Diversity of Life (II)

Diversity of Life: The
major Animal Phyla
Module HBio 102

Reading Material
by Caston Makaka
1 Caston Makaka (in prep)

PART 1
CHAPTER 1 Introduction
CHAPTER 2 Basic biological classification and criteria
PART 2- THE PARAZOA
CHAPTER 3 Phylum Porifera (Sponges)
PART 3-THE RADIATA
CHAPTER 4 The Cnidaria
PART 4-THE BILATERIA I
CHAPTER 5 The Protostome Phyla.
PART 5-THE BILATERIA II
CHAPTER 6 The Deuterostome Phyla
PART 6-THE MOST SUCCESSFUL
CHAPTER 7 The insects: the champions of the metazoan world
CHAPTER 8 Fishes dominate the Vertebrate world.
CHAPTER 9 Birds are the present day champions of the land Tetrapods

PART A
CHAPTER 1
Introduction
This module focuses on the diversity of animal life, a subject dreaded by many students of
biology. This dread is often reflected when the same students take up positions as lecturers,
when the course is often thrown from one lecturer to another. Many years of teaching this
course has shown that one need not know all the animal groups and the often times difficult
names to be comfortable in the area. For beginners, a basic understanding of the unifying
features and differentiating characteristics will go a long way in making the student find
his/her way through the myriad of animal groups. It is our hope that this module will help
students in appreciating the fact that in the myriad of animal life forms, there are basic
unifying features, which can be used to better understand the diversity of animals.
In this module the student is introduced to the concept of biodiversity, the nature of
metazoans, basic classification; the features important in uniting and separating the major
animal groups; and lastly a survey of the major animal phyla are given. It is impossible to
delve into animal diversity without making reference to evolution. The concept of evolution
will come out as the basis for all the diversity of animals present today. Evolutionary trends
will be highlighted in the major animal groups and the evidence for this will become
apparent as some of the diagnostic features of the animals will be reviewed. The dictum
that ontogeny recapitulates phylogeny will also be explored in illustrating the similarities
and differences between animal life. It is hoped that the selection of the animal groups used
in this module will not be misconstrued by the student as prioritising groups on the basis of
the author’s areas of interests but for purposes of illustrating evolutionary patterns.
What are metazoans/Animals?
Metazoans are multicellular, motile, heterotrophic organisms with a blastula stage in their
life cycle. Although motility may not always be an exhibited feature of metazoans in all life
stages, other features are normally quite evident even to the casual observer. Being

heterotrophs, metazoans take in (ingest) ready made food through openings (usually one) in
the body wall, which in the larger forms is referred as to as a mouth. The food is digested in
the gut (in some lower forms it is engulfed by cells and digested intracellulary) and taken up
(absorbed) into the cells lining the gut, enter a circulatory system with a fluid medium
usually referred to as blood in the more advanced forms. The food is assimilated to bring
about growth and some is respired to provide energy required for various life processes.
Oxygen required for respiration is usually taken into cells either directly through simple
diffusion or it enters the body through a system of tubes referred to as the respiratory
system. Waste products of metabolism are excreted either directly to the outside in animals
whose cells are directly exposed to the outside or channelled through a system of ducts
called the excretory system. Reproduction is either asexual or sexual, with both forms being
predominant in the lower animals and the latter in the higher animals. Sexual reproduction
involves the fertilization of the female sex cell (the ovum) by the male sex cell
(spermatozoa). During early embryonic development the newly formed zygote divides into
two then four, eight, 16, 32, 64 cells (blastomeres), doubling at each cell division without an
increase in size. A ball of cells called a morulla then forms. Its inside become vacuolated and
becomes fluid-filled. This is called a blastula. All metazoans pass through this stage in their
embryonic development. For the very primitive forms, this is the terminal stage in
embryonic development and for some others, further development occurs through a
process of gastrulation and, still in others, organogenesis.
A number of theories have been put forward to explain the origins of metazoans (see
Rapport 1992) but the colonial theory seems to be the most plausible one. According to this
theory, metazoans are thought to have evolved from colonial flagellated protozoans,
probably the choanoflagellates or colonial ciliates. The individual cells in the colony are
thought to have been held together by an extracellular matrix (ECM). These protozoans,
living as a colony (Figure 1.1), are thought to have developed some interdependence which
made the colony more successful than the individual solitary forms. This colony is
envisioned to have evolved some form of differentiation, with an anterior end and a
posterior end and some cells specialised for locomotion and sensation, others for feeding
and still others for reproduction.

Figure 1.1: Origin of Metazoa: choanoflagellate colonies and metazoan evolution. A and
B, different forms of the choanoflagellate colony Proterospongia haeckeli. C, A
hypothetical premetazoan based on the choanoflagellates in A and B. D, A hypotheitical
protometazoan showing cellular speciliazation along an anterior-posterior axis. Surface
cells are bound together and in mutual contact, thus allowing physiological regulation of
the anterior of the body
The hypothetical multicellular metazoan or protometazoan is called a blastea and is similar

to the blastula, a developmental stage that all metazoans pass through in their embryonic
development today. As multicellularity established itself, the ability of the individual cells to
recognize each other as self became central to the formation of multicellular organisms.
This was made possible by the evolution of cell surface recognition receptors and receptor
sites which could recognise each other and thereby allowing for adhesion of the individual
cells to each other.
The new evolutionary invention (the blastea), taking place some 650million years ago in the
Cambrian is thought to have been highly successful and further evolution gave rise to
numerous multicellular metazoans. It is important to note, though that, not all evolutionary
lines were successful. Many blind ending lines in the fossil record are a testimony to the
millions of species that could not make it in the struggle for survival. The evolution of
animals, therefore, like the evolution of any other life forms, was not as it were, like a
guided form of branching, each with all the necessary tools for a successful lineage. We
need not emphasise the fact that all extant animal forms are a result of successful lineages
that have managed to make it to this day since some 3.5 billion years ago when the first life
forms emerged. Their existence today is however not an assurance of their continued
survival in the ever unending repertoire of life's demands. Many species are getting extinct
each day that passes by and still many more are on their way out. The African elephant
(Loxodonta africana) has been cited as one such species whose heydays are long past and is
on its way out.

The complex nature of biodiversity
Biodiversity or biological diversity is the variety of life in all its forms, levels and combinations,
including ecosystem diversity, species diversity and genetic diversity. Just as there are many
different ways to define biodiversity, there are many different measures of biodiversity. Most
measures quantify the number of traits, individuals, or species in a given area while taking into
account their degree of dissimilarity. Some measure biodiversity on a genetic level while others
measure within a single habitat or between ecosystems.
Traditionally there are two levels at which biodiversity has been described. In effect it uses
genetic diversity as a basis for valuing both species diversity (for their relative richness in
different genes) and ecosystem diversity (for the relative richness in the different processes to
which the genes ultimately contribute) (Foram Mehta, 2013)
Measuring Genetic diversity

It is a genetic diversity which causes tulips to be different colors and different heights. Typically,
researchers measure genetic diversity by counting how often certain genetic patterns occur.
Measuring biodiversity on the genetic level requires gene map and then compare them to the
genetic make-up of the larger population. Another method of measuring genetic diversity works
in the reverse by evaluating the differences in physical appearance between individuals then
attributing these traits to the most likely genetic roots. Mapping diversity at the genetic level is
currently the most accurate measure of biodiversity, although it can be costly and time
consuming and, thus, impractical for evaluating large ecosystems. It is most often used to
examine managed populations or agricultural crops which can allow for selective breeding of
the most desirable traits (Foram Mehta, 2013).
Measuring Ecosystem diversity
In its simplest form Biodiversity refers to the totality of life in its various forms. When
taxonomic diversity is the point of reference, commonly, the species is the unit used as a
surrogate for assessing this biodiversity. Total biodiversity in the biosphere may never be
known. Biologists have approached the issue of coming up with total global diversity from
two angles: use of estimates and real inventories. Estimates of total biodiversity given by
various authorities have been precipitated by the knowledge that the number of newly
discovered organisms is quite large (about 13 000 species per year); pointing to the fact that
whatever total diversity is known to science is a miniscule amount of the total in the
biosphere. Thus, estimates have been obtained by inferring total diversity from well known
groups and the simplest of these approaches to calculating biodiversity is to compare the
ratio of known to unknown faunas. In 1985, Peter Raven postulated that for birds, mammals
and other well known animals, there are roughly twice as many tropical species than
temperate climate species. If this were true of all organisms, then of the 1.5 million species

already described, and two-thirds of these being temperate zone species, the world total
would be 3 million.
It is now known that this approach is inaccurate because birds and mammals comprise only
a tiny fraction of all life forms and are not at all indicative of species such as insects, which
are often 10 times as numerous in a given tropical location as for a temperate area.
Erwin used the "bug bomb" method in which he fogged the tree canopy with insecticide and
collected thousands of dying insects as they fell into a number of big funnels placed on the
forest floor beneath. From these samples gathered over three seasons from the same spot,
tentative numbers were then available, from which Erwin drew his conclusions. He
calculated that there are 163 species of beetles living in the canopy of a single species of
tree. In turn, there are about 50,000 species of tropical trees worldwide, so this comes out
to 8,150,000 species of beetles. Then, assuming that beetles represent 40% of all arthropod
species, and finally that there are twice as many arthropods (mostly insects) in the canopy
than on the ground, came up with a rounded estimate of 30 million species for the world's
rainforests.
The total global biodiversity was extrapolated using butterfly population which are a
relatively well known group of organisms. Butterflies are easy to collect and are relatively
conspicuous and thus easy to identify and collect in any survey and were used by others to
calculate global biodiversity
There is a maximum of 17,500 species of butterflies.

At both the continental and global level, butterflies comprise 6% of the order Lepidoptera.
The order Lepidoptera comprises 12% of all insects.
Therefore, there are approximately 2,430,000 species of insects.

One more step; insects comprise approximately 67% of all living species;
In only three simple steps, total biodiversity is calculated:
1). 17,500/.06 = 291,667 species of Lepidoptera
2). 291,667/.12 = 2,430,556 species of Insects
3). 2,430,556/.67 = 3,627,695 total biodiversity
Using such estimates the total biodiversity of all living organisms is projected to be
3,628,361 species, minor rounding errors aside.
Finally, there is mere opinion and pure guesswork, such as the 13.6 million species "working
estimate" developed by a United Nations-sponsored "consensus" effort. The data used: a
survey of the world's leading experts on each group of living organisms, simply asking each
what they felt the speculative total was. In a big contrast to Erwin's pumped up estimates,
he discredits the diminutive 13.6 million total as the product of "armchair biology", because
it is not based on any real experimental data .
Until someone comes forward with the hard evidence to produce a conclusive number, the
global species count, will continue to be the subject of wild speculation.
The parallel use of species inventories utilised figures of species of the different life forms
found in the literature. Conservatively there are 1.75 million species (Hammond, 1992)

assigned to 5 kingdoms (Whittaker, 1969; Marguilis and Shartz, 1982), 64 phyla, 146 classes,
869 orders and about 7000 families (Parker, 1982) all of which are a result of evolution.
Others have put this estimate at 1.4 million (Wilson, 1992).
Rana (2005), estimates that in the animal world there are 1 032 092 animal species in 35
phyla. These estimates are claimed to have been arrived at by taking an inventory of all
animal species known to science as cited in the literature. Rana’s inventory of the major
animal groups is depicted in Table 1.1. Earlier, Brusca et al (2003), by taking an inventory of
all animal phyla presented in the literature had come up with a total of 1 335 188 animal
species.
Table 1.1: Inventory of the major animal groups (After Rana, 2005)
Animal group Total in group Number estimated

Insects and other arthropods 874 161 30m species believed to be
present most in Tropical forest.
Invertebrates (excluding arthropods) 116 873 May number millions.
Fish 19 056 Estimated at 21 000.
Reptiles and Amphibians 8 962 Known species account 95% of
known species.
Birds 9 040 Known species account 98% of

known species.
Mammals 4 000 Known species account 95% of
known species.
Total 1 032 092
The number of species in a given area is known as its species richness (S). The species being
the basic unit of classification is the most practical and commonly used currency when
referring to biodiversity. Species richness can be considered at various spatial levels; local,
regional and global. It is thus important to note that species richness is highly dependent on
the size of the area considered or sampled, as any increase in land area, results in an
increase in species richness. In fact for every 10-fold increase in area there is a doubling in
species richness. Rana’s inventory shown in the table above is at the global scale.
The number of species given by many authorities remains estimates and the completeness
and accuracy of the global animal diversity cannot be ascertained. The number of species
that still remain unknown and to be incorporated into the inventory is not known. A number
of reasons can be cited to account for this deficiency. The animal species record is not
adequate in part, because, many species have not been described. This is largely so because
of the daunting task of learning and familiarising with the already described fauna. In
addition one can only recognize the yet unidentified species if he/she knows those that have
been described.
Although the larger animal groups are relatively better known to science, the smaller ones
are hazily known. It is an acknowledged fact that there are by far fewer taxonomists for
invertebrates and particularly the Hexapoda (insects), which form the bulk of metazoans
(about 85%) than for the larger animal groups. The species inventory is also largely
inaccurate because of errors of judgement. As pointed out by Gaston and Mound (1993),
the existing inventory contains large amounts of hidden synonymy. Synonymy arises when a
single species is given two or more names as a result of being erroneously recognized by
two observers as such. Thus whilst new discoveries and descriptions tend to increase the
species inventory list, many of these are later referred to synonymy for arbitration. For
example in 1979, 2116 species of beetles were newly described, but some 426 species were
later found to have been described before. For the years 1986-1987 and 1988-1989, the
rates of synonymy in the major insect orders were of the rates of 1/3 of the number of
newly described species in those years.
Although synonymy tends to pull down the numbers (once discovered) it is important to
note that there are certain groups of animals where very little is known. A case in point is
the terrestrial arthropods. In this group, already, close to a million species are recognised,
still a lot is anticipated from this phylum. Estimates put the undescribed at between 1 and
100 million, but conservative estimates are between 2-3 and 20 million, and for insects
alone, 10 million (Wilson, 1992). A large proportion of the undescribed species are thought
to be in the tropics and especially in third world countries.
Alpha (α), beta (β) and gamma (γ) diversity
We have referred to the total number of organisms in a locality demarcated at any scale as
the species richness (S). Species richness is a common way of measuring biodiversity and
involves counting the number of individuals - or even families – within a given area. This is also
expressed as Alpha diversity (α-diversity). This can be measured by counting the number of
taxa (distinct groups of organisms) within the ecosystem. For example, in a study of avian
communities in Hwange Main Camp, Hwange National Park, Zimbabwe, the author,
together with Mazire Samantha (2014) assessed alpha diversity by considering the number
species and noted that there were a total of 41 bird species. This is the α diversity for the
area. For birds, α diversity can simply be determined by undertaking a bird count as you
traverse the area. As one moves from one locality into another undertaking the same bird
count, one will encounter, in addition to those species already encountered, other new
species. As one goes through this second locality he/she will realize that not only had he/she
encountered new species, but some species encountered in the first were not encountered
in the second. The species encountered in the second locality also constitute the α diversity
for this second locality but the total number of species peculiar to either locality is called the
β diversity. At the ecosystem-level, measures of β diversity are often used to compare two
ecosystems or to determine changes over time in a given region. Beta diversity measures the
present and changes of species diversity between ecosystems; this involves comparing the
number of taxa that are unique to each of the ecosystems. In simpler terms, it calculates the
number of species that are not the same in two different environments. The resulting number
indicates to researchers whether there is any overlap in the species found in each group. Beta
diversity is given by the formula:
β = (S1 − c) + (S2 − c)
where S1 are number of species found in the first locality (community),
S2 are number of species found in the second locality (community) and C is
the number of species found in both communities.

High values of beta diversity indicate high dissimilarity and low beta diversity indicates high
similarity between communities or localities. Table 1.2 below shows α, β, and and another
form of diversity called γ diversity (we shall to come to this shortly) as one passes through
three different habitat types; Grassland, Miombo woodland and Acacia woodland. The
highest β diversity (7) is between the Miombo woodlands and the Acacia woodland
indicating highest dissimilarity between these two communities
Table 1.2: Hypothetical species, alpha (α), beta (β) and gamma (γ) diversities for three
communities.
Diversity Grassland Miombo Acacia woodland

woodland
A A
B B
C C
D D
E E E
F F F
G G
H H H
I
J
K
α 8 6 8
β 4 (i.e. species A, B,G and K)
7 (i.e. A,B,C,D,G I, J and K)
4 ( i.e. C, D, I and J
γ 11 (i.e. species A,B,C,D,E,F,G,H,I,J, and K
Beta (β) diversity is more often represented by the similarities and not differences between
samples or communities. Two major indices of similarities are commonly used. These are
Jacquards’ coefficient of similarity and Soerensen similarity ( ) index. QS is given by the
following formula:
Where QS is the quotient of similarity;

S1 and S2 are the number of species in samples (communities) S1 and S2; and
C is the number of species shared by the two samples (communities).
Using Soerensen similarity index, the similarities between the Grassland and Miombo, the
Grassland and Acacia woodland, Miombo woodland and Acacia woodland would be 0.71, 0.75
and 0.43 respectively.

The total number of species found throughout the region is called the gamma (γ) diversity.
Gamma diversity is thus the species richness for a region with many localities or communities.
Gamma diversity for a region with two communities can be expressed in terms of the species
richness of the communities as follows:
γ = S1 + S2 – c
Where, S = the total number of species recorded in the first locality (community)
1
S = the total number of species recorded in the second locality (community)

2
c= the number of species common to both communities.

Thus gamma diversity for a region covering the Grassland and the Miombo woodland would be
8+6-5=9.
Diversity indices
Although many authors are contended with the use of the concept of species richness as a
good representation of diversity, some advocate for the use of some other indices that factor
in species abundance. One such index is the Shannon Wiener index of diversity. The Shannon
Wiener diversity index ( ) is calculated using the following formula:
Where is the proportion of individuals belonging to the ith species in the data set.
Using this index, a community with relatively high proportions of each species will have a
high diversity index indicating high diversity in the species than one in which a few species
are predominant. This is illustrated by the two communities shown below each with a
species richness of 3 but different Shannon diversity indices.
Figure 1.2: Two communities with the same species richness (S=3) but having different
Shannon diversity indices, being higher in the second community due to equal
representation.

Now if we go back to our three hypothetical communities and assume that the total
numbers of each individual species for the three communities were also assessed (Table1.3),
the H' for the first community, the Grassland community, will be calculated as follows:
H'=-∑ (185/438In234/438) + (87/438In87/438) + (54/438In54/438) + (44/438In44/438) +

(35/438In35/438)+ (25/438In25/438) + (7/438In6/438) + (1/438In2/438)
= -∑ -0.364-0.321-0.258-0.231-0.202-0.163-0.066-0.025 = 1.63
The Shannon diversity indices for the other two communities have also been calculated and
are shown in the Table 1.3 below.
Table 1.3: Hypothetical species, numbers and diversities for three communities
Diversity Grassland Inividuals Miombo Individuals Acacia Individuals

woodland woodland
A 185 A 55
B 87 B 58
C 54 C 345
D 44 D 102
E 35 E 88 E 51
F 25 F 54 F 61
G 7 G 55
H 1 H 36 H 49
I 51
J 58
K 10
Total 8 438 6 671 8 438
H' 1.63 1.32 2.10
λ 0.251 0.313 0.296
If we make comparisons amongst the three communities with regards to total individuals,
the Miombo woodland, though having the least species richess (S=6), has the highest total
population of 671. On the other hand, although having the highest population, it has the
lowest Shannon diversity index (H'=1.32) because of the uneveness of species populations in
this community. A comparison of Grassland and Acacia communinities with the same
species richness (S=8) and the same total populations (438), diversity, as expressed by the
Shannon diversity index, is higher (H'=2.10 in the Accacia woodland because of the near
equality of the species populations as compared to the Grassland community (H'=1.63).
Shannon’s equitability (E H') can be calculated by dividing H' by H'max (here H'max =InS)
H'max is the maximum diversity of a community (plot/site/sample) when all species are
equally abundant and this is equivalent to In S. If there is complete evenness of species
abundances in the grassland community then each species would be represented by 438/8
or 54.75 individuals. Then
H'max=-∑ (54.75/438In54.75/438) + (54.75/438In54.75/438) + (54.75/438In54.75/438) +
(54.75/438In54.75/438) + (54.75/438In54.75/438)+ (54.75/438In54.75/438) +
(54.75/438In54.75/438) + (54.75/438In54.75/438) =2.079
And dividing H' by H'max we have 1.63/2.079= 0.78
And since H'max = InS we can still arrive at the same answer by the formula:
For the grassland E H'=1.63/In8 =1.63/2.079=0,78
Equitability (evenness) assumes a value between 0 and 1 with 1 being complete evenness.
Another index which takes into consideration the abundances of species is the Simpson
diversity ( ) index. It measures the probability that two individuals randomly selected from a
sample will belong to the same species. Simpson’s diversity index is then obtained by summing
up these probabilities.
The Simpson diversity index is calculated from the formula:
Where is the proportion of individuals belonging to

the ith species in the data set.
In everyday calculations that involve real or finite populations, the following formula is used
λ=
n = the total number of organisms of a particular species

N = the total number of organisms of all species
For the Grassland community the Simpson diversity index would be calculated as follows
λ =∑n (n-1)/N (N-1)
=185(184)/438(437) +87(86)/438(437) +54(53)/438(437) +44(43)/438(437) +35(34)/4389437)
+25(24)/438(437) +7(6)/438(437) +1(0)/438(437)
=0.178+0.039+0.0149+0.0099+0.0062+0.00313+0.000156+0.0
=0.251
The Simpson diversity index is also sometimes called the dominance index on the basis that
the contribution of the dorminant species in the community overrides the rest and those
with very few individuals contributing very little if any. Notice the very high contribution
made by species A (0.178) in the overall figure and the zero (0.0) contribution made by
species H. The Simpson diversity indices for the three hypothetical communities are shown
in Table 1.3 above. The Miombo woodland has the highest Simpson diversity index because
of the predominace of a few species (species C) in the community.

The dormance of a particular species is effectively highlighted by calculating the dorminance
index (d). Dominance (d): expresses the proportional importance of the most abundant
species and is calculated as:
For the Grassland community d is given by

185/438 100 = 42,
And for the Miombo and Acaccia woodland they are 51 and 14 respectively. The dorminant
species in these communities are A, C and F respectively.
The dominance of a single species is also visually evident when we consider the rank
abundance graph for a community which conforms to the quite common phenomenon
observed for many communities where there is usually a single dominant species with other
species contributing fewer and fewer individuals. The rank abundances of avian Orders
found in Main Camp Hwange is in consonance with this pattern (Fig 1.3).

