.,
2015; Dadebo, 2000, 2009; Dadebo et al., 2014; Eyayu, (2019); Teka & Admassu, 2016; Tekle-
Giorgis et al., 2016). In the Ribb Reservoir, while a few studies on fish diversity and abundance
(Mequanent et al., 2022) and the feeding ecology of the Nile tilapia (Oreochromis niloticus)
(Tesfahun & Alebachew, 2023) have been conducted, there has so far been no study on the diet
and feeding habits of C. gariepinus, despite its importance to the commercial fishery of the area.
Such area-specific information on the food and feeding habits of this commercially and ecologi­
cally important species is significant for management (protecting its natural prey types and
understanding its feed demand in fish culture systems) to sustain fish resources. Therefore, the
current study was conducted to study diet composition and feeding habits of the African sharp­
tooth catfish Clarias gariepinus (Burchell, 1822) from Ribb Reservoir, South Gondar, Ethiopia.

2. Materials and methods

2.1. Study area


The Ribb Reservoir was built at the midpoint of the two districts namely Farta and Ebinat for the
purposes of irrigation (Bezabih, 2021; Mequanent et al., 2022) (Figure 1). The Ribb Reservoir is
located in the northeastern part of the Lake Tana sub-basin, and was formed by the damming of
the Ribb River in 2018. The longitudinal catchment of the Ribb River covers about 130 km and
comprises a drainage area of approximately 1790 km2 with an annual average discharge of
14.6331 m3s−1. The four administrative kebeles such as Medeb Gubda, Jarashikra, and Ayvaniva
(Farta district), and Amstya (Ebnat district) neighbored the Ribb Reservoir. It is located at 11°59’0”

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Figure 1. The study area map


reveals the sites of specimen
collection of the Ribb Reservoir,
Ethiopia.

N to 12°2’0“N Latitude and 38°0’0“E to 38°2’59“E Longitude (Figure 1). The Ribb watershed is
experienced with a higher rate of irrigation practice particularly at Ribb Reservoir which increasing
time to time at an alarming rate (Mequanent et al., 2021). Concerning the climate condition May to
July is characterized as the rainy season. The monthly rainfall fluctuated from 65 mm in May to
411 mm in July. The mean precipitation was (1,400 mm annum−1) and the minimum (1,200 mm
annum−1). The weather condition of Ribb Reservoir is categorized as Woina Dega (moderate
climate) (Ezezew, (2019)). The annual temperature ranges from 22.5°C to 26.2°C (Mequanent
et al., 2022). The Microcystis (blue-green algae were found along the littoral side of the reservoir
(Tesfahun & Alebachew, 2023). Fourteen zooplankton species were found in the reservoir. The Nile
tilapia O. niloticus, the African catfish C. gariepinus, and Labeobarbus species were commercially
important in the reservoir (Mequanent et al., 2022). Fishing in Ribb reservoirs is a common practice.
Fishers established an association with the support of the government and began fishing in 2018.
Thus, annual production in 2018 was 148.5 tons, 1028.3 tons and 1034.1 tons, respectively, and
shows an increasing trend (Mequanent et al., 2022).

2.2. Fish sampling and extraction of stomach contents


The fish sampling sites (S1, S2 and S3) were selected based on the availability of fish species. Fish
specimen collection was conducted between February and April 2021 and between June and
July 2021 in dry and wet seasons respectively. Handlines, longlines, and single filament gillnets
were used with a mesh size of 4, 5, 6, 8, 10 and 12 cm and 25 m in length. Gillnets and longlines
were secured in the evening time (5:30 p.m.) and inspected the following morning (7:30 a.m.).
Length of fishes, such as total length (TL) and standard length (SL), were measured to the nearest
millimeter using a measuring board. The weights of the smallest fish (TW) were measured using
a sensitive balance with a sensitivity of 0.1 g while the larger fish were measured by non-digital
balance. Following the morphological and weight measurements, the fish were immediately
dissected by a dissection kit. Subsequently, the entire stomach of C. gariepinus was isolated and
exerted pressure to extract the contents of the stomach. In addition, the contents of the stomach

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(food) were immediately stored in a 5% formalin solution for further analysis. Finally, all samples
were marked (date of sampling, measurement of length and weight, location of sampling, fish
species, etc.) and transferred to the University of Debre Tabor for further analysis.

