813 1308 1 PB
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*Corresponding author: YEMDJIE MANE Divine Doriane, University of Douala Cameroon; Email:
[email protected]
Article history; Received: October 5th 2020; Revised: October 25 th 2020; Accepted: December 25th 2020
Abstract
The feed preferences of Conraua goliath, an endogenous frog in central Africa were assessed in the present
study. A total of 65 frogs (22 males, 22 females and 21 unspecified frogs) were obtained from hunters in 3
localities namely: Loum, Penja, and Nlonako in the Moungo division, littoral region of Cameroon. The methods
of dissection, measurement, and stomach content analysis were used. The results revealed a large diversity of
preys in the stomach namely: myriapods (25%), plants leaves and steems (21.19%), insects (19.56%),
indigestible matters (19.56%), arachnids (4.89%), crustaceans (4.34%), mollusks (3.26%), amphibians (0.54%),
annelids (0.54%) and reptiles (0.54%). The numerical percentage and the frequency of myriapods occurrence
were significantly (P < 0.05) higher in males captured in the locality of Loum (100% and 35.55%) compared to
the females (28.57% and 18.18%). Comparing the state of maturity, the numerical percentage and the frequency
of occurrence of insects (100% and 42.85%) and indigestible matters (100% and 42.85%) were significantly
higher in the youngs frogs (61.11% and 20.89%) captured in Loum compared to the adults (22.22% and 7.46%).
Frogs of high weight (˃1000g) recorded significantly (P < 0.05) higher occurrence and numerical frequency of
myriapods (100 and 70%), indigestible and detritus materials (100 and 40%) in low-weight frogs (<400g) and
plants leaves and steems (50 and 47.36%) in medium-weight frogs (400 – 1000g). In conclusion, goliath frog is
an omnivore fed on a variety of preys and plants according to their availability in the environment.
Key words: Conraua goliath, feed preferences, frog, stomacal conten
Introduction
The world is facing a massive decline in its biodiversity of the amphibians (IUCN, 2020), due to over
exploitation and the fragmentation of their habitats (Collins and Storfer, 2003; Bellard et al., 2016). In
2006, among the 4035 amphibian species that depended on water during their life cycle, 1356
approximately 33.6% were considered threatened (Maxwell et al., 2016). Emerging infectious
diseases, such as chytridiomycosis, (Harp et al., 2006; Voordouw et al., 2010; Cunningham et al.,
2015), climate change (Walther et al., 2002; Parmesan and Yohe, 2003; Foden et al., 2013; Maxwell et
al., 2016), chemical contaminants (Blaustein et al., 2002) and the introduction of predatory species
(Kats and Ferrer, 2003) are factors that affect the dynamism of the populations, causing a drop in their
number. However, it must be noted that most of the overfished amphibians in the world are edible
frogs (Neveu, 2004; Casimir et al., 2016). This is the case of Conraua goliath, a giant African frog
listed as endangered species by IUCN (2004).
In Cameroon, goliath frogs is endemic to the Moungo and Nkam regions where its constituted an
integral part of the eating habits of the indigenous population. The conservation of this species through
its domestication remains the best prospect to ensure its sustainability (Teletchea, 2017). Data on the
diet of a species can help to identify the environmental conditions favorable to its development as well
as the consequences induced by the degradation of its living environment (Parker and Goldstein,
2004).
© 2021 The Author(s). This open access article is distributed under a Creative Commons Attribution (CC-BY) 3.0 license.
Monthe Moumegni et al, (2021) Gen. Biodiv. J: 5 (1): 136-146
Thus, explaining the fluctuation of their population (Lips et al., 2005) and defined conservation
strategies (Batista et al., 2011). Studies on bioecology and feeding habit of the goliath frog are limited.
Only few studies have been carried out on neighboring species, in Europe, Asia, America and Africa
(Bellakhal et al., 2010; Benício et al., 2011; Camera et al., 2014). The objective of this study is to
contribute to the preservation and enhancement of endogenous species through a fairly extensive
analysis of its feeding regime in the natural environment.
Material and Methods
Area of Study
This study was carried out in the Moungo division, Littoral region of Cameroon (Figure 1), precisely
in Loum (LN: 4o41 'and LE: 009o43'), Penja (LN: 4o45 'and LE: 009o 45') and Nlonako (LN: 4o40' and
LE: 009o43') with an average altitude of 665m.
Relief, Pedology, Climate and Hydrography
The study area is characterized by three main types of relief, namely the plains along the Moungo river
and the surroundings of Penja, the scattered plateaus, and mountain ranges. The soils are dominated by
inceptisols (volcanic soils) on its northern and western part. The southern and eastern parts are
dominated with sandy soil, silty-sandy and clay-sandy textures. The climate is of equatorial type,
strongly influenced by the Guinean monsoon. It is characterized by a short dry season (November to
March) and a long rainy season (April to October). The average annual precipitation is 2700 mm, the
temperature varies between 20 and 27oC and a relative humidity of 75%. The hydrographic network is
dense and includes three main rivers namely the Dibombè, Mabombè, Nkam river, multitude of rivers
and streams. These rivers are permanent and their rate of flow varies with the season.
