CHAPTER ONE

1.0 INTRODUCTION

Ecological surveys identify the habitats and/or species that exist within an area at the time of

the survey. Ecological surveys are carried out with the intention of identifying particular

species, the interactions which exists between organisms in the habitat, the abundance and

distribution of these species as well as to help identify the factors that may affect the survival

of these species. Ecological surveys also help keep track of changes and patterns in the

habitat/environment under study.

Ecology is the scientific study of how organisms interact with each other and with their

environment. This includes relationships between individuals of the same species, between

different species, and between organisms and their physical and chemical environments.

Aquatic ecology is the study of these relationships in all aquatic environments, including

oceans, estuaries, lakes, ponds, wetlands, rivers, and streams.

Freshwater snails are gastropod mollusks which live in freshwater. There are many different

families. They are found throughout the world in various habitats, ranging from ephemeral

pools to the largest lakes, and from small seeps and springs to major rivers. The great

majority of freshwater gastropods have a shell, with very few exceptions.

Some groups of snails that live in freshwater respire using gills, whereas other groups need to

reach the surface to breathe air. In addition, some are amphibious and have both gills and a

lung (e.g. Ampullariidae). Most feed on algae, but many are detritivores and some are filter

feeders. According to a 2008 review of the taxonomy, there are about 4,000 species of

freshwater gastropods (Strong et al., 2008). Snail habitats include almost all types of

freshwater bodies ranging from small temporary ponds and streams to large lakes and rivers.

Within each habitat, snail distribution may be patchy and detection requires examination of

1
different sites. Moreover, snail densities vary significantly with the season. They live on

water plants and mud that is rich in decaying organic matter. They can also be found on

rocks, stones or concrete covered with algae or on various types of debris. They are most

common in waters where water plants are abundant and in water moderately polluted with

organic matter, such as faeces and urine, as is often the case near human habitations. Plants

serve as substrates for feeding and oviposition (laying of eggs) as well as providing

protection from high water velocities and predators such as fish and birds. Most aquatic snail

species die when stranded on dry land in the dry season. However, a proportion of some snail

species are able to withstand desiccation for months while buried in the mud bottom by

sealing their shell opening with a layer of mucus. Most species can survive outside water for

short periods (WHO, 1997).

1.1 JUSTIFICATION

Freshwater molluscs help in the assessment of the ecological status of water bodies and also

play significant roles in public and veterinary health. Hence, scientific studies pertaining to

their diversity, distribution and abundance becomes essential (Oyedibu et al., 2017).

Investigations of freshwater snails have been carried out by several authors, among which

are: Hira (1966) and Asumu (1975) for Ibadan City and its environs, Okwuosa (1979) for

Lagos, Ogun and Oyo States, Ukoli and Asumu (1979) for the New Federal Capital Territory,

Ebele (1981) for Zaria City, Dupont and Leveque (1968), Leveque (1967 and 1972) and

Betterton et al., (1983) for the Lake Chad area and Thomas and Tait (1984) for the Ibadan

area.

There is need for precise species identification to help aquatic snail vector control,

conservation and management initiatives. Snail shell morphology has previously proved

useful in taxonomic studies. Very little is known about freshwater snails in general (apart

2
from what is known about them as human disease vectors). The findings of this research

work should prove to be a useful source of primary data for improved freshwater snail

identification, sensitization of the general public and future researchers on its importance,

diversity, morphometry / morphology and distribution of freshwater snail species at Owan.

1.2 Aim and Objectives

The major aim of this study is to ascertain the current status of the aquatic snail species in

Ovia River at Owan. The specific objectives are to determine:

1) the species composition of freshwater snails in the water body in order to ascertain

their relative abundance.

2) the morphological features and variations of each available species and between the

sexes where possible

3) the distribution of freshwater snails in the water body

3
 CHAPTER TWO

2.0 LITERATURE REVIEW

Fresh water, which has less than 0.5 g per liter of dissolved salts, exists in many ecosystems

both above and below ground. These freshwater ecosystems are highly diverse: temporary or

permanent, large or small, stationary or flowing, intermittent or continuous, hot or cold,

surface or subterranean. As noted above, there are also transitional systems that link fresh

water with terrestrial and marine environments. In the latter case, where fresh water mixes

with seawater, for example in estuaries and coastal marshes, the water will be higher in salts

and hence “brackish” (Balian et al., 2010). Freshwater snails belong to a larger group of

shelled animals called molluscs. Freshwater, marine and terrestrial snails number is well over

50,000 individual species. Most species of freshwater snails originally evolved from saltwater

habitats, although, several species evolved from terrestrial habitats to freshwater. Snails

usually play a dominant role in the ecology of freshwaters by providing food for many other

animals and by grazing on vast amounts of algae and detritus (Hayes, 2009). Snail

populations are known to vary in seasons and are under the control of climate. This macro

distribution of gastropods in freshwater systems has been the subject of many surveys. In

most habitats, distribution has been found to be limited by climatic factors such as rain and

drought. For this reason, species will probably fail to colonize some habitats while in some

other similar habitats, they may be abundant. It is therefore very important that factors of the

environment that affects macro distribution of gastropods in any environment are investigated

and used to understand the biological interaction between the habitats (Igbinosa et al., 2015).

Freshwater gastropods (snails) are an important and diverse component of aquatic ecosystems

worldwide (Johnson et al., 2013). Gastropods have diversified into every conceivable natural

aquatic habitat including aquifers, springs, creeks, rivers, lakes and wetlands, and are

routinely found in ephemeral and man-made water bodies. Most graze on periphytic or

4
epiphytic algae and biofilms, while some are suspension or deposit feeders. Gastropods often

dominate stream benthos in both mass and numbers often exceeding 50% of the invertebrate

benthic biomass and are the principal grazers in many aquatic habitats. As such, gastropods

have a profound impact on algal primary productivity playing a pivotal role in nutrient

cycling. Gastropods were important dietary components of several animals, including fishes

(like Redear, Sunfish, River Redhorse) and turtles (like Stinkpot). Freshwater snails are

among the most rapidly declining groups of organisms on Earth (Lydeard et al., 2004;

Johnson et al., 2013).

2.1 ECOLOGY

Snail habitats include almost all types of freshwater bodies ranging from small temporary

ponds and streams to large lakes and rivers. Within each habitat, snail distribution may be

patchy and detection requires examination of different sites. Moreover, snail densities vary

significantly with the season. They are most common in waters where water plants are

abundant and in water moderately polluted with organic matter, such as faeces and urine, as

is often the case near human habitations. Plants serve as substrates for feeding and

oviposition as well as providing protection from high water velocities and predators such as

fish and birds (WHO, 1997).

Aquatic macrophytes have been shown to play vital roles in the distribution of snails in

different parts of Africa (Ofoezie, 1999). The macrophytes are believed to provide both food

and shade and to provide breeding sites for the snails. However, the importance of different

ecological factors vary significantly from one ecological zone to the other and even from one

water body to the other, suggesting local investigations to identify important factors in each

5
zone or water body (Dazo et al., 1966; Klumpp and Chu, 1977; 1980; Imevbore et al., 1988;

Ofoezie, 1999).

The snail species found at Guapimirim displayed a marked habitat preference, especially in

relation to the physical characteristics of each environment. P. marmorata, B. tenagophila,

and M. tuberculatus were associated to lotic environments, with a low percentage of aquatic

vegetation and variable degrees of domestic sewage input. The multiple regression analysis

showed that the high concentration of faecal coliforms and chloride were the most important

factors for explaining the abundance of M. tuberculatus in lotic environments. The increase

of the chloride concentration is expected in environments contaminated by sewage, due to the

inflow of urea-rich effluents (Feema, 1981). In lotic environments, the abundance of P.

marmorata was correlated with the alkalinity and the mean velocity. High alkalinity values

may be related to organic pollution, due to the increase in phosphate and ammonium salts

(Feema, 1981). Ndifon and Ukoli (1989) verified that M. tuberculatus and P. waterloti

(Germain, 1911) were most frequently encountered in water bodies polluted by high amounts

of human and animal waste and domestic sewage. The abundance of organic matter increases

the concentration of detritus and possibly aids in the proliferation of epiphytic algae. The diet

of both planorbid and prosobranch snails includes those items (Madsen, 1992; Lombardo and

Cooke, 2002). Another limiting factor for snail distribution in the studied lotic environments

was the water current velocity and the occurrence of spates. All the three species encountered

at those environments were found mostly on drainage channels, which have slower water

flow than those from streams (Giovanelli et al., 2005).

2.2 DISTRIBUTION AND EFFECT OF SEASONAL VARIATION

In the survey carried out by Igbinosa et al. (2013), at four different water bodies in Ovia

Southwest local government area of Edo state, a total of 579 individual snails were collected.

6
The snail species collected were Lanistes varicus Melanoides tuberculata 82(14.2%),

Gabbiella humerosa, Bulimulus teniussimus, Monacha cantiana Drymaetus multilineatus and

a bivalve species Aspatharia subreniformis. The monthly distribution and abundance of the

snail species indicated that Lanistes varicus (258) increasingly dominated throughout the

study period, while Bulimulus teniussimus had the least number throughout the study. The

survey also showed that there was a sharp decline the abundance of freshwater snails with the

onset of the dry season in September through October and November.