2.3. Stomach content analysis


During laboratory analysis, the stomach contents were placed into a Petri dish and investigated
(Hyslop, 1980). Therefore, the smallest food is analyzed using the XSZ-70DN dissection microscope
model and the ST-30-2 L stereo microscope (100 to 400 magnification). The identified food items
were classified to the lowest taxonomic level by using descriptions, keys, and literature (Vuuren
et al., 2006, Carling et al., 2004). The contents of fish’s stomachs were analyzed by relative
measures of prey quantities (RMPQ) (Hyslop, 1980). The relative importance of each food item in
the total food content of the stomach was analyzed based on percentage frequency (%Q) and
percentage volume (%V) (Assis, 1996).

In the frequency of occurrence, the number of stomach samples containing one or more of
a given food item was expressed as a percentage of all nonempty stomachs examined (Hyslop,
1980).

The proportion of fish that consumed certain foods is estimated and the frequency of occurrence
is calculated (Bowen, 1983; Hyslop, 1980). The proportion of the African sharptooth catfish that
feed on certain food items was estimated by this method.

The frequency of occurrence was calculated as:

where %Oi = frequency of occurrence of the i food item in the sample; ni = number of stomachs in
which the i item is found; n = total number of food contained stomachs in the sample (Bowen,
1983).

Food items were sorted into different taxonomic categories, and the water displaced by a group
of items in each category was measured in a partially filled graduated cylinder (Bowen, 1983). The
volume of water displaced by each category of food items was expressed as a percentage of the
total volume of the stomach contents (Bowen, 1983).

The volumetric measurement (%Vi) was estimated as:

Where %Vi is the percentage of volume.

The stomach analysis parameters such as frequency of occurrence (%Qi) and volumetric con­
tribution (%Vi, ml) were used to estimate the index of food preponderance (PIi) and geometric
importance of index (GIIi). To determine the importance of each food item the index of prepon­
derance (PIi) (Tomojiri et al., 2019) was calculated as:PIi ¼ ð%ViÞð%OiÞ

Where Qi is the frequency of occurrence of species i and %Vi is the percent composition by volume
of species i. To facilitate comparisons among species, PIi was converted into percent PIi (%PIi).
Furthermore, to estimate the relative importance of food items and species-level dietary differ­
ences, the Geometric Index of Importance (GIIi) (Assis, 1996) was estimated as:

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Where RMPQi = percentage of volume and frequency of occurrence (as a percentage of total
occurrences) and n = total number of RMPQ. The index of GIIi value ranges from 0 to 1 (1–
100%), with values close to 0 indicating feeding specialization and values close to 1.0 representing
generalization (Hurlbert, 1978).

2.4. Data analysis


Food and feeding habits in relation to season and size class of C. gariepinus in the Ribb Reservoir
were studied using the percentage of volumetric contribution, frequency occurrence, preponder­
ance importance index (PIi), and geometric importance of index (GIIi) of the five size classes
(<37.0, 37.0–42.0, 42.5–47.0, 47.5–52.0 and ≥52.5 cm SL), and seasons (the dry and the wet) of
the year (Tomojiri et al., 2019).

Descriptive statistics were interpreted and analyzed by Microsoft Excel (Windows 10). IBM SPSS
version 20 was also used to analyze the diet composition and feeding habits differences within the
size classes of the C. gariepinus through analysis of variance (ANOVA) at 95% confidence level.
Whereas, the t-test was used to compute the seasonal variation in the diet of the fish.

3. Results

3.1. Overall diet composition and feeding habits of Clarias gariepinus


From this study, 525 specimens of Clarias gariepinus were sampled. Out of these specimens, 379
(72.2%) consisted of different foods, whereas 146 (27.8%) had empty stomachs. The diet composi­
tion of C. gariepinus was comprised of numerous prey items such as zooplankton, fish, phytoplank­
ton, plant detritus, insects, ostracods, sand particles, macrophytes and plant seeds (Table 1).
Zooplankton was the most consumed food type followed by fish prey, phytoplankton, and detritus
in the diet of C. gariepinus. However, insects, sand particles, ostracods, and macrophytes contrib­
uted less to the diet of the fish. The percentage of the geometric importance index (%GIIi)
revealed that zooplankton was the main prey type fed by the fish (Figure 2). Based on this index,
fish prey, phytoplankton, and detritus were the second, third and fourth most important preys in
C. gariepinus diet respectively, whereas sand grains, ostracods, and insects are occasionally
ingested.