Figure 1.3: Rank-abundance graph of bird Orders according to number of individuals in Main
Camp, Hwange National Park, Zimbabwe (Source: Makaka Caston and Mazire S, 2014, in
press).
Rarity or commonness of individual species (taxon) can be evaluated by measuring the

frequency of occurrence (%). This is determined by dividing the number of plots
(sites/commuinites) where a particular species (taxon) was present by the total number of
plots and multiplying by 100. If we consider our three communities (Grassland, Miombo
woodland and Accacia woodland), species E, F, and H show a 100% frequency of occurrence,
and species A, B, C and D a 50% frequency whilst species I, J, and K exhibit a 33% frequency
of occurrence. Species can be ranked as to whether they are rare, uncommon, common or
supper-common.
In a study of bird species in a national park in Ghana, Wiafe et al (2010) set the relative
status for each species based on frequency of occurrence as follows:
• Super-common: species occurring within more than 50% of the census plots/sites
• Common: species found in between 20–49% of the census plots/sites
• Uncommon: species occurring within between 11–19% of the census plots

• Rare: species found in between 1–10% of the census plots/sites.
Weighted diversity
Some biologists are of the opinion that diversity based on purely richness and abundance
figures does not capture the real diversity and value of species and commuinties. In essence
they maintain that species are not equal. Some are more equal than others. To this end they
advocate for some way of placing value to those more valuable species when coming up
with diversity indices. Of note are those that advocate for the incorporation of a weighting
factor in the diversity index with the result that a community with many important species
(keystone species or rarer species) is considered richer than one with a fewer such species.
Still others place more weight on diversity at higher taxonomic level with the result that if
two communities have the same number of species, the one having the species from
different genera or families is considered richer than the one with all the species belonging
to the same genus or family.
The placing of more weight on taxa that do not have close relatives is also considered
worthwhile. The tautara lizards (Sphenodon) of New Zealand has sometimes been equated
to all the other reptilian groups (totalling 6 000 species) simply on the basis that it is quite
isolated in terms of kinship. This genus has remained quite unmodified and has given rise to
very few species ever since the age of reptiles, Mesozoic era, some 65 to 248million years
ago.
If more weight is placed on species which are contributing very little to rapid diversification
(speciation), what value should be placed on those that are really contributing immensely to
this divergence? This was a question that others thought about seriously and concluded
that instead of giving credit where it is not due, the rapidly diversifying taxa should be given
more weight in indices of diversity. However the placing of some form of weighting is
sometimes quite prudent when priorities of communities to be conserved are being made.
Otherwise for all other purposes of gaining insights into the diversity of species, species
richness is considered suffice. It is also important to note that for all other indices, the
starting point is to know the different forms and species and the taxa to which they belong.
We will begin with that aspect in this module.
Functional diversity
Before we leave measures and assesssements of diversity let us consider another form of
diversity which, like weighted diversity, is imported from an ecological vview point. This is
functional diversity. Functional diversity considers the varieties of organism from a
functional point of view. An ecosystem which contains primary and secondary consumers
only is less diverse than one which contains primary producers, primary consumers and
secondary consumers which in turn is less rich than another which contains primary
producers, primary consumers, secondary consumers and tertiary consumers. In the same
line of thinking, an aquatic system which contains shredders only is less diverse than one
containing shredders and scrappers and detritus feeders.

Is biodiversity evenly distributed? – Hotspots of
biodiversity.
Biodiversity is not evenly distributed. There are some areas with high numbers of species
and these are recognized at global, regional and local scales. A total of 34 areas of high
biodiversity have been recognized the world over, the major ones are depicted in Fig 1.4.
Fig 1.4: Biodiversity hotspots (Source: http://www.bbc.co.uk/2/science/nature)
Mittermeir and Werner (1990) introduced the term megadiversity centres to refer to those
areas where biodiversity is comparatively very high. Using this concept, a number of
regions/countries have been classified as megadiversity centres. These are Brazil, Zaire,
Madagascar, Columbia, Eucador, Peru, China, India, Malaysia, Indonesia, and Australia.
In a 1999 analysis, published in the book Hotspots: Earth’s Biologically Richest and Most
Endangered Terrestrial Ecoregions, and a year later in the scientific journal Nature (Myers,
et al., 2000), 25 biodiversity hotspots were identified. Collectively, these areas held as
endemics no less than 35 percent of terrestrial vertebrates in an area that formerly covered
only 11.8 percent of the planet’s land surface. The habitat extent of this land area had been
reduced by 87.8 percent of its original extent, such that this wealth of biodiversity was

restricted to only 1.4 percent of Earth’s land surface (http://www.conservation.org/- date
24-03-11).
Biodiversity gradients
A closer look at the geographical positions of the megadiversity centres/countries proposed
by Mittermneir and Werner (1990) shows that most of them are located along or within the
tropics and very few if any from the temperate regions. This leads us to the concept of
biodiversity gradients. The one already alluded to here is the tropical- temperate
biodiversity gradient. Generally, there are more species within the tropics and fewer in the
temperate regions, thus there is a gradual decrease from the tropics to the temperate
regions. This has partly been attributed to the high primary productivity in tropical plants as
a result of the high sunlight intensities and rainfall regimes. This high primary productivity in
turn supports a rich fauna, which, with increased competition, leads to rapid speciation and
hence diversity. Another gradient which has been noted is altitudinal. It has been noted that
in some cases as altitude increases species richness decreases.This has been attributed to, in
part, to a decrease in temperature and oxygen concentrations with altitude.
In the major aquatic environments such as the seas and oceans, biodiversity, apart from
showing the latitudinal gradient shown by terrestrial life also shows variation with depth.
Most life forms occur in the top layers of waters which constitute the photosynthetic zone
because of high light intensities and sufficient dissolved oxygen. With increase in depths the
amount of light penetrating to these depths markedly decreases and life forms also
decrease as only carnivorous and suspension or detritus feeders become the only thriving
life forms.
Endemism and biodiversity

Endemics are organisms that occupy limited geographical ranges, i.e. have limited extent of
occurrence and also have limited areas of occupation within their geographical range. At a
global scale examples of endemics that quickly come to mind are the African Antelopes (e.g.
bushbucks, water bucks, kudus etc), the African elephant (Loxodonta africana) and the
Tsetse flies (Glossina spp) which are endemic to Africa and the tuatara (Sphenodon) which is
endemic to New Zealand. At a local scale (country scale), good examples are the butterfly
Amauris echeria which is endemic to the Chirinda and Vhumba Forest, Zimbabwe, and the
butterfly Acraea igola which is endemic to the Chirinda Forest in Mt Selinda, Zimbabwe
(Cooper 1991). The Black maize beetle (BMB) Heteronychus licus is endemic to the South-
east low-veld of Zimbabwe where it a notorious pest in sugarcane plantations (Cackett,
1994 and Makaka, 2008). The Killifish Nothbranchius furzeri is endemic to the Gonarezhou
National Park, Zimbabwe (Minshull and Bill-cross, 1988).
Endemism is often found to be associated with certain biological features such as gigantism,
dwarfism, and low levels of self incompatibility, greater tendency towards asexual
reproduction, lower overall reproductive effort and poor dispersal abilities than widespread
species in the same area.

Knowledge of endemic species may help greatly in conservation efforts as those species that
are known to be found in restricted areas may require greater conservation because once
they are eliminated in their areas they cannot be replaced form anywhere else.

CHAPTER 2
Basic biological classification

and criteria
As alluded to in the preceding chapter, the daunting task of learning and understanding the
already described species impedes progress in making a comprehensive inventory of animal
species. Realising this impediment, humankind, from time immemorial has tried to lessen
the problem by way of devising methods of making inferences and retrieval of information
about the animal groups much easier. One great effort to do this was to classify the
organisms. Humankind has recognized the importance of classifying things ever since he
developed the mental capabilities that characterises him as human. The earliest forms of
classification were rather rudimentary and mainly based on the utility of the organism.
Organisms were thus classified as to whether they were of value to humans or not;
medicinal versus non-medicinal; edible versus non-edible etc. With the passage of time, as
biology (then natural history) evolved and gradually separated from philosophy, more
scientific criteria gained precedence over earlier criteria. With this, morphological and
anatomical features become commonly used to group and hence classify organisms. Thus
classification criteria based on shared morphological and anatomical features become
prominent.
To date, classification in biology, apart from using the same morphological and anatomical
features has moved on to look at embryonic features, histology, cytology, ecological,
behavioural and now very commonly, biochemistry. Organisms hitherto grouped together
using the old criteria are being re-examined using these new methods, in some cases
confirming the earlier relationships and in some cases (a lot of them) unveiling completely
new kinships. As these new revelations come to light, new phylogenetic trees are being
reconstructed.
The importance of classification need not be over-emphasized. In everyday life we see how
life is made much easier by simply packing grocery items of similar use in the same shelves
in supermarkets, and books of same disciplines in the same shelves in libraries. Classification
in biology serves a number of functions: it simplifies the study of organisms by grouping
organisms sharing a number of features in common; it helps make inferences; and finally it
shows phylogenetic relations between organisms. The importance of making inferences
dawned to this author when he was asked a question on the reproductive biology of
chameleons as an undergraduate by a high school student who had a very high regard for

my knowledge in biology. I want to profess that by that time I had not read anything about
chameleons per se but upon a quick mental storming I realized that the object of the
question posed was a reptile and I avoided the potentially embarrassing moment by simply
narrating reptilian reproductive biology. Thus, not only can we infer on the morphological
and anatomical characteristics, but on the biology as well.
Whilst different schools of classification have immerged, they all seem, in the end to point
to some form of relationships between the organisms concerned. The phenetic school of
classification emphasizes the importance of taking into consideration as many shared
features as possible and working out where the strongest relationships lay by finding those
that share the greatest number of features in common. This has been made easy by the use
of computers that can handle large quanties of data. The Phylogenetic school on the other
hand argues that not all features are of evolutionary importance and not all of them should
be lumped together to find evolutionary relationships- only those that are of some
evolutionary importance should be considered, Not doing so, they argue, would result in
finding relationships in distantly related species showing convergent evolution. Whilst in
cladistic classification, emphasis is put on the branching patterns. (See Mayr, 1981 for a
review of these schools of thought).
Categories used in Biological Classification.

The science which deals with biological classification is called taxonomy. Taxonomy is
branch of biology which deals with describing, naming and classification of organisms on the
basis of shared characteristics. Sytematics goes on further to deal with relationships
between organisms and to derive thereof phylogenetic relations. Classification of organisms
therefore inherently shows relationships with regards to common ancestry.
The simplest unit or category of classification, as earlier indicated, is the species. So many
definitions have been muted as to what a species is and the definition given here is a
composite of all these definitions. It defines a species as a group of organisms of common
ancestry, sharing many characteristics in common, occupying the same habitat and niche,
and capable of interbreeding to produce viable offspring. This can be regarded as an all
encompassing definition put together to try and pin down the very elusive concept of a
species. Using one part of the definition may not be adequate as experience has shown. For
example defining a species as a group of organisms capable of interbreeding (the biological
species concept), though, applicable to many species may not be adequate in others. Slight
variations in behaviour patterns and biological clocks may guard against interbreeding in a
group of organisms of the same species temporarily separated by a barrier for a
considerable length of time.
Two or more species may seem quite related but do not satisfy the criteria to be classified
as a single species. These are grouped together in bigger grouping (category) referred to as
a Genus. Related genera constitute a Family and closely related Families form an Order.
Related Orders make a Class which in turn constitute a Phylum, and ultimately a Kingdom.
Often times an even higher category, Super kingdom, is included at the top of the hierarchy
(Figure 2.1). This classification is called Natural Biological Classification and it envisions
natural relations of decreasing affinities as we go up the ladder. By grouping a number of

organisms into a single species are implied stronger relationships than those uniting species
into a single genus. It also follows that a genus is made up of more closely related members
than a family, and so on. The entire system of classification, based on interpretation and
gudgement, thus aims to express the degrees of relationship between various organisms. All
categories above the species level are referred to as the higher categories. This distinguish
them from the species group category.
Figure 2.1: Categories used in the hierarch of biological classification
Very often some intermediate categories are also included in the hierarchy and these tend
to complicate the whole hierarchy. These include subspecies, strains and varieties to refer
to subtypes of the same species and super species to refer to closely related species within a
genus. Other categories include sub- and super genus, sub- and super family, sub- and super
order, sub- and super class, sub- and super phylum etc. For our purposes here we restrict
ourselves to the use of the larger categories in the hierarchy.
As we have already alluded to, in biological classification, animals are grouped basing on
shared characteristics.The actual animal group that is placed in each category is called the
taxon. Thus the taxon Echinodermata is placed at the hierarchical level corresponding to the
category phylum: Echinodermata is the taxon; phylum is category. If we consider the
classification of humans the following categories and taxons are used.
Category Taxon
Phylum Chordata
Subphylum Vertebrata (Craniata)
Class Mammalia
Order Primate

Family Homidae
Genus Homo
Species Homo sapiens (Carolus Linneaus, 1758)
By convention the species name is followed by the authors name and a year, the person
who first described the organism Homo sapiens and gave it its name, and the year the
description was publised. In the case of humans it was Carl Linneaus in the year 1758.
The hierarchical classification of organisms is characterised by two inherent features. The

first aspect of this hierarchical classification is that as we go up the hierarchy, each category
becomes more and more inclusive. In other words the number of taxa in each successive
higher rank (category) increases as we move up the hierarchy. Thus, whilst, extant
humankind is the only representative in the species Homo sapiens; in the genus Homo are
included our closely related forms (fossil ancestors) like Homo erectus, Homo habilis etc and
in the family Hominidae the list expands to include the distantly related forms like the
Australopithecines and Pithecanthropus. In the order Primate the list even expands to
include monkeys, gibbons, orangtangs, baboons, macaques, bush babies, lemurs, tarsiers,
lorrisies etc. The Mammalia, which is next in the hierarchy, is quite a large grouping
including all those animals bearing hairs/furs, mammary glands and being endothermic. The
mammals are the present day dominant vertebrate group in the terrestrial environment,
coming second only to birds. Included here are diverse groups ranging from the very small
moles (Insectivores) to the very large elephants (Proboscidae). Others like the sea cows
(Sirenia) and whales (Cetacea) have gone back to the waters. In the highest grouping,
Animalia, all animals are included. Table 2.2 shows the classification of selected animals,
highlighting the more and more inclusive nature of the hierarchical classification. It is
important to note here that the only real tangible units of this classification are the species,
and every other higher order grouping combines varying numbers of species of varying
degrees of similarity.
The second feature is that the number of shared (unifying) characteristics become fewer as
we go up the hierarchy. Thus, whilst all extant humans (Homo sapiens) share many
characteristics in common {e.g. sweat glands and pores on almost entire skin surface, very
short fine hair giving a naked look, a prominent chin, receding jaw and brow bones, cranial
capacity of around 1600cm3 , complex brain with high intelligence, large brain to mass ratio,
fingers with nails, prehensile hand with opposable thumb capable of both precision and
power grip and highly manipulative with intricate co-ordination of hands with brains, ,
upright walking with striding gait, ability to communicate using speech and symbols, ability
to use and make tools, unusually extended parental care for offspring, a complex social
system based on sharing and cooperation, purposefulness, the ability to build in the brain,
to scheme and plot with a vision of the outcome (creativity) and many others too many to
exhaust here}, the number of features we share with closely related forms in the genus
Homo are considerably fewer, more so in the family Hominidae that we belong together
with our distant fossil relatives like the Australopithecines and Pithecanthropus This is
markedly so in the Order primate and up the hierarchy.

Table 2.2: The more inclusive nature of the hierarchy of biological
Classification
Kingdom Animalia Animalia Animalia Animalia Animalia Animalia Animalia Animalia
Phylum Chordata Chordata Chordata Chordata Chordata Chordata Chordata Arthropo
da
Subphylu Vertebrat Veretbrat Vertebrat Vertebrat Vertebrat Vertebrat Vertebrat Pterygota
m a a a a a a a
Class Mammali Mammali Mammali Mammali Mammali Mammali Mammali Insecta
a a a a a a a
Order Carnivora Primate Primate Primate Primate Primate Canivora Hymenop
tera
Family Canidae Cercopith Pongidae Hominida Hominda Cercopith Fellidae Apidae
ecidae e e edidae
Genus Canis Cercopith pan Homo Homo Popio Panthera Apis
ecus
Species familiaris manis troglodyt erectus sapiens arsinus leo mellifera
es
Common Common Blue Common Java/Beiji Humanki Chackma lion Honey
name dog monkey or robust ng man nd Baboon bee
chimpanz
ee
Role of the International Code on

Zoological Nomenclature (ICZN)
Systematists have devised codes of biological nomenclature which are slightly different for
different major groups of organisms and all these codes provide a mechanism for publishing
a new name for a newly recognized taxon, for fixing a name to a particular organism by
citing a type and arbitrating between synonyms where a taxon has accidentally been named
more than once, or where two taxa have been united into one (Biodiversity Assessment).
The type of a species name is a particular cited specimen in a particular collection
(museum), the type specimen.
How a species is defined will have bearing on the overall number of organisms described by
man. As we have already noted various species concepts have been put forward to try and
define what a species is for the various forms of life and varying reproductive and biological
traits. But whatever definition is used to pin down what a species is, a set of rules have to be
followed upon recognition of a new species so that it becomes known to science. The
International Code on Zoological Nomenclature (ICZN) is a set of laid down rules,
recommendations and procedures to be followed by all zoologists to make new species
names available to science. The International Code on Zoological Nomenclature was
adopted by the International Commission on Zoological Nomenclature in 1901 and declared

1st January 1758 as the first operational day of the Code. In 1961 the ICZN was authorized
by the International Congress of Zoology and printed for publication.
Having discovered a new species, and, having satisfied oneself that it is indeed a new
species through consultation of available literature, or comparing it to related types in a
collection, a taxonomist has to follow, among other things the following: He/she will have to
give it a name following the binomial nomenclature. This is a system of naming a species
first proposed and used by Carl von Linne, the great naturalist of the 18th century. Carl
himself was able to describe and name more than 12 400 species (8 000 plant species and 4
400 animals including Homo sapiens) during his lifetime and thus is often credited as the
father of taxonomy. Binomial nomenclature involves giving the species a two name name-
binomen (meaning two names). The first name is the genus name and the second is the
specific name. The specific name should always be in Latin or be Latinized by adding a suffix
which makes it sound Latin. To emphasize this aspect, the originator of this system, Carl
himself, Latinized his own name, thus he become known as Carolus von Linnaeus. Thus,
following this rule, the familiar canine (familiar dog) is called Canis familliaris. The first name
being the generic name and the second name is the specific name (trivial name). The genus
name is the genus to which the species belong. The genus name should always be initialled
with a capital letter and the specific name with a small letter. In writing, the whole species
name should always be underlined and in printing it should always be italicized. Thus for
humankind and the migratory locusts the names are cited as Homo sapiens and Locusta
migratoria, respectively. The subfamily name is formed by adding the suffix -inae to the
stem of the genus name and the family is coined by adding the suffix -idea. Thus the
subfamily and family to which humans belong are Hominae and Hominidae respectively. The
superfamily standard ending is –oidea.
During the times of Carl, it is important to mention that each species used to have a long
name (a polynomial) which was descriptive in nature. Those who read “Hard Times” by
Charles Dickens would recall the Victorian era definition given by young Tom when asked to
define a horse in class by his teacher, Mr Mchokmchild. He gave a long definition of a horse
sounding like “.....a quadrupled, graminivorous, nonruminant...” That indeed was a
polynomial. Such was the nature of the names that they were mouthfilling and cumbersome
in saying them out and writing, and, in one of his treaties or essays, Carl decided to use
short hand to denote each species. It is this short hand, because it had two names (a genus
name and an epithet or specific name) that was later to be adopted and become the usual
way of referring to a species.
To make the species name available to science the author should publish it. For a name to
be said to have been published, the author should write an article and make it accessible in
large enough quantities to be distributed to other stalk holders (biologists and in particular
zoologists). Today, this usually involves publishing in a scientific journal. The article must
detail the morphology and/ anatomy, and if needs be , even the biochemistry, histology and
other aspects of the new find, highlighting the aspects that are significantly different from
other closely related species and hence warranting it to deserve being placed in its own
taxon. The description can be in the form of diagrams, descriptive statements, pictures and
sometimes acoustic recordings, behaviour patterns or any such evidence that will be of help
in separating it from other closely related forms. The location the new species was found,

the nature of habitat and method/s used in its eventual discovery or capture should also be
presented. This description should ideally be accompanied by the deposition of the
specimen (a voucher) in a collection, usually a national/county or university museum. This
becomes the type specimen which becomes available for all concerned to see and to be
used in future for purposes of reference. It is important to note that almost all zoological
species have type specimens kept somewhere in a collection. The only species without a
type specimen kept anywhere in the world is humankind himself! Perhaps this is due to
moral and ethical issues, and of course having a single type specimen for all the human
races would raise a lot or racial issues – which race would have a representative as a type
specimen anyway?
Although the process of identifying new species and fixing of names appear meticulous and
thus not prone to error, many errors of judgement do occur and when they are eventually
discovered, are brought to a panel of Zoologists who arbitrate on such issues. Most cases
have to do with synonymy and homonymy. As we have already pointed out, synonymy
occurs when a species has accidentally been named more than once. The effect of this is to
inflate the species list and hence species richness. This may occur as a result of two authors
observing two developmental stages of the same organism and hence naming the two
morphological stages as different species. Once this type of error is discovered, whether by
one of the authors or by a third party, it is brought to the attention of the international
Panel of Zoologists for arbitration. In such cases, the senior name has priority over the junior
name. Thus, although both names are available to science, the senior name is considered
the valid name for the species and the other is discarded completely. The senior name is the
name which was given by the first author, i.e. the one who described and named it earliest.
In some cases the junior name may not necessarily be thrown away as in cases where the
junior name, because of popularity arising otherwise from extensive use or the stature of
the author. Discarding such a popularly known and used name may not do justice to many
who were very familiar with the species as associated with the name. In such cases,
although the senior name has priority over the junior name, the latter is retained and is
usually included after the senior name but in parenthis.
Homonymy occurs when two taxa have been united into one. Arbitration here involves
separating these into two taxa of which they really are. In such cases the taxon which was
described first retains the name and the second is given another name
As alluded to earlier in this chapter, although many species are described and named, a
significantly large proportion is later referred for arbitration because of hidden synonymy
and homonymy.
Classification Criteria.
In an introductory book of this nature our classification would rather remain at the coarse or
higher levels of classification such as Phyla (and in some cases Classes). Classification at such
higher ranks (i.e. class and phylum) should basically be broad based. That is, the features
used should be all inclusive. These features are highly conserved over generations. As
classification cascades down to lower and lower ranks more exact features become useful to
separate closely related forms. Such classification becomes based on less conserved
features, characteristics controlled by more or highly mutable genes. The criteria considered
here is thus at very high levels and highly conserved features will be used. Most of such
highly conserved features are controlled by highly conserved genes called hox genes.Thus,
for example, genes which regulate growth rates and patterns (hox genes) also determine
the symmetry of the resultant animal. These appear to be highly conserved genes and are
represented quite extensively in the animal world as their effects permeate across wide
groups and determine underlining unifying features. This is ease to grasp when we note that
despite the differences in form, the phenomena of bilateral or radial symmetry transcends
across phyla. Despite chasms of differences certain underlying developmental patterns
separate and unite animal groups. It is such highly conserved features which we will use for
the purposes of separating and unifying the varied metazoan groups.
Although modern relationships in taxonomy are now largely based on molecular features
and biochemistry, these, to a large extent serve to confirm or dispel previously held kinships
(especially at species level) and we will make reference to them as they crop up.
Classification based on cell number

We have already alluded to the metazoans as multicellular animals. This distinguishes them
from any other animal-like unicellular protozoans. In addition to being multicellular
metazoans are diploid. As depicted in the Figure 2.2 below, the protozoans are separated
from the other metazoans in that they are unicellular and the rest multicellular. In
protozoans, the single cell is the whole organism, which, though less complex than whole
metazoan organisms, is more complex when compared to a single metazoan cell by having
all the apparati for feeding, locomotion, excretion and sensation, all housed in a single
organism (cell).
Figure 2.2: Multicellularity as criterion for classification

Classification based on levels of organisation
The level of organisation exhibited by an animal forms a major basis for classification.
Animals exhibit three major levels of organisation; the tissue level of organisation, the organ
level and the organ system. Originating from a single eukaryotic cell, the metazoan cells
evolved into multicellular organisms as already pointed out. It is now widely accepted that
the primitive multicellular animals evolved through various lines, with specialisation of cells
to form tissues then organs and organ systems. Each of these levels has representatives in
which they are presently highly conserved. The Porifera, though having cells specialized for
particular functions, the specialized cells are scattered all over the body and many biologists
still contest their tissue status. Campbell aptly described the condition in sponges as not
being at the tissue level of organisation but “lose federations”. On this basis the Porifera are
separated from the others (Figure 2.3 below). The rest of the higher animals are
characterised by tissues, organs and organ systems.
Figure 2.3: Multicellularity and level of organisation as criteria for classification
Classification based on body symmetry.