Table 2.1: Showing abundance and distribution of snails at Eleyele Dam

Subclass Family Species Sampling points

Pulmonata Planorbidae B. p A B C D

1426 511

(63.95%) (22.91%)

7
Source: Oyedibu et al., (2016)

8
The spatial distribution of snails shows that sampling point A had the highest percentage of

mollusc abundance of 44.71% and richness with 8 species; sampling point B had 36.23%

with 8 species; sampling point C had 14.30% with 7 species, while sampling point D with

4.76% had the poorest composition with 3 species. (Oyedibu et al., 2016).

In another survey carried out in three parts of Jakara dam, Kano State Nigeria, it was

discovered that all the Bulinus species except Bulinus globosus were found in the Yadakunya

part of the dam. Fako has the highest number of snails (1230) followed by Yadakunya (911)

and Kwata (695) had the least but Yadakunya has all the snail species except Lymnea

natalensis. During the months of November, December and January, the rains have

completely receded and there is more access to the dam for collection of snails and new

species have also emerged. Some of the snails were more during the rainy season while some

were completely absent (Duwa, 2017).

2.3 STRUCTURE, APPEARANCE AND BEHAVIOUR

Freshwater snails generally are dull coloured, especially in comparison to their marine and

terrestrial cousins. Their colour generally varies from yellow-brown to black, but some

species have stripes and others have distinctive markings. Freshwater snails come in a variety

of shapes and sizes. Some snails are almost completely round in shape, but most species are

generally conical. Adults of some species are smaller than the head of a pin and others are

larger than a baseball. The snail’s soft body is divided into three distinct sections. A well-

defined head is connected to a large muscular foot. The foot is the most visible external

feature of the snail’s body and is the animal’s mode of locomotion. A snail propels itself with

this single foot, which is controlled by hydrostatic action inside the snail’s body.

Additionally, the foot surface is covered with tiny projections (cilia) that assist the gliding

motion. Snails feed along the bottom as they move, partially clearing a path for the foot. As

9
the foot passes over the substrate, it leaves a “trail” behind the snail. These trails are

commonly visible in soft sediments or across hard surfaces such as rocks or submerged trees

(Johnson, 2009).

The third distinctive section, the internal organs, is concealed inside the shell. The internal

organs of a snail include a heart for circulating “blood;” a complete digestive system with a

distinct mouth, stomach, and intestine; a reproductive system; and a gill or other respiratory

surface used for oxygen exchange. When a snail is threatened by a predator, the head is

retracted into the shell first, followed by the foot; thus the entire body can retreat inside the

shell. Some snails even have a hard “lid” on their tails called an operculum (oh-pûr´-ky-lum)

that is made of protein. The hard operculum covers most soft tissue of the foot exposed to a

predator when the snail has retracted into its shell (Johnson, 2009).

All freshwater snails have a shell, a hard structure covering most of the body, providing

protection. That shell normally consists of several layers, one to several mineral (calcium

carbonate, usually aragonite or calcite) and one outermost (periostracum), composed mostly

or entirely of organic material and usually pigmented. The shell is basically a tube coiled

variously around a central pillar, termed an axis. Each complete turn is called a whorl:

generally, the first 1 or 2 whorls are formed before the snail egg hatches and are termed

embryonic, or nuclear, whorls. Collectively, these make up the protoconch. Often, coiling and

surface morphology of these whorls are different from those that follow, termed

postembryonic or teleoconch whorls (Frest and Johannes, 1999).

While most mollusks do not have determinant growth, strictly speaking, the total number of

whorls and shell size are pretty stable features of sexually mature individuals of each species.

Adults seem to put much of their physiological efforts into gamete production instead of

growth. The whole shell is termed the conch; all whorls except the last one make up the spire.

10
The final (last, ultimate) whorl often differs in morphology in some way from those

preceding, most often in terms of aperture (mouth; shell opening proper: the rim or border

itself is termed the peristome) features. This is mostly the case if the animal is mature;

descriptions without specification always refer to mature (adult; sexually capable)

individuals. The whorl preceding may also be discriminated as the penultimate whorl. The

line of contact between adjacent whorls is the suture. Usually, it is sunken to some degree,

i.e. impressed. Whorl profile is important, also. Generally, a whorl in side view is rounded to

some extent; if very evenly so, it is convex; often, whorl shape is flattened to some degree.

The whorl periphery may be even, so that the outermost point is at midwhorl; or it may be

shouldered to varying extent, with the outermost point displaced adapically or abapically

(Frest and Johannes, 1999).

Some freshwater snails exhibit shell differences between the sexes (e.g. Melanoides

tuberculata which broods its young in a modified cephalic ‘pouch’, some species of

Lavigeria which brood in the pallial oviduct and Vinundu westae which is an egg-layer) while

others such as Tarebia granifera do not (Brande, Turner and Heller, 1996; Michel, 2004).

Another group where sexual dimorphism has been explored in detail is the genus Viviparus, a

holarctic group of freshwater brooding snails of the family Viviparidae. Sexual dimorphism

in Viviparus has been documented as differences in size (Van Cleave and Chambers, 1935;

Chiu et al., 2002). Female Viviparus are significantly larger than males due to differences in

reproductive age (females reproduce later in ontogeny) and selection of larger females for

increased brood size and brood viability (Van Cleave and Lederer, 1932; Jokinen, Guerette

and Kortmann, 1982; Brown, Varza and Richardson, 1989; Jakubik, 2006).

Sexual dimorphism in snails is traditionally quantified through straight-line shell

measurements and ratios (Son and Hughes, 2000; Velecka´ and Ju¨ttner, 2000). Recently,

geometric morphometrics have been employed for examinations of snail shell dimorphism,

11
both to provide descriptive analyses of shell shape and to answer broader evolutionary

questions (Hollander et. al., 2006).

2.4 REPRODUCTION

Coelho et al., (2012), observed that after the first 20 days of keeping the adults, courtship and

copulation behavior began to be observed. The male usually approaches the female from

behind, crawls over her shell and when at the last whorl searches for the genital aperture,

grasping with his penial sheath, and inserts it. Copulation can last for 1-5 hours and during

this period the female usually crawls around feeding, while the male retracts inside his shell.

The pair may be lifted out of water without interrupting copulation, showing that the male is

firmly attached to the female during the entire copulation period. Oviposition was difficult to

observe since it usually occurs at night. It was observed only once. The female crawls from

the water and deposits eggs on the tank wall. The eggs slide, one by one, from the mantle to

the lower edge of the clutch.

All species of Biomphalaria and Bulinus are hermaphrodite, possessing both male and female

organs and being capable of self- or cross-fertilization. A single specimen can invade and

populate a new habitat. The eggs are laid at intervals in batches of 5–40, each batch being

enclosed in a mass of jelly-like material. The young snails hatch after 6–8 days and reach

maturity in 4–7 weeks, depending on the species and environmental conditions. Temperature

and food availability are among the most important limiting factors. A snail lays up to 1000

eggs during its life, which may last more than a year. The amphibious Oncomelania snails,

which may live for several years, have separate sexes. The female lays its eggs singly near

the water margin (WHO, 1997).

12
13
FIG 2.1: Copulation behaviour in Pomacea bridgesii. Male inserting penial sheath in female
genital aperture (Coelho et al., 2012)

14
2.5 ECONOMIC IMPORTANCE

The two main areas of snail consumption in the world are western Africa and Western

Europe (Owolabi, 2006). In spite of the considerable external and local demand, commercial

snail farms such as those in Europe, South-East Asia and the America do not exist in West

Africa. In Nigeria, Ghana and Cote d’Iviore where snail meat is particularly popular, snails

are gathered from the forests (Owolabi, 2006). One of the important alternative sources of

animal protein, which has received relatively scanty attention in Nigeria (Enugu State in

particular) is the snail. Elsewhere, the potential of snail as source of high quality protein has

long been recognized. For instance, snail breeding started as far back as the beginning of 20th

century (Ayodele and Asimalowo, 1991) and the Romans raised snails on farms and fed them

with special herbs to improve their taste and increase overall snail meat availability (Odiabo,

1997).

From earliest times, humans have used many snail species as food. Periwinkles (Littorina) in

Europe and South Africa, queen conchs (Strombus gigas) in the West Indies, abalones

(Haliotis) in California and Japan, and turban shells (Turbo) in the Pacific are the most

frequently eaten marine snails. Occasionally limpets and whelks are used for food, but they

are more commonly used as fish bait (Solem, 2019). Freshwater snails rarely are eaten. Shells

of certain snails are highly prized by collectors. The operculum of some Turbo species is used

in making earrings; cameos are cut from the shell of the Red Sea snail Cassis rufa. Serious

medical problems are caused by the few freshwater snails (Pomatiopsis, Bulinus,

Biomphalaria) that serve as intermediate hosts for flatworms that parasitize humans.