Table 1. Proportions of different food items in the diet of C. gariepinus (n = 379) in Ribb
Reservoir, Ethiopia
Food items %Oi %Vi %IP %GIIi
Zooplankton 100 21.8 32.95 46.0
Cladocera 55.7 3.8 3.2 22.7
Copepod 73.1 17 18.8 34.4
Rotifer 10 0.6 0.1 4.0
Fish prey 36.1 48.8 26.6 32.0
Phytoplankton 55.7 5.9 4.96 23.3
Blue-green algae 45.6 5.4 3.7 19.5
Green algae 5.6 0.3 0.025 2.3
Euglenoids 4.5 0.2 0.01 1.8
Detritus 35.6 17.1 9.2 19.9
Insect 17.1 1.5 0.4 7.0
Ostracods 7.1 0.2 0.02 2.8
Sand particles 6 0.2 0.018 2.3
Macrophytes 3.7 0.4 0.01 1.5
Note: percentage frequency of occurrence (%Qi), percentage of volumetric contribution (%Vi), percentage index of
preponderance (%IP) and percentage Geometric Index of Importance (%GIIi).

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Figure 2. The percentage con­ 50


tribution of geometric index 45 Se c onda r y foods
importance (%GIIi) in the diet 40

Primary prey
of C. gariepinus (n = 379) from
35
Ribb Reservoir.
30

GIIi (%)
PKT—Phytoplankton; ZPK— 25
Zooplankton; DET—detritus;
20
SAG-sand grain; and OST-
ostracod. Dotted vertical lines 15 Incident foods
separate the different grades 10
of favorite food items. 5
0
ZPK Fi sh PKT DET INST OST SAG
Prey types

Zooplankton (Rotifers, Copepods, and Cladocera) collectively occurred in 100% of all stomachs
analyzed and comprised 21.8% of the stomach contents by volume. Fish prey occurred in 36.1% of
all stomachs analyzed in the diet and comprised 48.8% of the stomach contents by volume.
Phytoplankton (blue-green algae, green algae, and euglenoids) together consisted of 55.7% of
the total occurrences and accounted for 5.9% of the total volume. The index of preponderance (%
PI) of food types such as zooplankton, fish prey, phytoplankton, and detritus were 32.95, 26.6, 4.96,
and 9.2 respectively. The index of preponderance (%PI = 32.95%) and geometric importance index
(%GIIi = 46.0%) revealed that zooplankton was the dominant food type in the diet of C. gariepinus.
Fish prey (%GIIi = 32.0%), phytoplankton (%GIIi = 23.0%), and detritus (%GIIi = 19.8%) exhibited
the second favorite food types in the diet. However, the remaining food items such as insects (%PI
= 0.4%; %GIIi = 7.0%), ostracods (%PI = 0.02%; %GIIi = 2.8%), sand particles (%PI = 0.02%; %GIIi =
2.3%), and macrophytes (%PI = 0.01%; %GIIi = 1.5%) contributed less in the diet of C. gariepinus
(Table 1 and Figure 2).

3.2. Food and feeding habits in relation to the size class


The food and feeding habit of C. gariepinus was significantly varied in relation to the size classes
(ANOVA, p < 0.05). Detritus was the foremost important food item followed by mud and zooplank­
ton for the size class below 37.0 cm (SL) (Figure 3). Their volumetric contributions were 41.2%,
29.3%, and 19.8% of the total volume in the diet of this size class respectively. But, other food
items such as phytoplankton (%V = 4.7) and fish prey (%V = 5.8) had a slight contribution to the
diet of the fish this size class. According to the index of preponderance (PIi%) and Geometric Index
of importance (GIIi%), zooplankton was the most preferred food item (PIi% = 29.3 and GIIi% =

Figure 3. The size class varia­ 100%


Volumetric contribution of food items

tion in the diet of C. gariepinus 90%


in Ribb Reservoir, Ethiopia. 80%
70%
60%
50%
(%)

40%
30%
20%
10%
0%
<37.0 37.0-42.0 42.5-47.0 47.5-52.0 ≥52.5
Size classes (cm SL)
Detritus Mud Zooplankton Phytoplankton Fish Insects Ostracods

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Figure 4. The relative impor­ 120