Metazoans may also be classified according to their general body form. Most animals exhibit
one of two major types of body symmetry; radial and bilateral. At the extreme left are the
irregular forms without any symmetry. An animal is said to have some form of symmetry if,
when cut through the centre, the two halves produced are mirror images of each other.
Radially symmetrical animals are those whose external can be divided into two
approximately mirror images of each other by many incisions that pass through the centre
of the organism. These include some Ctenophora, and Cnidaria. These are generally refered
to as the Radiata. They have similar parts similarly arranged about a central axis. Radially

symmetrical animals can further be classified into biradially, tetramerously, pentamerously
and hexamerously symmetrical depending on whether they can be divided into two equal
halves through two, four, five or six planes on a single axis. Radial symmetry is very often
associated with sessility and hence the need to receive environmental stimulation from all
sides equally. Radially symmetrical animals therefore tend to have a diffuse nervous system.
Even in the motile Cnidarians the nervous system still retains the diffuse architecture.
Bilaterally symmetrical animals posses right and left sides that are approximate mirror
images of each other. In such animals, there is usually an anterior and posterior end, and
sense organs, to a more or less extent concentrated in the anterior portion of the animal.
This is referred to as cephalisation. This allows the animal, because it moves with the
anterior portion forwards, to be able to explore the new environment it is moving into
before the whole body moves on. The concentration of nervous tissues and sense organs in
the anterior portion is in consonance with the principle of neurobiotaxis which insists that a
concentration of nervous material evolves in the area of maximum environmental
stimulation (Parker and Haswell, 2003).
Examples of bilaterally symmetrical animals include the Platyhelminthes, Nematoda,

Annellida, Arthropoda and Vertebrata. These are commonly called the Bilateria.
Bilateral symmetry has become synonymous with animals to the extent that many people
fail to appreciate any other form of symmetry in the animal world, and this is one reason
why many people fail to recognize animals with other forms of symmetry like the sponges,
cnidarians and echinoderms as metazoans in pictures and videos on TV sets.
From an evolutionary view point, bilateral animals are thought to be more evolved than
radially symmetrical animals. Using this line of thinking the echinoderms (like the sea
urchins) would be placed very close to the Cnidarians, for having radial symmetry, however
studies have shown that the former are even more advanced to warrant being placed
ahead of even such groups like the nematodes and annelids. Investigations have shown
that the embryo of these animals show typical bilateral symmetry, implying that the adult
pentaradial symmetry is a secondary evolutionary development.
It would appear that the genes for these two major types of symmetry (radial and bilateral),
evolved quite early in metazoan life and have tended to be passed down through evolution
without any modifications in these two major metazoan lines.
Using the classification based on symmetry we can add another notch onto our classification
based on levels of organisation depicted in figure 2.3. This is shown in figure 2.4. below.

Figure 2.4: Classification based on multicellularity, level of organisation and body
symmetry
Classification based on body plans
Metazoans have for long time been divided into groups based upon the number of germ
layers formed during embryogenesis. These germ layers are groups of cells that behave as
distinct units during the early stages of embryonic development and give rise to distinctly
different tissues and /or organ systems in the adult. Thus metazoans can be classified as
either diploblastic, or triploblastic. Diploblastic (diplo=double) animals have only two
distinct layers following gastrulation (movement of cells into the interior of the embryo).
The outer layer is called the ectoderm and the inner layer is the endoderm. Examples of
diploblastic animals are the Cnidaria and Ctenophora.
Triploblastic (Triplo- triple) animals, which are the majority of metazoans have three germ
layers; endoderm, mesoderm and ectoderm. The mesoderm arises from either the
endodermal tissues or from a line of cells that form quite early in embryonic development
through incorporation of mesodermalizing factors in the early dividing zygote. The
mesoderm is thus a middle layer between the endoderm and ectoderm.

Further classification of triploblastic animals can be done using the criterion of the presence
or absence of an internal body cavity independent of the gut and hence not connected to
the outside (Fig 2.5 below). This internal body cavity is referred to as a coelom and is
entirely surrounded by mesodermal tissue. Those triploblastic animals devoid of a coelom
are said to be acoelmate. In these animals, characteristically, the region between the
ectoderm and the endoderm is filled by the proliferation of mesodermal tissues and there is
no trace of an internal body cavity. The Platyhelminthes are such animals. In the second
group of triploblastic animals, the mesoderm does not proliferate to fill the whole space
between the outer body wall and the gut, and a cavity remains, which is in fact a persistent
blastocoel (a cavity which develops in the embryo during the blastula stage prior to
gastrulation). This type of internal body cavity is called a pseudocoelom (false coelom) and
animals having this condition are called Pseudocoelomates. Some authors have coined the
name blastoceolom to refer to the pseudoceolom since there is nothing false about it. Good
examples of pseudoceolmates (blastoceolmates) are Nematodes and Rotifers. In the third
group of triploblastic animals, there is an internal body cavity which develops embryonically
within the mesoderm and is completely surrounded by mesodermal tissue. These animals
are said to have a true coelom and thus referred to as Coelomates or Eucoelomates, The
Annelids, Molluscs Echinoderms, Arthropods and Chordates are examples of animals
bearing such a condition. Thus adding another two notches to our scheme of classification
we have the scheme shown in figure 2.6 below. Figure 2.7 depicts an evolutionary tree
constructed on the bases of the criteria discussed this far.
Figure 2.5: The body plans of metazoans based on the presence and/nature of coelom.
Source:http://image.wistatutor.com/content/animal-kingdom/body-cavity-
conditions.jpeg

Figure 2.6: Scheme of classification based on multicellularity, level of organisation, body
symmetry and presence and nature body cavity

Figure 2.7: Schematic diagram showing the positions of the major animal groups in the
evolutionary tree
Classification based on early embryonic

development: The
protostome/deuterostomes dichotomy

We live in a world where evolutionary theory is the reigning paradigm in biology to the
extent of being an unquestioned fact to many biologists. That organisms evolved, and are
still evolving, has become an underlying theme in biology. And as Watson (2007) aptly puts
it in his forward to the book: Darwin, the Indelible Stamp, “Evolution is Law (with several
components) only disputed by those who choose to ignore the evidence, put their common
sense on hold, and believe that unchanging knowledge and wisdom can be reached only by
revelation”. He goes on to say “Darwin’s insights remain as profound today as it was 150
years ago and continues to shape the way we think about, not only the living world and our
place in it, but about existence”.
One of the pieces of evidence for evolution first put forward by biologists were the
similarities between embryos of different animal groups which was used as part of the
evidence of common ancestry and popularised in the dictum 'ontogeny recapitulates
phylogeny'. According to Heackel who proposed this theory, animals, during their
embryonic development (ontogeny) retrace the morphologies and anotomies of their past
ancestral adults (phylogeny). In other words, in their embryonic development, animals pass
through the stages resembling their past ancestral forms. The exact consequence of this is
that goups of animals whose embryoes pass through a similar stage in their embryonic
devepments are deemed to have an ancestor of that form and hence the same ancestor at
some distant past. If the embryonic resemblences are transient, and, in the initial stages of
embryonic development, then they shared the same ancestor in the very long distant past
but if they are drawn out in time right up to the very late stages of development then they
are deemed to have had the same ancestor in the near distant past. Hence the more drawn
out the resemblances, the more related the groups are and Vis versa.
Although this dictum has fallen out of favour with some in the biological sciences it gives
some plausible explanations to the many relationships between the metazoans. The fact
that all metazoans pass through a blastula stage early in their embryonic development is not
a disputed fact. It is reasonable to suggest that this convergence is not by mere coincidence
and this definitely points to common ancestry in the distant past. That, later embryonic
developmental stages do vary to varying degrees in the major groups is also an uncontested
fact. It is rational to suggest that this points to later divergence of the lineages in the groups
with the passage of time.
It is usually assumed that similarity among early embryonic stages is the result of
evolutionary conservation (Seidel, 1960; Sander, 1983; Buss, 1987; Raff, 1996; and Hall,
1996 cited in Galis et al 2002). Even though von Baer (1828) did not believe in evolution by
descent, he was the first to propose that early changes in ontogeny were more likely to have
cascading consequences than later changes when most of development has already taken
place (Galis et al., 2002). The early stages of cleavage and gastrulation are often remarkably
similar among metazoans (Gilbert and Raunio, 1997). The succeeding stage, organogenesis,
is far more diverse among metazoans than cleavage and gastrulation (Gilbert and Raunio,
1997). This, as already alluded to, points to a common ancestry and subsequent divergence.
It is important to emphasize that although the similarity of cleavage and gastrulation is
largely due to conserved similarity, it is also, to an important extent due to convergence. For
a book at this level we are going to dwell on the former and students interested in the later
twist can quench their thirst in the works of Galis and Sinervo (2002).

A survey of the general trends in early metazoan development is quite revealing on the
major unifying aspects of this seemingly highly heterogeneous group of animals. This survey
brings to light the fact that distantly related groups have fewer and transient similarities
with each other while more closely related ones tend to have extensive and more drawn out
similarities in their embryonic development. Thus whilst groups like the Porifera have
transient similarities with the rest of the metazoans as seen in the blastula stage, slightly
higher groups like the cnidarians have even longer drawn out similarities up to the gastrula
stage, producing a diploblastic embryo. Consistent with this, are the even greater
similarities shared by the Bilateria, having many similarities in later embryonic stages but
also showing a strong divergence into two major clades- the protostomes and
deuterostomes.
As will become clear later, these two groups of bilaterans show marked divergence in
developmental patterns but strong similarities within each of the two groups. They have
marked differences in cleavage patterns, the stage at which cell fate is determined, the
nature of gastrulation, the fate of the blastopore and origins of the mouth or anus, origins
of the third germ layer (mesoderm) and hence coelom formation and the type of larvae
which finally develop from the embryo. It is important to emphasize at this juncture that
although one set of developmental patterns are predominantly characteristic of one group
of animals, they nonetheless are not a preserve for that animal group only. Exceptions do
occur here and there and this is the nature with many aspects in biology. This will be quite
evident when we consider the developmental stages. In summary it is suffice to say that
protostomes are characterised by the following: spiral cleavage of blastomeres, determinate
development- cell fate being determined quite early in embryonic development with the
result that twinning is not possible; mouth forming from the blastopore opening and anus
developing elsewhere.; the mesoderm derived from a mesodermal cell whose origins are
quite early in embryonic development, schizocoelic coelom formation- a coelom which
develops from the splitting of mesodermal tissue; and finally the embryo developing into a
trocophore larva. In addition to these characteristics there is a trend in the major
protostomes phyla for the presence of ventral nerve cord and a dorsal heart as shown in the
Annelid-Mollusc-Arthropod Line.
On the other hand, deuterostomes predominantly exhibit the following: radial cleavage of
blastomeres; indeterminate development -fate of cell being determined quite late in
embryonic development (at least after a few cell divisions) and hence making production of
twins quite possible in this group; gastrulation by invagination; the blastopore forming the
anus and mouth forming elsewhere; mesoderm forming through outpocketing of endoderm
and thus coelom forming through pinching off of coalesced, spherical, fluid-filled
endodermal tissue- a process called enterocoely. In addition to these characteristics there is
a trend in the major deuterostomes phyla for the presence of dorsal nerve cord and a
ventral heart as shown in the Echinoderm-Chordate Line. For us to grasp the differences
between these two groups of Bilateria we need to explore their early embryonic
development. Now, here, we make a detour and explore the nature of early embryonic
development in the Bilateria.

Early stages of embryonic development
We have already alluded to the fact that metazoans are diploid, thus meiosis is restricted to
the formation of gametes. At fertilization, which may either be internal or external, the
diploid condition is restored once again. The so formed zygote is a polarized cell with
distinctly different poles, one referred to as the animal pole and the other the vegetal pole.
With development the zygote divides into two, four then eight, 16, 32, 64, etc, as the
number of cells double after each cell division cycle. The result is a ball of cells called a
morula. This ball of cell becomes vacuolated to form a fluid filled blastula (Figure 2.8). The
fluid-filled cavity is called a blastocoel and is typically surrounded by a single layer of cells.
Certain aspects of the early cleavage patterns are determined by the amount and
distribution of yolk, while other features are determined by the genetic programming of
that particular organism. The first two divisions of the zygote have planes running parallel to
the polar axis of the cell and the cleavage leads to 2 then 4 cells (blastomeres). Because this
cleavage is complete (cleavage planes pass completely through the cell, producing
blastomeres that are completely separated from one another by thin membranes), it is
referred to as holoblastic cleavage (Fig 2.8 A, B). Holobastic cleavage occurs in isolecithal
(evenly distributed) and weakly to moderately telocithal eggs ova. In some cases cleavage is
incomplete. This is known as incomplete cleavage or meroblastic cleavage. Meroblastic
cleavage occurs whenever very large amounts of yolk are present (as in strongly telolecithal
eggs). In this type of cleavage, cleavage planes do not pass readily through the dense yolk,
so the blastomeres are not fully separated from one another by cell membranes (Fig 2.8 C).

Figure 2.8: Summary of stages involved in formation of blastula (blastocyst): metazoan
eggs, early cleavage stages in relation to yoke content, and blastulae. A, A small
microlecithal egg with a uniform distribution of yolk (showing equal cleavage). B, A medium
size mesolecithal egg with yolk concentrated in vegetal half (showing unequal cleavage). C,
A large macrolecithal egg with only a small apical blastodisc free of yolk (Source:
http://www.paleo.pan.pl/people/Dzik/Publications/origins.pdf)
In holoblastic cleavage, after the second cleavage the amount and distribution of yolk in the
egg determines the nature of further cleavage patterns resulting in two patterns i.e.
complete and equal cleavage and complete but unequal cleavage.
Complete and equal cleavage is characteristic of eggs with relatively small amounts of yolk
that is also evenly distributed. Such eggs are also referred to as homolecithal or isolecithal
eggs. Cleavage in such eggs results in blastomeres above and below the plane of cleavage
being of equal size because of the uniform distribution of yolk. Continued repeated divisions
result in a sphere of small blastomeres, the blastula. In such cases a single layer of
blastomeres constitute the blastula with the fluid filled blastocoel (Fig 2.8 A).
The other type of cleavage, that is, complete but unequal cleavage occurs when there are
moderate amounts of yolk restricted to the vegetal hemisphere (telelecithal eggs). In such
eggs division occurs in such a way that the blastomeres at the animal hemisphere are
relatively smaller than those in the vegetal hemisphere (Figure 2.8 B). These are called

micromeres and macromeres respectively. Generous amounts of yolk in the vegetal
hemisphere can cause the blastocoel to be pushed to the vegetal half or to be completely
filled so that a solid sphere results. This is called a stereoblastula.
It is important to note, at this point, that the least evolved of the Metazoa, the Porifera (also
called Parazoa= besides the animals) show complete cleavage and the larvae are usually at
the blastula stage of development. In this group, further development occurs well after the
larva has escaped from the parent. The majority of sponges possess a parenchymaula larva,
in which monociliated cells occur on the outer surface, save for the posterior pole (Figure
2.9). This is an example of stereoblastula as it is composed of solid mass of cells. In some
porifera however, the blastula is said to be a coeloblastula. Further larval development will
involve the turning of the larva inside out so that the ciliated cells which were once outside
are on the inside -the typical sponge condition.
Figure 2.9. Diagram of a parenchymula larva of a sponge.
In the true Metazoans (Eumetazoa) embryonic development proceeds from a blastula

through gastrulation. These are a series of stages which converts the blastula from
spherical hollow or solid blastula into a bilayered embryo or gastrula, bearing the symmetry
of the adult. Like cleavage pattern, the gastrulation pattern is significantly influenced by the
levels of yolk. It usually occurs through three major patterns; invagination, epiboly and
ingression.
Invagination involves cells of the vegetal half of the blastula infolding into the interior
forming a new cavity called an archenteron. This is also sometimes referred to as a primitive
gut. The blastocoel surrounds this new cavity which opens to the outside by an opening
called a blastopore (Fig 2.10).

Figure 2.10. Progression of events during invagination. (Source: The Science of Biology,
Purves et al, 1998 (http://biology.kenyon.edu/courses)
In the second type of gastrulation, i.e. epiboly, cells of the animal hemisphere of the blastula
overgrow those on the vegetal half. (Figure 2.11). This is a characteristic typical of
protostomes.
Figure 2.11; Gastrulating by epiboly in the zebrafish embryo, Note the enveloping layer

(EVL) spreading to enclose the yolk synctyial layer (YSL). Source:
http//en.wikepedia.org/wiki/Epiboly
In ingression, cells of the blastula wall multiply and fill the blasocoel (Figure 2.12).
Figure 2.12: Progression of events during ingression.(Source:

http://biology.kenyon.edu/courses/biol114/Chap14/Chapter_14A.html)
Cleavage patterns and early development in Bilaterals
The description we have just given is greatly simplified and devoid of the details that
distinguish protostomes from deuterostomes. In Bilateria the late embryo (postgastrula) is
bilaterally symmetrical but is usually preceded by radial symmetry, indicating the original
metazoan pattern of symmetry. The initial radial symmetry results from one of two
predominant cleavage patterns: radial and spiral (Fig 2.13). On the basis of the type of
cleavage and other features, the Bilateria are classified as either Protostomes or
Deuterostomes.
Radial Cleavage.
Radial cleavage occurs in Deuterostomes. In this type of cleavage, cleavage planes are
oriented parallel or at right angles to the polar axis of the egg. The tiers (layers) of
blastomeres arrange themselves radially around the polar axis, and corresponding
blastomeres of different tiers are located directly above and below each other (Figure 2.13).
Spiral cleavage.
This occurs in a group of organisms referred to as protostomes. The cleavage planes are
oriented obliquely to the polar axis of the egg. Successive tiers of blastomeres arrange
themselves radially about the polar axis, but corresponding blastomeres of different tiers
are offset with respect to those above and below. This results in a blastomere in an upper
tier resting in the furrows between two blastomeres in the next lower tier (Figure 2.13).
After gastrulation the radial symmetry is replaced by bilateral symmetry.

Figure 2.13. Radial and Spiral cleavage patterns (Source:
http://forums.studentdoctor.net/showthread.php?t=647250)
Determinate and indeterminate development.

As the embryo develops, at some stage the embryo’s cell fate is determined. Whatever a
cell is going to be is rigidly cast so that a whole organism is produced. In some embryos the
fate of the individual cells is determined quite early in embryonic development and in some,
at a later stage. The stage at which this occurs is largely dependent on whether the
organism is protosome or deuterostome.
Determinate (mosaic) development

This is predominant in embryos showing spiral cleavage, that is protostomes in general
(Annelids, Molluscs etc). In these embryos cell fate is established early in development (at
the first cell division) and results mainly from cell factors that are unevenly distributed in the
uncleaved egg (zygote). By the sixth division the ancestors of all the three germ layers can
be recognized. The fate of every cell in the embryo is determined and for many species is
known. Endoderm develops from the macromeres and three micromeres. Mesoderm
develops entirely from the 4d micromere (=mesentoblast) and ectoderm develops from the
remaining micromeres. The development of the mesoderm is thought to be due to a yet
unidentified factor which segregates into one of the cells (the 4d or mesentoblast) and
causes it to differentiate and produce mesodermal tissue. It is the inheritance of this
mesodermalizing factor, early in embryonic development, that will result in all the cells
derived from it forming mesodermal tissue (Details of mesoderm formation will be looked at
later). The cells derived from the first division are therefore not totipotent. Any lose of cells
at this early stage will result in an embryo lacking certain tissues and hence failing to
develop further. It is for this reason that production of twins is not possible in protostomes.
Indeterminate (regulated) development

This is the predominant form of development in deuterostomes (animals that exhibit) radial
cleavage). Cell fates are determined much later in embryonic development (at least after
the first few cell divisions). A network of cellular communication determines what each cell
will eventually develop into and thus the position of the cell in the network is crucial in
determining what will eventually become of it. All cell that are formed early in embryonic
development have all the instructions to develop into full embryos and hence adults. It is for
this reason that twinning is possible in humans (we are deuterostomes of cause).
Further development and fate of gastrula

After blastula formation, gastrulation proceeds, resulting first in the formation of a
bilayered embryo, the gastrula. We have noted that the gastrula which is formed has an
opening which leads to the outside called the blastopore. The blastopore will form the
mouth or anus depending on whether the embryo is of a protostome or deuterostomes. In
protostomes the blastopore will form the mouth and the anus forming elsewhere and in
deuterostomes it will form the anus and the mouth forming elsewhere. In the higher
Metazoa the archenteron will form the gut which will connect to the outside through the
mouth and anus. Following gastrulation a third germ layer will form making the embryo
three layered. Thus all bilateral embryos are Triploblastic, composed of the original
endoderm and ectoderm and in between, a third layer, the mesoderm. Each of the germ
layers will differentiate to give adult epithelium. The ectoderm will give rise to the
epidermis; the mesoderm will give rise to the mesothelium and the endoderm will develop
into the gastrodermis. We shall look at the functions of these three germ layers but first, the
formation of the third germ layer, the mesoderm
Mesoderm formation
The mesoderm has either of two origins: endodermal or from the mesentoblast (Figure
2.14). Its origins from the former is characterised by the evagination of the endodermal wall
into the blastoceol of the gastrula. As the embryo develops the evaginated pouches pinch
off from the endoderm and proliferate in the blastocoel forming mesodermal tissue which
either completely fills the blastocoel or partially filling it (Figure 2.14). Such mesodermal
formation occurs in deuterostomes (excluding the vertebrates). Mesoderm formation from
the mesentoblast involves, first the division of the mesentoblast into teloblast cells which
occupy some ventral-lateral positions in the blastocoel. These in turn will proliferate to give
mesodermal tissue which may or may not entirely fill the blastocoel. This type of mesoderm
formation occurs in Protostomes.
Coelom formation
In animals where the mesoderm is formed from the out pocketing of the endoderm
(deuterostomes), the coelom is a consequence of this process, as the pinched off spherical
mesodermal tissue becomes fluid-filled upon formation (Fig 2.14). This is referred to as
enterocoelous coelom formation and the coelom is sometimes referred to as an enterocoel.
On the other hand, where the mesoderm is derived from the proliferation of the
mesentoblast (protostomes), a slit may occur in it resulting in the formation of a coelomic
cavity. This process is called schizoceoly and the coelom so formed is called a schizocoel
(Figure 2.14). As we have already pointed out, bilaterans bearing a coelom are called
coelomates or eucoelomates. Fig 2.14 below summarises develpmental patterns after

gastrulation and Table 2.1 summarises differences protostomes and deuterostomes. We
shall look at the broader picture of the kinships in metazoans but for now let us pursue and
see what later become of the three germ layers in the adult triploblastic metazoan.
Figure 2.14: Developmental patterns after gastrulation in protostomes and

deuterostomes

Table 2.1: Differences between protostomes and deuterostomes
Characteristic Protostomes Deuterostomes