Schistosomiasis is a disease caused by minute blood flukes (schistosomes). Both snails and

flukes are most common in areas where fields are irrigated. Schistosomes also parasitize birds

and mammals. A skin rash called swimmer’s itch results from bird schistosomes trying, only

partly successfully, to penetrate human skin. They die in the upper skin layers, and their

15
decomposition causes local infection. Freshwater snails also have aesthetic use in aquaria

(Solem, 2019).

2.6 FEEDING HABITS/BEHAVIOUR

Hydrobioids snails have a taenioglossate radula (i.e., seven teeth per row) comprising

numerous rows of cuspate teeth, each of which includes a typically squarish or trapezoidal

central tooth flanked on each side by lateral, inner marginal, and outer marginal teeth. Teeth

near the anterior end of the radula are often worn or broken, whereas the proximal portion of

the ribbon has many rows of poorly differentiated or incompletely formed teeth. To observe

some characters, teeth must be isolated because overlapping teeth obscure details when whole

mounts are photographed (Hershler and Ponder, 1998).

Many species of freshwater snail live in the wild and can find their way into home aquariums.

Different snails have a variety of diets.

2.6.1 Algae Eaters

Many species of freshwater snail eat algae. However, if something goes wrong with your

water parameters, excessive algae can result in a plague of snails. Malaysian trumpet snails

(Melanoides tuberculata) are notorious for this. Additionally, if you have too many snails and

they exhaust their food supply, the starving snails can die and further foul the water. Keeping

only a few algae-eating snails and keeping up on aquarium maintenance can help prevent this

sort of problem (Boumis, 2019).

2.6.2 Scavengers

Many freshwater snails act as scavengers. In aquariums, this trait can prove useful. Most

freshwater snails will eat extra fish food, as well as dead and dying fish. Such snails can help

keep a tank clean. However, if there is excessive food in the tank, you could see a population

16
explosion among your snails, which detracts from the look of the tank and triggers more

serious problems with your water chemistry (Boumis, 2019).

2.6.3 Aquarium Plants

Some species of freshwater snail eat plants. Even a few common aquarium species, like

mystery snails and apple snails from the family Ampullariidae, may devour aquarium plants.

Most snails with vegetarian inclinations will also devour algae, so they may make a great

addition to aquariums with plastic or silk aquarium plants. However, any live aquarium plant

is fair game for these species, and few can grow fast enough to survive their browsing.

(Boumis, 2019).

2.6.4 Meat

A few freshwater snails will even eat live prey. The assassin snail (Clea Helena) is probably

the best known example for aquarium hobbyists. You can actually use assassin snails to

control and possibly eradicate other species of snails. These snails will rapidly decimate other

species. Once they eat all of the other snails, they act as scavengers and usually breed too

slowly to become a pest in their own right. They may eat small decorative shrimp, but usually

can't harm fish or plants -- unlike a few saltwater snails that actually capture fish (Boumis,

2019).

CHAPTER 3

3.0 MATERIALS AND METHODS

17
3.1 Description of Study Area

The study is to be carried out along a stretch of Ovia River at Owan (also known as Owan

River) in Ovia-Northeast Local Government Area, Edo State (Lat. 6o16'0" N; 6o45'24" N and

Long. 5o43'54"; E5o46'30"E). The region is notable for its intensive cocoa farming and

plantain markets. A vast number of these farms are located along the river bank. These farms

are also heavily treated with Agrochemicals to protect and boost the crops yield. The

sampling point for this study is at Owan (Omoigberale et. al., 2014; Akinbo and Okaka,

2010).

Fig 3.1: Drawn by: Ukrakpor Ajiri, Department of Geography & Regional Planning, University
of Benin, Benin City. (2014)

The survey carried out at the above specified location resulted in the discovery of 3 species of

freshwater snail, Pila Africana, Pila ampullaceal and Lanistes libycus.

18
3.2 Method of Sampling/Collection of Samples

The survey will be carried out with the assistance of the local fisher folks. The freshwater

snail samples are mainly collected by hand picking or with the use of small scoop nets and

baskets at marginal areas and submerged vegetation. The collected samples will be washed

with the river water to remove debris from sand, organic waste, algae and vegetation.

Basketfuls of freshwater snails will be purchased at the river side and counted. Identification

of the samples will be done at the lab with the aid of the necessary keys.

3.3 Morphometric Measurements

Morphometric measurements such as shell width, shell length, whorl length, aperture width

and spire length will be taken with the aid of digital vernier caliper and measurements will be

taken in millimeters (mm). Number of whorls, height of spire will also be taken into

consideration. Also the shape of the shells - whether discoidal or conical, will be considered

(Sangeeta et al., 2015).

Fig 3.2: Morphometric traits of Lymnaea

acuminata shell

Abbreviation: (ASL=animal shell length,

ASW=animal shell width,

ApL=aperture length, ApW=aperture width.)

(Priyanka et al., 2016).

3.4 Materials to Be Used For Study

19
The apparatus and materials needed for the study include venire caliper for taking inner or

internal measurements of the shell width, electrical digital weighing balance and transparent

rule will be used for measuring the length in centimeters, writing materials for recording

values, small scoop nets, gloves and Identification keys.

3.5 Statistical Analysis

The data obtained from the field during survey will be analysed using the SPSS (Statistical

Package for Social Sciences Students) version 24.0 software. Pie charts, bar charts and graphs

will be used to descriptively show relationship. The data obtained was subjected to analysis

of variance (ANOVA) to determine significant differences among treatments and the

treatment means were separated by Duncan’s Multiple Range Tests (DMRT) at 5%

probability level.

3.6 Condition Factor

The condition factor for the snails will be computed using the formula below

CF = W * 100
L3

The condition factor (CF) is an estimation of the general well-being of the species (Oribhabor

et al., 2011). The condition factor of 1.0 or greater indicates the good condition of the animal

species while less than 1.0 shows bad condition (Abobi, 2015).

3.7 Experimental Design

20
The experimental design that will be used is a factorial design which involves composite

samples: three species that have been found at Owan (Pila Africana, Pila ampullacea,

Lanistes libycus), one season (wet), three sizes (big, medium and small) replicated twice. The

experiment will be carried out for a period of 6 weeks (three months, from September to

November).

3.8 Sex Identification

Selected samples of the species obtained from the site will be dissected to examine the

internal structure of snails to ascertain sexual differences (if any) in order to confirm whether

or not the species have separate sexes or hermaphroditic.

CHAPTER 4

4.0 RESULTS

4.1 INTRODUCTORY STATEMENT

During the course of the field work, 3 species where identified at the study station – Pila

ampullacea, Pila africana, and Lanistes libycus. From the Table below and overall

comparison can be made between the three species (irrespective of size). From table below

(Table 4.1), it can be inferred that there exist no significant difference (p>0.05) between P.

africana and P. ampullacea, but that significant difference exist between Lanistes libycus and

the other two species. Significance level was determined using F-test and means were

separated using the Duncan Multiple Range Test.

4.2 SPECIES COMPOSITION AND RELATIVE ABUNDANCE

The the river was sampled every two weeks for a period of three months as seen in Table 4.2.

A total of 658 snails were worked with during the sampling period. Three different species of

freshwater snails where identified in the river at Owan river, which were – Pila ampullacea,

21
Pila africana and Lanistes libycus. From week 1 to week 3 only two out of the 3 species were

observed (P. ampullacea and P. africana) while the third species (Lanistes libycus recorded

occurrence from week 4 to week 6).

The highest number of snails was sampled in the second week (18.9%). Pila ampullacea was

the most abundant snail species (44.5%), followed by Pila africana (38.1%) and then

Lanistes libycus (17.5%). The distribution of snails shows that week 2 had the highest

percentage of the freshwater snail abundance of 18.9% with Pila ampullacea being 10.6%

and Pila africana being 8.4%. At the first sampling Pila ampullacea had percentage

abundance of 9.1%, while Pila africana was 8.8%. In 3, Pila ampullacea had 6.4% and Pila

africana was 4.3%. In week 4 there was occurrence of Lanistes libycus and it had the highest

abundance in that week with 7.3%, while P. ampullacea and P. africana recorded 4.9% and

6.2% respectively. In week 5 P. ampullacea was 6.4% P. africana – 5.8% and L. libycus

5.3% while the abundance for week 6 for P. ampullacea, P. africana and L. libycus were

7.1%, 4.6% and 4.9% respectively. Overall, P. ampullacea had the highest abundance during

the period of study.

22
Table 4.1: Table showing comparison of the means and standard deviations of the morphometric
parameters across the three species of freshwater snails found at Owan.

Parameters Pila Pila Lanistes F P

ampullacea africana libycus

Weight (Grams) 48.22±27.76a 56.64±38.30a 29.28±15.97b 16.456 0.000

Total Length (CM) 5.22±1.63a 5.65±1.79a 4.26±1.30b 15.107 0.000

Apperture Width (CM) 2.58±0.86a 2.76±0.95a 1.96±0.63b 18.102 0.000

Apperture Length 4.16±1.23a 4.30±1.30a 3.41±1.17b 11.097 0.000

(CM)

Shell Width (CM) 5.06±1.70a 5.19±1.60 a 4.39±1.23 b 5.576 0.004

Condition Factor 34.21±13.36a 32.17±14.54a 41.74±30.33b 6.273 0.002

NOTE: The different superscripts within rows indicate significant differences (p< 0.05).
Rows with similar superscripts are not significantly different (p>0.05).