Proportion of PIi (%) and GIIi%


tance of prey items (PIi% and 100
GIIi%) with respect to size
80
classes in the diet of
C. gariepinus. 60
40
20
0
PIi% GIIi% PIi% GIIi% PIi% GIIi% PIi% GIIi% PIi% GIIi%
<37.0 37.0-42.0 42.5-47.0 47.5-52 52.5-93
Size classes (cm SL)

Detritus Mud Zooplankton Phytoplankton Fish Insects Ostracods

76.2) followed by detritus (PIi% = 44.34 and GIIi% = 65.5), phytoplankton (PIi% = 5.06 and GIIi% =
51.8), and mud (PIi% = 19.7 and GIIi% = 42.4) to the diet of this size class (Figure 4).

Zooplankton was the most important food type for size interval between 37.0 and 42.0 cm (SL).
It accounted for 31.2% of the total volume of the stomachs. In addition, detritus, fish prey,
phytoplankton, and mud were the following most preferred food items in this size class and
their volumetric contributions were 25.6%, 14.0%, 12.4%, and 10.5% respectively. While, remaining
food items such as ostracods, and insects had the least importance in the diet of the fish.
Considering the index of preponderance (PIi%) and Geometric Index of importance (GIIi%), zoo­
plankton was the most preferred food item (PIi% = 55.2 and GIIi% = 96.2) followed by detritus (PIi
% = 21.5 and GIIi% = 49.76), phytoplankton (PIi% = 14.1 and GIIi% = 58.9), and fish prey (PIi% = 5.8
and GIIi% = 25.1) to the diet of this size class (Figure 4).

Fish prey was the most frequent food item for the size class between 42.5 and 47.0 cm (SL). It
consisted of 31.1% of the total volume in the diet of this size class. Detritus and zooplankton were
the second most important food items in this size class and their volumetric contributions were
24.1%, and 21.3% respectively. However, other food items such as ostracods, phytoplankton, and
insects had less contribution to the diet of the fish this size class. As per the index of preponder­
ance (PIi%) and Geometric Index of importance (GIIi%), zooplankton was the most preferred food
item (PIi% = 44.8 and GIIi% = 83.1) followed by phytoplankton (PIi% = 9.7 and GIIi% = 54.8), detri­
tus (PIi% = 23.0 and GIIi% = 43.2), and fish prey (PIi% = 18.8 and GIIi% = 33.4) to the diet of this
size class (Figure 4).

Fish prey was the most important food item for the size class between 47.5 and 52.0 cm (SL). It
comprised 50.3% of the total volume of the diet. Detritus was the second most favored food item
in this size class and its corresponding volumetric contribution was 22.9%. While, the remaining
food items such as ostracods, phytoplankton, insects, mud, and zooplankton contributed less
importance to the diet of the fish this size class. As per the index of preponderance (PIi%) and
Geometric Index of importance (GIIi%), zooplankton was the most preferred food item (PIi% = 25.9
and GIIi% = 87.2) followed by fish prey (PIi% = 51.0 and GIIi% = 60.9), detritus (PIi% = 18.8 and GIIi
% = 42.5) and phytoplankton (PIi% = 3.0 and GIIi% = 40.1) to the diet of this size class (Figure 4).

Fish prey was by far the most important food item for the size class ≥52.5 cm (SL). It accounted
for 74.1% of the total volume of diets. Zooplankton was the following chosen food item in this size
class and its volumetric contribution was 11.2%. However, the remaining food items such as
detritus, mud, ostracods, phytoplankton, and insects were ingested in the least amount in the
diet of the fish in this size class. The volumetric contribution of fish prey revealed an increasing
trend with the size of the fish increases. However, the volumetric contribution of detritus, mud,
phytoplankton, and zooplankton exhibited decreasing trend as the fish size increased (ANOVA, p <

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0.05 (Figure 4). Furthermore, as per the index of preponderance (PIi%) and Geometric Index of
importance (GIIi%), fish prey (PIi% = 79.9 and GIIi% = 89.2) was the most favorite food item
followed by zooplankton (PIi% = 14.3 and GIIi% = 76.6), phytoplankton (PIi% = 1.9 and GIIi% =
37.4) and detritus (PIi% = 3.4 and GIIi% = 33.4) to the diet of this size class (Figure 4).

