Chapter 1

Introduction

Anurans are generalist foragers who do not only feed on insects but are known to also feed on a

wide diversity of active invertebrates and vertebrates such as Chilopoda, Annelida, Diplopoda

and molluscs (Yap and Jaafar, 2011; Amadi, 2021). Anurans are essential to the stability of the

trophic communities of most ecosystems, as they are responsible for the consumption of various

species so are they also consumed by diverse predatory species (Hirschfeld and Rodel, 2011;

Federico et al., 2012; Amadi, 2021). Understanding their dietary habits is not only for natural

history studies, but also to comprehend their mode of interactions and patterns of nutrient cycling

within host communities. Studies that entails the killing and dissection of multiple species of

anurans for their stomach contents have been common in the past (Yap and Jaafar, 2011; Jamdar

and Shinde, 2013). However, faecal content and stomach flushing techniques (Sole et al., 2005;

Mahan and Johnson, 2007; Hirschfeld and Rodel, 2011) have greatly reduced the rate at which

reptiles and amphibians die from research related exposures. This study aims to ascertain as

much as possible the dietary habits of the less studied galam lipped frog in Rivers State, Nigeria.

Thus, the galam lipped frogs are members of the Ranidae family, and are found in central,

western and eastern parts of the African continent (Rodel et al., 2004). Although, various studies

on the parasitic contents of the galam lipped frog (A. galamensis) have been conducted in

southern Nigeria by parasitologists (e.g. Aisien and Amuzie, 2017; Aisien et al., 2017), but there

seems to be a paucity of data on the dietary behaviour of this nocturnal frog. Multiple studies

have been conducted on the trophic behaviour of most anurans, but the galam lipped frogs

(Amnirana galamensis) which have mostly been found in freshwater swamps, wetlands, rural

and urban gardens, ponds, ditches, rivers, shrub lands and moist savannahs (Rodel et al., 2004),
have almost been ignored by trophic ecologists. Therefore, this study hopes to employ the novel

dietary preference test method in ascertaining the foraging habits of the galam lipped frogs as

well as their preferred prey in Rivers State, Nigeria.

 Chapter 3

Materials and Methods

The study on the dietary behaviour of the gallam lipped frog was carried out in the premises of

the Rivers State University Campus, and in Rumuagholu community of Obio/Akpor Local Area

of Rivers State, Nigeria.

Research protocol

Ten (10) individuals of the Galam lipped frog were randomly captured from their foraging sites

at night, and then stored in perforated plastic containers. The individuals were randomly captured

between 7-8pm from Rumuagholu (Latitude: 040 53.028’ and Longitude: 0060 58.1135’E) Town

of Obio/Akpor Local Government Area. With the aid of a flashlight, all frogs were captured by

hand while wearing a latex glove. Considerable care was taken in order to reduce the general

possibility of injuring or infecting the frogs with human pathogens. Although, the dietary

preference test was conducted outside, the frogs were kept in the darkest part of the laboratory in

order to prevent their exposure to the harsh effects of sunlight. In order to mimic their

environments and make them comfortable, water for rehydration and soil (sourced from their

habitats) for anchorage were provided for the frogs. The dietary preference test for the Gallam

lipped frogs was conducted outside so as to mimic as much as possible the natural conditions that

governs the foraging behaviour of the frogs.

In order to test for dietary preference, various invertebrates (Apoda, Lepidoptera, Coleoptera,

Hymenoptera, Isoptera, Orthoptera, Hemiptera, Blatodea, slugs, Chilopoda, Aranea, etc), which

were originally captured from the locality of the Gallam lipped frogs, were presented to the

confined predators for food. Although, only the prey whose photoperiod coincided with the
nocturnal activity patterns of the predators was presented to the frogs. The abundance and

taxonomic clade of the captured prey was noted.

The 10 frogs were kept in pairs of 5 perforated plastic containers, i.e. 2 individuals per container.

Live and mobile prey of various kinds were intermittently presented to the contained frogs and

then allowed to feed for 20 minutes without disturbance from light and noise. After 20 minutes

of foraging activities, the cages were searched for concealed prey, and the previously

unconsumed prey was removed. Thus, the prey taxa that was neither removed nor in

concealment was recorded to be preferred (foraged upon) by the frogs.

The dietary assessment of the Gallam lipped frog (Amnirana galamensis) was ascertained by

using the dietary preference test method (Amadi, 2021).

Statistical analysis

Simpson index was used to measure the diversity of the prey taxa, as well as their evenness

which collectively measures the quantitative assortment of the prey taxa used in the test D =

∑ n (n−1)/ N ( N −1 ) (Hill, 1973; Dendi et al., 2019).