23
Total snails collected during the study was 658 in number, represented as P. ampullacea 293,
P. africana 250 and L. libycus 115.

Table 4.2: Table showing the relative species composition and abundance of freshwater
snails in Ovia river at Owan.

Number of Species/Percentage Abundance

Species found Week 1 Week Week 3 Week Week 5 Week 6 Total

2 4
Pila 60(9.1% 70(10. 42(6.4 32(4.9 42(6.4% 47(7.1% 293(44.5%
ampullacea ) 6%) %) %) ) ) )
Pila africana 58(8.8% 55(8.4 28(4.3 41(6.2 38(5.8% 30(4.6% 250(38.1%
) %) %) %) ) ) )
Lanistes - - - 48(7.3 35(5.3% 32(4.9% 116(17.5%
libycus %) ) ) )
Total 118(17. 125(18 70(12.3 121(18. 115(17. 109(16. 658(100%)
9%) .9%) %) 4%) 5%) 6%)

24
 17%

44%

Pila ampullacea
Pila africana
Lanistes libycus

38%

FIG 4.1: Pie chart showing the relative species composition of the freshwater snails found
in Ovia River at Owan

25
 80

70

60

50

Pila ampullacea
40
Pila africana
Lanistes libycus
30

20

10

0
Week 1 Week 2 Week 3 Week 4 Week 5 Week 6

FIG 4.2: Bar Chart showing the relative abundance of the different species found in Ovia
River at Owan

4.2 MORPHOMETRIC CHARACTERISTICS OF THE SPECIES

The different species found were measured in three categories – large, medium and small

sizes. The morphometric parameters of concern include – aperture width, aperture length,

total shell length, total shell width and the weight.

4.2.1 Weight

On the basis of size comparison amongst the species, the analysis of the samples collected

showed that there was significant difference (p<0.05) for the large/big sizes of P. africana

and P. ampullacea. The mean and standard deviation of weight for P. ampullacea was

80.56±16.44 while Pila africana was 104.14±32.14. For the medium size samples the mean

and standard deviation values for P. ampullacea, P. africana and L. libycus were 46.87±9.01,

46.02±9.22, 43.77±7.19 respectively. While for the small size samples 16.21±5.45,

20.33±6.34, 14.80±5.76 means and standard deviations were recorded for P. ampullacea, P.

26
africana and L. libycus respectively. From these values it was observed that from the samples

collected Pila africana had higher weight ratio in comparison to the other two species.

4.2.2 Total Length

The statistical analysis revealed that the mean and standard deviation for the total shell length

of the small sized specimen of Pila ampullacea was 3.47±0.43, Pila africana was 3.71±0.67

and L. libycus was 3.13±0.62. Significant difference (p<0.05) was not observed in terms of

total length for all three species for this size (Table 4.3).

The means and standard deviation for the medium sized specimen was as follows: Pila

ampullacea - 5.06±0.73; Pila africana - 5.35±0.75 and Lanistes libycus - 5.38±0.64. While

for large size the means for total length were 7.09±1.05 and 7.81±1.03 for P. ampullacea, P.

africana and L. libycus respectively (Table 4.3).

4.2.3 Aperture Width

Significant difference (p>0.05) was not really observed between P. ampullacea and P.

africana large sizes with 3.38±0.59cm and 3.82±0.75cm respectively. The medium sizes for

P. ampullacea and P. africana and L. libycus recorded the following mean measurements for

the aperture width 2.71±0.42cm, 2.59±0.46cm and 2.42±0.39.cm while for the small size

1.59±0.44cm, 1.84±0.48cm and 1.50±0.47cm respectively (Table 4.3).

4.2.4 Aperture Length

It was observed that the aperture length for both P. ampullacea and P. africana were

observed to be nearly twice or one and a half times as long as the width. The readings for

large size specimen were 5.43±0.72cm and 5.84±0.79 (in comparison to 3.38±0.59cm and

3.82±0.75cm for aperture width) for P. ampullacea and P. africana.

27
The medium size specimens had mean and standard deviation readings for aperture length of

4.26±0.59, 4.10±0.61 and 4.42±0.58 for P. ampullacea, P. africana and L. libycus while the

readings for small size respectively each of the species were 3.28±0.41cm, 3.44±0.67cm and

3.32±0.66cm (Table 4.3).

4.2.5 Shell Width

Lanistes libycus shell width was consistently greater than the total length of the shell for both

medium and small sizes having mean and standard deviation readings of 5.46±0.53cm and

3.32±0.66cm respectively for medium and small sizes. In P. ampullacea, the mean and

standard deviation readings were 3.28±0.41cm, 4.89±0.67cm and 6.96±1.31cm for small,

medium and large size of the species, while P. africana recorded 7.10±0.86cm, 4.97±0.72cm

and 3.44±0.67cm for large, medium and small sizes respectively (Table 4.3).

4.2.6 Condition Factor

The overall mean K-value for Pila ampullacea during the sampling period in the study area

was found to be 34.21, while that of Pila africana was 32.17 and Lanistes libycus 41.74

(Table 4.1), which indicates that the species are doing well in their environment.

Lanistes libycus had the highest mean and standard deviation condition factor value for the

small size among the three species with 54.47±38.61, P. ampullacea with 39.29±12.38 and P.

africana with 42.32±14.94. For the large size P. ampullacea was 24.16±8.11 while P.

africana was 22.39±6.37 while the medium size was 38.52±13.13, 32.59±13.91 and

29.01±6.73 for P. ampullacea, P. africana and L. libycus.

28
Table 4.3: Comparison and variations of morphometric measurements of the three species of
freshwater snails found in Owan.

LARGE/BIG Pila ampullacea Pila africana Lanistes libycus F P

WEIGHT (GRAMS) 80.56±16.44 104.14±32.14 20.752 0.000

TOTAL LENGTH (CM) 7.09±1.05 7.81±1.03 11.224 0.001

APPERTURE WIDTH (CM) 3.38±0.59 3.82±0.75 9.933 0.002

APPERTURE LENGTH
(CM) 5.43±0.72 5.84±0.79 7.011 0.010

SHELL WIDTH (CM) 6.96±1.31 7.10±0.86 0.382 0.538

CONDITION FACTOR 24.16±8.11 22.39±6.37 1.360 0.247

MEDIUM

WEIGHT (GRAMS) 46.87±9.01 46.02±9.22 43.77±7.19 1.258 0.287

TOTAL LENGTH (CM) 5.06±0.73 5.35±0.75 5.38±0.64 3.110 0.048

APPERTURE WIDTH (CM) 2.71±0.42 2.59±0.46 2.42±0.39 4.720 0.010

APPERTURE LENGTH
(CM) 4.26±0.59 4.10±0.61 4.42±0.58 3.084 0.049

SHELL WIDTH (CM) 4.89±0.67 4.97±0.72 5.46±0.53 7.991 0.001

CONDITION FACTOR 38.52±13.13 32.59±13.91 29.01±6.73 6.693 0.002

SMALL

WEIGHT (GRAMS) 16.21±5.45 20.33±6.34 14.80±5.76 8.945 0.000

TOTAL LENGTH (CM) 3.47±0.43 3.71±0.67 3.13±0.62 8.705 0.000

APPERTURE WIDTH (CM) 1.59±0.44 1.84±0.48 1.50±0.47 5.284 0.006

APPERTURE LENGTH
(CM) 2.72±0.53 2.91±0.48 2.41±0.60 7.652 0.001

SHELL WIDTH (CM) 3.28±0.41 3.44±0.67 3.32±0.66 0.891 0.413

CONDITION FACTOR 39.29±12.38 42.32±14.94 54.47±38.61 4.340 0.015

29
4.3 MORPHOLOGICAL FEATURES/VARIATIONS OF THE SPECIES

Freshwater snails during the study were observed to have separate sexes. The dissection

carried out at the main lab Faculty of Agriculture, UNIBEN, showed that in comparison with

the land snails which have a double hermaphroditic duct carrying gametes from each of the

two genital organs present within the ovotestis, the freshwater snail samples collected showed

when dissected, only one reproductive duct which signifies the presence of one genital organ

meaning that the snails were not hermaphroditic. Externally, sex differences are subtle and

are not easily noticeable. During the course of the study, the differences in the shell structure

of both genders were not however clearly identified.

In terms of morphological differences, significant difference (p<0.05) was not observed

between P. ampullacea and P. africana. Lanistes libycus showed significant difference

(p<0.05) from both P. ampullacea and P. africana (Table 4.1). Smaller sized snails were

more easily encountered than large or medium sized ones for Lanistes libycus and Pila

ampullacea.

30
PLATE 4.1 (Size comparison): From left to right – Medium sized Pila ampullacea, Large
sized Pila ampullacea and Medium sized Pila africana.