3.3. Seasonal variation of the diet


The seasonal differences in the diet of C. gariepinus were presented in (Table 2) from Ribb
Reservoir. In this study, the results revealed that seasonal variations in the diet of the fish were
observed (t-test, p < 0.05). During the dry season, fish prey (%IP = 64.5 and %GIIi = 40.5) was the
most important food item accounted for 66.1% of the total stomachs, and comprised 63.6% of the
total volume. Zooplankton (%IP = 7.3 and %GIIi = 32.6), and phytoplankton (%IP = 1.2 and %GIIi =
20.6) also secondly consumed food items occurred in 99.5%, and 64.6% of the total stomach
contents and their volumetric contributions were 4.8%, and 1.2% of the total volume in the diet of
fish during the dry season respectively (Table 2). The remaining food types such as mud (%IP = 2.4
and %GIIi = 8.8), insect (%IP = 0.1 and %GIIi = 7.9), ostracod (%IP = 0.04 and %GIIi = 4.3), sand
particles (%IP = 0.04 and %GIIi = 3.7), macrophytes (%IP = 0.01 and %GIIi = 2.4), and plant seeds
(%IP = 0.01 and %GIIi = 0.5) had less contribution to the diet of the fish.

In the wet season, zooplankton and phytoplankton were by far the most important prey type in the
stomachs of C. gariepinus. These prey types occurred in 100% and 96.8% of the total stomachs and their
respective volumetric contributions were 71.8% and 22.2% of the total volume respectively. The %IP and
%GIIi indexes also exhibited that zooplankton (%IP = 40.3 and %GIIi = 71.6) was by far the most
important food items in the diet during the wet season. Phytoplankton (%IP = 12.1 and %GIIi = 49.6)
was the second most important food item in the diet of the fish. However, detritus (%IP = 0.08 and %GIIi
= 4.0), insects (%IP = 0.08 and %GIIi = 3.9), fish prey (%IP = 0.11 and %GIIi = 3.8), and mud (%IP = 0.003
and %GIIi = 0.8) had less contribution to the diet of the fish during the wet season (Table 2).

4. Discussion

4.1. Diet composition and feeding habits


In this study, 525 specimens of Clarias gariepinus were sampled. Out of these specimens, 379
(72.2%) were found with prey items in their stomachs, whereas 146 (27.8%) had empty stomachs.
Most empty stomachs may be related to the digestive process after the fish has been caught.
Similarly, inappropriate sampling of fish also leads to an accelerated digestion process (Tesfahun &
Alebachew, 2023). The authors confirmed that a great proportion of fishes with empty stomachs
were recorded in various Ethiopian water bodies when fishes were sampled with gillnets (Dadebo
et al., 2014; Temesgen et al., 2022; Wagaw et al., 2022). This can be due to the fact that food in
their stomach may be vomited or assimilated as fish fight to escape the fishing gear during fishing.

In this study, different food types were documented in the stomachs of C. gariepinus including
zooplankton, fish prey (the Nile tilapia and Labeobarbus spp.), phytoplankton, detritus, insects,
ostracods, macrophyte, and sand particles in Ribb Reservoir. Several studies also confirmed the
presence of the aforementioned prey types in the diet of C. gariepinus in Lake Hawassa (Dadebo,
2000); Lake Langano (Teka & Admassu, 2016); Lake Babogaya (Admassu et al., 2015); Lake Hayq
(Dereba, 2019); Lake Koka (Dadebo et al., 2014), Lake Chamo (Dadebo, 2009), Egbe Reservoir
(Adewumi et al., 2014), Lake Alau (Wakil et al., 2014), Tungan Kawo Reservoir (Habiba, 2021)
and Lake Hawassa and Shallo swamp (Tekle-Giorgis et al., 2016).