Figure 1: Localization of the area of study and hydrographic network (ORSTOM, 1973)
Flora and Fauna S
The flora along the rivers is very diverse and dominated by a dense rain forest. We note the presence
of industrial plantations (pepper, cocoa, plantain cultivation), Puereria pubescens species as a ground
cover plant and a diversity of plant species. The fauna is terrestrial (Potamochoerus porcus, Manis
tricuspis, Cercocebus sp., hinds, antelopes, hares, and Arthropods), aerial (Acedinidae, Anatidae,
Myliobatidae), and aquatic (Clariidae, Cyprinidae, Cichlidae, Channidae, Potnidae, Channidae, sea
turtles, Conraua goliath, Rana esculenta, Conraua robusta, Crocodilia sp., Varanus niloticus). Many
predatory snakes and aquatic invertebrates are also encountered there.
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Monthe Moumegni et al, (2021) Gen. Biodiv. J: 5 (1): 136-146
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Monthe Moumegni et al, (2021) Gen. Biodiv. J: 5 (1): 136-146
- Complementary prey of the first order (CPFO) and of the second order (CPSO): Q˂10 and F˃10%
and Q˂10 and F<10%.
Where F is the frequency index of a prey with F = Number of stomachs containing the item i × 100 /
Total number of full stomachs, Q is the food quotient with: Q =%Cw ×%Cn or Cn being the percentage
by number and Cw is the percentage by weight of a prey. %Cn = 100 × Number of individuals of each
type of prey / Total number of prey ingested per individual and %Cw = 100 × Weight of each type of
prey consumed / Total weight of prey ingested per individual.
Statistical Analysis
The data collected were subjected to analysis of variance (ANOVA). Where there was significant
differences between the means, Duncan's Multiple Range test was applied to separate them at 5%
significance level. The statistical software SPSS 20.0 (Statistical Package for Social Sciences) was
used for these analysis and the map location of the capture sites with the MAPINFO 7.8 software.
Results
The different categories of prey consumed by Conraua goliath in the localities of Loum, Penja and
Nlonako are summarized in Table 1. It appears that the preys were divided into 5 categories, namely
preferential primary prey (diplopod, indigestible matters and detritus), frequent secondary prey
(decapod, beetle and plant materials), accessory secondary prey (gastropod), first-order
complementary prey (aranea, chilopod, orthoptera) and second order complementary prey (anoure,
squamata, odonate, hemiptera, hymenoptera, dermaptera, annelid, diptera). These include the presence
of terrestrial, aquatic and semi-aquatic prey. The diplopods and plant matters were more numerous and
abundant in the stomachs. The percentage weight and relative index of importance were respectively
higher. Indigestible matter and litter have a high frequency of occurrence in the stomach.
Table 1: Category of food (taxon) consume by Conraua goliath
Categories of feed
N(%) W(%) V(%) RII(%) O(%) Q(%) Classification
Annelid 1.08 0.01 0.01 3.92 3.57 0.02 Q<10% and %O < 10% : (CPSO)
Aranea * 4.89 0.29 0.11 80.45 16.07 1.43 Q<10% and %O ˃10% : (CPFO)
Chilopod 3.80 1.77 0.93 59.22 12.5 6.74 Q<10% and %O ˃10% : (CPFO)
Coleoptera* 11.41 1.91 0.46 403.00 33.92 21.89 10% < Q< 100 % and %O ˃ 10% : (FSP)
Decapode * 4.34 10.65 23 390.75 14.28 46.32 10% < Q< 100 % and %O ˃ 10% : (FSP)
Dermaptera 2.17 0.03 0,05 11.96 5.35 0.08 Q<10% and t %O < 10% : (CPSO)
Indigestibles matter
and detritus 20.10 7.56 1.58 1045.71 48.21 152.11 Q ˃ 100% and %O ˃ 30% : (PMP)
Diplopod 20.65 59.81 10.62 1396.18 44.64 1235.29 Q ˃ 100% and %O ˃ 30% : (PMP)
Diptera 1.08 0.02 0.0049 3.89 3.57 0.03 Q<10% and %O < 10% : (CPSO)
Gasteropod * 3.260 8.06 1.99 37.51 7.14 26.28 10% < Q< 100 % and %O < 10% : (PSA)
Hemiptera * 0.54 1.42 0.60 2.04 1.78 0.77 Q<10% and %O < 10% : (CPSO)
Hymenoptera 0.54 0.003 0.004 0.97 1.78 0.001 Q<10% and %O < 10% : (CPSO)
Plant matters 20.65 1.55 47.60 1950.25 28.57 32.04 10% < Q< 100 and %O ˃ 10% : (FSP)
Odonate 0.54 0.09 0.016 0.99 1.78 0.05 Q<10% and %O < 10% : (CPSO)
Orthoptera 3.80 1.73 0.41 52.794 12,5 6.60 Q<10% and %O ˃10% : (CPFO)
Anoure 0.54 3.96 2.53 5.50 1,78 2.15 Q<10% and %O < 10% : (CPSO)
Squamata 0.54 1.05 10.02 18.87 1.78 0.57 Q<10% and %O < 10% : (CPSO)
N: numerical percentage; W: weight percentage; V: volumetric percentage; RII: relative index of importance; O: frequency
of occurrence; Q: food quotient; PMP: Preferred main prey; OMP: Occasional main prey; FSP: Frequent secondary prey;
PSA: Secondary accessory prey; CPFO: First order complementary prey; CPSO: Second order complementary prey. *:
Aquatic and semi-aquatic prey; the largest clue is in bold.