 Chapter 4
Results
The dietary preference test results of the gallam lipped frog showed that a total of 145

individuals of both invertebrates and vertebrates were presented to the frogs for food, but only

29.27% (n=60) of the introduced prey was actually foraged upon by the frogs (Figure 4.1). With

a feeding frequency of 85%, the insects were the most consumed prey taxa of the anuran

(Figure 4.2). Generally, The Hymenopterans which were mostly dominated by the ants, were

the most commonly introduced (n = 38) and foraged prey (n = 23). With an introduction rate of

21, the prey preference was followed by the Isopterans which had a 21.67% consumption rate

by the predatory frogs. With the exception of Diptera, Hemidactylus sp, Chilopoda,

Afrotyphlops sp. and Arhynchobdellida, the least foraged prey clade were Coleoptera and

Opisthopora with the two having a cumulative consumption frequency of 6.66% (Table 4.1). In

terms of consumption rate, only 38.24% of the introduced Isopterans were consumed by the

frogs. More so, 11.76% of the Coleopterans were foraged by the frogs. Similarly, 38.89% of the

spiders in the order Araneae were consumed, while 37.50% of Lepidopterans were foraged by

the anurans. Also, 18.75% of the presented Orthopterans were actually consumed by the

predators, whereas, only 28.57% of the Opisthoporans were actually consumed by the

predatory frogs, more so, 19.05% of the introduced cockroaches in the order Blatella were

actually consumed by the frogs. Three individuals of the vertebrates Hemidactylus sp and

Afrotyphlops sp with a cumulative frequency of 2.07% were presented to the frogs but were

totally ignored. While in captivity, we observed that the adult frogs were very selective and

reluctant to feed and were only able to feed on 18.33% of all consumed prey, whereas, the

juveniles and sub adults easily adapted and foraged on 81.67% of their preferred prey.
In their natural habitats, we observed that they often commence their foraging activities from

6-30 to 7pm of each night, and are often spotted near the entrance of ant holes where they

prey on emerging ants, but would stop if distracted or disturbed by noise and bright light which

triggers them to leap away. In terms of their retrieve site, we observed that they often

disappear into burrows, under logs and the darkest part of their habitats before sunrise.
Table 4.1. Dietary Preference and percentage frequency of the Gallam lipped Frog Amnirana
galamensis in Rivers State, Nigeria
No. of introduced % No. of %
S/N Prey taxa prey Frequency consumed prey Frequency
1 Isoptera 21 14.48 13 21.67
2 Coleoptera 15 10.34 2 3.33
3 Diptera 5 3.45 0 0.00
4 Araneae 11 7.59 7 11.67
5 Lepidoptera 10 6.90 6 10.00
6 Orthoptera 13 8.97 3 5.00
7 Opisthopora 5 3.45 2 3.33
8 Hymenoptera 38 26.21 23 38.33
9 Blattodea 17 11.72 4 6.67
10 Hemidactylus sp 2 1.38 0 0.00
11 Chilopoda 4 2.76 0 0.00
12 Afrotyphlops 1 0.69 0 0.00
13 Arhynchobdellida 3 2.07 0 0.00
Total 145 100.00% 60 100.00%
 145, 70.73%
160
140
120
100 60, 29.27%
80
60
40
20
0
Total no. of introduced prey Total no. of consumed prey

Figure 4.1. Relative abundance of the total number of introduced prey and the quantitative
abundance of consumed prey.
 Aran Ophisthopora; 2; 3%
eae;
7;
12%

Insects; 51; 85%

Figure 4.2. Quantitative and percentage abundance of the preferred prey clades of the galam
lipped frog in Rivers State.
 Chapter 5

Discussion

Compared to the gut content analysis method, which entails the killing of anurans (Yap and

Jaafar, 2011; Jamdar and Shinde, 2013), traditional stomach flushing method and the analysis of

faecal contents (Hirschfeld and Rodel, 2011), we believe that the dietary preference test

method is far better than the aforementioned techniques. The dietary preference test method

is better than others because (1) it does not lead to the death of the amphibians; (2) the stress

of going through the faecal contents of the frogs which often leads to the omission of the

heavily digested prey is removed; (3) it gives the researcher the opportunity to record the

preferred prey of a predator in a short period of time. Although, there are disadvantages to this

method, for instance, when predatory ants are kept in the same containers with other animals,

the ants often kill the other prey animals before they are ready to be introduced to the frogs.

The use of just 10 individuals of the galam lipped frog in the conduction of the dietary

preference test experiment shows that only small percentage of the test frogs were required in

getting a desired result with zero anuran mortality. Although a larger proportion of the frogs

could have been used with a high chance of success, thus, the result shows how advantageous

the method could be.

More so, the fact that 85% of their preys were insects showed that they are largely

insectivorous, and it correlates with the insectivorous dietary habits of most amphibians

(Hirschfeld and Rodel, 2011; Chanda, 1993). For instance, a large Percentage of their preferred

prey were mostly invertebrates, whereas the vertebrates, though nocturnal could have been

out of their reach due to the arboreal nature of the geckos or even the fossorial nature of the
blind snakes. We observed that they had high foraging preference for nocturnal species, this

behaviour could be linked to their nocturnal habits.

The result showed the zero consumption of the introduced vertebrates by the frogs, Although, the

adults did not feed on any of the introduced vertebrates, but there is the possibility of large sized

individuals feeding on small frogs and snakes in the wild. For instance, Hirschfeld and Rodel,

(2011) reported the consumption of fishes and amphibians by the African tiger frog

(Hoplobatrachus occipitalis) in northern Benin. Just like the African tiger frog, the gallam lipped

frog preys on invertebrates such as moths, ant’s spiders, earthworms with no evidence for

immediate vertebrate consumption (Hirschfeld and Rodel, 2011). The prey selectiveness

observed in the adult galam lipped frogs suggests that as they age, they only require succulent

and fairly large sized prey such as cockroaches, earthworms and possibly small frogs and snakes

that could satisfy their dietary requirements.

Recommendations

Studies on the dietary behaviour of the Amnirana tadpoles should as well be carried out in order

to ascertain their specific feeding behaviour.

There is need to study the life cycle of the galam lipped frogs in order to determine their life

span.

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