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 2

LITERATURE REVIEW

2.1 Feeding Behaviour of Selected Frog Species.

Frog refers to any of the various tailless amphibians belonging to the order Anura. Used strictly,

the term “may be” limited to any member of the family Ranidae (true frogs), but more broadly the

name frog is often used to distinguish the smooth-skinned, leaping anurans from squat, warty,

hopping ones, which are called toads (Add source).

In general, frogs have protruding eyes, no tail, and strong webbed hind feet that are adapted for

leaping and swimming. They also possess smooth, moist skins. Many are predominantly aquatic,

but some live on land, in burrows, or in trees.

The snout-vent length of frogs ranges from 9.8 mm (0.4 inch) in the Brazilian Psyllophryne

didactyla to 30 cm (12 inches) in the West African Goliath frog (Conraua goliath). The male

anuran is generally smaller than the female (Add source).

During feeding, frogs extend their legs, lunge toward the prey and rapidly open their mouths.

Rapid depression of the lower jaw (9–24 m s –1) propels the tongue out of the mouth and onto the

prey, after which the tongue is retracted with the prey into the mouth. Movement amplitudes

including lunge, gape and tongue protrusion distance were not especially sensitive to temperature.

However, the durations and speed of those movements were, for example, the mean velocity of

mouth closing at 10°C was ∼0.1 m s–1 and at 25°C it was ∼0.3 m s–1. (Add source). But not all

durations and speeds were equally sensitive to temperature. Velocities and accelerations associated

with mouth opening had Q10 values less than 1.25, indicating a relatively low sensitivity to

temperature, while those associated with tongue projection were much higher (Add source). The

higher Q10 values indicate the relative importance of muscle contraction in driving the movement

because we know muscle contractile performance depends heavily on temperature. Thus, while
mouth opening appears to be driven largely by elastic energy release (i.e. is not especially sensitive

to temperature); tongue projection presumably relies more heavily on muscle contraction for its

power. This is quite distinct from what has been found for tongue protrusion in toads, lungless

salamanders and chameleons, where elastic energy is the main driver (Add source).

2.2 Feeding Behaviour of the African Tiger frog (Hoplobatrachus occipitalis)

The African Tiger Frog (Hoplobatrachus occipitalis) is an amphibian species in the

family Dicroglossidae. They are associated with freshwater habitats. The frog is a large and flat

anuran containing protruding eyes, numerous dorsal warts, and a minute inner metatarsal tubercle

(Add source). It has complete webbing between the toes and fingers. A large dorsoventrally

flattened frog with a very warty skin whose eyes and nostrils are dorsally positioned (when viewed

from above the eyes are contained within the outline of the head). Due to numerous glands, the

skin is very slippery (Add source).

The African Tiger frog (H. occipitalis) is a common and widespread West African frog occurring

in both savannah and disturbed forest habitats. It also inhabits diverse water habitats including

small, large, fast flowing, stagnant, temporary and or permanent water bodies (Rödel, 2000). It has

also been found in forested regions (Hughes, 2008; Gruschwitz et al., 2001). It is native to Nigeria

according to IUCN (2014) and in view of the wide distribution; it is tolerant to a wide variety of

habitats. The presumed large population of H. occipitalis led to its categorization as “Least

Concern” by the International Union for Conservation of Nature (IUCN) probably because it is

unlikely to be declining fast enough to qualify for listing in a more threatened category. Its

population however, is on the decline (IUCN, 2014).

Detailed studies concerning the amount and composition of the diet of H. occipitalis in the tropics

have until now been limited to countries like Senegal (Lescure, 2017), Ivory Coast (Tohé, 2009;

Tohé et al., 2013), Kenya (Bwong and Measey, 2010) Congo (Mady-Goma et al., 2012). Though
Hirschfeld and Rödel (2011) carried out their research mainly in the Republic of Benin; they

sampled specimens from Burkina Faso, Niger Republic and Nigeria. The study of the food

preference and feeding habits of frogs is also important considering the concerns regarding global

declines of amphibian populations (Pechmann and Wilbur, 2004).