Type of cleavage Spiral Radial
Type of development Determinate Indeterminate
Fate of blastopore Forms mouth Forms anus
Origins of mesoderm mesontoblast endoderm
Coelom formation schizocoely enteroceoly
Position of nerve cord if ventral dorsal
present
Position of heart if present Dorsal ventral
Examples Platyheminthes Echinodermata
Nematodes Chordates
Anellida
Mollusca
Arthropoda
Functions of the three germ layers and coelomic cavity

In the adult, the ectoderm forms the epidermis. The epidermis covers the body and thus
constitutes the skin. It is responsible for the secretion of the exoskeleton -the shell in
gastropods, the cuticle in arthropods, the scales in fish and reptiles, and other derivatives
like enamel of teeth, hairs, fur, claws, nails horns and antlers in mammals. It is also
responsible for formation of nerve tissue, glandular cells, and absorptive cells. The
mesoderm lines the coelom, forms internal organs, forms the musculature that moves body
organs; it forms septa, gonads and gametes, connective tissues and blood capsules, and
mesenteries that hold the gut. The endoderm forms the lining of the gut and contains
ciliated, secretory, absorptive and storage cells. The coelomic cavity within the mesoderm
houses the internal organs like excretory, reproductive and circulatory systems; functions as
a hydrostatic skeleton and allows for movement in animals like annelids and nematodes. It
also allows for independent movement of the gut and the body wall. Were it not of this
shock absorber, any slight movement of food in the body would result in the distension of
the body wall.
Classification based on metameric segmentation

Another feature which has become of major biological importance is the phenomenon of
metameric segmentation. This has occurred parallel in both protostomes and
deuterostomes. Segmentation is characterised by a sequential repetition of body segments
along the anterior- posterior axis of the animal's body (Figure 2.15.). This phenomenon, in
its primitive form, shows the faithful duplication of segments and the associated body
organs and systems. The earthworm body plan shows this quite well. In these animals there
is typically a prostomium anteriorly and a pygdium posteriorly and a chain of serially
repeated segments separated by thin peritoneum called septa. Typically in each segment,
the coelom persists and is provided with an excretory organ, the metanephridia, opening
into a nephrostome, segmental longitudinal and circular muscles and pairs of parapodia and

or cheata protruding from the body surface. Although the alimentary canal and nervous
system are continuous, the latter also becomes segmental, in the sense that the ventral
Figure 2.15 Metameric segmentation in earthworms
nerve cord becomes knotted with ganglia in each segment from which segmental nerves
enervate segmental viscera and body wall.
Metameric segmentation allows for specialization as a number of body segments become

functionally united into single unit- a condition called tagmatization. This is pronounced in
the insectan arthropods where we see a number of segments become functionally united
into head, thorax and abdomen (Figure 2.16). The head becomes a region concerned with
feeding and sensation, the thorax becoming specialized in locomotion and the abdomen
specializing in digestion and reproduction. In crustaceans and chelicerates the body is
divided into cephalothoraxes (prosoma) and abdomen (opsithosoma). In the Mryiapoda the
body is divided into head and trunk. Tagmatization has also been accompanied by fusion of
segments in these functional units, making them more compact and even more specialized,
and, at the same time, hiding the underlying segmentation. The primitive insectan head,
thorax and abdomen for example, are composed of six, three and up to 11 segments
respectively.
Figure 2.16 Tagmatism in Hexapod arthropods (Source:

http://threes.com/index.php?option=com_content&view=article&id=1887:ant&catid=7
3:nature&Itemid=51)

Tagmatization is quite evident in Arthropoda as already alluded to, as well as in more
advanced phyla like the Hemichordates and Chordates. In advanced Chordates, the
Vertebrates the body is divided into head, trunk and tail. The segmental vertebrae column
in this group shows the hidden underlying segmentation.
The occurrence of segmentation (Figure 2.17) in highly divergent groups like the arthropods
(protostomes) and chordates (deuterostomes) points to the evolution of this feature more
than once in the animal kingdom and serves to illustrate the highly adaptive nature of this
phenomenon.
Figure 2.17: Relationships between the Bilateria, note the occurrence of segmentation in
both protostomes and deuterostomes
Separation based on jointed body appendages

Co-evolving together with segmentation was the trait of jointed body appendages. This
phenomenon took root in the arthropods and it is from this trait, as we will see later, that
the name Arthropoda was inspired (arthros- jointed, poda- leg). Jointed appendages also
evolved in Vertebrates, a subplum of the Chordates. The evolution of the jointed
appendages was a milestone achievement and allowed the bearer to use the limbs to gather
food as well as get traction on the substratum and allowing for locomotion (Fig 2.18). As
Raven et al (2009) points out, it is difficult to imagine locomotion in humans without hips,
knees, elbows and wrists. These serve as the hallmark with regards to appendage
segmentation in the animal world. Jointed appendages are thus found in both arthropods
and chordates and although not quite obvious in the latter, they serve to allow for the
agility characteristic of both groups and separate them from other animal phyla. Appendage
segmentation, like body segmentation, thus occurred more than once in the animal world

(in arthropods and Vertebrates). It is thus possible that this convergent evolution was
necessitated by the same circumstances of having to move on land.
Figure 2.18: Jointed appendages in Arthropods and Vertebrates (Modified from

http://www.nvo.com/jin/homepage8/)
Separation based on internal skeleton.

Within the chordate group emerged one major feature which was to distinguish this group
from the rest of the other groups. This will also be highlighted when we consider the
broader picture of metazoan kinships. In the chordates an internal skeleton evolved to
support the body so that it doesn’t collapse under its own weight or upon contraction of
body muscles. This feature is called a notochord. This is a rod like structure dorsal to the gut
running from anterior to posterior end. It is composed of fluid-filled vacuolated cells
enclosed within a sheath that acts as a hydrostatic skeleton. In one chordate group (the
Vertebrata); there emerged a vertebral column to replace the notochord. The vertebral
column is made of a series of vertebra units (Figure 2.19), and it is from this structure that
the the name Vertebrata was inspired. This structure serves for muscle attachment and to
support the animal as it moves in water and more so on land where there is no buoying
effect of water as found in the aquatic environment. The vertebral column is largely made of
bone but in the primitive vertebrates it is composed of cartilage, a relatively weaker
material than bone. Anteriorly, the vertebrae column is constructed around the brain to
form a cranium to protect the later from damage. The presence of the cranium inspires the
name Craniata, the other name for the vertebrates. In these animals, the brain is relatively
enlarged and allows the bearers considerable powers of learning from past experiences.

Figure 2.19: The vertebrate vertebral column (Source:
https://www.google.co.zw/search?q=Vertebral+column+of+rabbit&biw=1366&bih=551
)
The Broader picture of kinships in

metazoans.
The link between the first and higher metazoans
One major subject for zoology students is to try to bridge the gap between the simplest
multicellular and the higher multicellular animals. A question which usually arises is- which
one of the simplest metazoan gave rise to the higher forms of metazoans? Up to this stage,
we have tended to imply that there is linear evolution from the unicellular choanoflagellates
to the sponges and giving off cnidarians and Platyhelminthes and the rest of the higher
forms. Biologists are however still not in agreement as to the true transition from the
simplest multicellular to higher forms. Thus, debate is still present as to whether the
sponges gave rise to the other multicellular metazoa. What seem to be clear is that sponges
could have been a very successful, blind-ending experiment, which could not be modified
any further to bring about further novelties (Barnes et al., 1993). The lack of any form of
coordination between cells, the total lack of symmetry and any form of tissue organisation
has led many biologists to exclude them in the list of possible contenders for the title of
higher metazoan forbearer. Although the presence of flagellated cells in their bodies which
are also ubiquitous in the higher metazoans, (in the form of sperm cells) had favoured their
candidacy, the fossil record says otherwise. It has been observed that fossils of hard shells
predate the fossils of sponges (Barnes et al., 1993). This, now, as it stands, exclude the
sponges as the exact progenitors of the other higher multicellular organisms. This leaves the
Cnidarians and Platyhelminthes as the contenders for that title. Is it possible that the
diploblastic condition of the cnidarians, later gave rise to a third germ layer which is

ubiquitous in the Metazoa or it is the triploblastic Platyhelminthes that gave rise to the
other Metazoa?
Some authors are of the opinion that the cnidarians, bearing a diploblastic organisation and
having no organ at all to show, seem too distant to be capable of giving rise to metazoans
with organs and organ systems. Fossil evidence also disqualifies them on the same grounds
that saw the sponges falling by the wayside. This leaves the triploblastic Platyhelminthes in
the race. These, have ciliated bodies and Willimer (1990) and Barnes et al (1993) favour the
possibility of these worms being the progenitors of the rest of the Metazoa. These, they
argue, because they have bodies covered by cilia, they could probably have arose from a
ciliated protozoan, probably a Cilliophora. They contend that these Cilliophora could have
become colonial, giving rise to a primitive Platyhelminthes of the same ancestry with the
Acoela. Thus if the sponges are ruled out of the line of contenders of higher metazoan
ancestry, then choanoflagellates also fall out as the possible progenitors of the primitive
multicellular animals themselves.
The Acoela, though now significantly diverged from the progenitor, are still close to the
ancestral stalk with their non-permanent synctyial gut and lack of bilateral symmetry and a
loose association of cells. From the ancestral Acoela-like form evolved the other worms (like
the nematodes, Rotifers, kinorhynks etc), the annelid ancestors, the lophophorates and the
deuterostomes. Thus instead of envisioning a phylogenetic tree as has been the tradition,
they see the kinships as akin to a field of grass.
The broader picture

Having considered the criteria used in unifying and separating the major animal groups, and
having revisited the link between the first metazoans and higher forms, it is prudent, at this
stage, to consider the broader picture. Considering the whole Metazoan group, the Porifera
are taken as an odd group on account of lacking true tissues and hence placed separately
as the Parazoa (=besides the animals) and the rest, the Eumetazoa. The latter is separated
into the Radiata, those with radial symmetry (e.g. Cnidaria and Ctenophora), and Bilateria,
those with bilateral symmetry. The Bilateria are essentially triploblastic and constitute the
rest of the animals (Platyhelminthes, Nematoda, Rotifer, Mollusca, Annelida, Arthropoda,
Echinodermata and Chordata, and many others). The Bilateria are also divided into
acoelomate, pseudocoelomates and coelomates (eucoelomates) (Figure 1.5) depending on
whether they have a solid body, i.e. without any internal body cavity (coelom); or whether
the embryonic blastocoels persist and occupy the space between the gut and mesoderm or
there is a cavity completely enclosed within the mesoderm. Using this classification, the
Platyhelminthes fall under the acoelomate, the nematodes and associated groups like the
rotifers constitute the pseudocoelomates and the Annelida, molluscs, arthropods,
echinoderms and chordates forming the eucoelomates.
The Coelomates are further divided into Protostomes and Deuterostomes. The Protostomes
(= mouth first) as we have seen are characterized by a mouth which develops from the
blastopore, spiral determinate cleavage, a stereoblastula and hence gastrulation through
epiboly, mesoderm forming from a 4d blastomere and schizocoelic coelom formation. This
group is also further divided into those that undergo moulting (ecdysis), the Ecdysozoa
(Arthropods) and those that do not, the Lophophotracha (Annelida and Mollusca).

The deuterostomes on the other hand are characterized by radial and indeterminate
cleavage, an anus which forms from the blastopore, a mesoderm which forms from the out-
pocketing of the endoderm and hence derived through enterocoely. The deuterostomes
include such seemingly distantly related animals like the starfish (Echinodermata) and man
(Chordata). Figure 2.20 shows the Phylogenetic tree of the major metazoan phyla. Note that
segmentation has occurred twice and independently (in the Protostomes and
deuterostomes).
So far, our construction of the phylogenetic relationships has been based on morphological
and anatomical features. The use of genetic methods has recently been employed to
analyse the relationships. Small subunit rRNA (SSU rRNA or 18S rRNA) phylogenies generally
have supported the Protostome–Deuterostome (P/D) branch point although they have
suggested reassignment of some taxa from one of these clades to the other, (e.g.,
pogonophorans not shown here) have been reassigned from Deuterostomia to Protostomia
(Valentine, 1997). SSU rRNA data have also confirmed the fact that the Protostomes may
comprise two major branches the Ecdysozoa, including arthropods and the Lophotrochozoa,
including annelids. Aschelminths appear to be a polyphyletic assemblage of Protostomes,
some of which are more closely allied to Ecdysozoa with others allied to the
Lophotrochozoa. Moreover, some molecular data suggest that at least some of the
acoelomate flatworms are lophotrochozoan Protostomes (Valentine, 1997).
The invertebrate/vertebrate dichotomy

Having looked at the kinships in the metazoan group, students of zoology are often
confronted with another classification criterion which, in essence, is purely subjective. This
groups the metazoans into two groups, the Invertebrates and Vertebrates. This is a
classification, which on the one hand employs real biological traits as references for
categorising into a single group, and on the other hand disregards any such unifying trait in
coming up with the other grouping. Thus whilst the vertebrates are a collection of all
animals with a vertebral column, the invertebrates are a bunched up collection of all the
animal phyla whose members lack a vertebral column. The vertebrates are thus a very small
group unified by a tangible biological trait and thus form a real taxonomic group as opposed
to the invertebrates whose relationships are as varied as the number of groups involved.
There are 35 animal phyla, of which 34 are purely invertebrate and the 35th phylum contains
two invertebrate subphyla and one vertebrate subphylum, the Vertebrata. According to
Brusca at al (2003), of the 1 335 188 estimated described species of Animalia, 1 288 518
(96%) are invertebrates.
The Vertebrata are thus a subphylum in the Phylum Chordata and thus constitute a very
small proportion of the metazoans (4%) (Fig 2.21). As we have already alluded to, the
vertebral column is a firm supporting girdle lying dorsally inside the vertebrate's body. It is
composed by a series of ring-like structures (either bone or cartilage) linked together in a
chain made firm by articulating protuberances anterior and posterior to each unit. Thus,
although the vertebrates are in essence a biological grouping, the invertebrates are a loose
collection of heterogeneous animal forms, embracing the primitive sponges (phylum
Porifera) to the advanced arthropods, echinoderms and the urochordates and
cephalochordates. The latter two groups, like the vertebrates, are also chordates, and, to
differentiate them from Vertebrates, they are called protochordates or Invertebrate
Chordates.
Fig 2.21: Proportion of Invertebrates and Vertebrates

The classification of metazoans into Invertebrates and Vertebrates has been cited by some
biologists as strong evidence for humankinds' strong bias towards life forms that are closer
to him- humankind are Vertebrata. A lot is known about the Vertebrates and very little is
known about the invertebrates, which, ironically, constitutes the bulk (96%) of the
metazoans.
Criticism of the present classification

The criteria for separating the Metazoa into different groups, have received a thorough
bashing from critics who insist that separating metazoans into groups on the basis of a few
'key features' is not consistent with present thought in biology. They insist that the
presence, of say, nematocysts in the cnidarians for example, cannot be used as a basis for
their separation into their own phyla because such key features may develop when
organisms of different phylogenies are faced with the same challenge in their environment.
Thus they point out that certain protoctists having faced the same predicament in their
evolution also evolved nematocysts. They also point out the presence of muscle tissue
which 'innervates' nerve chords in the nematodes as not being a typical nematode feature
as these also occur in certain cephalochordates. In this regard, Barnes et al (1993) insists
that the presence of a metameric segmentation, for example should not result in
relationship being attached between different groups bearing the same as this could have
been a result of convergent evolution.
The theory of 'ontogeny recapitulating phylogeny’ as mooted by Haeckel has also had its
fare share of criticism. But here, we insist that these had formed a rational basis for
relationships and the relationships being established today using other criteria like
biochemistry together with cladistics are not resulting in the complete over hall or
rewriting of the kinships, but slight modifications here and there as we have seen in the
sequence analysis of the 18S r RNA which has resulted in very little alteration in the overall
picture in the Protostomes/ deuterostomes dichotomy. As a beginning we thus advocate
for their use, and, as the student gets his/her feet firm in the subject, can then learn the
excerptions. We thus serve to bridge the gap.
What has been the picture like through

time – a summary?
It is important to note that by the turn of the Cambrian (545million years ago) all the major
animal phyla (Porifera, Cnidaria, Platyhelminthes, Nematoda, Annelida, Mollusca,
Echinodermata, Arthropoda, and Chordata) and all the others had evolved. Although no
definite explanation has been given for this sudden boom of life, it is thought to have been a
result of the evolution of novel characteristics that opened up new adaptive zones. The
emergence of such characteristics like, the shell of the mollusc, the cuticle and wings of
insects and the endoskeleton of chordates among others had such profound impact on the

competition and other interactions between organisms and the result was a complete
rewriting of the rules of interaction. These led to intense diversification sometimes
popularized as the Cambrian explosion. The Cambrian explosion resulted in the
macroevolution (evolution of higher taxa) that has never been repeated again in the history
of life on this planet.
Although the great extinction events of the late Permian (250 million years ago) and the
Late Mesozoic (some 150 million years ago) resulted in massive extermination of animal life,
the diversification which were to follow these events, though marked, were not anything
comparable to those of the Cambrian explosion. These two extinction events resulted in the
wiping off of about 90% and 50% of all aquatic and land Families respectively, but the
rebound of life as the remaining species filled in the gaps left out by the victims were not
anything near the Cambrian explosion. The latter was just of great seismic proportions.
Thus, although new families and orders evolved after these extinction events, there was no
evolution at the phylum level. It would thus appear that there have been no new
evolutionary inventions to alter on a grand scale the fortunes of the Metazoa as did
happened in the Cambrian.
Of the 35 phyla of extant metazoans, there are nine major ones, based on their high
representation in terms of both species numbers and populations. These are the Porifera,
Cnidaria, Platyhelminthes, Nematoda, Annelida, Mollusca, Arthropoda, Echinodermata and
Chordata (Table 2.2). These make up 99.2% of all animal species. We will consider these
phyla one after the other, looking at the distinguishing and the phylogenetic trends that
they exhibit. Although kinships linking the primitive multicellular animals with the other
metazoans are still an area of hot controversy, here, for now, we follow tradition and start
with the sponges.
Table 2.2: The major animal phyla and their estimates (extracted from Brusca, 2003)
Phylum Estimated no of species examples
Porifera 5 500 Freshwater sponge
Cnidaria 10 000 Portuguese man of war
Platyhelminthes 20 000 tapeworm
Nematoda 25 000 hookworm
Anellida 16 500 Garden earthworm
Mollusca 93 000 Garden snail (Helix)
Echinodermata 7 000 Sea cucumber
Arthropoda 1 097 631 crab
Chordata 49 693 Tunicate
Total 1 324 324

PART 2- The Parazoa
CHAPTER 3
Phylum Porifera (Sponges)
The Porifera (pore-bearing organisms) are considered the most primitive of all metazoans.
Although multicellular, like all the other metazoans, the cells are not truly specialized into
tissues; they are loose associations of cells. They are thus sometimes referred to as the
Parazoa (meaning beside the animals). In addition to the lack of tissues, they differ from
other metazoans in having no nervous tissues and hence having no coordinated activities.
Unlike other animals also, their principle opening, the osculum is exhalent and not inhalent.
The body of sponges is constructed on a system of water canals that bring water, nutrients
and dissolved gases into the sessile animal. They live a sedentary life, attached onto rock,
cement blocks or pieces of timber
Sponges are largely marine with very few species (about 150) out of the 10 000 described
species being freshwater. They live in the intertidal zones attached to rocks, or any other
had substrata.
Figure 17. The three

grades of organisation in the sponges: asconoid, syconoid and leuconoid grade. (Source:
http://en.wikipedia.org/wiki/Sponge)
They vary in size from a few cm to over 2metres in length, occurring as tubular or vase-like
structures (Fig 17) and very often as branched encrustations. Usually brightly coloured, with
colourations of orange, red, yellow and purple being quite common and exhibiting three
levels of organisation: Asconoid, Syconoid and Leuconoid. In the Asconoid forms, size is
generally limited to small vase-like encrustations and the body wall is not folded as in the
other forms. In the Syconoid forms the body wall becomes folded and this is accompanied
by an increase in size, and finally in the Leuconoid forms, very large sizes of up to 2metres
are attained (Figure 17).

Figure 18: Common shapes of sponge skeletal material
The sponge body is generally supported by some skeletal material in the form of inorganic
or organic structures called spicules and spongin fibres respectively. The spicule skeleton
varies in structure and composition. The inorganic material can either be siliceous (made
from silicon) or calcareous (calcium carbonate). Sponges can have a skeleton made up of
calcareous spicules, siliceous spicules or protein spongin fibres or a combination of the last
two. The spicules occur in various shapes (Fig 18). The simplest are needle or rod shaped
and can be straight or curved. These are referred to as monaxon spicules (or just
monaxons). Some are triaxons, bearing three rays. Others are tetraxons, hexaxons, and
polyaxons, bearing 4, 6 and many rays respectively. These skeletal materials are secreted by
amoeboid cells in the mesohyl layer of the sponges called sclerocytes and spongocytes. The
sclerocytes are responsible for siliceous and calcareous spicules whilst the spongocytes
produce sponging fibres. The nature of the skeletal elements is important in the
classification of sponges. We shall return to the classification of sponges, but first, let us
take a detour and look at the different sponge organisations.

Sponge grades
The Asconoid sponges

The typical Asconoid sponge body form is vase-like (Fig 17). They are among the smallest in
the sponge world with sizes ranging from 1cm to 10cm in height. The body wall is
perforated by small openings called ostia (singular -ostium) which allow water to be drawn
in to the central cavity called an atrium and escape through the osculum- an opening at the
tip of the organism. The movement of water into the animal brings with it particulate food
in the form of bacteria and protozoa that are engulfed by the cells lining the atrium. Gases,
likewise also move with the water in and out as the water flows.
The body wall (Figure 19) is composed of three cell layers- the outer pinacoderm, a middle
mesohyl, and inner choanocyte layer lining the atrium. The pinacoderm layer is composed of
flattened cells called pinacocytes. These form an outer sheath of cells perforated at intervals
by the ostia, openings which traverse the whole body wall into the atrium. Each ostium is an
opening through an elongated pinacocyte cell which traverses the whole breath of the body
wall. These cells are thus referred to as porocytes because of the bore or lumen which they
form, presumably as a result of cell invagination. The pinacoderm cells have at their bases
some contractile fibres which they alternatively contract and relax thereby opening or close
the pores in the porocytes.
The mesohyl layer is gelatinous layer containing amoeboid cells and skeletal structures (the
spicules and spongin fibres). As we have already noted the skeletal material occurs in
various shapes and compositions. Amoeboid cells of various types occur in the mesohyl
layer. There are amoeboid archeocytes which are totipotent and thus responsible for
production of other cell types including eggs and sperms. These are large nucleated cells
capable of phagocytic activity and digestion of engulfed food particles. Amoeboid
scleroctyes and spongocytes are responsible for the secretion of the spicule and protein
spongin fibres respectively.

Figure 19: Section through the body wall of an Asconoid sponge. (Source:
http://www.esu.edu/~milewski/intro_biol_two)
The inner-most layer, the choanocyte, lines the atrium and abuts with the mesohyl. It is
composed of flagellated cells with a microvillus collar (just like the choanoflagellate cells
from which the sponges are thought to have evolved). The choanocyte cells are responsible
for creating a current of water moving into the atrium from the outside, and out through
the osculum through the asynchronous beating of their flagella. They are also responsible
for engulfing food particles and gaseous exchange.
As pointed out earlier, the Asconoid sponges are small in size. They have failed to attain
larger sizes by the restriction which is brought about by a none corresponding increase in
surface area as size increases. This tends to limit sufficient uptake of food material and
gaseous exchange through the choanocyte layer.
Other species of sponges have attained relatively larger sizes by folding the body wall to
varying degrees. This folding of body wall resulted in the extensive increase in surface area
for food and gaseous exchange. The Syconoid body structure, to which we turn now, has
been accompanied by a manifold increase in body size. Some amongst this group have
attained sizes of up to a metre in height and diameter.