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Monthe Moumegni et al, (2021) Gen. Biodiv. J: 5 (1): 136-146
The categories of food illustrated in photo 1 was composed of detritus, indigestibles matters, plants
(plant matter), invertebrate (insects, crustacean, molluscs, myriapod, arachnid and annelid) and
vertebrate (amphibians and reptile).
A B C
D E F
G H I
J
Figure 1: Galery of food categories consumed by Conraua goliath (a) Diplopode; (b) beetle; (c)
amphibian; (d) decapod (e); gastropod (f); plant matter (g) indigestible matter; (h); hemiptera; (i)
odonate; (j) chilopod.
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The frequency of occurrence of prey according to the gender and state of maturity as illustrated in
Figures 2 and 3 shows that, irrespective of the factors considered, organic matter of animal origin was
significantly higher (P <0.05) in the stomachs of Conraua goliath. Their frequency of occurrence was
significantly represented (P <0.05) in the stomachs of males (95.23%). The values obtained in adults
(86.48%) and youngs (89.47%) frogs were comparables. The frequency of occurrence (42.85%) of
plant organic matter recorded in males were significantly higher compared to the frequency of
indigestible and detritus materials (19.04 %).
90 90 86.48 78.94
78.4 80
80
70 70
60
60 50
50 42.85 40.06 38.9
40 38.62 40
28.32
30
30 19.04 20 15.78
20 10
10 0
0 Animal Plant Indigestible
organic organic matter and
matter matter detritus
Figure 2: Frequency of occurrence of the type of Figure 3: Frequency of occurrence of the type of
organic matter consumed by Conraua goliath organic matter consumed by Conraua goliath
according to the gender according to the state of maturity
Food Preferences by Locality and Gender of Conraua goliath
Frequency of occurrence
Numerical percentage
90 85.71
80
70
Myriapods
60
50 42.85
37.5
40
30
18.18
20
10
0
Male Female
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Monthe Moumegni et al, (2021) Gen. Biodiv. J: 5 (1): 136-146
The frequency of occurrence and the numerical percentage of myriapods (figure 4) recorded
significantly higher values in males (42.85 and 85.71%) compared to females (18.18 and 37.50%).
The prey was more diversified in the stomachs of frogs captured at Loum, followed by Penja and
Nlonako respectively. Reptiles and annelids were absent irrespective of the location and gender. With
equal diversity, the myriapods were significantly (P <0.05) more frequent and numerous regardless of
the locality considered. Depending on the localities (figure 5), myriapods and insects were
significantly less frequent (25 and 0%) in Nlonako compared to the frogs from Penja (55 and 40%)
and Loum (60 and 55%). Food Preferences According to Locality and State of Maturity of Conraua
goliath
Food preferences of Conraua goliath with respect to the locality and state of maturity (Figures 6)
indicate that regardless of the localities and states of maturity, frogs mainly consumed myriapods
(25%), plant materials (21.19%), insects (19.56%), indigestible materials and detritus (19.56 %),
arachnids (4.89%), crustaceans (4.34%), gastropods (3.26%), annelids (0.54%), amphibians (0.54%)
and reptiles (0.54%). Depending on the locality, insects were more frequent in the stomachs of adults
frogs found in Loum (table 2). In this particular locality, the frequency of occurrence, of insects (100
and 42.85%) and indigestible matter (100 and 42.85%) were significantly higher in youngs frogs
captured in comparison to the adults frogs respectively (61.11 and 20.89%) and (22.22 and
7.46%).