Hoplobatrachus is quite an evasive species; this made it difficult to make meaningful observations

of their feeding behaviour in their natural environment especially during daytime. H. occipitalis

were usually observed along the edges of the drainage and remained properly camouflaged with

most of their body hidden in the vegetation waiting for prey to move. Deductions could be made

from this that they are“sit-and-wait” predators (Add source).

The mouth of the frog opens directly into the oesophagus which is a continuous tube with the

stomach. The frog's stomach is usually found on the right side of the body cavity i.e. the frog's left

side and is continuous with the small intestine. They have relatively short intestines in relation to

their body size; this indicates that they are carnivores.

The presence of a wide variety of prey items suggests an opportunistic feeding behavior. This frog

captures all the moving preys which have a suitable size for consumption (Balint et al., 2010). H.

occipitalis is a generalist feeder based on the wide food diversity in its diet. The presence of about

fifteen prey categories and plant parts indicates an intense feeding based on the availability of a

rich trophic base.

2. 3 Feeding Behaviour of Rana galamensis

Rana galamensis, commonly called galam white-lipped frog belongs to the family Ranidae, which

has widest distribution of any frog family, and the class amphibian. It is a strongly aquatic species,

where it lives in and around permanent lakes, rivers, ponds and swamps. Rana galamensis is

abundant throughout most of the continents except Antarctica. In Africa, they are found in

savannah region of West Africa, South Africa and East Africa (Rodel et al., 2004). In Nigeria, they
are found in many states such as Lagos, Ogun, Oyo, Kwara, Osun, Ondo, Ekiti, Kaduna and Benin

City (Walker, 2007). The fresh Rana galamensis could be salted and sundried to make them into

stocks for transportation into various parts of the country. Consumption of Rana galamensis in

various parts of Nigeria and other West Africa countries had been reported (Mohneke et al., 2010),

however, information on its nutritive value is still very scanty in literature.

Table 1: Proximate analysis of Rana galamensis in g /100g

Parameters Rana galamensis

Ash 6.10 ± 1.08
Lipid content 9.52 ± 0.31
Fibre 1.60 ± 0.30
Protein 53.74 ± 0.89
Carbohydrate (by differences) 29.04 ± 0.01

Each value is a mean of three determinations ± SEM

The proximate compositions of R. galamensis are shown in Table 1. R. galamensis contains a high

protein and understandably, low fibre content. The protein content of R. galamensis (53.73g/100g)

is higher than those reported in moon fish (39.3g/100g) and cat fish (43.70g/100g) (Abdullahi,

2000 Abdullahi et al., 2001), Haddock (17g/100g), Sardine (20g/100g), Mackerel (12g/100g) and

Oyster (11g/100g) (Brain and Allan, 2007b; Pearson, 2001), and beef (18g/100g), lamb (16g/100g)

and pork (10g/100g) (Bhulyan et al., 2003). Therefore R. galamensis can serve as a better source

of protein especially, with its high levels of indispensable (essential) amino acids.
Table 2: Amino acid profile of Rana galamensis (g / 100g protein), each value in column 2 is a
mean of three determinations ± SEM

Amino acids Rana galamensis FAO/WHO (2000)

Lysine 6.93± 0.02 5.80
Histidine 3.13± 0.01 2.5
Arginine 6.55± 0.02 5.2
Asparatic acid 9.66±0.21 7.7
Threonine 4.33±0.05 3.40
Serine 5.15±0.03 7.7
Glutamic acid 13.24 ±0.25 14.7
Proline 5.10 ±0.03 10.7
Glycine 5.01±0.02 2.2
Alanine 6.10±0.11 6.1
Cystine 1.06±0.01 3.0
Valine 4.82±0.04 5.00
Methionine 3.00±0.34 2.50
Isoleucine 4.00±0.02 2.80
Leucine 7.05±0.02 6.60
Tyrosine 4.02±0.01 1.10
Phenylalanine 5.02±0.02 6.30
(Table 2). The carbohydrate content of the R. galamensis was higher than those reported in moon

and cat fishes (Abdullahi, 2000; Abdullahi et al., 2001). The carbohydrate content, if digestible

could serve as a source of energy. The relatively low lipid content (9.52 ± 0.31 %) of R.