The Syconoid sponges
Sponges exhibiting the Syconoid body structure were the first to venture into massive
increases in body sizes (Figure 20) through body wall folding. In these sponges, the body
wall is folded (convoluted) so that a series of invaginations and evaginations alternate with
each other (Fig 17 and 21). The choanocyte cells are no longer in the region bordering the
atrium but are now restricted to the evaginations which now form chambers referred to as
flagellated chambers (choanocyte chambers). The invaginations from outside form incurrent
(inhalant) canals. Water now courses in through the inhalant canals, into the flagellated
chambers through prosopyles (the equivalence of ostia in the Asconoid sponges) and in the
atrium through apopyles and finally to the outside through the osculum (Fig 21).
In some more specialized Syconoid sponges, pinacocytes proliferate and sort of plug the
incurrent canals leaving smaller dermal pores which lead into subdermal spaces before
connecting to the incurrent canals. Water now flows through the dermal pores into
subdermal pores then into incurrent canals leading into prosopyles, into flagellated
chambers to the atrium through prosopyles, and out through the osculum
Figure 20: A Syconoid sponge, Scypha. Though it isn’t evident from the outside, Scypha has a
Syconoid body plan. (Source: http://www.freethought-forum.com)

Figure 21: Drawing of a section of the body wall of a Syconoid sponge. (Source:
http://webs.lander.edu/rsfox/invertebrates)
Leuconoid structure.
This is thought to be the most evolved in the sponge world. Its efficiency is evidenced by the
occurrence of some of the largest sponges (up to 2metres high and 2metres in breadth). In
these sponges the body wall is extensively folded with the flagellated chambers also
becoming folded (Figure 22). Water now moves in through dermal pores, subdermal spaces,
incurrent canals, prosopyles, flagellated chambers, and out
through apopyles and excurrent channels which finally unite
to form larger channels leading out through the osculum.
Figure 22: Construction of a Leuconoid sponge. Note the
extensive folding which has taken place in these sponges
The Leuconoid sponges form the giants (Figure 23) in the

sponge world, and as have been alluded to already this is

evident enough to show that this grade of organisation is efficient in collection of food and
oxygen for the body needs.
Figure 23: A variety of Leuconoid sponges
Sponge classes.
There are 10 000 described species of sponges exhibiting various levels of organisation.
Some exhibit the Asconoid structure only, some the Syconoid and others the Leuconoid
structure and still others have body forms that traverse through all the body forms, passing
from Asconoid to Syconoid to Leuconoid in their life cycles. Sponge structure is thus not a
criterion used in their classification, but as we have already alluded to, nature and chemistry
of skeletal material become quite handy in sponge classification into classes. The 10 000
described species of sponges are grouped into 4 classes: Calcarea, Hexactinellida,
Desmospongia and Sclerospongia.
The Calcarea sponges (Calcispongia) are characterised by a skeleton made up of calcareous

(calcium carbonate) spicules which are monaxons. They exhibit all three grades of structure;

Asconoid, Syconoid and Leuconoid. Examples include Leucosolonia, Sycon and Grantia (Figs
23a, b and c)
The Hexactinellida or Hyalospongia are commonly known as the glass sponges from the
appearance of their skeletal spicules. The spicules are hexaxons and often fused to form a
lattice like structure. They are made of siliceous material, hence the glass appearance. Their
bodies are usually cup or vase-shaped and height averages from 10 to 30cm. They lack a
pinacoderm but have an epidermis consisting of a net-like synctium formed by pseudopodia
of amoebocytes that form interconnecting structures. The Syconoid structures dominate in
these sponges and are entirely marine, occurring in deep waters. Figure 24 a and b shows
common deepwater Hexactinellida and Euplectella.
The Desmospongia have a variable skeleton consisting of either siliceous spicules or spongin
fibres or a combination of both. When both siliceous and spongin fibres exist, the spicules
are usually connected to or completely embedded in the spongin fibres. All Desmospongia
are leuconoid and they constitute the largest class with over 90% of all the classes. The
Leuconoid structure allows them to attain very large sizes with some, like the tropical
loggerhead sponges (Spheciospongia) forming masses more than a meter in height and
diameter. Pigment granules present in amoebocytes in many species make this group
frequently brilliantly coloured. They are distributed from great depths to shallow water.
Examples include the common bath sponges (family Spongidae), like Spongia and
Hypospongia.
The Sclerospongia are a small group of sponges with a small number of species found in
grottoes and tunnels associated with coral reefs. They are leuconoid with siliceous spicules
and spongin fibres and an outer encasement of carbonate.

PART 3 -The Radiata
CHAPTER 4
The Cnidarians. (Hydras, jellyfish, sea anemones and

corals)
There are about 10 000 species of Cnidarians most of which are marine with very few
freshwater species. They are diploblastic animals with two body layers, an ectoderm and a
gastrodermis enclosing a gastrovascular cavity. In between the two layers (ectoderm and
gastrodermis) is a gelatinous layer called a mesoglea. The gastrovascular cavity opens to the
outside through a mouth which also serves as the anus. The mouth is surrounded by
tentacles which are used during feeding (prey capture).
The Cnidaria are typically dimorphic (exhibit two body forms); the
polyp and the medusa (Figures 24 and 25).
Figure 24: The Polyploidy form of cnidarians.
(Sourcehttp://www.mbgnet.net/salt/coral/animals/cnidar.htm)
Figure 25: The medusoid form of cnidarians. (Source:

http://www.mbgnet.net/salt/coral/animals/cnidar.htm)
The polyp is tubular with the oral end up and the aboral end used for sticking on rocks.
Polyps may occur as solitary individuals as in Hydras or colonial like in Obelia. The medusa is
the motile form and umbrella like. It is an inverted version of the polyploidy form, with
tentacles hanging down from the mouth which faces downwards from the medusa. The top
surface is referred to as the exumbrella and the oral surface is called the subumbrella. The
mesoglea in the medusa is often quite massive, giving the whole animal a jelly-like
appearance, hence the term jellyfish which is used to refer to the medusae of some

Cnidarians. Contraction of the muscles in the subumbrella margins pushes water down or
backwards and the organism moves with exumbrella surface forwards.
Some Cnidaria only exhibit the polypoid form and others the medusoid form and still others
pass from one form to another in the life cycles. It is important to note that the two morphs
do not represent alternation of generations as in plants. In fact, the two forms are both
diploid. In forms that alternate between polypoid and medusoid morphs, one form is usually
predominant and the other less conspicuous. For examples in the hydrozoans, Obelia exhibit
both forms, but the polypoid form is the dominant form. In hydra only the Polypoid form
exist.
Figure 26: Section through the body wall of Hydra showing the body layers and associated
structures. (Source: http://cas.bellarmine.edu/tietjen/images/cnidarians.htm)
As already noted, the body wall is composed of two germ layers; the epidermis and an inner
gastrodermis, and in between, a gelatinous mesoglea which is usually cellular (Figure 26).
The epidermis is composed of five major cell types; the epitheliomuscular cells, interstitial
cells, cnidocytes, mucous secreting cells and sensory nerve cells. The epitheliomuscular cells
are columnar cells that line the outside. At their basis are strands of contractile filaments
which contract and relax when the animal is disturbed thereby bringing about bending. In
between these cells are interstitial cells that are totipotent and give rise to either the
epitheliomuscular cells or germinal cells like eggs and sperms.
The third cell types in the epidermal layer are the cnidocytes. These are located throughout
the body wall, interspaced between the epitheliomuscular cells and they are particularly
abundant on the tentacles. They contain organelles capable of eversion (protrusion by
turning inside out) called cnidae. Typically a cnidocyte is an ovoid cell with a basal nucleus
(Figure 27).
B
Figure 27: (A) A cnidocyte containing a nematocyst and (B) A nematocyst being discharged
upon stimulation by pre or predator. (Source:
http://www.tutorvista.com.biology/polymorphism-coelenterata)
One end of the cell interfacing with the environment contains a short bristle-like process
called a cnidocil in hydras (hydrozoans) and jellyfish (scyphozoans). This, in ultrastructure, is
basically a flagellum.
Cnidae include the nematocysts, spirocytes and ptychocytes. The nematocysts (Figure 27)
are stinging organelles that are varied in form. They are used in prey capture and defence.
They are capable of penetration and delivering a toxin. The nematocyst is like a capsule
which contains a coiled, usually pleated tube. The end of the capsule directed to the exterior
is covered with a thread-like flap called an operculum. Stimulation of the cnidocil by a prey
or predator readily elicits the discharge of the nematocyst. This is due to rapid entry of
water into the cnidocyte caused by a change in the permeability of the cell as a result of the
stimulation. The discharged nematocyst is of varying length, consisting of a thread and
equipped with spines at its base. It penetrates the skin of the victim, get anchored by the
spines and discharge the toxins to kill or immobilize its victim.
The spirocyst is an adhesive structure found in some anthozoans. Upon discharge the thread
gelatinizes to form an adhesive net that is used in prey capture. Like the spirocyst, the
ptychocyst is an adhesive structure but is elaborately pleated and devoid of spines. This is
the typical cnidae in some sea anemones.
All the three types of cnidae, once used, like the sting in bees, are discarded. New
cnidocytes can be regenerated from interstitial cells and the developing cnidocyte is called a
cnidoblast.
Some cnidarians are notorious for their very painful stings and, with others; the toxins can
be lethal to animals as large as the size of an adult man. It is quite common to see along
some beaches with warnings such as ‘Beware of sea wasps” a warning which keeps off all
who have experienced the excruciating pain of these stings.
The fourth cell types, the mucous secreting cells are abundant in the basal disc of the sessile
polyps. They secrete mucous for prey capture, protection and adhesion. The last types of
cell, the sensory nerve cells are receptors. They are elongated cells perpendicular to the
epidermal cells. Their bases give rise to neuron processes and their distal ends terminate in
a sphere or sensory bristle. These receptor cells are particularly abundant in the tentacles.
The gastrodermis is also composed of a number of cell types. One type, the nutritive
muscular cells are oriented perpendicularly to the long axis of the body stalk and thus form
a circular layer bordering the gastrovascular cavity. They serve to engulf particulate food
from the gastro vascular cavity for intracellular digestion. Being flagellate, they also help to
continuously stir the contents of the gastrovascular cavity. Enzymatic gland cells are
interspaced between the nutritive muscular cells, where their tipped ends directed towards
the mesoglea are wedged. They are responsible for extracellular digestion by secreting
enzymes into the gastrovascular cavity. The gastrodermis, like the epidermis, also contains
mucous secreting cells, these being particularly abundant in the region around the mouth
where they serve to secrete mucous which aids in lubricating the food and thus facilitate
ease swallowing of prey. Also present in the gastrodermis of some cnidarians like the
cubozoans and scyphozoans are nematocysts.

Movement in Cnidarians is limited in the polypoid forms but can be quite active in the
medusoid forms. In the polyploidy forms, body contractions in the musculoepithelia results
in bending in the hydras resulting in slow movement involving looping or somersaulting.
This is brought about detaching and shifting location by somersaulting or floating. In the
medusa, the extensive muscles in the subumbrella margins contract and relax thereby
producing movement in the oral /aboral plane.
Feeding is brought about by the activity of the tentacles. Stimulation of cnidocil in the
tentacles results in discharge, entangling and paralysing of prey. The prey is then brought
into the mouth region by the tentacles and the prey is swallowed. Mucous in the mouth
region aids swallowing and once the food is in the gastro vascular cavity, enzymatic gland
cells secrete proteolytic enzymes which initiate the digestion. This results in a soupy broth.
The nutritive muscle cells continuously agitate the contents of the gut by the beating of
their flagella. And also, through their phagocytic activity they take up the particulate food in
the soup broth and digestion continues intracellulary.
In the whole Cnidarian group, there are no specialised organs for gaseous exchange and
nitrogenous waste excretion. These occur throughout the body surface.
Classes of Cnidaria
The cnidarians are divided into 4 classes: Hydrozoa, Scphozoa, Cubozoa and Anthozoa,
mainly on the basis of whether a polyp or medusa is the dominant feature in the life cycle.
The nature of the medusae, in medusoid forms is also used as criteria for classification.
In the Hydrozoans, the polyp (Figure 28). Is the dominant feature and the medusae are less
conspicuous.
Figure 28: A polyploidy hydrozoan:

The Pink Hearted hydroids. They look like delicate plants but they are animals that sting and
capture food (Source: http://www.oceanicresearch.org/education/wonders/cnidarian.html)
The most common hydrazoan is the freshwater Hydra which exhibits the polyploidy form
only. It is a common laboratory animal used on investigations into behaviour patterns in the
Cnidaria. Hydra is a small tubular organism just visible to the eye. Its body consists of a

stalk, with a basal end which is used for attachment to the substrate and the other terminal
end bearing a mouth surrounded by tentacles. It reproduces asexually, budding off small
polyps form its axial body. Sexually is it reproduces by way of producing eggs and sperms in
ovaries and testis which form from interstitial cells in the epidermal walls. The testis is
usually formed above the ovaries on the body stalk to facilitate ease movement of sperm
from testis to ovary to fertilize the egg.
Obelia, another common hydrazoan, exhibits both the medusoid and polypoid form (Figure
29 and 30). The Polyp, which is colonial basically consist of a vegetative structure built on a
series of highly branched stalks. To the unwary observer it thus appears plant-like. Each
stalk is an individual contributing to the built up of the whole colony. Each stalk is like a tube
within a tube. The outer tube, called the perisarc, is none living and is the product of the
secretion of the inner tube, the coenosarc. It is annulated prior to each branching to allow
for bending as water currents move the different individuals relative to each other. The
coenosarc, which is living, is built on the basic cindarian structure- the diploblastic
condition- common in all cnidarians. Within the coenosarc is the gastrovascular cavity
which is continuous throughout the whole body of the colony. The vegetative body consists
of a horizontal stem or stolon called a hydrorhiza. From the hydrorhiza arise vertical
Figure 29: A colonial polyploidy form of Obelia.

(Source http://www.tutorvista.com/biology/polymorphism-coelenterata)

Figure 30: A magnified form showing
a single polyp of Obelia attached to a hydranth.
(Source:http://www.tutorvista.com/biology/polymorphism-coelenterata)
stems called hydrocaulus. Each hydrocaulus terminates in a feeding individual called a

hydranth. The hydranth has a bulbous terminal bearing a mouth situated on a mound, the
hypostome. The hypostome is surrounded by tentacles. The expanded region of the
hydranth houses an equally expanded gastrovascular cavity called the gastric region. The
hydranth is enclosed by an expanded part of the perisarc called the hydrotheca. Tentacles
surrounding the mouth are responsible for prey capture and hence feeding. In the axial
region of the hydrocaulus are branches of reproductive individuals called blastostyles. Each
blastostyle is also surrounded by an expanded perisarc called the gonotheca. The blastostyle
buds off young transient medusae which temporarily become pelagic but soon settles to
grow and give another mature polypoid which branches off to form a new colony.
In the jellyfish (Cubozoa and Scyphozoa) the medusoid form is the dominant morph and the
polypoid forms, being restricted to small larval stage, are very inconspicuous and transient.
They are however distinguished on the bases of the shape of the bell (umbrella or bell-like
medusa), the nature of the bell margin and number of tentacles or tentacle clusters.
Scyphozomedusae and cubomedusae are generally larger than hydromedusae (Figure 31

and 32). The majority of their medusae have bell diameters ranging from 2 to 40cm,
although some species are even larger. In the Cubozoa the bell is roughly cuboidal shaped
and the bell margin is not indented and they typically bear 4 tentacles or tentacle clusters.

In the Scyphozoa (the true jellyfish), on the other hand, the bell varies in shape from being a
shallow saucer to a deep helmet, with deeply indented or scalloped margins to form lappets
and the tentacles around the bell margin ranging from four to many. The scyphomedusea
are generally larger than cubomedusae.
Figure 31; The scyphozoan medusa.
Their gonads are gastrodermal like hydromedusae and like hydromedusae release their
gametes into the water. Upon fertilization the zygotes develops into a planura larva that
settles and grow into a small inconspicuous polyp that buds juvenile medusae that will grow
into adults. Examples include Cyanea capillata, Chrysaona, Stomolophus and Chironex
fleckers.

Figure 32. A cubomedusa
of Chiropsalmus sp. (Source:http://www.ucmp.berkeley.edu/cnidaria/Chiropsalmus_sp.html)
The Anthozoans (corals, sea anemones, as well as sea fans and sea pens) constitute the
largest class in the cnidarians with about 6 000 species. They occur as polyps only and their
polyps differ from those of the hydrozoans in that they are generally larger with most sea
anemones polyps ranging from 1.5 to 10cm in length and 1 to 5cm in diameter. They have a
flattened oral disc bearing tentacles and a slit-like mouth leading into laterally flattened
tubular pharynx that extent more than halfway into the gastrovascular cavity. Their
gastrovascular cavities are divided into radiating compartments by longitudinal mesenteries
or septa, with the edges of the mesenteries bearing nematocysts. Their gonads are
gastrodermal and the mesoglea is fibrous structures and bearing cells.
The sea anemones (Figure 33) are solitary polyps that occur throughout the world in coastal
waters. They commonly live attached to rocks, shells and submerged timbers.

Figure 33: A sea anemone oral end, note the tentacle cluster surrounding the mouth in the
oral end (Source: http://library.thinkquest.org/J001418/anemone.html)
The other anthozoan group, the corals live as solitary or colonial forms and secrete hard
external skeletons of calcium carbonate (Figure 34). Each polpy generation builds on the
skeletal remains of the previous generations to 'construct' rocks which will develop into
coral reefs.
Figure 34: A coral colony, a coral

colony consists of hundreds or thousands of tiny polyps. Each polyp is an individual animal
(basically a small anemone) but they live together as a group.
(Source:http://www.oceanicresearch.org/education/wonders/cnidarian.html)

PART 4-THE BILATERIA
CHAPTER 5
The Protostome Phyla.
The Platyhelminthes (flatworms-turbellarians, fukes

and cestodes).
The platyhelminthes are typically triploblastic, bilateral animals with moderate cephalisation
(Figure 35). They have an anterior end and a posterior end, with the former bearing sensory
structures. The group is composed of both free-living and parasitic forms. They are typically
worm like and flattened- belt or leaf-like. Their body walls are made up of three germ layers,
their being, in addition to the two germ layers present in Cnidarians, a third germ layer, the
mesoderm. The presence of a third germ initiates in these animals (and in higher
metazoans) an ability of the body to attain organ and organ systems level of organisation.
Figure 35. Whole mount of planarian Platyhelminthes. Note the two protuberances in the
head region which function as mechanoreceptors.

The mesodermal tissues however fill the whole of the embryonic blastocoel and thus there
is no internal body cavity- the coelom (Figure 36). They are thus called acoelomate animals.
The mesodermal tissue proliferates as parenchyma tissue which fills the whole space
between the body wall and the gut.
Figure 36; The acoelomate condition as obtained in Platyhelminthes
Their nervous system, unlike the nerve-net present in Cnidarians is slightly more elaborate
with an anterior nerve mass (cerebrum) aggregating in the head region and usually two
ventral nerve cords that run posteriorly (Figure 37). This may be in the form of a ladder like
structure as in the turbellarians due to transverse nerves connecting the two nerve cords.
Free-living forms have also developed some sensory structures in the head region but these
have been lost in parasitic forms. These sensory structures occur as mechanical or light
sensitive structures in the planarians. The free-living forms and some parasitic forms have
primitively developed a true alimentary canal with a mouth and gut but no anus. The gut is
thus said to be blind ending. In these groups with a digestive system, the mouth is
primitively midventral to terminal. In the more derived (advanced parasitic) forms the gut
has completely disappeared.
The body wall is composed of an outer epidermis followed below by basal membrane then
circular and longitudinal muscles. The epidermis may be ciliated or thrown into folds like
microvilli and microtriches, adaptations to either a free-living or parasitic existence. In the
free-living forms the cilia is used as gliding structures in bringing about locomotion and in
the parasitic forms the cilia and microtriches serve to increase surface area for absorption of
food materials from the fluids of their hosts in which they are resident.
Figure 37; Typical nervous system of a Platyhelminthes

Parasitic Platyhelminthes also bear attachment or adhesive structures to cling on their
hosts. These attachment organs are in the form of suckers and hooks.
Figure 37: Anterior end (head region of Cestoda (Taenia; sagitum)
There are four classes of Platyhelminthes, the free-living Turbellaria, the Monogenea,
Trematoda, Opisthobothridae and Cestoda.
Turbellaria
The turbellarians are mainly free-living with most of them being aquatic, living in freshwater
and marine environments. Very few are terrestrial, living in very dumb habitats. The
woodlice are one example of a terrestrial turbellarian. The turbellarians vary in size from a
few centimetres to the giant planarians. Their bodies are typically leaf like (Figure 38). Being
free-living they show most of the primitive feature. Anteriorly, in the head region are,
usually tentacle- like structures, the auriles which are sensitive to a variety of stimuli
including light to which they respond by negative taxis when exposed to high intensity light.

Figure 38. A typical turbellarian (Pseudoceros dimidiatus). Note the leaf-like body (Source:
http://upload.wikimedia.org/wikipedia/commons/thumb/1/1e/Pseudoceros_dimidiatus.jpg/
800px-Pseudoceros_dimidiatus.jpg)
External morphology and body wall.

Externally, turbellarians are leaf like with a head region equipped with sensory structures
responsive to light. These structures are called auriles. The mouth is typically ventral, and in
the advanced forms it is borne on an eversible pharynx. The turbellarian body wall is
covered by cilia which they use for locomotion. Cilia are either restricted to the ventral
surface or may cover the entire body surface. The ciliated epithelium is either composed of
individual cells linked by gap junction or is synctyial. Just below the epidermis is the basal
lamina. Below the basal lamina are the circular and longitudinal muscles respectively.
Between the body wall and the gut are parenchyma cells of mesenchymal origins. These can
either completely fill the space or leave small spaces that act as a pseudocoelom. Some of
the parenchyma cells serve as gland cells for production of mucous to lubricate the body
surface and hence enable the animal to glide forward. Other parenchyma cells are
totipotent and give rise to other cell types. In between the cells is the ground connective
tissue interrupted by branches of the gut and dorsoventral muscles (Fig 39).

Figure 39: Section through the body of a turbelarian (Planaria).
(Source:http://cas.bellarmine.edu/tietjen/Laboratories/Bio%20Pix%204%20U/Image38.gif
The gut
Turbellarians like all other flatworms do not have an entire alimentary canal. At the anterior
end is a mouth which is usually borne at the end of an eversible pharynx. The pharynx leads
into a short oesophagus which bears a muscular bulb which is used in pumping food into the
intestines. The nature of the intestines varies from group to group depending on the size of
the worms. In the very small Acoela there is no permanent gut. The intestine is composed of
a cynctyial mass into which ingested food is digested intracellularlly. In relatively bigger
cutenellids, macrostomids and rhabdocoels the gut is a simple sac. In the triclads the
intestines is triradiate with one anterior and two anterior ceaca and in the polyclads, to
which the popular planarians belong, the gut is thrown into a number of diverticula which in
turn also have diverticula. The result is the complete ramification of the entire space
between the gut and body wall and hence eases distribution of nutrients in these animals
which are devoid of a circulatory system. The pharynx also shows the same trend in
complexity with the simple turbellarians having a simple pharynx and the complex ones
having complex pharynx that can be everted and swallow the pray outside the body cavity.
The nervous system

Unlike the simple diffuse nervous system present in the cnidarians, turbellarians nervous
system show marked advance in consonance with bilateral symmetry and directional
movement shown by the animals. Anteriorly in the head region is cerebrum which gives off
nerves to the sensory structures in the head as well as circumoesophageal commissures that

connect to a suboesophageal ganglion from which two ventral nerve cords run posteriorly
with transverse nerves connecting the two to form a ladder-like structure. In primitive
turbellarians like Acoela, however the nervous system is still very primitive, with a
subepidermal nerve plexus and the brain totally absent. This is one of the reasons why many
authorities have downgraded the Acoela to a group separate from all other Platyhelminthes.
Figure 39. Typical nervous system of turbellarians (Source:

http://cas.bellarmine.edu/tietjen/images/platyh10_small.jpg
Excretory system
Excretion is typically through protonephridial tubes. Flame cells with nephridial tubes
terminate in two lateral collecting tubes which run anteriorly (Figure 40) and open to the
exterior in the anterior region.