Zooplankton was by far the most important food type in the diet of C. gariepinus from Ribb
Reservoir. This may be a result of an abundance of zooplankton owing to suitable physico-chemical
conditions in the Ribb Reservoir (Bhuyan et al., 2020; Mequanent et al., 2022). The diversity of
zooplankton also varies depending on many factors, including geographical regions, feeding type,
and morphometric characteristics of water (Vijayakumari et al., 2018). The abundance and diver­
sity of zooplankton also varied with seasonal fluctuations. The average zooplankton biodiversity

Page 8 of 14
Table 2. Relative contribution percentage frequency of occurrence (%Qi), volumetric contribution (%Vi), index of preponderance (%PIi) and geometric index of
importance (%GIIi) of different food items in the diet of Clarias gariepinus in Ribb Reservoir during the dry and wet seasons
https://doi.org/10.1080/23311932.2023.2284228

Food items %Oi %Vi %PIi %GIIi


Dry Wet Dry Wet Dry Wet Dry Wet
Fish prey 66.1 5.4 63.6 3.7 64.5 0.11 40.5 3.8
Zooplankton 99.5 100 4.8 71.8 7.3 40.3 32.6 71.6
Tesfahun & Alebachew, Cogent Food & Agriculture (2023), 9: 2284228

Cladocera 36.5 62.9 1.6 10.5 0.89 3.7 11.9 30.3


Copepod 48.4 94.1 2.4 61.1 1.8 32.3 15.9 64.7
Rotifer 14.6 3.2 0.8 0.2 0.2 0.003 4.8 1.4
Phytoplankton 64.6 96.8 1.2 22.2 1.2 12.1 20.6 49.6
Blue-green algae 49.5 92.5 0.7 21.8 0.5 11.3 15.7 47.6
Green algae 10.4 - 0.4 - 0.1 - 3.4 -
Euglenoids 4.5 3.8 0.2 0.4 0.02 0.01 1.5 1.8
Mud 20.8 1.6 7.4 0.4 2.4 0.003 8.8 0.8
Insect 25.0 7.6 0.2 1.8 0.1 0.08 7.9 3.9
Ostracods 13.5 - 0.2 - 0.04 - 4.3 -
Sand particles 11.5 - 0.2 - 0.04 - 3.7 -
Macrophytes 7.3 - 0.5 - 0.01 - 2.4 -
Plant seeds 1.6 - 0.03 - 0.001 - 0.5 -

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index values of the Ribb reservoir, such as the dominance index (0.902) and the Shannon diversity
index (H′) (2.47) indicate that the reservoir is in a good state (Mequanent et al., 2022). Besides, the
abundance of zooplankton in the diet of C. gariepinus might be associated with a high production
of zooplankton due to high temperature and nutrient availability this makes C. gariepinus con­
sumed high bulk of zooplankton (Wakil et al., 2014). In agreement with the present study, Lakes
Langano (Teka & Admassu, 2016) and Chamo (Dadebo, 2009) conveyed zooplankton as the main
food source in the diet of C. gariepinus. However, the food items such as detritus, insects, macro­
phytes, and fish prey were reported as the main food component from Lakes Koka (Dadebo et al.,
2014), Hayq (Dereba, 2019), Hawassa and Shallo swamp (Tekle-Giorgis et al., 2016), Alau Nigeria
(Wakil et al., 2014), Babogaya (Admassu et al., 2015) and Tungan Kawo reservoir Nigeria (Habiba,
2021). Fish preys (O. niloticus and Labeobarbus spp.) were the second most important food items in
the diet of the fish. The piscivorous feeding behavior of the C. gariepinus was also reported in Lakes
Koka (Dadebo et al., 2014), Alau Nigeria (Wakil et al., 2014), Hayq (Dereba, 2019), Langano (Teka &
Admassu, 2016), Hawassa (Desta et al., 2006; Dadebo, 2000). However, phytoplankton was
reported as the major food type in Egbe reservoir (Adewumi et al., 2014). The variability of food
and feeding habits of fish may be due to the different prey availability in different waters. The
variations in prey availability is likewise linked with nutrient availability and the characteristics of
the physicochemical of waters. Moreover, variations in prey availability may be linked to changes in
the environment and biological factors of the freshwater, which impact the food type in the diet of
this fish (Temesgen et al., 2022). Information on food and feeding habits of freshwater fish species
is a subject of continuous research. Because it makes up a basis for the development of
a successful management program on fish culture and capture (Temesgen et al., 2022).
Furthermore, studies on the natural feeding habit of fish enable us to identify the trophic relation­
ships present in aquatic ecosystems, identifying feeding composition, structure and stability of
food webs (Teka & Admassu, 2016). Now a days the natural water bodies of Ethiopia experiencing
pollution at alarming rate. This phenomenon makes declining the wild capture fishery. The aqua­
culture is the alternative option of fish production for this fast-growing population. Therefore,
identifying the natural food and feeding habits of this fish species is important for in view of
aquaculture applications as well as climate change.