galamensis (Table 1), compared to that of moon fish (27.7 ± 1.06g/100g) and cat fish (32.30

±1.32g/100g) (Abdullahi, 2000; Abdullahi et al., 2001), would probably make it a useful

ingredient in poultry feeds than the fish meals. This is because very high fat content in feed

ingredients would cause difficulty in mixing the feed and could even predispose the feed to

oxidative rancidity (Ewing, 2001; Atteh, 2002). The roughage (fibre) quantity of R. galamensis

may due to its tough skin. This plays an important role in preventing colon cancer and constipation

(Ahmed, 2005; Bingham et al., 2003; Park et al., 2005). The amino acid profiles of R. galamensis

are shown in Table 2. Seventeen amino acids were observed; out of which nine were indispensable

with leucine having the highest concentration. The seventeen amino acids present in R. galamensis

were also reportedly present in moon and cat fishes (Abdullahi, 2000; Abdullahi et al., 2001). The

indispensable amino acids (lysine, histidine, arginine, threonine, valine, methionine, isoleucine,

leucine, and phenylalanine) are within FAO/ WHO 1990 reference values for daily intake. These

amino acids serve as raw materials for the synthesis of many other cellular products, including

hormones, enzymes and pigments. In addition, several of these amino acids are key intermediates

in cellular metabolism (Murray et al., 2000).

2.2 Amphibian Diversity of Rivers State

Amphibians are consumed in many parts of the world either as whole organisms or as frog-legs

(Akani and Luiselli, 2002; Anosike and Keke, 2002; Onadeko, 2009; Onadeko et al., 2011;

Imkongwapang et al., 2012). Akani and Luiselli (2002) reported that frog legs of big amphibian

species, such as Hoplobatrachus occipitalis and all members of the family Pipidae, when treated

with condiments cost several dollars in groceries in Europe where they were considered as a
delicacy. Anosike and Keke (2002) noted that amphibians were considered as therapy for

hypertension, tuberculosis, conjunctivitis, abortion and Kwashiorkor, and Onadeko (2009) reported

the commercial sale of sun-dried frogs in Lolo market in Northern Nigeria and at Badagry and

Oyingbo markets in south-western Nigeria. Onadeko et al. (2011) reported that Ptychadena

pumilio, Hoplobatrachus occipitalis and Xenopus muelleri were commonly consumed by the local

inhabitants of south-western Nigeria, noting that frog meat served as a good source of protein for

them, with P. pumilio having a protein content of 19.79% followed by X. muelleri (19.53%) and H.

occipitalis (19.46%).

Some researchers have reported on the amphibian species endemic in parts of Rivers State,

Nigeria. For instance, Akani et al. (2011) reported the presence of Amietophrynus maculatus,

Ptychadena aequiplicata and Hoplobatrachus occipitalis from Azuabie, Rumuokwurusi, Woji

Housing Estate, and Rumunduru. Reuben (2014) and Ukawu (2014) reported on the diversity of

amphibians in Rumuomoi and Ahoada, repectively, and found the following species: Rana sp;

Silurana tropicalis; Hoplobatrachus occipitalis; Ptychadena oxyrhynchus; P. mascareniensis; P.

pumilio; Amietophrynus regularis; A. maculatus; Hymenochirus sp., Afrixalus dorsalis; A.

fulvovittatus; Phrynobatrachus francisci; Hyperolius guttulatus; and H. fusciventris. Omereji

(2014) also found Leptopelis hyloides, Ptychadena bibroni, and Ptychadena schubotzi in parts of

Emohua. The amphibian species encountered in this research have been reported in earlier studies

from neighbouring communities (Akani et al., 2011; Omereji, 2014; Reuben, 2014; Ukawu, 2014).