31
PLATE 4.2s: Morphological difference in terms of shell orientation and structure
between small sized Lanistes libycus (on the left) and Pila ampullacea (on the right), with
L. libycus having a sinistral shell and P. africana having a dextral shell.

32
PLATE 4.3: Presence of a single reproductive duct (indicated by the red arrow)
connected to the bright yellow sac-like albumen gland which in turn connected to the
reproductive organs, indicate that the organism posses a single reproductive organ.

33
 CHAPTER 5

5.0 DISCUSSION

Freshwater snails are important components of aquatic ecosystems, and many species are

excellent water-quality indicators. Approximately three-fourths of these freshwater snail

species are threatened, endangered, or of conservation concern or are already thought to be

extinct (Johnson, 2019).

Freshwater snails are an important food source for many fish, turtles, and other species of

wildlife and because of their sensitivity to certain chemicals, many species are excellent

water-quality indicators (Johnson, 2019).

Three freshwater snail species were found at Owan community, Ovia-North East local

government area, Edo state namely – Pila ampullacea, Pila africana and Lanistes libycus.

Sampling was done over a period of six weeks (three months), twice a month. The snails are

commonly referred to as “kere kere” by the native people.

Snail samples were obtained either by hand picking from the river shore or from the marginal

vegetation of the water or with the use of hand-made local traps (which are primarily made

for catching fish). Local fishermen in area commonly make use of fresh water snails as baits

in their traps for catching fish.

5.1 GENERAL MORPHOLOGY OF AMPLURIIDAE (APPLE SNAILS)

All species identified in the area (– Pila ampullacea, Pila africana and Lanistes libycus) fall

under the family Ampullaridae. There are general features which are common to all snails in

this family, such as possession of hard bony plate-like operculum which act as a covering or

closure over the aperture opening.

34
The Ampullariidae are unusual because they have both a gill and a lung, with the mantle

cavity being divided in order to separate the two types of respiratory structures. This

adaptation allows these snails to be amphibious (Bouchet & Rocroi, 2005).

5.2 MORPHOMETRY/MORPHOLOGY AND IDENTIFICATION OF THE

SPECIES

Pila ampullacea is a large to medium sized freshwater snail and from the study carried out,

can reach total length of 10.5cm (that is over 100 millimeters). This agrees with the findings

of Ponder et al., 2016. Both Pila ampullacea and Pila africana come from the same genus

and share the similarity of having a dextral shell (shell opening at the right), but Lanistes

libycus was observed to have a sinistral shell (shell opening at the left). However, P. africana

was noticed to have a more protruded or longer spire than P. ampullacea whose spire were

shorter or more reduced or flattened.

From the measurements taken, the shell width for L. libycus was usually or more frequently

larger than its shell lengths (wider than longer) while P. africana and P. ampullacea had

lengths that were more frequently greater than their widths. Although this was not always the

case with every sample measured, there were some exceptions to this observation.

Measurements carried out by Nwe Nwe San (2015) in Natyaekan Fish Culture Ponds and

Taungthaman Lake in Amarapura Township, Singapore revealed that the mean total length

and total width for P. ampullacea were 3.4cm and 2.2cm respectively, while mean total

length and mean total width for P. africana were 3.94cm and 4.67cm respectively. This

implies that larger sizes of P. africana were obtained compared to P. ampullacea. This bears

similarity with the results obtained from Ovia river at Owan during this study, as the results

show that larger sizes of P. aficana were obtained than P. ampullacea since mean total length

35
and width for P. africana were 5.65cm and 5.19cm respectively while P. ampullacea was

5.22cm and 5.06cm (Table 4.1).

The identification of Pila ampullacea was carried out based on diagnosis made by W. S. S.

van Benthem Jutting (1956), and Brandt (1974) as follows - Shell sub-globose, often

somewhat reversely conic; surface smooth, usually with fine axial lines; to about 100 mm in

shell height; spire low, conical to almost flat, suture distinct, but not deeply impressed; body

whorl large, rounded or indistinctly angular at the shoulder, especially in juveniles; peristome

not continuous except in large shells, which is joined by a thin parietal callus; columellar and

basal margin of aperture slightly reflexed; umbilicus open, small and partly hidden by

apertural margin.

Pila africana has a relatively short spire with the tip of the spire having a more pointed look

than P. ampullacea, has globe-like curved shell, shell opening is rounded at the bottom but

sharper at the top. The umbilicus is ranges from medium to wide and deep, the colour of the

shell is dark brown or sometimes light yellow with faint spiral bands.

Lanistes has a unique anatomy among the Ampullariidae: it has a "hyperstrophic" sinistral

shell. This means that the body of the snail is dextral (as in all other ampullariids), but the

shell appears to be sinistral. However the sinistral appearance stems from the fact that the

rotation of the shell as it grows is in an upward direction rather than the usual downward

direction (Shell 2019).

5.3 COMPOSITION, ABUNDANCE AND DISTRIBUTION

From the study carried out it was discovered that the composition of the snails at the site

include P. ampullacea, L. libycus and P. africana. Calculations on the abundance of the

freshwater snails revealed that P. ampullacea had the highest percentage abundance over P.

36
africana and L. libycus with about 44% while the others were 38% and 18% respectively.

This is however in contrast to the findings of Edegbene, A.O. and Arimoro, F.O. (2014) who

identified four species which include; Potadoma moerchi, Melanoides tubaculata and

Melanoides moerchi of the family Thiaridae and Pseudospatha sp. of the family Unionidae in

Owan river at 4 sample locations (Evbiamen – Emai, Okpokhumi- Emai, Ojavun and Ogute

community, in Edo state). Lanistes libycus had an occurrence and percentage abundance of

30.8% at Ifaki-Ekiti, Iworoko-Ekiti and Are-Ekiti in Ekiti State, Nigeria (Olorunniyi and

Olofintoye, 2017).

Hayes et al., (2015), suggested that members of the Pila genus (including P. ampullacea and

P. africana) could be found across Africa from West Africa (Liberia) to East and Southern

Africa (Tanzania). There is reported occurrence of Pila ampullacea in River Benue by

Obande et al., (2013) but the percentage relative abundance was not stated. Danladi et al.,

(2019), reported the occurrence of Lanistes libycus at Dadinkowa man-made reservoir,

Gombe State, Nigeria, percentage relative abundance not stated. Ndifon and Ukoli (1989)

reported the occurrence of Lanistes libycus across 404 freshwater habitats in south-western

Nigeria with a percentage relative abundance of 17.6%.

The study was carried out during the rainy season, from September to November twice a

month with two preliminary surveys done in august, a total of 6 weeks from September to

November. It was observed that towards late November, there was higher occurrence of

smaller sized snails than large ones, as the fishermen reported greater difficulty in obtaining

the samples in this period. From the results of the sampling, the reduction or receding of the

water due to reduced rainfall and approaching dry season, the assumption can be made that

increased diversity of freshwater snail species occurred as the water level fell. This shows a

directly proportional relationship with falling water levels or receding water due to reduced

rainfall. This is because Lanistes libycus which was not previously found in the area as at the

37
period of more frequent rainfall (early september to mid-october, that is from weeks 1 to 3)

was observed during the period the river began to recede, that is from early November. This

assertion is corroborated by Appleton (1978), who stated that during the rainy season, rainfall

affects water movement and temperature thereby affecting the distribution and density of

aquatic snails. During the rainy season most of the snail species and their eggs are carried

away by high water current resulting in lesser number of snail species caught during this

period (Kenneth, 2002).

5.4 SEX DIFFERENCES

Although the external sexual differences were not clearly defined in this study, the

dissections of some specimens at Main Lab, Faculty of Agriculture University of Benin with

assistance from Dr. Omoyakhi and Dr. Okhale, confirmed that the snails indeed have separate

sexes as a single reproductive tract was seen connected to the bright yellow sac-like albumen

gland which in turn is connected to the reproductive organs, indicate that the organism posses

a single reproductive organ. This is in direct contrast to the internal structure of land snails

which possess double reproductive tracts within the hermaphroditic duct and are separately

connected to the ovaries and testes both contained within the ovotesties (Duncan, 1958).

38
 CHAPTER 6

6.0 CONCLUSION

Pila africana is a species of freshwater snail that is currently on the IUCN (International

Union for Conservation of Nature) Red List as threatened (Jørgensen et al., 2010). There is

difficulty in accessing enough detailed information on the ecology and distribution of

freshwater snails from the study location. This is one of the major issues this study was aimed

at resolving. The species found at the project site (Owan river) are in good condition as the

human interference by pollution (deposition of human and domestic waste) seem not be at a

level where the health and survival of the snails are adversely affected.

6.1 RECOMMENDATION

I. Efforts should be made in order to preserve the species of freshwater snails found in the

area.

II. More studies should be carried out regularly in order to provide current data on the

freshwater snail status from the area.