4.2. Food and feeding habits in relation to the size class


In this study, size class variations in the diet C. gariepinus were observed. The smallest size class (<37
cm SL) primarily fed on detritus, zooplankton, and mud. Dadebo et al. (2014) also reported detritus and
insects as the main food component for the smallest fish in Lake Koka. Admassu et al. (2015) and
Tekle-Giorgis et al. (2016) also reported that the smallest fish consumed more insects and zooplankton
whereas, adult fish ingested fish prey in Lakes Babogaya and Hawassa and Shallo swamp respectively.
Tesfahun and Temesgen (2018) also reported the dominance of insects and zooplankton in the diet of
the smallest fish while the larger fish had more fish prey and zooplankton as well in Ethiopian waters.
This might be due to, the smallest fish requiring a high amount of protein to improve their growth rate
and metabolism (Temesgen et al., 2022). Other researchers also confirmed that C. gariepinus exhibited
diet shifts because of the change in habitat use in different water bodies (Dadebo, 2000, 2009). Dadebo
(2000) studied size-based differences in feeding habits of the smallest C. gariepinus fed more on
insects. In the present study, larger C. gariepinus with size classes 37.0–42.0 cm SL, 42.5–47.0 cm SL,
47.5–52.0 cm SL and ≥52.5 cm SL primarily fed on zooplankton and fish prey. The importance of fish
prey progressively increased, while the contributions of detritus, zooplankton, and mud decreased with
fish size. The fish shifted its feeding habit from detritus, zooplankton, and mud to fish prey from Ribb
reservoir as the fish length increased. The ontogenetic dietary shift may be due to the productivity of
the water and habitat differences of the fish (Tekle-Giorgis et al., 2016)). Furthermore, this could be
because large-sized C. gariepinus living deeper waters, whereas small ones exist in shallow waters
among macrophytes as a refuge where bulks of benthic macroinvertebrates are usually high. In
addition, the life history of fish depends on habitat use, and their feeding habits differ significantly
(Njiru et al., 2004; Temesgen et al., 2022; Wagaw et al., 2022).

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4.3. Seasonal variations in the diet of Clarias gariepinus


There was a seasonal variation in the diet of C. gariepinus from Ribb Reservoir. In the present study,
fish prey was by far the most important food item followed by zooplankton in the diet of
C. gariepinus during the dry season. Many of the larger fish species consumed small fishes as
source of food. This might be due to C. gariepinus is the top predator in the reservoir it feeds on
small fishes. The other reason may be the piscivory nature of C. gariepinus associate with water
level fluctuation. For instance, when the water level decreases the prey fishes aggregate some­
where this creates good opportunity for the predator fish (Tekle-Giorgis et al., 2016). Alebachew
et al. (2022) also confirmed that the water level of the Ribb Reservoir decreases in the dry season
when the water needed for irrigation purposes. According to Spataru et al. (1987) piscivory is
common in fishes with presence of oral teeth, tough stomach and stomach acidity for digesting of
large preys. During the dry season, zooplankton was the second most important food item in the
diet of C. gariepinus. This might be due to high nutrient availability and good water quality that
supports production of zooplankton biomass in the reservoir. During the dry season, the contribu­
tion of phytoplankton was less in the diet of C. gariepinus. This may be due to the turbidity that
might have cause a low light penetration in the reservoir and reduced photosynthetic rate. The
high turbidity of the reservoir may be due to the water abstraction and water release for irrigation
purposes (personal observation). The seasonal variation in the diet of the fish may be due to the
opportunistic habits of the fish which has the ability of shifting from one food item to another food
item. Besides, the seasonal variation of the food and feeding habits of C. gariepinus also may be
due to the seasonal production difference of the preys in the reservoir. The food items such as
mud, insects, ostracods, sand particles, macrophytes and plant seeds had minor contribution in
the diet of the fish. This is may be due to the food and feeding habits of the fish depending on the
availability of preys in the reservoir. For instance, the contribution of macrophyte in the diet of the
fish was minor, this may be due to the less importance of macrophytes in the diet of C. gariepinus
during the dry season in Ribb Reservoir might be due to decrease of nutrients that facilitate the
growth of macrophytes. The most important factor that determine the availability and emergence
of various food items in Ribb Reservoir could be the seasonal changes in water level.