Figure 41: Protonephridia tube with associated flame cell (Source:modified from
http://cas.bellarmine.edu/tietjen/images/platyh12.jpg)
Reproduction.
Reproduction is typically both sexual and asexual. In many worms, a cut of the worm into
two halves results in the regeneration of the other half, and with many species serial cuts
along the body results in each segment regenerating the missing portions.
Turbellarians are typically hermaphroditic (Figure 41) although cross fertilization is the
norm. Variations do occur in the turbellarians with regards to the nature and disposition of
the sexual organs, but typically, the female reproductive system consists of ovaries
anteriorly located and connected via oviducts to germinal chamber posteriorly from which a
single tube connects to the genital pore ventrally. The genita pore is a common pore and
serves both the female and male system. A copulatory sac connects to the genital chamber.
Viteline glands distributed within the body are connected to the oviducts and provide
nourishment to the developing egg. In other turbellarians there may be numerous pairs of
ovaries, but only a single pair of oviducts. The male reproductive system consists of two
tubes running anteriorly and receiving tubes (Vas deferens) from numerous testes lying in
the parenchymatous tissues. The tubes lead posteriorly before uniting to give a single tube
which terminates in a penis armed with stylets which in turn opens in the common genital
aperture. In some species, instead of a penis a cirrus is present. Sperm transfer is mutual,
the penis of each partner being inserted into the copulatory sac of the other as they abbut
to each other oppositely. Sperm transfer occurs and sperms are stored in the copulatory
sacs from which they will subsequently move and enter the oviducts enroute to the ovaries
where fertilization occurs as soon as the eggs are released.

Feeding.
Most turbellarians are free living and utilize microscopic as well as planktonic organisms as
food. They are typically either scavengers or carnivorous. Prey is captured by pharynx
(eversible or non-eversible) which pumps the food material into the intestines. Digestion is
initially extratracellularly but finally intracellularly.
Figure 41. The hermaphroditic condition of turbellarians, (O-Ovary, Ov-oviduct, yg york

glads, , Gc -genital chamber, T- testis, vd-vasa deferens, P- penis, Cs -Copulatory sac)

(Source:http://www.path.cam.ac.uk/~schisto/general_parasitology/parasitology_platyhelmi
nthes_reproductive.html)
The Monogenea.
These are mainly parasitic on the external surfaces of aquatic organism like fish, frogs and
reptiles and some have been reported on hippos. Monogeneans, as the name implies, are
typically single host/generation parasites suitably adapted to ectoparasitism through
development of architecturally modified structures to fit into specific host sites.
Like the turbellarians they are flattened and thin (Figure 42). They bear surface attachment
organs in the head region (haptor) and more conspicuous ones in the posterior region called
opisthohaptors. The opisthohaptors are well sculptured structures with anchors (hamuli),
hooks and clamps. The anchors and hooks are elongated keratinized structures that pierce
into host tissue to secure the parasite in place. The clamps help secure the parasites in its
specific host site by clamping onto the skin of the host just like clothespins. All these
structures serve to allow the bearer to cling tenaciously to the outer surface of its host and
thus not lose hold as the host moves against water currents.
Figure 42: Structure of a typical Monogenean. Note

the conspicuous opisthohaptor equipped with hooks
Body wall
Like the turbellarian body, the monogenean body wall is covered by epithelia, but unlike
that of the former, it is not ciliated. The epidermis is made of either a layer of cells or a
synctyium which is continuous with cells originating deeper in the parenchyma cells. A basal
lamina follows below, and then a layer of longitudinal and circular muscles respectively.

The gut
Terminally is mouth leading into a prepharyngeal tube which then leads to a muscular,
bulbous pharynx which is continuous with oesophagus? The oesophagus bifurcates to give
two blind ending ceaca which may bear some fenestrations (Bush et al 2003).
The excretory/osmoregulatory systems.

Excretion and osmoregulation are achieved by protonephridia. Flame cells and their
protonephridial tubes terminate into two longitudinal tubes which swing posteriorly then
anteriorly to open in the head region through a single pore.
The nervous systems.

Monogeneans are capable of simple responses to light and chemicals. They are equipped
with photosensitive as well as chemoreceptive structures in the head region. Internally in
the head region is a pair of cerebral ganglia which link with the head receptors as well as
developed commissures linking to a suboesophageal ganglion. Two ventral longitudinal
nerve chords from the suboesophageal ganglion lead posteriorly and are linked by
transverse nerve to form a ladder-like nervous system.
Reproductive system
The reproductive systems of monogeneans are very similar to that of the Trematoda except
that one or two vaginas may be present. Monogeneans are typically hermaphroditic with
each worm bearing one set of either reproductive system. The female reproductive system
typically consists of a single ovary connected to an ootype by an oviduct. Before the oviduct
reaches the ootype it is joined by three tubes; a vitelline tube from the numerous vitelline
glands, a tube from the seminal receptacle and another from the vagina. From the ootype
which, is surrounded by a Mehli's gland, the uterus continuous anteriorly to the outside
through a common genital opening. The male reproductive organs consist of a pair of testis
(there can be many testis but they are usually two), from which vas efferentia lead to a vas
deferens which in turn lead to a cirrus pouch containing seminal vesicles and prostate
glands. From the cirrus pouch emerge a copulatory organ with spines at its distal end (the
cirrus) which leads to the outside during copulation through the common genital atrium and
pore.
Monogeneans are typically one host parasites, producing eggs that hatch into free
swimming onchomiracidia which attach onto new hosts. The onchomiracida have some
hooks at the posterior which are retained in the adult as the attachment hooks of the
opistohaptor. In some Monogeneans, like the gyrodactyllidae a single larva, viviporously
gives rise to another young one which soon after being produced also gives birth to another
larva which does the same until a host is located and entry is achieved. Mucous produced by
host are thought to elicit a chemotactic response in the onchomiracidium which aids host
location and attachment.
The Trematoda

Figure 43: Body plan of a trematode (the amphistome condition) (Source:
http://www.path.cam.ac.uk/~schisto/general_parasitology/Platy_RepSys.female.gif
This are typically leaf-like and e very thin (Figure 43). They occur mainly as parasites in the
guts and associated organs of the gut in their vertebrate and invertebrate hosts. Anteriorly,
like in all Platyhelminthes, is a head-like organ, though less defined than that of the
turbellarians. Primitively they have an anterior sucker and ventral sucker (the diastome
condition), or posterior sucker (the amphistme condition) or a single posterior sucker. The
mouth is typically ventral. Their bodies are covered by synctyial epithelia whose roots are
embedded in the deeper seated parenchyma cells. Below the epithelia are basal laminas
below which are circular and longitudinal muscles. Like all platyhelmithes the gut is blind
ending. The ventral mouth opens into a buccal opening which leads into a muscular
pharynx. After the pharynx a short tube leads into two blind ending ceaca, which are usually
thrown into numerous diverticula ramifying the whole body.
The excretory/osmoregulatory system

Excretion and osmoregulation are effected by protonephridia that have tubes leading into
two collecting duct that run laterally to the posterior end of the body where they unite into
a single bulbous bladder which opens to the outside through a nephridiopore.
The nervous system

In the head region is a pair of cerebral ganglia which sent at least two ventral nerve cords
running to the posterior. These are connected at intervals by transverse nerves to form the
typical ladder-like nervous system characteristic of platyhelmithes.
The reproductive system

Trematods are typically monocieous with individuals bearing both male and female
reproductive systems. The female reproductive system consist of, usually a single ovary
from which a single oviduct leads towards the ootype. Before it reaches the ootye it is
joined by a number of tubes; one of this is a tube from the seminal receptacle and another
one, the vitelline duct, from the vitelline glands. From the ootype (surrounded by Mehli's
gland) the uterus courses its way anteriorly leading into a vagina which opens into a genital
atrium or Laurel's organ which opens dorsally. The latter opens to the exterior through a
gonopore.

Figure 42: Female reproductive
system of a Trematode (O- ovary, Ut-Uterus, E- egg, Oo0 ootype, M- Mehl's gland, vt-
vitelline glands, vt.d- viteeline glands ducts, v or l vagina or Laurels organs) (Source:
http://www.path.cam.ac.uk/~schisto/general_parasitology/Platy_RepSys.female.gif
The male system is composed of two testes whose vas eferentia unite anteriorly to form the
sperm duct (vas deferens). The vas deferens runs towards the anterior and opens into a
pouch, the cirrus sac, equipped with seminal vesicles and prostate glands. From the cirrus
pouch an eversible copulatory organ (cirrus) terminates in the common genital atrium.
Figure 43: Male

reproductive system of a Trematode (T- testis, Vd- vasa deferens, cs- cirrus sac, CGA-
common genital atrium, SV-seminal vescle, pr-- prostate gland, and c – cirrus
(Source:http://www.path.cam.ac.uk/~schisto/general_parasitology/Platy_RepSys.male.gif)

The Opisthobothridae
These are thought to be the most primitive of all the parasitic platyhelmithes. They are
mainly parasitic on aquatic molluscs or aquatic ectothermic vertebrates like fish. Their
bodies, like those of the trematoda are leaf like but unlike trematodes with distinct suckers,
those in opisthobothridae are usually in the form of continuous ventral suckers divided by
muscular septal ridges or a series of ventral suckers. And, still, in other Opisthobothridae,
the suckers occur as ridged structures called rugae that clamp the surface of the host.
The opisthobothrid bodies are covered by a synctyial mass as in the trematodes. Below this
is basal lamina as in the trematodes then circular and longitudinal muscles. They have the
typical platyhelminthes nervous system with, an anterior cerebral ganglion linked to two
ventral nerve cords that run posteriorly with cross commissures at intervals. Excretion is
effected by protonephridia that collect into two lateral collecting tubes that open into a
bladder in the posterior region before opening to the outside through a nephridiopore.
Their reproductive systems are similar to those of the trematodes, but, unlike the
trematodes their life cycles are simple and direct. The eggs, which are voided together with
the host's excreta, hatch to give rise to ciliated larvae called cotylocidium which enters a
new host (if present) to develop into the adult. In those species that are parasitic on
ectothermic vertebrate, the larvae have to be ingested first by a molluscan host which then
passes it to the definitive vertebrate host as it is ingested in turn. The opisthobothrids are
not very specific in their host selection, with some species having been identified in up to
twenty different hosts.

Figure 45; The dog tapeworm Echinococus granulosum
(Source:http://animaldiversity.ummz.umich.edu/site/resources/Grzimek_inverts/Cestoda
/Echinococcus_granulosus.jpg/medium.jpg)

Figure 44. The Broad fish tapeworm Diphyllobothrium latum
(Source:http://animaldiversity.ummz.umich.edu/site/resources/Grzimek_inverts/Cestoda
/Echinococcus_granulosus.jpg/medium.jpg)
The Eucestoda
This is a group comprising highly evolved internal parasites with two to three or four hosts.
The eucestodes or true tapeworms, with a few exceptions, are internal parasites living, at
one time in their life cycles, in the gut of their vertebrate hosts and their associated tubes.
They have lost most of the features that their free-living and less derived parasitic relatives
possess. They do not possess a gut and all the light sensitive structures associated with their
relatives. They occur as parasites in the gut lumens, blood vessels, the coelom, heamocoel
and internal organs and muscles. The use of multiple hosts (both invertebrate and
vertebrate) and the alteration of host behaviour are some of the hallmarks of parasitic
evolution in this group.

Figure 46; Whole mount of the scolex of
the beef tapeworm (Taenia saginata) showing suckers and hooks.
They are very thin, flattened and typically ribbon like (Figures 44 and 45). Their bodies
appear segmented form the outside but the division is only superficial as the muscle tissues
inside are not in any way segmental. Anteriorly is a head region called the scolex (Figure 37,
46 and 47), followed by a short nonsegmented region called the neck, then a series of
segments (proglottids) making up the body (strobilus) of the organism. The head (Figure 46)
or scolex is equipped with a number of host attachment structures, including suckers,
hooks, spines and tentacle-like structures. The suckers are of three types, viz; the bothria,
the bothridia (diphyllidia) and acetebula. The bothria are slit-like grooved structures whilst
the diabothria are muscular earlike structures with thin margins suitably designed to fit into
particular sites on the host. The acetebula are simple suction sacs or cups. In some groups
like the teanids the anterior most part of the scolex is produced into mound called the
rostrum which surrounds the mouth and is also equipped with circular rings of small hooks
or spines.
The neck region is short non segmented section of the worm, the posterior end of which
gives off new proglottids by transverse division. The young proglottids of the strobilus are
those close to the neck region and the youngest one abbuts against the neck. The oldest
proglottids are those at the terminus of the strobilus. The strobilus is composed of a series
of proglottids each with a full complement of reproductive organs (both male and female
reproductive systems). The proglottids mature as they traverse to the end in the production
line and the oldest are at the end. Here they release free eggs to the exterior (anapolysis) or
bud off as bags full of eggs (apolysis. These, together with the excreta or faeces of the host,
are passed to the outside.

Figure 47: Life Cycle of a
Cestoda- Hymenolepsis nana. Note the Head (scolex, neck and strobilus) (Source:
http://www.parasitecleanse.com/images/tapeworm04.jpg)
Body wall
The Eucestoda body is covered by epithelia which is typically a cynctial mass composed of
cells whose roots are embedded in the parenchyma tissues. The outer wall is thrown into
many folds of microvilli called microtriches. These serve to increase the surface area for
nutrient absorption from the host fluids. The synctial mass contains amongst other
organelles, numerous mitochondria. The mitochondria contain very few cristae probably
signifying the heavy reliance on anaerobic respiration (Bush et al., 2003)). These epithelia is
covered by a glycocalyx, which is a carbohydrate rich mucous layer responsible for buffering
the epidermal layer from the host enzymatic activity as well as allowing for absorption of
mineral salts and bile salts. Below the epithelial layer is the basal lamina below which are
circular and then longitudinal muscles. Dorso-ventral and oblique muscles are also present.

Eucestodes are monocieous (Figures 48 and 49) except for a few like the Schistosomes that
are dioceous. Among the monocious species, copulation and hence sperm transfer is
reciprocal. The male reproductive system is composed of numerous testis connected to
small vas eferentia which then unite to form the vas deferens (sperm duct). The sperm ducts
leads laterally or ventrally to an expanded chamber, the cirrus pouch bearing accessory
glands, the seminal vesicle and prostate glands. From the cirrus pouch an eversible
copulatory structure, the cirrus extrude to the exterior through a common genital atrium
and gonopore.

Figure 48. The
hermaphroditic reproductive system of Cyclophyllidean Cestoda, P- proglottid, LEC -lateral
excretory canal, LGA-Lateral genital aperture (Female system (in red)= Ov- ovary, vt- vitelline
glands, Vt.d- vitelline gland ducts, Oo- ootype, S-seminal receptacle, V- vagina; Male system
(blue)= testis, V -vasa deferens, C-cirrus),(Source:
http://www.path.cam.ac.uk/~schisto/general_parasitology/Pseudo.pro.gif)
The female reproductive system is composed of, usually a single bilocular ovary from which
leads, anteriorly, a short oviduct to an ootype which is surrounded by a Mehli's gland.
Before reaching the ootype the tube receives a tube from the seminal receptacle and
ovivitelline duct. Thus sperm from the seminal receptacle fertilizes the egg enroute to the
ootype. From the ootype, two tubes take root, a blind ending uterus leads anteriorly and a
vagina courses its way laterally or ventrally to the genital atrium and opens to the outside
through the gonopore. A lot of variability occurs with regards to the precise number of
reproductive systems in a single strobila. In some forms there is a single pair, in others there
are two sets of male and female systems, and still in others, there are two sets for the male
and one set of female system. Exchange of sperms can and do occur between two
proglottids of the same individual or between two adjacent individuals, but the later is
naturally favoured. Self fertilization does occur but cross fertilization is thus the norm.
Protandry (maturation of male sex organs before maturation of female organs and
gyndantry (maturation of female sexual organs before male sexual organs) normally guard
against self fertilization.

Figure 49. The
hermaphroditic reproductive system of Pseudophyllidean Cestoda, LEC- lateral excretory
canal, GA-Genital aperture (Female system (in red) = Ov- ovary, vt- vitelline glands, Vt.d-
viteline gland ducts, Oo- ootype, M-Mehl's gland, V- vagina, aperture; Male system (in blue)
T= testis, V -vas deferens, C-cirrus),. (Source:
http://www.path.cam.ac.uk/~schisto/general_parasitology/Pseudo.pro.gif)
The excretory/osmoregulatory system

Osmoregulation and excretion are effected by the protonephridial system. These are
horizontal tubes terminating in flame cells and have their contents drained into four
collecting tubes. Two of the lateral tubules are ventro-lateral and another two are dorso-
lateral. These tubes unite anteriorly in the scolex and open to the outside through two
pores.
The nervous systems.

Two pairs of cerebral ganglia occur in the scolex and these are linked to two ventral nerve
cords that run posteriorly, and like that in the monogeneans, there are transverse nerves
linking the two major nerve cords.
The Nematoda (the thread worms).

There are between 16 000 and 20 000 species of nematodes making them the third most
speciose phylum after the arthropods and molluscs. They are both aquatic and terrestrial.
Aquatic forms occur in both marine and freshwater. The terrestrial forms inhabit all habitats
where there is a thin film of moisture. They occur as both free-living and parasitic form.
Free-living forms are either carnivorous feeding on other small invertebrates and
microscopic forms like bacteria or detritus feeders feeding on remains of plants and
animals. Parasitic nematodes feed on plant and animal tissues and fluids. Many feed on
insects as entomopathogens (Bush et al, 2001)
Nematodes (nemas =threadlike), as their name suggest, are typically threadlike (Figure 50)
with both ends tapering (fusiform) or rounded (filiform). In some, sexual dimorphism is the
norm, with the male being filiform or fusiform and the female, especially when mature
attains some grotesque forms due to high fecundity. These (like Heterodera, Tetramere and
Meloidogyne species) may attain pear shapes (succate) or spherical forms as the inside
become engorged with developing eggs. Characteristic of all these aberrant variations is the
appearance of a constricted neck like region after the head region. Nematode sizes of range
from 0.3mm to 8m long. The largest nematode on record is Placentonema gigantisim
isolated from the placenta of the blue sperm whale.
Figure 50: The characteristic threadlike form of

nematodes (Source:
http://t3.gstatic.com/images?q=tbn:ANd9GcTH4qV583PF1KFNGC2AIZBxpKxNOB_SRokifa-
5srKU0FptEdSl)
Figure 51: Anterior end of nematode with lip-like structures

(Source:http://biodidac.bio.uottawa.ca/thumbnails/images/NEMA004B.gif0
The body of the nematodes is covered by a cuticle which may be variously ornamented or
sculptured with annular rings. Anteriorly is the mouth which is surrounded by lip-like
structures varying in number between zero and six? In the primitive aquatic forms, the six
lipped condition is quite common and in others there is reduction as a result of fusion or
lose (Figure 51). The anterior is also equipped with sensila, numbering 18 in the primitive
aquatic form. These sensila are in the form of amphids, which are openings into the body,
below which are hidden a glands. These, together with the papilla, cephalic alae and bristle-
like structures serve for chemosensation and mechanoreception. Posteriorly the body
tapers and there is a subterminal anal or cloacal opening. The cloaca (present in males) is
usually provided with a gubernuculum which guide the spicule out of the cloaca during
copulation and a bursa which tenaciously holds onto the females during activities preceding
procreation. In some nematodes (the Phasmida) there is an opening in the terminal region
which opens into a gland below the surface called the phasmida. This is also chemosensory
like the anterior amphids and is of great taxonomic importance at higher ranks of
classification. The anus like the mouth is also surrounded by sensory structures like anal or
cloacal papilla and caudal alae.

The mouth, surrounded by the lips lead into a buccal cavity, then a short tube leads into a
bulbous pharynx used to pump food into the intestine of the animal (Figure 56 and 58). The
intestine, which is single layered and often synctyial leads to a subterminal anus. The
mouth, in plant parasitic nematodes is equipped with a hollow spear like structure, the
stylet, which is used during feeding to pierce plant tissue and direct the fluids into the
buccal cavity like a hypodermic needle. In animal parasitic nematodes like hookworm
(Anklostoma), the buccal cavity is provided with a tooth-like structure called an odontostylet
by which the bearer uses to hang onto the surface of its host (Figure 52). The foregut
(stomodeaum), buccal cavity, pharynx and oesophagus and terminal (proctodeaum), rectal
and annual regions are covered by cuticle which is continuous with the outside cuticle and
hence shed during ecdysis.
Figure 52: The odontostylet of the

dog nematode (Source:http://www.astrographics.com/GalleryPrints/Display/GP2097.jpg)
Body wall
The nematode body wall is covered, as alluded to earlier, by a cuticle overlying a synctyal
epidermis. The nature of cuticle layers and their numbers vary from group to group, and can
be from two to nine, but basically it is composed of an outer epicuticle followed by a median
zone and finally an inner collagen fibrous layer overlying the synctyial epidermis. The
callogenous fibres are made up of three layers whose fibres are crossed so as to resist any
stretching due to the high hydrostatic pressure inside. Below the body wall are longitudinal
muscle fibres responsible for the adulatory locomotion of nematodes. There are four muscle
bands, two ventro-laterals and two dorso-laterals (Figure 53). The muscles are constructed
in manner unique to the Nematoda, terminating in U-shaped cells which abut with the
epidermis and elongated arms that directly 'enervate' (link to) the nerve chords (Figure 54).
The muscle fibres are arranged in bands inside the body wall to correspond with areas not
accommodating the nerve chords.

Figure 53: Section of a nematode in the Pharyngeal area showing the
four muscle bands in between the nerve cords (Source:
http://www.ucmp.berkeley.edu/phyla/ecdysozoa/nematodexs.gif
Between the body wall and the gut there are no tissues, giving the characteristic
pseudocoelomate condition of the nemas. The pseudocoelom is fluid-filled and contains,
very rarely some coelomocytes, usually not numbering more than four. The fluid in the
pseudocoelom acts as a hydrostatic skeleton as its acts against the muscular activities of the
longitudinal muscles. The gut is single cell layered and its contents are under the high
hydrostatic pressure of the fluid in the pseudocoelom, only the two valves between the
anterior gut and the posterior gut serve to prevent the gut contents from being forced out
anteriorly or posteriorly to the exterior.
Figure 54: The characteristic link between muscle

cells and the nerve cord in nematode. (Source: http://1.bp.blogspot.com/-
Twtn6oPvUNs/TbBAtukFyOI/AAAAAAAAANo/UKIKq3mCjEI/s1600/NematodeMuscleNerve.gi
f)

The nervous system
Figure
56: Nematode Nervous system and associated structures. (Source: http://sharon-
taxonomy2009-p2.wikispaces.com/file/view/nematode.gif/99304909/nematode.gif)
An anterior nerve ring (ganglion) around the oesophagus (Figure 56) is all that the nematode
has to show for a brain (Bush et al 2001). A number of nerve chords course anteriorly from
the nerve ring to enervate the sensory structures in the head region (the papilla and
amphids). Also from the nerve ring, two major nerve chords (ventrally and dorsally) and two
minor nerve chords (laterally) run to the posterior. In the anal region, another nerve ring,
the anal nerve ring connects the longitudinal nerves and from this ring emerge nerves that
supply the phasmida and the caudal alae and anal papilla (if present).
The excretory system.
The nemas have an excretory system peculiar to them (Figures 56 and 57). It consists of two
rennete cells which are medially situated in an H-shaped system of tubes. Each of the two
arms of the H system runs subepidermally posteriorly and anteriorly along the length of the
worm. The cross of the H system occurs subterminally anterior in the body and a tube
originating meridonially on the cross leads to the outside to release the excreta.