39
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44
 APPENDIX

WEIGHT (GRAMS) TOTAL LENGTH (CM) APPERTURE WIDTH (CM) APPERTURE

LENGTH (CM) SHELL WIDTH (CM) Condition factor * Species * Week
TOTAL APPERTURE SHELL
WEIGHT LENGTH APPERTURE LENGTH WIDTH Condition
Species Week (GRAMS) (CM) WIDTH (CM) (CM) (CM) factor
Pilaampullace 1 Mean 49.7667 5.2767 2.6877 4.1650 5.3720 33.3325
a Std. 31.65404 1.61527 .94673 1.17807 1.78244 14.51123
Deviation
Minimum 10.00 3.00 1.20 2.50 3.20 17.14
Maximum 122.00 8.11 4.72 6.95 8.90 73.44
N 30 30 30 30 30 30
2 Mean 49.2500 5.7121 2.5796 4.3461 5.6779 28.9821
Std. 28.43658 2.10561 .83107 1.32861 2.35893 13.60238
Deviation
Minimum 8.00 3.04 .90 1.40 2.70 7.60
Maximum 102.00 10.50 4.20 6.82 10.55 67.64
N 28 28 28 28 28 28
3 Mean 60.1500 5.8690 3.2685 4.7320 5.4055 32.4200
Std. 17.85696 1.11810 .52250 .91058 .89939 12.73144
Deviation
Minimum 40.00 4.00 2.70 3.30 4.00 16.12
Maximum 121.00 8.20 4.50 6.70 7.40 64.06
N 20 20 20 20 20 20
4 Mean 46.5333 4.9407 2.4383 4.0580 4.6063 38.2305
Std. 28.63171 1.51522 .79769 1.11978 1.40924 15.42614
Deviation
Minimum 12.00 3.20 1.20 2.48 2.94 19.87
Maximum 98.00 7.82 3.80 6.08 7.30 71.07
N 30 30 30 30 30 30
5 Mean 50.5667 5.2387 2.5830 4.3337 5.1130 33.4024
Std. 29.83019 1.59940 .85779 1.28084 1.57290 9.95289
Deviation
Minimum 13.00 2.77 1.23 2.30 2.56 19.87
Maximum 98.00 7.82 3.92 6.44 7.68 61.17
N 30 30 30 30 30 30
6 Mean 31.5000 4.2005 1.9505 3.2395 3.9480 39.8289
Std. 17.32507 1.00401 .69448 1.16304 .96218 10.42006
Deviation
Minimum 7.00 2.73 .62 1.23 2.51 26.47
Maximum 58.00 5.66 2.85 4.73 5.36 59.32
N 20 20 20 20 20 20
Tota Mean 48.2152 5.2216 2.5815 4.1634 5.0556 34.2116
l Std. 27.76387 1.63068 .86377 1.23376 1.69612 13.35626
Deviation
Minimum 7.00 2.73 .62 1.23 2.51 7.60
Maximum 122.00 10.50 4.72 6.95 10.55 73.44
N 158 158 158 158 158 158
Pilaafricana 1 Mean 54.7000 6.4617 3.1050 4.4470 5.7740 19.8669
Std. 35.85041 1.91863 .73048 1.20671 1.65114 6.47160
Deviation
Minimum 13.00 3.50 2.08 2.91 3.10 10.27
Maximum 153.00 9.63 4.70 6.90 8.92 37.36

45
 N 30 30 30 30 30 30
2 Mean 54.9000 5.4363 2.6203 4.0230 5.0697 30.1694
Std. 44.80483 1.86620 1.20756 1.52286 1.66415 9.32770
Deviation
Minimum 10.00 2.70 1.10 1.90 2.30 19.73
Maximum 172.00 8.78 5.92 7.24 7.62 66.05
N 30 30 30 30 30 30
3 Mean 64.4375 5.8306 2.8006 4.4825 5.3869 33.1544
Std. 24.21836 1.13052 .61279 .75763 1.04773 9.01439
Deviation
Minimum 38.00 4.10 2.02 3.55 3.80 20.93
Maximum 110.00 8.07 3.70 5.70 7.22 56.59
N 16 16 16 16 16 16
4 Mean 57.6786 5.5200 2.7929 4.4925 5.0700 33.1842
Std. 46.34177 1.93368 1.04366 1.46441 1.81807 12.86620
Deviation
Minimum 13.00 2.97 1.21 2.44 2.50 15.32
Maximum 177.00 10.25 5.20 7.84 9.37 59.27
N 28 28 28 28 28 28
5 Mean 58.6333 5.2930 2.6103 4.2137 4.8900 41.2004
Std. 39.01059 1.77278 1.01589 1.37977 1.63088 16.06456
Deviation
Minimum 15.00 3.00 1.22 1.90 2.87 17.69
Maximum 172.00 8.78 4.92 6.75 7.62 70.49
N 30 30 30 30 30 30
6 Mean 46.3000 4.9620 2.4000 4.1200 4.7880 43.6370
Std. 10.52035 .99173 .39944 .81279 .94283 23.18390
Deviation
Minimum 33.00 3.43 1.80 3.18 3.39 23.83
Maximum 61.00 6.02 3.00 5.40 5.81 94.17
N 10 10 10 10 10 10
Tota Mean 56.6389 5.6472 2.7575 4.3001 5.1947 32.1743
l Std. 38.30273 1.79072 .95474 1.30147 1.60031 14.53579
Deviation
Minimum 10.00 2.70 1.10 1.90 2.30 10.27
Maximum 177.00 10.25 5.92 7.84 9.37 94.17
N 144 144 144 144 144 144
Lanisteslibycus 4 Mean 27.0000 4.0795 1.8915 3.1335 4.2800 38.6901
Std. 14.30863 1.13156 .54821 1.09420 1.00613 10.57065
Deviation
Minimum 6.00 2.30 1.20 1.60 2.60 23.36
Maximum 47.00 5.86 2.93 5.03 5.93 58.00
N 20 20 20 20 20 20
5 Mean 30.9000 4.3905 2.1280 3.6320 4.5535 36.5379
Std. 14.00714 1.04844 .49406 .91518 1.01469 9.51379
Deviation
Minimum 12.00 3.01 1.23 1.93 2.56 23.96
Maximum 52.00 5.81 2.90 4.92 6.03 52.92
N 20 20 20 20 20 20
6 Mean 29.9500 4.2965 1.8580 3.4715 4.3395 49.9988
Std. 19.54879 1.66609 .79527 1.44803 1.62700 50.43684
Deviation
Minimum 6.00 1.45 .68 1.23 1.78 19.66
Maximum 60.00 6.58 3.63 5.85 6.77 196.81
N 20 20 20 20 20 20
Tota Mean 29.2833 4.2555 1.9592 3.4123 4.3910 41.7423
l Std. 15.96829 1.29518 .62749 1.17235 1.23452 30.32810
Deviation
Minimum 6.00 1.45 .68 1.23 1.78 19.66
Maximum 60.00 6.58 3.63 5.85 6.77 196.81
N 60 60 60 60 60 60
Total 1 Mean 52.2333 5.8692 2.8963 4.3060 5.5730 26.5997

46
 Std. 33.62171 1.85710 .86435 1.19085 1.71544 13.04560
Deviation
Minimum 10.00 3.00 1.20 2.50 3.10 10.27
Maximum 153.00 9.63 4.72 6.95 8.92 73.44
N 60 60 60 60 60 60
2 Mean 52.1724 5.5695 2.6007 4.1790 5.3633 29.5962
Std. 37.58319 1.97265 1.03416 1.42918 2.03441 11.50077
Deviation
Minimum 8.00 2.70 .90 1.40 2.30 7.60
Maximum 172.00 10.50 5.92 7.24 10.55 67.64
N 58 58 58 58 58 58
3 Mean 62.0556 5.8519 3.0606 4.6211 5.3972 32.7464
Std. 20.71569 1.10760 .60393 .84376 .95377 11.08846
Deviation
Minimum 38.00 4.00 2.02 3.30 3.80 16.12
Maximum 121.00 8.20 4.50 6.70 7.40 64.06
N 36 36 36 36 36 36
4 Mean 45.5256 4.9278 2.4254 3.9769 4.6891 36.5369
Std. 35.43376 1.67519 .90496 1.34300 1.50189 13.47779
Deviation
Minimum 6.00 2.30 1.20 1.60 2.50 15.32
Maximum 177.00 10.25 5.20 7.84 9.37 71.07
N 78 78 78 78 78 78
5 Mean 48.6750 5.0470 2.4795 4.1133 4.8895 37.1105
Std. 32.42666 1.58213 .86574 1.25834 1.47639 12.82668
Deviation
Minimum 12.00 2.77 1.22 1.90 2.56 17.69
Maximum 172.00 8.78 4.92 6.75 7.68 70.49
N 80 80 80 80 80 80
6 Mean 33.8400 4.3912 2.0034 3.5084 4.2726 44.6585
Std. 18.02760 1.31638 .70953 1.25111 1.28364 33.89099
Deviation
Minimum 6.00 1.45 .62 1.23 1.78 19.66
Maximum 61.00 6.58 3.63 5.85 6.77 196.81
N 50 50 50 50 50 50
Tota Mean 48.4282 5.2308 2.5484 4.0933 5.0008 34.6494
l Std. 32.34024 1.71119 .90859 1.28603 1.61022 17.95812
Deviation
Minimum 6.00 1.45 .62 1.23 1.78 7.60
Maximum 177.00 10.50 5.92 7.84 10.55 196.81
N 362 362 362 362 362 362