Ptychadena species and H. occipitalis were more abundant at Agbada while the bufonids were

more abundant at Rumuesara

2.3 Global Amphibian Decline

Declines and losses of amphibian populations are a global problem with complex local causes.

These may include ultraviolet radiation, predation, habitat modification, environmental acidity and
toxicants, diseases, changes in climate or weather patterns, and interactions among these factors.

Understanding the extent of the problem and its nature requires an understanding of how local

factors affect the dynamics of local populations.

Amphibian population declines were first recognized as a global phenomenon in the early 1990s

(Wake, 2001). We now know that current extinction rates for amphibians may be as much as 200

times higher than background (Roelants et al., 2007). With 32% of the world's nearly 6600

amphibian species threatened with extinction, 43% experiencing declines and another 22% with

insufficient data (Stuart et al., 2004), this phenomenon represents the Earth's sixth mass extinction

(Wake and Vredenburg, 2008).

A number of causes have been suggested to explain global amphibian declines. Habitat

modification (loss) has played a significant role in many declines and is likely the single most

important cause of local declines (Adams, 2000; Becker et al., 2007; Davidson et al.,

2002; Davidson et al., 2001; Delis et al., 2006; Eigenbrod et al., 2008; Harper et al., 2008; Smith

et al., 2009). Other factors may explain many local declines including the use of road salt,

catastrophic events, etc. The global phenomenon particularly declines in apparently physically

undisturbed habitats especially so-called ‘enigmatic declines’ which represent 48% of threatened

amphibian species according to Stuart and colleagues (Stuart et al., 2004), begs other explanations,

however. Identifying the most urgent issues and those factors that are truly global is vital to

developing plans to halt declines or to provide whatever remediation is possible.

2.3.1 A Hierarchical Approach to Defining the Causes

Here, we propose that ultimately there is no single cause of global amphibian declines, but rather

interactions between several factors are important. Some local declines may primarily have single

causes, but we propose that even local declines and extinctions are likely (and most often) caused

by interactions between two or more factors. Individual factors that interact likely differ from
species to species and even population to population, within a species. In addition, several factors

may be more detrimental when combined, and result in emergent properties that render individual

factors more severe when combined with other factors.

Robert May recently addressed the ‘likely causes and consequences of the manifest acceleration in

extinction rates over the past few centuries’ (May, 2010), with a focus on current extinction rates

in general. According to May, ‘the ultimate causes are habitat destruction, alien introductions,

overexploitation and climate change’. Young and colleagues reported that declines in amphibian

populations due to overexploitation are concentrated in East and South East Asia (Stuart et al.,

2004); thus, overexploitation is seemingly not a global concern for amphibians. This leaves habitat

destruction, alien introductions and climate change from May's list. Two additional factors are

important in global amphibian declines, however: environmental pollutants and pathogens. We

propose that none of these factors act alone, but that interactions between five ultimate factors

(atmospheric change, environmental pollutants, habitat modification, invasive species and

pathogens) are the cause of amphibian declines. This review is not the first to propose that there is

more than one cause of amphibian declines (Blaustein and Kiesecker, 2002; Blaustein and Wake,

1990; Kiesecker et al., 2001; Wake and Vredenburg, 2008), nor is it the first to suggest that

causative factors interact (Boone and James, 2003; Boone and Semlitsch, 2002; Boone et al.,

2007; Davidson et al., 2007; Davidson et al., 2002; Davidson et al., 2001; Garcia et al.,

2003; Jancovich et al., 2005; Kiesecker and Blaustein, 1995b; Kiesecker and Blaustein,

1999; Marcogliese et al., 2009; Pounds et al., 2006; Pounds and Crump, 1994; Relyea,

2003; Relyea, 2004b; Relyea and Hoverman, 2008; Sodhi et al., 2008; Wake and Vredenburg,

2008). As such, the goal here is not to present an exhaustive review of all of these factors and

interactions.