Figure 57; Excretory system of nematode. (Source:
http://www.wormatlas.org/ver1/handbook/fig.s/exc1.jpg)

Nemas are typically dioecious and the female is usually larger than the male (Figure 58). The
male reproductive system consists of testis, which lie anteriorly in the coelom. From the
testis emerge a vas eferentia which courses posteriorly, merging into a seminal vesicle
before continuing as the vas deferens (sperm duct) which posteriorly joins the rectum to
form the cloaca. The latter is provided with a one or two spicules used to open the valva of
the female during copulation against the high hydrostatic pressure of the pseudocoelom. A
gubernuculum and bursa are also present in some, and help, as alluded earlier, to guide the
spicule out and hold the female respectively. The female reproductive system is usually
double. A vulva, in the posterior half of the worm opens to the outside to the ventral side.
This leads into a short vagina, from which a bifurcation leads to two uteri, seminal
receptacles, oviducts and ovaries. In some nemas the two loops of the two sets are directed
either side of the vulva (the didelphic condition), in others both are directed anteriorly (the
prodelphic condition). In still others, both are directed posteriorly, (the opisthodelphic
condition).

Figure
58: The diocieous condition of most nematodes- male and female separate (Source:
http://t0.gstatic.com/images?q=tbn:ANd9GcT5C_ErKTmtW6AKvlY1wKwbkYQgtBbOIN2u1Yp
ltvyUZlDBVXuKcw)
Basic life cycles
The life cycle of nematodes typically consists of an egg, four larval (juvenile stages) and an
adult (Figure 59). In the simple life cycles as that exhibited by the parasitic Strongyllides,
eggs produced by the adults in the gut of a vertebrate host and are voided together with
faeces. These hatch to produce a larval stage 1 (juvenile stage 1=J1) which is free living. This
molts to give rise to a J2 and the J3 which is the infective stage. This penetrates a host,
circulate within the blood stream and lodges in the lungs and molts into J4. From the lungs
they enter the trachea from where they are coughed and brought up the pharynx only to be
swallowed into the gut. Once in the gut, the J4 molts into the adults which mate and lay
eggs for the next generation. In the Anklostoma, there is either a free living cycle as well as a
parasitic life cycle. Eggs are voided together with faeces and the hatch in the soil to give rise
to J1 which in turn gives rise to the J2 and J3. The J3 can either reinter the host or enters a
free-living cycle

Figure 59: Basic life cycle of a
nematode. (Source: http://www.personal.psu.edu/ncj111/images/Lifecycle.gif)
Classification of nematodes.
There are about 1600 to 2000 described nematodes, with up to 1million yet to be described
(Barnes, 1993). These nematodes are classified into two classes depending on whether they
bear or do not bear a phasmida. The two classes are the Secenentea and Adenophorea.
The Annelida: The Segmented worms.

The Annelida are a medium sized phylum of more than 9,000 species of worms. Most
species prefer aquatic environments, but there are also a number of well known terrestrial
species. Only a few species of annelids are commonly known by laymen, these include the
Earthworms that work so hard to make our soils healthy, the Rag worms and Lugworms
used by marine fishermen and the much smaller Tubifex or Red worms used by aquarists to
feed their fish. In many countries people are still familiar with Medicinal leeches, and people
who live closer to nature are naturally more familiar with a much wider range of Annelids
than those who live in cities.
The majority of annelids are free-living although a relatively high proportion also occurs as
parasites. They occur in both terrestrial and aquatic environments. In the aquatic
environment, the marine forms are predominant and only very few are freshwater. The
terrestrial forms tend to inhabit the moist environments in mud and very wet swampy
areas.

Annelid sizes vary. The giant in the annelid world is the Giant Earthworms, of which
Michrochaetus rappi (Michrochaetus michrochaetus) is the largest, with an average length
of 1.36 m. A new find has recently been recorded that measured 6.7 meters (22 ft) in length,
and 2cm in diameter. Larger worms have been reported but not scientifically proven. The
smallest Annelid known to science is Chaetogaster annandalai which is full grown at 0.5 mm
(http://www.earthlife.net/inverts/annelida.html).
Annelids are coelomate (Figure 60), bilateral, true segmented worms. Their bodies are
typically vermiform and segmented, being composed of a series of repeated body units
(Figure 61). Anteriorly is a head region, the prostomium, and posteriorly is the terminal
pygidium and intervening, are a series of segments of varying numbers. Growth occurs both
laterally, by enlargement of the segments during the juvenile stages, and through the
addition of new segments just anterior to the pygidium.
Figure 60: The coelomate condition found in annelids. (Source:

http://www.utm.edu/departments/cens/biology/rirwin/coelom.GIF)
Segment production occurs throughout life in some annelids, and others production stops
once a certain size has been attained. They are characterized by the following features: an
entire gut (figure 61) with mouth at the anterior and terminating posteriorly in an anus,
segments typically separated by septa, moderate cephalisation, with head bearing sensory
organs and internally a supraoesophageal ganglion connected to a suboesophageal ganglion
by circumoesopahageal commissures. The suboesophageal ganglion sends, posteriorly, a
ventral nerve chord with segmental ganglia and nerves in each segment (Figure 62). A
closed blood circulatory system with dorsal blood vessels carrying blood anteriorly and a
ventral blood vessel carrying blood posteriorly serve to move materials in these animals.
Each segment has its supply of blood vessels branching from the dorsal blood vessel. The
gut, nerve chord and blood vessels run though the septa (Figure 63). Body wall typically
bearing protruding chitinous cheata and leg like appendages or parapodia.
Their bodies are covered by cuticle overlying an epidermis. Below the epidermis are circular
muscles followed by longitudinal muscles which are, usually in association with the body
appendages. The body cavity is a true coelom, and being fluid filled it plays a central role in
locomotion. Within the coelom, are housed the excretory system (metanephridia and
coelomoducts) and the reproductive system (testis, sperm ducts ovaries and oviducts). The
Annelida are typically hermaphroditic.
Figure 61; Digestive system of L. terretris is typical of many annelids.
Classification of annelids.
Annelids are classified into three classes, The Polycheata, the Clitella (subclasses
Oligochaeta, Hirudinea and Branchiobdella) and the Aleometamorpha.

Figure 62: Nervous system of an annelid. (Source:
http://cronodon.com/files/earthworm_CNS_lateral.gif
Figure 63: Model of Circulatory system of annelids and associated structures (Source:
http://cronodon.com/files/Lumbricus_circulation_POV.gif)

Class Clitella.
These are the Annelida close to the primitive annelids with typical metameric segmentation
and having a clitellum, a group of segments that secrete mucous used in cocoon moulding.
Subclass Oligochaeta.
These bear most of the primitive structures. They are a characterized by few cheata or few
groups of cheata. Amongst these is Lumbricus terrestris, an extensively studied laboratory
animal which bears many of the features in this group and will be presented here because it
is typical of the group.
L. terrestris is typically terrestrial. Its body is covered by a cuticle which is kept moist (Figure
64) by mucous produced by gland cells in the epidermis. Each segment bears four pairs of
cheata in the lateral and ventro-lateral sides of the worm. Each segment bears
metanephridia which open into the coelom through a nephrostome and to the outside
through a nephridiopore. In the prostomium is a suproesophageal ganglion connected to
the suboesophageal ganglion by circumoesophageal commissures and the typical
ganglinated ventral nerve chord run posteriorly (Figure 62). The dorsal blood vessel which
runs anteriorly gives of branches which run ventrally as semi loops which become swollen in
the pharyngeal area to give rise to lateral hearts (Figure 63). There is an elaborate gut, with
mouth, buccal cavity, oesophagus, crop, gizzard, intestine and anus (Figure 61).
Figure 64: L. terretris. Note the

moist glistening body.
(Source:http://upload.wikimedia.org/wikipedia/commons/3/30/Regenwurm1.jpg)

L. terretris is hermaphroditic with a pairs of female and male gonopores opening ventrally to
the outside near to the anterior half of the worm. The female system consists of ovaries and
oviducts that lead to the outside through paired gonopores. The male system consists of
testes, seminal vesicles, spermotheca and sperm ducts which lead to the outside through
the paired male gonopores. There is swollen glandular region called a clitellum close to the
gonopores which is responsible for mucous production. The mucous is responsible for
holding two worms during sperm transfer as well as formation of egg cocoon.
Prior to sperm transfer, which is mutual, two worms pair out and get involved in a complex
presperm transfer behaviour involving massive secretion of mucous with which the pair
smears itself with. The two worms then lay antiparallel with ventral sides abutting against
each other (Figure 65). The copious mucous secreted by the clitellums of each worm hold
the two together. Each of the pair then releases sperms which move externally towards the
seminal receptacle of the other for temporal storage. After sperm transfer the two separate
and each release eggs which move externally to the clitellum region where they are
temporarily stored under the mucous. The mucous, together with the eggs then slides
forwards passing through the spermotheca where the eggs are fertilized before being shed
enclosed in the cocoon which sloughs off the head of the worm (Figure 66). The eggs hatch
to give young ones similar in all aspects to the adults

Fi
gure 65: Sperm transfer in L. terrestris. (Source:
http://upload.wikimedia.org/wikipedia/commons/thumb/f/f6/Mating_earthworms.jpg/800
px-Mating_earthworms.jpg)
L. terestries is valued by many farmers who associate its presence with high soil fertility. This
is largely due to its feeding behaviour which involves feeding on the remains of plant
remains together with the soil particles. It does literally eats its way as it burrows into the
soil anteriorly and egests through the other terminus casts of soil. The soil casts which
constitute the faeces of the worm are mainly soil and much of the ingested organic material.
The organic material has however been greatly made more particulate and hence easier
for microbial decomposition. These worms thus make the soil more fertile by partially
degrading plant debris as well as creating burrows which help aerate the soil and thus
favouring microbial proliferation and hence nutrient release to crops.

Figure 66: Cacoons of L. terretris
(Source:http://upload.wikimedia.org/wikipedia/commons/0/01/Earthworm_-
_L._terrestris_cocoons.jpg)
Subclass Hirudinea
The leeches, as they are commonly called have largely been modified for a parasitic mode of
life, although some are still free-living. They have secondarily lost the internal segmentation
as there are no longer any septa internally. Segmentation is still quite evident as shown by
the segmental ganglia in the ventral nerve chord. The associated segmental nerves and the
segmental vessels of the closed blood circulatory system also betray the typical annelid
metameric segmentation. The surface annulations which occur in this group which may be
misconstrued as evidence of segmentation do not, however, correspond to the internal
segmentation.
Figure 67: A hirudina worm- note the two

suckers used for host attachment.
They typically have 33 segmental ganglia, evidence that they have the same number of
segments though now obscured. Their parasitic mode of living has seen them evolving
attachment organs in the form of suckers (Figure 67). The two suckers., one anterior
(covering segments 1-4) and another posterior (covering segments 25-33), serve to attach
them to their vertebrate hosts from where they obtain copious blood meals after slashing
the flesh with their mandibulate-like jaws. The victim is oblivious of the damage being
effected as the leech, apart from producing saliva which prevents the blood from
coagulating, also produces an anaesthetic to soothe the pain. Once engorged some of these
leeches can go for up to a year without getting another blood meal as digestion is very slow.

Figure 68: Internal structure of
Hirudina -digestive and reproductive systems. (Source:
http://bio.classes.ucsc.edu/bio136/annelids/hirudinea/hirudinea2.gifhttp
Leeches are typically hermaphroditic. A series of testes are connected to short vas eferentia
which in turn lead to sperm ducts that open into an ejaculatory organ, the penis which
protrudes through the common genital opening during copulation. The female reproductive
system consists of a pair d ovaries which lead, through an oviduct, to a vagina which also
open to the outside through the common genital opening. Sperm exchange is mutual and
fertilization is internal. Fertilized eggs are released into a cocoon and hatch in the soil.
Class Polycheata
These are largely marine although there are also many freshwater forms. In marine
environments the numerous species of Polychaetes play a fundamentally important role in
the maintenance of food chains and the whole ecological balance of the seas, thus
supporting the seemingly endless stocks of fish we like to eat.

Figure 69; A Polycheata (Eurythoe
complanata- an errant polycheate) (Source:
http://animaldiversity.ummz.umich.edu/site/resources/Grzimek_inverts/Polychaeta/Eurytho
e_complanata.jpg/badge.jpg)
Polycheates are characterized by bearing the typical biramous parapodia with groups of
chaetae and a head region typically bearing sensory structures (Figure 69). Supporting the
parapodia are proteinecious rods called aciculums. The detailed structure of parapodia
varies from group to group. The polychaetes have secondarily lost internal septa, and
segmentation is still quite evident from the outside. They are commonly grouped into two,
the motile or errant polychaetes and the sedentary ones. The errant polychaetes are in turn
divided into two easily identifiable orders on the basis of the nature of the mouth and
proboscis. The two orders are Phyllidacida and the Eunicida.
The Phyllodocidae (Figure 70) are characterized by a highly agile or eversible proboscis. The
mouth typically borne at the tip of the proboscis bears jaw like structures and teeth like
structures which are used to stab prey and predator. The Eunicida bear a protrusible mouth
bearing a sclerotized jaws, spine like mandibles and maxilla of varying numbers. (Fig?)

Figure 70: A Phylodocidae polycheate
(Source:http://content5.eol.org/content/2011/02/16/03/18479_large.jpg)
The Polycheata exhibits epitoky, which is characterized by metamorphoses and swarming

upon reaching sexual maturity. Epitoky manifests itself in two ways; epigamy and
schizogamy. In epigamy, upon reaching sexual maturity, the posterior becomes modified by
way of having its parapodia elongating and becoming fleshy. In schizogamy, the posterior
end becomes modified into bag full of eggs which eventually are pinched off from the rest
of the body. These can regenerate complete worms.
The sedentary worms are, unlike the errant ones, typically sessile, leaving in mud or
variously modified burrows from which they emerge, usually at night to mate and feed.
Their mouths are typically surrounded by tentacle-like structures used in filter feeding.
Some of these tentacles are prehensile and are used to get hold of food and bring it to the
mouth. The thoracic regions also typically bear a lot of appendages.

Figure 71: Eunicidae polycheaate
(Source:http://polychaetes.info/sites/polychaetes.info/files/images/Onuphidae-
Eunicidae%20juv.%20sp.%20A.jpg)
Phylum Mollusca
The mollusc are the soft bodied animals thought to have evolved from the flatworm but
underwent wide adaptive radiation to be become the second largest animal phyla after the
Arthropoda. They occur in both terrestrial and aquatic environments. In the aquatic
environment, many are marine and others have moved into freshwater. One group, the
Gastropoda have taken the challenge to make the terrestrial environment their second
home after the marine environment.
Molluscs, as the name implies are soft bodied animals. Because of the very unifying features
or defining features (synapomorphies) occurring in this group we follow the tradition of
described a primitive hypothetical (Archetype or supposed ancestral) mollusc (Figure 72)
with most of the features present and then as we move on to the different groups we look
at the excerptions.

Mollusca are bilateral, coelomate animals that are primitively elongated in the dorso-ventral
axis. Primitively their bodies are divided into three major regions; a head, foot and visceral
mass. The latter houses the internal organs (the stomach, intestines, heart, gonads and
nepridial system). The head bears the sense organs, like the tentacles and eyes as well as
the mouth opening. The visceral mass is covered by a flap of tissue, the mantle which
primitively secretes a protective shell that covers the soft tissue dorsally. The shell is
composed of a conchiolin periostracum layer outside, followed by a columnar calcite layer,
then laminated calcite that is often prismatic nacreous. Some shells are made of aragonite.
The shell is typical conical and is secreted gradually by the margins of the mantle cavity.
Some have secondarily lost the shell, reduced, or internal.
Figure 72: Hypothetical (Archetype) mollusc. (Source:http://www.sciencewithme.com/wp-

content/uploads/mollusks2C.jpg
The reproductive system is composed of two gonads next to the coelom that surrounds the
heart, shedding their ova or sperm into the coelom from where they are collected by the
nephridia which releases them out into the mantle cavity. Fertilization is thus primitively
external and molluscs using this method remain typically monocieous, although some have
become hermaphroditic and exhibit internal fertilization. After fertilization, the egg hatches
a simple trochophore or veliger or a miniature adult.
The nervous system is primitively simple with a circumoesophageal ring from which two
pairs of nerve chords run posteriorly as the visceral and pedal nerves, enervating the viscera
and foot respectively. The nervous system as we will see in some molluscs like the
octopuses (Class Cephalopoda) is highly developed and aids learning capabilities.
The gut is entire, with a mouth anteriorly which leads into an oesophagus. The mouth is
provided with a rasping tongue or ribbon-like organ, the radula, which sits on a collagenous
like pedal, the odontophore. The radula (Fig 73) is equipped with chitinous teeth-like
structures arranged in transverse rows which are used to rasp food materials and thus take
them into to the buccal cavity. The odontophore together with the radula can be protruded
through the mouth by protractor muscles. With radula teeth held against the surface to be
rasped, the retractor muscles attached to the odontophore then pull the radula, thereby
scrapping the surface with the hooked teeth tearing off fragments which are swallowed
with the aid of copious mucus. Although mineralized with Ca 2PO4 and Fe3O4, the radula

teeth wear off with time, and the radula moves new teeth to the active portion by
continuously moving slightly forwards.
The radula has allowed molluscs to be efficient herbivores feeding on plant material, but in
others the teeth have become elongated and fewer to allow for a carnivorous life. These are
used to subdue prey by stabbing. Others are even provided with in sunken poison glands to
be injected hypodermically into the prey.
The oesophagus leads to the stomach which is peculiar in having some diverticula which
terminate in digestive glands where digestion takes place. The stomach has a style region
which is ciliated and hence responsible for propelling food from the oesophagus by the
continual beating of its cilia. Once the food is in the stomach, it is sorted according to size by
a sorting region which is not chitinous as opposed to much of the stomach which is
chitinous. The smaller particles are diverted into the digestive glands through the diverticula
ducts and the larger ones are led to the intestines towards the rectum and anus for
egestion. Mollusca are eucoelomates although the coelom is greatly reduced so as to only
accommodate the heart in the pericardium and the nephridia. The gonads open into the
coelom as well. Posteriorly, between the overlapping mantle and the foot is mantle cavity
into which opens the anus and nephridial system. Also housed in this cavity is usually a pair
of a special kind of gills found in the mollusc, the ctenidia. These are constructed on a
ridged frame of supporting rods from which, on either side, arise flaps of triangular
filaments in rows. This is the typical bipectinate ctenidia (Fig 75). The ctenidia are highly
vascularised, with blood in the filaments flowing counter current to the water which flashes
from below and escapes dorsally. Blood from the ctenidia is channelled directly to the heart
for delivery into the open circulatory system composed of open sinuses between the
tissues.
The foot of the mollusc is usually muscular and is used in creeping on the surface. The foot
has been variously modified with the result that the molluscs are amongst the slowest and
fastest moving invertebrates. In others it has been modified into tentacles with suckers for
prey capture.

Figure 73:
Structure of radula (Source: http://barnegatshellfish.org/images/snails/_radula_02.PNG
Figure 74: Radula in action during feeding. (Source:

http://barnegatshellfish.org/images/snails/_radula_03.PNG)

Figure 75; Bipectinate ctenidium (Source:
http://i.quizlet.net/i/7snFi2A2VMZfCiGboLbgQQ_m.jpg
Classification of the Mollusca.
The Mollusca are one such group which has been able to exploit all the potentialities in the
primitive body plan and this has paid off. To date about a 100 000 species of molluscs have
been described and these are placed into 7 classes, viz Cheatodermomorpha,
Neoniomomorpha, Monoplacophora, Polyplacophora, Gastropoda, Bivalvia and
Cephalopoda.
Class Cheatodermomorpha
These are thought to have retained only a few of the primitive molluscan features. They are
wormlike and have lost the shell, the eyes and tentacles (Fig 76). They live in mud burrows
with heads down, feeding on the organic matter in the soil as they burrow. The anterior
placed mouth is equipped with the radula housed in a radula sac. The foot is greatly
reduced and the mantle cavity open posteriorly usually protruding from the burrow. The
mantle cavity houses the opening to the anus and nephridia and is equipped with a single
pair of bipectinate ctenidia. Lose of shell has been compensated by development of scale-
like structures that cover the vermiform body. Fig ? shows the variety in this relatively very
small group of molluscs.

Figure 76: A Cheatodermomorpha mollusc
(Source:http://www.abdn.ac.uk/~nhi708/classify/animalia/mollusca/chaetodermomorpha/c
haeto.jpg
Class Neoniomomorpha.
These, like the Cheatodermomorpha, have lost most of the primitive molluscan features.
They are vermiform, elongated in the posterior anterior axis, but unlike the former, their
bodies are laterally compressed (Figure 76). They possess a reduced, ridged foot which
usually lay concealed in a ventral groove. The mantle overlies the animal dorsally and
overlaps in the lateral margins, usually concealing the much reduced foot. They have lost
the ctenidia, and in its place, posteriorly is a gill. The mouth opens ventrally anteriorly,
housing the radula inside a radula sac.