WEIGHT (GRAMS) TOTAL LENGTH (CM) APPERTURE WIDTH (CM) APPERTURE

LENGTH (CM) SHELL WIDTH (CM) Condition factor * Species * Size
TOTAL APPERTURE SHELL
WEIGHT LENGTH APPERTURE LENGTH WIDTH Condition
Species Size (GRAMS) (CM) WIDTH (CM) (CM) (CM) factor
Pilaampullace Big Mean 80.5600 7.0906 3.3830 5.4306 6.9586 24.1575
a Std. 16.44057 1.05336 .58600 .71672 1.30605 8.11029
Deviation
Minimum 61.00 5.00 2.35 3.40 5.18 7.60
Maximum 122.00 10.50 4.72 6.95 10.55 52.00
N 50 50 50 50 50 50
Medium Mean 46.8667 5.0627 2.7103 4.2602 4.8905 38.5244
Std. 9.00935 .73137 .42477 .58681 .66733 13.12947
Deviation
Minimum 30.00 3.50 2.00 3.10 3.20 17.29
Maximum 68.00 6.70 3.42 5.36 6.00 73.44
N 60 60 60 60 60 60
Small Mean 16.2083 3.4733 1.5856 2.7225 3.2798 39.2937

47
 Std. 5.44980 .42956 .43628 .52555 .40552 12.38008
Deviation
Minimum 7.00 2.73 .62 1.23 2.51 21.70
Maximum 29.00 4.80 2.60 3.67 4.40 67.64
N 48 48 48 48 48 48
Total Mean 48.2152 5.2216 2.5815 4.1634 5.0556 34.2116
Std. 27.76387 1.63068 .86377 1.23376 1.69612 13.35626
Deviation
Minimum 7.00 2.73 .62 1.23 2.51 7.60
Maximum 122.00 10.50 4.72 6.95 10.55 73.44
N 158 158 158 158 158 158
Pilaafricana Big Mean 104.1364 7.8120 3.8177 5.8418 7.1016 22.3862
Std. 32.14400 1.02846 .74928 .78896 .85980 6.37127
Deviation
Minimum 61.00 5.88 2.50 4.30 5.59 10.27
Maximum 177.00 10.25 5.92 7.84 9.37 38.24
N 44 44 44 44 44 44
Medium Mean 46.0167 5.3538 2.5945 4.0978 4.9663 32.5897
Std. 9.22137 .75064 .45564 .60776 .71908 13.91094
Deviation
Minimum 30.00 3.43 1.70 3.00 3.39 16.02
Maximum 61.00 6.90 3.74 5.42 6.50 94.17
N 60 60 60 60 60 60
Small Mean 20.3250 3.7060 1.8358 2.9078 3.4398 42.3182
Std. 6.33827 .67418 .47825 .48237 .67195 14.94408
Deviation
Minimum 10.00 2.70 1.10 1.90 2.30 19.92
Maximum 30.00 5.32 2.80 4.20 4.77 70.49
N 40 40 40 40 40 40
Total Mean 56.6389 5.6472 2.7575 4.3001 5.1947 32.1743
Std. 38.30273 1.79072 .95474 1.30147 1.60031 14.53579
Deviation
Minimum 10.00 2.70 1.10 1.90 2.30 10.27
Maximum 177.00 10.25 5.92 7.84 9.37 94.17
N 144 144 144 144 144 144
Lanisteslibycus Medium Mean 43.7667 5.3800 2.4157 4.4190 5.4647 29.0102
Std. 7.18563 .64402 .38611 .57774 .53448 6.72652
Deviation
Minimum 28.00 3.82 1.87 3.22 4.20 19.66
Maximum 60.00 6.58 3.63 5.85 6.77 50.23
N 30 30 30 30 30 30
Small Mean 14.8000 3.1310 1.5027 2.4057 3.3173 54.4743
Std. 5.75596 .61795 .46992 .60481 .65572 38.60915
Deviation
Minimum 6.00 1.45 .68 1.23 1.78 32.79
Maximum 30.00 3.98 2.47 3.44 4.45 196.81
N 30 30 30 30 30 30
Total Mean 29.2833 4.2555 1.9592 3.4123 4.3910 41.7423
Std. 15.96829 1.29518 .62749 1.17235 1.23452 30.32810
Deviation
Minimum 6.00 1.45 .68 1.23 1.78 19.66
Maximum 60.00 6.58 3.63 5.85 6.77 196.81
N 60 60 60 60 60 60
Total Big Mean 91.5957 7.4283 3.5865 5.6231 7.0255 23.3284
Std. 27.56861 1.09757 .69862 .77525 1.11610 7.36309
Deviation
Minimum 61.00 5.00 2.35 3.40 5.18 7.60
Maximum 177.00 10.50 5.92 7.84 10.55 52.00
N 94 94 94 94 94 94
Medium Mean 45.9067 5.2426 2.6051 4.2270 5.0357 34.2477
Std. 8.78402 .73312 .44091 .60180 .69582 12.94941
Deviation

48
 Minimum 28.00 3.43 1.70 3.00 3.20 16.02
Maximum 68.00 6.90 3.74 5.85 6.77 94.17
N 150 150 150 150 150 150
Small Mean 17.2458 3.4652 1.6493 2.7047 3.3436 44.1784
Std. 6.22503 .60726 .47592 .56244 .57284 23.31619
Deviation
Minimum 6.00 1.45 .62 1.23 1.78 19.92
Maximum 30.00 5.32 2.80 4.20 4.77 196.81
N 118 118 118 118 118 118
Total Mean 48.4282 5.2308 2.5484 4.0933 5.0008 34.6494
Std. 32.34024 1.71119 .90859 1.28603 1.61022 17.95812
Deviation
Minimum 6.00 1.45 .62 1.23 1.78 7.60
Maximum 177.00 10.50 5.92 7.84 10.55 196.81
N 362 362 362 362 362 362

SORT CASES BY Species.

SPLIT FILE SEPARATE BY Species.
ONEWAY Weight Totallengthaperwidthaperturelengthshellwidth K BY Size
/MISSING ANALYSIS
/POSTHOC=DUNCAN ALPHA(0.05).

Oneway

Species = Pilaampullacea
ANOVAa
Sum of
Squares df Mean Square F Sig.
WEIGHT (GRAMS) Between Groups 101591.514 2 50795.757 405.233 .000
Within Groups 19429.170 155 125.349
Total 121020.684 157
TOTAL LENGTH (CM) Between Groups 322.883 2 161.441 264.518 .000
Within Groups 94.600 155 .610
Total 417.483 157
APPERTURE WIDTH Between Groups 80.721 2 40.360 171.780 .000
(CM) Within Groups 36.418 155 .235
Total 117.139 157
APPERTURE LENGTH Between Groups 180.509 2 90.254 239.262 .000
(CM) Within Groups 58.469 155 .377
Total 238.978 157
SHELL WIDTH (CM) Between Groups 334.074 2 167.037 220.185 .000
Within Groups 117.586 155 .759
Total 451.660 157
Condition factor Between Groups 7410.005 2 3705.003 27.881 .000
Within Groups 20597.180 155 132.885
Total 28007.185 157
a. Species = Pilaampullacea

49
Post Hoc Tests

Multiple Comparisonsa

a. Species = Pilaampullacea

Homogeneous Subsets

WEIGHT (GRAMS)a
Duncanb,c
Subset for alpha = 0.05
Size N 1 2 3
Small 48 16.2083
Medium 60 46.8667
Big 50 80.5600
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaampullacea
b. Uses Harmonic Mean Sample Size = 52.174.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

TOTAL LENGTH (CM)a

b,c
Duncan
Subset for alpha = 0.05
Size N 1 2 3
Small 48 3.4733
Medium 60 5.0627
Big 50 7.0906
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaampullacea
b. Uses Harmonic Mean Sample Size = 52.174.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

APPERTURE WIDTH (CM)a

b,c
Duncan
Subset for alpha = 0.05
Size N 1 2 3
Small 48 1.5856
Medium 60 2.7103
Big 50 3.3830
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.

50
a. Species = Pilaampullacea
b. Uses Harmonic Mean Sample Size = 52.174.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

APPERTURE LENGTH (CM)a

b,c
Duncan
Subset for alpha = 0.05
Size N 1 2 3
Small 48 2.7225
Medium 60 4.2602
Big 50 5.4306
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaampullacea
b. Uses Harmonic Mean Sample Size = 52.174.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

SHELL WIDTH (CM)a

b,c
Duncan
Subset for alpha = 0.05
Size N 1 2 3
Small 48 3.2798
Medium 60 4.8905
Big 50 6.9586
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaampullacea
b. Uses Harmonic Mean Sample Size = 52.174.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

Condition factora
b,c
Duncan
Subset for alpha = 0.05
Size N 1 2
Big 50 24.1575
Medium 60 38.5244
Small 48 39.2937
Sig. 1.000 .734
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaampullacea
b. Uses Harmonic Mean Sample Size = 52.174.
c. The group sizes are unequal. The harmonic mean of the
group sizes is used. Type I error levels are not
guaranteed.