On the other hand, zooplankton and phytoplankton were the main prey type in the wet season.
During the wet season the contribution of zooplankton in the diet of C. gariepinus was high in
reservoir. According to Dadebo (2009) high productivity of the water bodies in terms of zooplank­
ton biomass that leads to shift the feeding habits of C. gariepinus to zooplankton filter feeding. In
the wet season, the high zooplankton production may be due to high water temperature in the
Ribb Reservoir. In addition, some reasons could be mentioned to elaborate this high consumption
of zooplankton. It is known that the Ribb Reservoir is tropical, may be eutrophic and have high
water temperature that promote high level of phytoplankton, and thus high zooplankton produc­
tion. As the biomass of zooplankton increases, the fish may use ram feeding by opening its mouth.
According to Dadebo (2009) C. gariepinus has the anatomical adaptations for filter feeding, which
allows the fish to feed on prey ranging from zooplankton to a fish half its own length. This may be
the reason it shifted from piscivorous feeding habit to ram feeding. The other reason may be the
increment of water level when the water level increases the prey fishes may be unevenly dis­
tributed which makes C. gariepinus may not be able to compete for the available prey fish in such
an environment. Phytoplankton was also the second most important food item in the wet season.
The high contributions of phytoplankton in the diet of C. gariepinus during the wet season in
reservoir might be due to increase of nutrients during the floods that promote the growth of
phytoplankton. The remaining food items such as insects, fish, macrophytes and mud had minor
contribution in the diet of C. gariepinus in the reservoir.

Many investigators also documented a seasonal variation of food items in the diet composition
of C. gariepinus in various water bodies (Adewumi et al., 2014; Admassu et al., 2015; Dadebo et al.,
2014; Dereba, 2019; Eyayu, (2019); Habiba, 2021; Teka & Admassu, 2016; Tekle-Giorgis et al., 2016;
Tesfahun & Temesgen, 2018; Wakil et al., 2014). However, phytoplankton, mud, insects, ostracods,
sand grains, macrophytes, and plant seeds have insignificant contributions to the bulk of the fish in

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the dry season. Comparable studies were also conducted on Lakes Koka (Dadebo et al., 2014) and
Hawassa (Tekle-Giorgis et al., 2016) fish prey and zooplankton as the main food sources in the dry
season. From this study, zooplankton and phytoplankton were important prey types in the wet
season. In contrast, detritus, macrophytes, gastropods, and insects were reported as the main
consumed food types in Lake Koka (Dadebo et al., 2014). Other investigators also found different
food items in the wet season from different lakes in Ethiopia. For instance, Teka and Admassu
(2016) investigated that fish prey was the most preferred food item from Lake Langano in the wet
season. Moreover, in Lake Hawassa (insects and fish eggs) and Shallo swamp (macrophyte and
detritus) were the most ingested food items in the wet season (Tekle-Giorgis et al., 2016). The most
important factors influencing the availability and appearance of different food in Ribb reservoirs
may be seasonal changes in water levels. The high contribution of zooplankton and phytoplankton
to the diet of C. gariepinus during the humid months of the Ribb Reservoir could be due to
increased nutrients during floods promoting phytoplankton.

5. Conclusions
This finding established zooplankton as the main food item, followed by fish prey, phytoplankton, and
detritus. The smallest-sized fish mainly fed on detritus followed by zooplankton and mud whereas the
adult fish fed majorly on fish prey followed by zooplankton. The prey types such as fish prey followed by
zooplankton and phytoplankton were the most favored food types during the dry season. Whereas
zooplankton followed by phytoplankton and detritus were the most significant food items in the diet
C. gariepinus in the wet season. C. gariepinus revealed omnivorous feeding habits in its diet from the
Ribb Reservoir. Generally, the diet composition and feeding habits of fish are affected by habitat
differences, seasons, and the size class of fish in the reservoir. Therefore, watershed management is
required to improve the food and feeding habits of this fish species for better sustainable use.

Acknowledgments 1822) from Ribb Reservoir, South Gondar, Ethiopia,


We thank the Guna Tana Integrated Field Research Agumassie Tesfahun & Sale Alebachew, Cogent Food &
Development Centre Debre Tabor University for helping Agriculture (2023), 9: 2284228.
financial provision and vehicle support. We also thank
the fishermen of Ribb Reservoir for the helping with the References
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