30
Food preferences (%)
25
25 21.19 19.56 19.56
20
15
10 4.89 3.26 4.34
5 0.54 0.54 0.54
0
Feed category
Figure 6: Food preferences of Conraua goliath regardless the locality and state of maturity
Table 2: Frequency of occurrence and numerical percentage of insects and indigestible matter by
state of maturity in Loum
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weight. Frogs of high weight (˃1000g) recorded significantly (P<0.05) higher occurrence and
numerical frequency of myriapods (100 and 70%), indigestible and detritus materials (100 and 40%)
in low-weight frogs (<400g) and plants leaves and steems (50 and 47.36%) in medium-weight frogs
(400 – 1000g).
Table 3: Food preferences with respect to weight of Conraua goliath
Discussion
The analysis of the stomach contents of Conraua goliath shows that this frog consumes a wide
varieties of feed. Feed from animal origin are predominant both in frequency of occurrence, and in
numerical percentage suggesting that this frog is omnivorous with carnivorous tendency. These preys
are terrestrial, aquatic and semi-aquatic in nature, indicating that frogs foraged in water and on land.
This result corroborates the results of the studies carried out in Tunisia on Rana saharica, in which
adults consume 96.32% of terrestrial preys and 3.68% of aquatic preys (Bellakhal et al., 2010). Study
on Rana ridibunda in Russia (Mordovia) revealed that this species fed mainly on aquatic preys
(Ruchin and Ryzhov’s, 2002). Other investigations on the diet of ranidae revealed that they fed
exclusively on terrestrial preys (Berry, 1965; Jenssen and Klimstra, 1966; Beschkov, 1970; Whitaker
et al., 1981; Hirai and Matsui, 1999; 2001a). Feed from plant origin is also frequent in the stomachs in
high number irrespective of the factor of variation considered. The ingestion of plant materials is
recurrent in many anurans (Batista et al., 2011; Maragno and Souza, 2011; Pazinato et al., 2011;
Sabagh et al., 2012). However, many authors reported that they are accidentally ingested during food
capture (Whitaker et al., 1977; Evans and Lampo, 1996; Anderson et al., 1999; Teixeira et al., 1999).
The variability in the frequency of occurrence and the number of feed with respect to locality, sex,
state of maturity, size and weight of Conraua goliath is attributed to the fact that the choice of preys is
above all it’s availability and also edaphic characteristics of the surrounding environment (Werner et
al., 1995; Das, 1996; Low and Török, 1998; Hirai and Matsui, 1999; 2001b), due to the dynamics of
prey populations (Hirai and Matsui, 2001a; Maneyro et al., 2004; Santos et al., 2004). This variability
can also be linked to the fact that in the same species, there would be disparities in the diet according
to gender and size. Studies on Leptodactylus latrans in Uruguay and Argentina shows great overlap in
diet that exists between males and females (Maneyro et al., 2004; Sanabria et al., 2005). However,
certain investigations attested that there is no difference in diet between the gender. This is the case of
Rhinella scitula in Brazil (Maragno and Souza, 2011) and Ptychadena mascareniensis in Madagascar
(Vences et al., 2004).
The composition of Conraua goliath diet is more diversified in youngs ones, thus reflecting
opportunistic and passive eating behaviors. Studies on the diet of genus Leptodactylus, show that there
is a large quantity of larvae and more sedentary groups, describing an opportunistic feeding behavior
(Maneyro et al., 2004; Rodrigues et al., 2004, Sanabria et al., 2005; De Carvalho et al., 2008; Solé et
al., 2009; Solé and Rödder, 2010). This can be explained by the fact the consumption of larger prey in
small numbers by frogs is sufficient to cover their needs, unlike small prey.
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Monthe Moumegni et al, (2021) Gen. Biodiv. J: 5 (1): 136-146
Conclusion
The goliath frog is an omnivore with a carnivorous tendency and fed on a variety of preys according to
their availability in the environment. Preys are more diversified in youngs frogs than adults ones with
the myriapods been the more frequent and numerous.
Acknowledgement
Grateful acknowledgement is expressed to the international fund for Science (IFS) for the financial
support of the present work. The authors wish to thank all participating fishermen and laboratory
technicians for their cooperation.
Author’s Contributions
MONTHE MOUMEGNI Liliane Fleurette, YEMDJIE MANE Divine Doriane, NGOUANA
TADJONG Ruben and MAFOUO SONHAFOUO Vanessa went to the field to carry out the research
and collect the samples. KANA Jean Raphael supervised the overall research work. MONTHE
MOUMEGNI Liliane Fleurette, YEMDJIE MANE Divine Doriane and KANA Jean Raphael wrote
the first draft before being revised by NGOUANA TADJONG Ruben, MAFOUO SONHAFOUO
Vanessa and TCHOUMBOUE Joseph and approved by all the authors.
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