Species = Pilaafricana

51
 ANOVAa
Sum of
Squares df Mean Square F Sig.
WEIGHT (GRAMS) Between Groups 158782.282 2 79391.141 219.437 .000
Within Groups 51012.940 141 361.794
Total 209795.222 143
TOTAL LENGTH (CM) Between Groups 362.103 2 181.051 264.671 .000
Within Groups 96.453 141 .684
Total 458.555 143
APPERTURE WIDTH Between Groups 85.039 2 42.519 132.314 .000
(CM) Within Groups 45.311 141 .321
Total 130.349 143
APPERTURE LENGTH Between Groups 184.584 2 92.292 225.793 .000
(CM) Within Groups 57.633 141 .409
Total 242.217 143
SHELL WIDTH (CM) Between Groups 286.317 2 143.159 252.618 .000
Within Groups 79.905 141 .567
Total 366.222 143
Condition factor Between Groups 8341.796 2 4170.898 26.887 .000
Within Groups 21872.540 141 155.124
Total 30214.337 143
a. Species = Pilaafricana

Post Hoc Tests

Multiple Comparisonsa

a. Species = Pilaafricana

Homogeneous Subsets

WEIGHT (GRAMS)a
Duncanb,c
Subset for alpha = 0.05
Size N 1 2 3
Small 40 20.3250
Medium 60 46.0167
Big 44 104.1364
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaafricana
b. Uses Harmonic Mean Sample Size = 46.588.

52
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

TOTAL LENGTH (CM)a

b,c
Duncan
Subset for alpha = 0.05
Size N 1 2 3
Small 40 3.7060
Medium 60 5.3538
Big 44 7.8120
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaafricana
b. Uses Harmonic Mean Sample Size = 46.588.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

APPERTURE WIDTH (CM)a

b,c
Duncan
Subset for alpha = 0.05
Size N 1 2 3
Small 40 1.8358
Medium 60 2.5945
Big 44 3.8177
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaafricana
b. Uses Harmonic Mean Sample Size = 46.588.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

APPERTURE LENGTH (CM)a

Duncanb,c
Subset for alpha = 0.05
Size N 1 2 3
Small 40 2.9078
Medium 60 4.0978
Big 44 5.8418
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaafricana
b. Uses Harmonic Mean Sample Size = 46.588.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

SHELL WIDTH (CM)a

Duncanb,c
Subset for alpha = 0.05
Size N 1 2 3
Small 40 3.4398
Medium 60 4.9663
Big 44 7.1016

53
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaafricana
b. Uses Harmonic Mean Sample Size = 46.588.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

Condition factora
b,c
Duncan
Subset for alpha = 0.05
Size N 1 2 3
Big 44 22.3862
Medium 60 32.5897
Small 40 42.3182
Sig. 1.000 1.000 1.000
Means for groups in homogeneous subsets are displayed.
a. Species = Pilaafricana
b. Uses Harmonic Mean Sample Size = 46.588.
c. The group sizes are unequal. The harmonic mean of the group sizes
is used. Type I error levels are not guaranteed.

Species = Lanisteslibycus

ANOVAa
Sum of
Squares df Mean Square F Sig.
WEIGHT (GRAMS) Between Groups 12586.017 1 12586.017 296.965 .000
Within Groups 2458.167 58 42.382
Total 15044.183 59
TOTAL LENGTH (CM) Between Groups 75.870 1 75.870 190.479 .000
Within Groups 23.102 58 .398
Total 98.972 59
APPERTURE WIDTH Between Groups 12.504 1 12.504 67.604 .000
(CM) Within Groups 10.727 58 .185
Total 23.231 59
APPERTURE LENGTH Between Groups 60.803 1 60.803 173.826 .000
(CM) Within Groups 20.288 58 .350
Total 81.090 59
SHELL WIDTH (CM) Between Groups 69.166 1 69.166 193.299 .000
Within Groups 20.753 58 .358
Total 89.919 59
Condition factor Between Groups 9726.374 1 9726.374 12.665 .001
Within Groups 44541.463 58 767.956
Total 54267.837 59
a. Species = Lanisteslibycus

SPLIT FILE OFF.

ONEWAY Weight Totallengthaperwidthaperturelengthshellwidth K BY Species

54
 /MISSING ANALYSIS
/POSTHOC=DUNCAN ALPHA(0.05).

Oneway

ANOVA
Sum of
Squares df Mean Square F Sig.
WEIGHT (GRAMS) Between Groups 31706.543 2 15853.272 16.456 .000
Within Groups 345860.089 359 963.399
Total 377566.633 361
TOTAL LENGTH (CM) Between Groups 82.057 2 41.028 15.107 .000
Within Groups 975.010 359 2.716
Total 1057.067 361
APPERTURE WIDTH Between Groups 27.301 2 13.651 18.102 .000
(CM) Within Groups 270.719 359 .754
Total 298.020 361
APPERTURE LENGTH Between Groups 34.760 2 17.380 11.097 .000
(CM) Within Groups 562.285 359 1.566
Total 597.045 361
SHELL WIDTH (CM) Between Groups 28.202 2 14.101 5.576 .004
Within Groups 907.801 359 2.529
Total 936.002 361
Condition factor Between Groups 3930.954 2 1965.477 6.273 .002
Within Groups 112489.359 359 313.341
Total 116420.313 361

Post Hoc Tests

Homogeneous Subsets

WEIGHT (GRAMS)
Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 29.2833
Pilaampullacea 158 48.2152
Pilaafricana 144 56.6389
Sig. 1.000 .056
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.

55
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

TOTAL LENGTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 4.2555
Pilaampullacea 158 5.2216
Pilaafricana 144 5.6472
Sig. 1.000 .068
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

APPERTURE WIDTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 1.9592
Pilaampullacea 158 2.5815
Pilaafricana 144 2.7575
Sig. 1.000 .152
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

APPERTURE LENGTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 3.4123
Pilaampullacea 158 4.1634
Pilaafricana 144 4.3001
Sig. 1.000 .440
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

SHELL WIDTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 4.3910
Pilaampullacea 158 5.0556
Pilaafricana 144 5.1947
Sig. 1.000 .536
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.

56
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

Condition factor
Duncana,b
Subset for alpha = 0.05
Species N 1 2
Pilaafricana 144 32.1743
Pilaampullacea 158 34.2116
Lanisteslibycus 60 41.7423
Sig. .416 1.000
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

ANOVA
Sum of
Squares df Mean Square F Sig.
WEIGHT (GRAMS) Between Groups 31706.543 2 15853.272 16.456 .000
Within Groups 345860.089 359 963.399
Total 377566.633 361
TOTAL LENGTH (CM) Between Groups 82.057 2 41.028 15.107 .000
Within Groups 975.010 359 2.716
Total 1057.067 361
APPERTURE WIDTH Between Groups 27.301 2 13.651 18.102 .000
(CM) Within Groups 270.719 359 .754
Total 298.020 361
APPERTURE LENGTH Between Groups 34.760 2 17.380 11.097 .000
(CM) Within Groups 562.285 359 1.566
Total 597.045 361
SHELL WIDTH (CM) Between Groups 28.202 2 14.101 5.576 .004
Within Groups 907.801 359 2.529
Total 936.002 361
Condition factor Between Groups 3930.954 2 1965.477 6.273 .002
Within Groups 112489.359 359 313.341
Total 116420.313 361

Post Hoc Tests

Homogeneous Subsets

57
 WEIGHT (GRAMS)
Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 29.2833
Pilaampullacea 158 48.2152
Pilaafricana 144 56.6389
Sig. 1.000 .056
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

TOTAL LENGTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 4.2555
Pilaampullacea 158 5.2216
Pilaafricana 144 5.6472
Sig. 1.000 .068
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

APPERTURE WIDTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 1.9592
Pilaampullacea 158 2.5815
Pilaafricana 144 2.7575
Sig. 1.000 .152
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

APPERTURE LENGTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 3.4123
Pilaampullacea 158 4.1634
Pilaafricana 144 4.3001
Sig. 1.000 .440
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.

58
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

SHELL WIDTH (CM)

Duncana,b
Subset for alpha = 0.05
Species N 1 2
Lanisteslibycus 60 4.3910
Pilaampullacea 158 5.0556
Pilaafricana 144 5.1947
Sig. 1.000 .536
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

Condition factor
Duncana,b
Subset for alpha = 0.05
Species N 1 2
Pilaafricana 144 32.1743
Pilaampullacea 158 34.2116
Lanisteslibycus 60 41.7423
Sig. .416 1.000
Means for groups in homogeneous subsets are displayed.
a. Uses Harmonic Mean Sample Size = 100.200.
b. The group sizes are unequal. The harmonic mean of the group
sizes is used. Type I error levels are not guaranteed.

59