Level 1: Proximate (Direct) Causes of Amphibian Declines

There are only two proximate (direct) causes of amphibian population declines: death (or removal)

of individuals from a population and reduced recruitment within a population. This observation

may seem to state the obvious, but it is important to point it out. Many population declines have

received attention because adults simply disappeared rapidly and/or massive die-offs were

documented (Crump et al., 1992; Pounds and Crump, 1994). Just because adults are present at a

locality and massive deaths are not observed in a population does not mean that a given population

is not in decline. Populations can decline slowly if otherwise healthy adults are not breeding or if

progeny do not develop properly.

These two causes of declines (death and recruitment failure), of course, interact in that the death of

individuals will ultimately result in decreased recruitment. Although many studies have examined

declines in amphibian populations due to loss of individuals (death), we are not aware of any

studies in amphibians that have examined population declines resulting from failed recruitment in

populations with otherwise physiologically healthy individuals.

Level 2: Factors Contributing to Death and Failed Recruitment

There are multiple factors that contribute to the death of local populations. We include five major

factors that contribute to local declines as a result of death: (1) increased disease rates can lead to

individual deaths and population declines; (2) decreased nutrition (loss of food base) can result in

declining numbers within a population; (3) predation can remove individuals from populations;

and (4) we consider human take (exploitation) separate from predation by other animals. Finally,

we have included a ‘catch all’ category, (5) ‘other mortality’, which represents everything from

death due to old age, to incidental deaths not due to the aforementioned factors, to catastrophic

events that may reduce or even eliminate entire populations. Failed recruitment, on the other hand,

can result directly from reproductive failure or as a result of some developmental failure not

directly related to reproductive function. Although important in local declines, the factors
presented at Level 2 are not likely to be universal factors that contribute to global amphibian

declines. In other words, it is not likely that global amphibian declines are universally caused by

human take, predation, incidental or catastrophic events leading to mortality, or decreased

nutrition.

What is mainly important with regards to recruitment is that a number of factors (causes) can

reduce recruitment in a population without otherwise causing harm (developmental or

physiological) to exposed individuals. Decreased recruitment can result directly from reproductive

failure or from developmental failures that lead to reproductive failure, i.e. failure of reproductive

organs or structures to develop properly. Developmental failures that do not lead to death can also

directly decrease recruitment (cause reproductive failure). For example, limb deformities or

decreased growth are developmental effects that would limit reproductive potential without killing

affected individuals. This scenario is especially true for short-lived species where failed

recruitment for one or two breeding seasons may be disastrous for populations (Harper et al.,

2008). Complete inhibition or delays in metamorphosis that prevent reproduction should also be

considered here: although some salamander species can reproduce in the larval form, anurans

cannot (Hayes, 2007).

Level 3: Ultimate Causes of Amphibian Declines

There are at least five ultimate factors that influence all seven factors at Level 2. These five (Level

3) represent the most likely factors to have truly global effects on amphibian population declines,

and include atmospheric change (temperature, rainfall, UV levels, etc.), environmental pollutants,

habitat loss, invasive species and pathogens (Davidson et al., 2002; LeNoir et al., 2001).

Atmospheric change is clearly a phenomenon that amphibian populations will experience globally.

Also, environmental pollutants are widespread enough to have global effects. For example, aerial

transport of pesticides into California's Sierra Nevada likely contributed to amphibian declines
there (Davidson et al., 2002; LeNoir et al., 2001), and pesticides, like the popular herbicide

atrazine, can travel over 1000 km in precipitation and contaminate otherwise pristine habitats

where they are not used (Fenelon and Moore, 2008; Lode et al., 2005; Mast et al., 2007; Müller et

al., 2007; Thurman and Cromwell, 2000; Vogel et al., 2008). Although direct habitat modification

(loss) may not be ubiquitous, it is indeed quite widespread, but furthermore other global impacts

(atmospheric changes, environmental pollutants and invasive species) all generate habitat

modifications.

Invasive species may be widespread, but are probably not as independently important as other

factors on this level. Finally, at least some pathogens may be widespread enough to have a global

impact, but in most cases this factor is likely dependent on many other factors. All five ultimate

factors at Level 3 affect all factors at Level 2 and, thereby, have the potential to affect both death

and recruitment.

2.4 Conservation Status of Amphibians

Amphibians play important roles in ecosystem services, medical research and as bio-indicators.

Yet, nearly 41% of all existing amphibians are threatened with extinction. Why this is so and how

local interventions can influence global conservation outcomes is illustrated in this review.

Nigeria boasts of a wide variety of ecosystems ranging from mangroves and rainforests in the

south, to different savannahs up north, to the Jos plateau in Central Nigeria and mountainous

habitats in Eastern Nigeria towards the Cameroon border (Onadeko and Rodel, 2009). About 115

species of amphibians have been recorded within the country, two of which are caecilians and one

endemic species (Amietophrynus perreti) has been described in the country. A high biodiversity

loss including steep decline in amphibian populations has necessitated assessment of the

conservation status of amphibians in Nigeria. The causes of amphibian population declines are
diverse and there appears to be no single cause for their decline (Blaustein et al., 2010), but the

following factors are incriminated:

Habitat Loss

Habitat is essential to the survival of species. According to Baillie et al. (2004), habitat loss is a

key threat to about 70% of amphibian species. Most habitat loss is often due to agricultural

expansion, logging, mining activities and land for human settlement. Nigeria has lost a vast

majority of its rainforests and most of its savannah as well (Onadeko and Rodel, 2009).

Environmental Pollution

In Nigeria, greater amphibian biodiversity is observed in areas around the coast. The Niger Delta is

a region notoriously polluted because of the booming oil industry. Amphibians can be expected to

be seriously affected by this type of pollution as their larvae feed off algae and continuously

process water for respiration. Runoffs from agricultural farmlands into water bodies and nearby

terrestrial habitats of amphibians also constitute a threat (Onadeko and Rodel, 2009)..

Global Climate Change

Climate change which is largely due to the emission of greenhouse gases is global and have many

implications for the survival of amphibians. These effects may be direct or indirect. Temperature

plays an important role in the life cycle of amphibians (Blaustein et al., 2010). In amphibians,

temperature acts as a controlling factor for many physiological processes, including rates of

oxygen uptake, heart rate, locomotion, water balance, digestion, developmental rate, sex

determination, and immune function.

Diseases and Pathogens

Currently, a major disease that has contributed to the decline in amphibian populations is the

chytrid fungus (Batrachochytrium dendrobatidis). Its spread is often enhanced by other factors

such as climate change. It works by infecting the skin of amphibians, a vital organ necessary for
respiration and exchange of materials (Blaustein et al., 2010). The disease is believed to have

originated in Africa. The international trade of the African clawed frog, an asymptomatic carrier of

the fungus, is believed to have resulted in its spread (Blaustein et al., 2010).

Overharvesting

Anurans have been exploited for both consumptive and non-consumptive uses. Some frog species

are edible and a delicacy to some ethnic communities in Nigeria. In Nigeria, dried, skinned meat of

toads and fresh frogs has been found useful in the feeding of catfish and poultry, either as raw

materials for processed feed or whole as food for catfish (Tran, 2015). International trade in frog

legs in 1998 was valued at approximately USD 48.7million (Teixeira et al., 2001). There has been

an increasing trend in the export of frogs, and edible anuran species are being removed from the

wild at a rate higher than replenished (Onadeko et al., 2012). In a study carried out by Mohneke et

al. (2010), it was inferred that 2,738,610 frogs were harvested from the wild annually. Most frogs

traded were reported to originate from the northern Nigeria and neighbouring countries such as

Benin, Niger and Chad.

2.5 Adaptive Behaviour of Nocturnal Amphibians

Amphibians left water, and emerged on the land for the first time somewhere around 360 million

years ago, and the fact that they continue to exist even today, while several other species with

which they shared habitats have gone extinct, indicates their high adaptability (Add source).

Life on land was pretty different from life in water for this species, as a result of which early

amphibians had to undergo several changes and adapt to the new environment to ensure survival

on the planet. In order to get a better understanding of adaptations that these amphibians resorted

to, we need to go back to square one i.e. their evolution, which can be traced back to the Devonian

period.

2.5.1 Evolution of Amphibians

Amphibians were the first vertebrates to adapt to the terrestrial life; something which occurred

somewhere around 400-360 million years ago. The first major group of amphibians to evolve on

the planet was armed with fins – which were quite similar to legs that helped them crawl out of the

water (Add source).

They had characteristic traits which were quite similar to that of the extant species like coelacanth

(Latimeria chalumnae) and lungfish. Even though they succeeded in crawling out of the water,

onto the land, they were not fully equipped to lead a terrestrial life, and hence had to return to the

water for every other life process including breeding (Add source).

These amphibians – which evolved during the Devonian Period, dominated the food chain during

the Carboniferous Period, before they were out-classed by other species which evolved during the

Triassic Period. While several amphibian species became extinct with time, the three major groups

of amphibians which are found on the planet today include the frogs, caecilians, and salamanders

(Add source).

2.5.2 Amphibian Adaptations

For amphibians, limbs and lungs were two of the most important adaptations as the former helped

them move around without having to depend on the buoyancy of water, and latter replaced the gills

to facilitate respiration. As time elapsed, amphibian species on the planet also evolved and came

up with several new adaptations which help them survive on a planet which has changed a lot over

the last 360 million years (Add source). Even though they still resort to water bodies for life

processes such as breeding, they have adapted to other conditions – and are even found in deserts

and other arid regions now.

While the bodies of various amphibian species are specially designed to help them swim, their

powerful hind legs help them take large leaps and change direction with utmost ease when it

comes to movement on the land. On the other hand, their typical colour pattern – with dark body
and light underside, plays a crucial role in camouflage in water as well as on the land ( Add

source).

The thin and moist skin that they sport is also an adaptation which facilitates processes like

cutaneous respiration and osmosis in these species. Even though amphibians usually inhabit moist

areas, the chances of water loss due to evaporation cannot be ruled out, and that’s where their skin

– which is specifically designed to minimize the amount of water loss, comes into play. Other than

inhabiting moist environment, some species also resort to nocturnal lifestyle to minimize water

loss.

As I mentioned earlier, amphibians are found in deserts as well; though the number of species is

very less. Species like the Sonoran Desert toad and desert spade foot excavate burrows and spend

entire day in them, only to step out in search of food at night. Inside these burrows they secrete

semi permeable membrane and cover their body to minimize the water loss (Add source).

As amphibians are cold blooded animals, they cannot withstand cold environment and hence go

into hibernation during the winter season (Add source).

Eyes positioned on the top of the head don’t just facilitate a wide angle view for these species, but

also help them stay in the water with their eyes just above the water surface.

Some amphibians release toxins from their skin which is yet another defence mechanism in some

species of amphibians meant to keep predators at bay (Add source). At the same time, they sport a

transparent eyelid which helps them see underwater with relative ease.

While their eyes are considerably large, their mouths are even larger in proportion to their size an

adaptation which helps them catch and eat large prey with relative ease (Add source). Even the

tongue of some amphibians is designed in such a manner that they can flick it out, grab the prey

and pull it back to their mouth when hunting.

Amphibians lay eggs in masses, as a result of which only the outer eggs are affected – while the

inner eggs are safe, in case of harm. Similarly, the gelatinous cover on these eggs makes sure that

they are safe from infection – chances of which exist in plenty in moist environment which these

creatures usually inhabit during the breeding season.

The probability factor also comes into play for this species, as laying large number of eggs ensures

that at least some will survive predation and facilitate the survival of species. In deserts,

amphibians lay eggs in temporary pools of water which can dry up at any moment.

Those amphibians which inhabit the deserts sport an amazing adaptation in form of accelerated

development rate wherein species – like the desert spadefoot, grow from an egg to toad let with a

period of two weeks (Add source).

Right from their body shape to eye position and from camouflage to release of toxins from skin,

every single characteristic trait of amphibians mentioned above plays a crucial role in their

survival.

Other than these general adaptations, every single species on the planet has some or other

adaptations of its own. If it were not for these adaptations, regions with harsh conditions, such as

the Arctic tundra or Sahara, would have been devoid of species (Add source).

2.5.3 Frog Adaptation

Some frogs have behaviour adaptations to conserve water, including becoming nocturnal and

resting in a water-conserving position. Some frogs may also rest in large groups with each frog

pressed against its neighbours. This reduces the amount of skin exposed to the air or a dry surface,

and thus reduces water loss (Add source).

Weather Adaptations

When frogs are cold, they can move into the sun or onto a warm surface; if they overheat they can

move into the shade or adopt a stance that exposes the minimum area of skin to the air. This
posture is also used to prevent water loss and involves the frog squatting close to the substrate with

its hands and feet tucked under its chin and body. The colour of a frog's skin is used for thermo-

regulation. In cool damp conditions the colour will be darker than on a hot dry day.

Camouflage Behaviour Adaptations

Camouflage is a common defensive mechanism in frogs. Most camouflaged frogs are nocturnal;

during the day, they seek out a position where they can blend into the background and remain

undetected. Some frogs have the ability to change colour, but this is usually restricted to a small

range of colours. For example, the White's tree frog varies between pale green and dull brown

according to the temperature, and the Pacific tree frog has green and brown morphs, plain or

spotted, and changes colour depending on the time of year and general background colour (Add

source).

Structural Adaptations

Frogs are carnivores which eat insects, spiders, worms, snails, fish, and even small rodents like

mice. When a frog spots a tasty meal, it flicks out its long, sticky tongue. The tongue lassos the

meal/creature and pulls it back into the frog's mouth. Frogs have teeth which are small and so not

really good for chewing. Frogs mostly use their teeth just to hold their prey in their mouths, and

then they swallow their prey whole (Add source).

Head

A frog's head has adaptations that help it to swallow prey. Unlike a human's eyes which are fixed

in our heads in bony circles called orbits, the orbits surrounding a frog's eyes do not have bottoms.

This means that when a frog swallows large prey, it can close its eyelids and drop its eyeballs

down into its mouth. Then the eyeballs help push the prey down the throat of the frog (Add

source).

Feet and Legs

Most frogs are proficient at jumping. Many frogs have webbed feet and the degree of webbing is

directly proportional to the amount of time the species spends in the water. Sometimes during the

tadpole stage, one of the developing rear legs is eaten by a predator such as a dragonfly nymph. In

some cases, the full leg still grows but in others it does not, although the frog may still live out its

normal lifespan with only three limbs (Add source).

Webbed Hind Foot of Common Frog

A frog's skin is protective with a respiratory function, can absorb water and helps in the regulation

of body temperature. It has many glands, particularly on the head and back, which often exude

distasteful and toxic substances. The secretion is often sticky and helps keep the skin moist,

protects against the entry of moulds and bacteria and make the animal slippery and more able to

escape from predators (Add source).

 Chapter 3
Materials and Methods
The study on the dietary habits 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).

 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%
Figure 4.1. Relative abundance of the total number of introduced prey and the quantitative
abundance of consumed prey.
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.

 Chapter 6

Conclusions and Recommendations

6.1 Conclusions

In conclusion, 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. 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.

Knowledge regarding amphibian trophic ecology in Nigeria must be expanded. This will require

partnerships between protected areas, local communities, and educational institutions. Monitoring

programmes should be put in place to ensure the survival of frogs around our homes. Frog

farming should be encouraged with sustainable use in mind, because these animals play very

important ecosystems services that are essential to the success of our immediate ecosystems.

6.2 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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