Figure 77: Neomenimorpha
molluscs (Source:
http://www.abdn.ac.uk/~nhi708/classify/animalia/mollusca/neomeniomorpha/neo.jpg
Class Monoplacophora.
These have retained the primitive shell overlying the animal in the dorsal position (Fig 78
and 79). The shell is usually conical or cup-shaped with a foot ventrally and a head
anteriorly. Between the mantle margin and the foot that is in the mantle cavity are five to
six pairs bipectinate ctenidia and six pairs of nephridia.
Figure 78: A monoplacophoran mollusc (Source:

http://t1.gstatic.com/images?q=tbn:ANd9GcQzLyTVffPBSpw-
cryM09JRWanHdDjar2xusLtWD06l8icgWnPG_g

Figure 79: Photograph of a monoplacophoran mollusc.Source:
http://t1.gstatic.com/images?q=tbn:ANd9GcQzLyTVffPBSpw-
cryM09JRWanHdDjar2xusLtWD06l8icgWnPG_g
Class Polyplacophora.
These are generally known as the chitons. They have lost the anterior posterior elongation.
They are elongate to oval in shape with the dorsal part covered by 8 overlapping shell plates
(Fig 80). The mantle overlaps at the margins of the plates as a muscular ridge. Ventrally is a
foot and in the mantle cavity are varying numbers of gills ranging from 6 to 88. They are
known for their ability to cling tenaciously to rocks and other hard substrata. This, they
achieve by using their muscular mantle margins as suction cups. If detached from the
substratum, which requires a lot of effort, they quickly roll into a ball like a millipede, until
they secure another surface for attachment. They are known for their grazing habits on
algae attached on rock using their robust radula.
FIGURE 80: A Chiton molusc-Tonicella lineata. (Source:

http://upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Tonicella-lineata.jpg/220px-
Tonicella-lineata.jpg
Class Gastropoda.
These is the largest molluscan class with up to 75 000 species of both aquatic and terrestrial
forms. These primitively show the dorsal shell which houses the whole organism although
some have secondarily lost it. Primitively they have a well developed foot, a visceral mass, a
head with eyes and tentacles and a mouth bearing a radula. They are united by exhibiting
torsion at some stage in their development with many retaining it as adults. Torsion is a
phenomenon in which the visceral mass has been turned through 180 o with respect to the
head and foot so that the mantle cavity now opens to the anterior over the head (Fig 81).
Torsion is thought to occur as result of the unequal development of foot retractor muscles.
Figure 81: How the phenomenon

of torsion is thought to have occurred. The rotation hypothesis sketched in dorsal (A) and
lateral (B) view. Rotation of the entire visceropallium relative to the head and foot moves the
mantle cavity to an anterior position, crosses the pleurovisceral nerve connectives, and
changes shell orientation from exogastric to endogastric. a, anus; f, foot; g, gill; h, head; mc,
mantle cavity; pvc, pleurovisceral connective; sh, shell. (A) from Naef (1913). (Source
http://icb.oxfordjournals.org/content/46/2/134/F1.medium.gif)
Consequent to this, the anus and nephridia now release their egesta and excreta over the
mouth and hence fouling the whole anterior. This has, however, been ameliorated in many
groups by directing the fouled water current which baths the ctenidia in such a way that it
now escapes through an opening in the apex of the shell or the degeneration of the right
ctenidia so that water enters through the left side and escapes through the right side
thereby directing the fouled water away from the mouth (Fig ?). In some cases siphon like
structures have evolved which direct the movement of water in the incurrent as well as the
excurrent in such a way that minimum fouling occurs (Fig ?). In still others, detortion has
occurred. This involves the rotation of the visceral mass through 90 o so that the mantle
cavity is now directed to the side (Fig ?)
The bringing of the mantle cavity though with it inherent problems had the advantage of
bringing the ctenidia or gill to the front where there is fresh, undisturbed water for aerating
them. It also has the advantage of bringing the chemosensory organ in the mantle cavity,
the asphridium, into contact with fresh water and hence effectively picking up the chemistry
of the water.
The shell (in groups where it is present) has been variously modified as a protective shield
against predators into which the whole animal can withdraw, or as water proofing roof into
which the animal can withdraw and the opening sealed by an operculum which evolved
dorsally at the back of the foot mainly in terrestrial forms. The shell has also been modified
into a spiral or helical structure with an internal rod, the collumella forming the central axis.
In many forms the shell has been lost as alluded to earlier. Two major groups have emerged
in the Gastropods: the Prosobaranchs and Heterobranchs.
The Prosobranchia (Fig ?) retained most of the primitive gastropod features of torsion, well
defined head with eyes and tentacles, muscular foot for locomotion a paired bipectinate
ctenidial gaseous exchange apparatus and a helico-spiral shell. However there was a
tendency towards reduction of the gaseous exchange apparatus, with an initial loss of the
right bipectinate ctenidia leaving left bipectinate ctenidia, then partial loss to leave
monopectinate ctenidia. This was also accompanied by the loss of the right kidney, the right
auricle of the heart and the right osphridia.
In the other group of the Heterobranchia (Fig ?) there remerged a tendency towards
reduction, internalization or loss of the shell; a reversal of torsion; loss of the ctenidial
gaseous exchange apparatus and its replacement with a lung or bodily folds and tendency
towards hermaphroditic condition. The first groups to emerge were the marine
Opisthobranchia then the Pulmonates. The Pulmonates (snails and slugs) become terrestrial
although some remained aquatic. These developed a gaseous exchange apparatus for the
terrestrial environment, the lung ((Figure 82). This was achieved by extensive folding of the
wall of the mantle cavity enclosed and with a contractile opening called the pneumostome.
Most of them retained the helical shell although with thinner constituents, but others, the
Symnomorpha completely lost the shell and become the land slugs (Figure 83). Also to
evolve from the Heterobranchia were the Germnomobranchia and the Alligastropoda. The
former retained most of the Opisthobranchia and pulmonate mollusc features and the latter
retained most of the Prosobranchia and Opstobranchia traits.

Figure 82. A Gastropod snail, the land snail (Helix pomatia (Source :
http://upload.wikimedia.org/wikipedia/commons/thumb/6/64/Grapevinesnail_01a.jpg/220
px-Grapevinesnail_01a.jpg0
Figure 83: The Banana slug-Ariolimax columbianus. (Source:

http://animaldiversity.ummz.umich.edu/site/resources/jack_burch/097.rjb1.jpg/badge.jpg
Class Scaphopoda.
These are the tusk shells (Fig 84), so called because of their elongated shells that resemble a
tusk. The bodies are elongate in the dorso-ventral axis and the visceral mass is completely
enclosed by the mantle which secrets the hollow tusk like shell open at either end. They live
in vertical burrows with their heads and foot downwards feeding off the detritus in the mud
burrows. Protruding downwards is the cylindrical foot which is modified as a burrowing
apparatus together with the head with a mouth anteriorly. The mouth is surrounded by
adhesive rod like clubed feeding structures called captacula. The mouth is equipped with
radula which is used to scrape on the detritus below. The mantle cavity is posterior and into
it drains the egesta and the excreta.

Figure 84: A Scaphopoda mollusc INTERNAL ANATOMY
(Source:http://t2.gstatic.com/images?q=tbn:ANd9GcQoBdMvqAdgME2NnEmebEu8-
BI8NwuBGtgXZZBAnDdUSYy-zi2l
Figure 85: A Scaphopoda mollusc EXTERNAL

MORHPOLOGY
(Source:http://t2.gstatic.com/images?q=tbn:ANd9GcQoBdMvqAdgME2NnEmebEu8-
BI8NwuBGtgXZZBAnDdUSYy-zi2l
Class Bivalvia.
The Bivalvia include the clams, scallops, mussels, oysters. These are laterally compressed
molluscs with bodies enclosed in a two shell valves at the lateral sides. The shell valves are
connected dorsally by an elastic ligament which tends to make the ventral edges of the
valves open slightly. The animal is enclosed within the valve and is restricted to the dorsal
aspect of the shells leaving a space ventrally and latero-ventral through which the muscular
foot may protrude in the ventral space between the valves. A small head appears anteriorly.
This bears the mouth which is equipped with the radula. The bipectinate gill is replaced by a
monopectinate gill. The elastic dorsal ligaments tend to pull the valves open and these are
countered by the action of a pair of abductor muscles which pull the valves together so that
the whole animal is enclosed. These almost sedentary animals may be attached to the rock
through their shells by byssal threads. Figure 86a and b show the external morphology and
internal anatomy of the bivalvia respectively.
Figure 86a: External morphology (shell, right half) of a freshwater Unionidae bivalve,
Ceolatura kunenensis, collected from Tokwe River, Zimbabwe. Magn x 8. (Photo by CM
Makaka)
Bivalvia classification into subclasses is mainly based on gill structure. There are three
subclasses, two major and a minor one. These are the Protobranchia and Lamellibranchia
and Septibranchia

Fig 86b: Bivalve internal anatomy- Source -
http://www.cbu.edu/~seisen/Mollusca/sld025.htm
.
Class Cephalopoda.
Amongst this group are the most agile, largest and intelligent of all the invertebrates. The
largest squid on record, the giant squid measured 20 meters and weighed 500kgs. Their
intelligence is also quite remarkable and this correlates with their highly developed brains.
One Octopus (named the Oracle Paul), kept in an aquaria in a Germany zoo, generated a lot
of interest during the 2010 FIFA World Cup Soccer Tournament in South Africa. It was able
to successfully predict all the score margins of the Germany team well in advance!
Cephalopoda are elongated in the dorso-ventral direction and they have completely lost the
foot so that embryonically the tissue which gives rise to the foot develops into tentacles or
arm like structures surrounding the mouth. They have closed circulatory system and ganglia
fused to form a miniature brain in a cartilaginous cranium. The mantle cavity opens
posteriorly and it is usually provided with a siphon or funnel which directs the water
entering anteriorly. The exhalent current is pushed out weakly or massively ejected
forwards thereby propelling the animal backward in typical jet-propulsion fashion as occurs
in the squids and octopuses. The heads are equipped with jaws and a radula which
macerate the food. In some groups like the octopuses the tentacles around the mouth have
sucker like structures to achieve a strong grip on prey. In some cases they are prehensile
and serve to bring forth food to the mouth.

The Cephalopoda are an aquatic group the majority being marine, though a few are
freshwater. There are about 800 species of described cephalopods. They occur at every
depth from the abyssal zone to the intertidal zones and they are particularly abundant along
the equator. Amongst these are those that have retained the shell like the Nautiloids (Figure
87), those that have reduced the shell and internalized it like the squids (Figure 88) and
those that have completely discarded the shell, the Octopoda (Figure 89).
Figure 89: The common Octopus Octopus vulgaris-

(Source:http://animaldiversity.ummz.umich.edu/site/resources/jo_okeefe/octopus_101_486
7.jpg/badge.jpg)
Figure 87: The chambered nautilus-Nautilus pompilius (Source:

http://animaldiversity.ummz.umich.edu/site/resources/Grzimek_inverts/Cephalopoda/Nauti
lus_pompilius.jpg/badge.jpg)
Figure 88: Longfin inshore squid- Loligo pealeii (Source:

http://animaldiversity.ummz.umich.edu/site/resources/Grzimek_inverts/Cephalopoda/Lolig
o_pealeii.jpg/badge.jpg)
The Cephalopoda are now only represented by two distantly related subclasses, the
Nautilodea (Nauttilus and Allonautilus) and the Coleoidea (octopuses, squids and cuttlefish).
In the Coleoidea the shell has been internalized or is absent and in the Nautiloidea the
external shell has been retained. The nautiloids have retained the shell which has become
planospiral and composed of a series of chambers (Fig 86). Only the youngest, anterior most
chamber is inhabited by the animal though a thin filament of tissue (siphuncle) runs back
through the other closed chambers. In this group, which is largely sessile, secretion of
chamber occurs anterior most in the planospiral shell and the animal moves into the new
chamber, and closes the one behind by secreting a shell wall. Water enters the mantle
cavity anteriorly and it is pushed out by muscular contraction of mantle cavity resulting in
movement.
Phylum Arthropoda
The arthropods are a loose grouping of metazoans of diverse characteristics unified by a few
shared characteristics. As a group, the arthropods are thought to be polyphyletic. From a
biological view point, they are a very successful group. They form the largest animal group
with over 85% of all animal species and have been around for a very long time. Fossil
records show that by the close of the Cambrian, some 505 million years ago, the arthropods
were already quite a massive group represented everywhere by forms which have changed
very little up to this day. Some authorities estimate that there are over one million extant
species of arthropods with a population estimated at 1018 at any given time. The insects
constitute the largest proportion of these arthropods, with close to a million insect species
and constituting about 80% of all metazoans.
Arthropods are characterized by bodies covered by cuticle exoskeleton and jointed

appendages. The exoskeleton (Fig 90) is chitinous and is usually hardened. The cuticle is
primitively divided into plates; dorsal (tergal), ventral (sternal) and lateral (pleural) plates
(Fig ?) with intervening section flexible and membranous. In terrestrial forms the cuticle is
largely sclerotized and in aquatic forms it is typically mineralized with calcium carbonate.
The skeleton serves largely as an impervious layer which limits water loss and hence
prevents desiccation in terrestrial forms. It also serves as a frame for muscle attachment
and a barrier to entry of pathogens. The cuticle is layered, with an outer epicuticle, a middle
exocuticle and an inner endocuticle. The epicuticle is usually covered with wax produced by
an epidermis lying below the endocuticle. The wax serves the critical role of waterproofing
the animal which otherwise would be prone to desiccation on account of the large surface
area to volume ratios obtaining in the majority of the arthropods The exocuticle is
mineralized through deposition of calcium carbonate in some aquatic arthropods.
The cuticle is a ridged inelastic covering and thus restricts growth once hardened. It is thus
periodically shed to allow for growth. Prior to shedding (moulting or ecdysis) a new cuticle is
formed just below the older cuticle by the underlying epidermis. The older cuticle, after
much of it has been digested by enzymes secreted by the epidermis and resorbed by the
epidermis, breaks along lines of weaknesses typically starting anteriorly and the arthropod
“craws” as it were, from the old cuticle. The new cuticle, which, soon after the old one has
been shed, is light in colour and still soft and allows for increase in the size of the animal.
This increase in size is achieved by either taking in water or air. Soon the new cuticle
hardens and the pale colour disappears as it becomes melanised.

Figure 90: The
exoskeleton of arthropods (Source:http://media-3.web.britannica.com/eb-media/99/5899-
004-DBA32515.gif)
The process of moulting is quite spectacular as one marvel at the animal literally crawling
out of its old cuticle and the sudden increase in size. It however, makes the animal
vulnerable to predators as the animal is less active. It has been noted that it is during
moulting that mortalities are high in the arthropods. Prior to moulting the animal thus tries
to secure a place where it is less prone to attacks by hiding in secluded places.
Arthropods as a group are also characterized by a metameric body, with each segment
primitively having a pair of appendages. Anteriorly is a legless segment, the acron and
posteriorly the pygidium, and in between a series of segments of variable number. This
construction, together with the structure of the nervous system, as we will note, has been
interpreted to suggest an annelid progenitor to the arthropods exhibiting itself in an
onychophoran-like animal.
Tagmatism is quite extensive in the Arthropoda. This is the organization of body segments
into functional units, such as a sensory and feeding region followed by a locomotory region
and finally a digestive and reproductive region. In many arthropods, these occur as the head
region, the thorax and abdomen, in others there is a head region highly fused to the thorax
to form a cephalothorax and then an abdomen. The anterior head which is made up of a
variable number of anterior segments bears sensory organs like antennae in the
Crustaceans and Uniramia. Housed in the head region is brain which is highly complex using
the standards obtaining in the invertebrate world. The dorsal brain is connected to
suboesophageal ganglia by commissures from where a ventral nerve cord runs posteriorly
with nerve ganglia in each segment issuing out segmental nerves.
The open circulatory system is a typical feature of the arthropods (Fig ?). Blood, more
correctly called heamolymph is pumped anteriorly into an artery usually by a dorsal heart or

a series of hearts and poured into the spaces between tissues to bath the cells. These
spaces, constituting the heamocoel collectively a result of a reduced coelom are used as a
vascular system. Gaseous exchange is effected through gills in aquatic arthropods, and in
terrestrial forms, book lungs and the tracheal system take charge of gaseous exchange. Gills
occur mostly in the aquatic crustaceans where they occur as modified structures on
appendages like legs (Fig?). Book lungs occur in the scorpions and spiders. They are
ectodermal in origin but they are sunk in pockets in the ventral aspect of the animal (Fig?).
The tracheal system of insects is composed of tubes which ramify the whole insect and open
to the outside through openings called spiracles which occur as paired pores, one on either
side of the insect body in the latero-ventral aspect (Fig?) Typically the tracheal system is
constructed on a system of tubes (trachea) which are invaginations form the outer body
wall. From each spiracle leads a short tube which links up with a lateral tube which runs
longitudinally inside the insect body. These are called lateral tracheal trunks. The lateral
tracheal trunks on either side are connected to two dorsal tracheal trunks and two by dorsal
and ventral trachea respectively. These also run longitudinally within the insect body. From
each lateral tracheal trunk also, in each segment, visceral trachea supply the gut.
The major tracheal tubes are at interval swollen into bulb like sacs which serve as oxygen
stores. The tracheal tubes issue out small branches, which, as the ramify the body, become
smaller and thinner, ending blindly as finger like tubes poking into and between cells
without disrupting their cell membranes. The insect tracheal system thus supply and
remove gases from tissues without the need of an internal fluid transport medium like
blood.
There is variability in organs used for excretion and osmoregulation. In the aquatic
arthropods (Crustacea) it is effected by coelomoducts which terminate in glands opening to
the outside, usually at the base of body appendages like legs and feeding appendages. In
land arthropods it is achieved by blind ending tubes which float in the heamocoel and open
into the hindgut. These tubes, called Malphigian tubules, actively pump mineral salts
together with nitrogenous wastes into their lumens and water follows secondarily from the
heamocoel due to osmosis. The Malphigian tubes empty their contents into the gut from
where water is reabsorbed and the rest are excreted together with faeces through the anus.
The million and odd Arthropods which are also united by their lack of ciliated cells in their
bodies are classified into three extant subphyla, the Chelicerata, Biramia and Uniramia.
Classes of Arthropods
There are five major classes of arthropods and each class can be separated based on shared
characters. These are the Arachnida, Crustacea, Diplopoda, Chilopoda and Insecta. The
Arachnida, together with minor classes like the Merostomata falls under the subphylum
Chelicerata. The Crustacea, Diplopoda, Chilopoda and Insecta are often grouped together as
the Mandibulata because of presence of mandubulate mouthparts. Many authors however
argue that the mandibles in the Crustacea are not homologous with those in the latter three
groups. The latter are also commonly grouped together as the Uniramia on the bases of
their having unbranched appendages unlike in the Crustacean where the appendages are
commonly branched and hence falling into their own subphylum the Biramia.

Subphyum Chelicerata.
The Chelicerata are bilateral, coelomate animals with both aquatic and terrestrial habits.
They are characterized by a metameric body which is elongate to spherical. The bodies are
typically divided into two regions; an anterior prosoma (cephalothorax) and posterior
opisthosoma (abdomen) (Figure 91). The prosoma bears appendages which are used for
feeding, walking and mechanoreception. These prosomal appendages are; anteriorly, a pair
of cheliceri, which are chelate or subchelate or spine like appendages, followed by a pair of
pedipalps which are chelate or leg like or feeler-like appendages then four pairs of walking
legs. Anteriordorsally on the prosoma are median and lateral osceli used in light
perception. A mass of segmental ganglion lies in the prosoma or may extend into the
opisthosoma. The mouth is anterio-ventral and is surrounded by the prosomal appendages.
The opisthosoma is appendageless and elongate to spherical. The anteriorventral mouth

leads into a through gut with an oesophagus which leads into a stomach typically provided
with digestive diverticula and opens posteriorly to the anus. Respiration, in aquatic forms is
effected by external gill books. In the terrestrial forms it is achieved by external lung books
housed in insunk ectodermal pockets or sieve trachea and tube trachea. Excretion is
through coxal glands which open at the base of prosomal appendages in aquatic
chelicerates and Malphigan tubules in terrestrial forms.
Chelicerates are typically gonochoristic with a single gonad opening ventrally in the second
opisthosomal segment. Although they engage in pseudo copulation, fertilization is external
in the marine forms, but internal in the terrestrial forms. In the latter, copulation or the use
of spermatophores is the norm. The young ones are exact replicas of the adults into which
they grow without any metamorphoses.
The Chelicerata stand out as a distinct group from other arthropods by their lack of
antennae and mandible. Their lack of mandibles deprives them of any opposable feeding
structures, as occur in the crustacean and Uniramia. Their carnivorous feeding behaviour is
consequent on the nature of the mouth. They use their chelicerae to pierce or rapture their
prey which they then spoon into the mouth using the pedipalps and other walking legs, or
flood it with digestive enzymes for predigestion to take place before they suck the partially
digested soupy materials with their buccal pumps. The lack of antennae is compensated in
some forms by the modification of the pedipalps into feelers. This lack of antennae is also
reflected in the nature of the brain which is devoid of the third ganglionic mass enervating
the antennae in the other arthropod groups.

Figure 91: Typical Chelicerate body (Source:
planhttp://wshs.wtvl.k12.me.us/~science/MrsV~Bio/Posting/arthropods3_files/image004.gi
f
Classification of the Chelicerata

The 63 000 species of Chelicerata are grouped into three classes; the Merostomata, the
Arachnida, and Pycnogona.
Class Merostomata.
This is a small group of about four species all classified into a single order Xiphesura. They
are all marine and characterized by a horseshoe shaped carapace covering the dorsal aspect
of the prosoma (Fig 92). The latter is composed of an acron and six segments. The carapace
is large and overlaps anteriorly so the mouth is ventral. It also overlaps laterally so that the
legs are invisible from above. Conspicuous on the carapace are paired compound eyes. All
the prosomal appendages are pincer like except for the last pair which bears spines which
are used for burrowing into the mud sands which constitute the greater proportion of the
coastal areas which form their home in their benthic mode of life. All the prosomal
appendages originate from the midventral line of the prosoma and the front ones are
radially arranged around the mouth and their bases from part of the oral region
(gnathobases) used in crushing the molluscan shells which form the major diet of these
animals.
The opisthosoma is flat and plate-like and wedges in a notch posteriorly in the horseshoe
shaped anterior carapace. Laterally, it is provided with spines, and quickly constricts to an
anal region from which a post anal caudal spines issues. Also characteristic of this group is

the presence of appendages on all opisthosomal segments, the first of which are tube-like
followed by six plate like appendages which are modified into gills, the first of which also
covers the genital aperture. Like all aquatic cheliceretes, fertilization is external. Excretion,
unlike in other aquatic Chelicerata is effected by Malphigan tubules.
The horseshoe crabs are nocturnal emerging at night from their burrows to feed and mate.
Figure 92: The horseshoe crab- Limulus polyphemus (Source:

http://animaldiversity.ummz.umich.edu/site/resources/jo_okeefe/horseshoe_crab_100_592
1.jpg/badge.jpg)
Class arachnida (scorpions, spiders, mites, ticks, daddy longlegs, etc.)
Mostly terrestrial and predatory, some secondarily aquatic and sap-sucking. Their bodies
are typically divided into two regions; prosoma and opisthosoma (cephalothorax and
abdomen). Six pairs of appendages are borne on the cephalothoraxes. These are chelicerae,
pedipalps and 4 pairs of walking legs. Abdomen with up to 3 pairs of appendages
(spinnerets)

Figure
93: Arachnids (Source:http://visual.merriam-webster.com/images/animal-kingdom/insects-
arachnids/examples-arachnids.jpg)

Subphylum
Biramia
Figure 94 A typical crustacean, a Decapod crab, Potamonautes warreni, collected from

Tokwe river, Zimbabwe
Class Crustacea (crabs, lobsters, shrimp, isopods, water fleas, copepods, etc.)
Primarily aquatic with most species being marine and few terrestrial representatives
Like the arachnids, their bodies are divided into 2 regions (cephalothorax and abdomen). As
alluded to earlier, they bear biramous appendages (forked segmentation). Borne on the
cephalothorax are 2 pairs of antennae, 3 pairs of mouthparts and at least 5 pairs of legs.
Most species are scavengers or predatory.
Subphylum Uniramia
Class Diplopoda (millipedes)

Figure 95: The Rhinocricid milliped (Source:
millipedhttp://www.discoverlife.org/nh/tx/Myriapoda/Diplopoda/images/Rhinocricid_Millip
ede, I_PA1.320.jpg)
The Diplopoda are primarily herbivorous and all are terrestrial. Their bodies are divided into
two regions; a head and trunk. The head bears one pair of antennae and often a pair of
compound eyes. Each compound eye is composed of a number of facets each functioning as
an individual visual structure so that many images of the some object are formed and send
to the brain where they are fused into a single image. Body segments are usually cylindrical
and each is composed of two fused segments (a condition referred to as
diplosegmentation). Typically two pairs of legs per segment. The reference to this group as
the millipedes (thousand legs) is thus a misnomer as there are no species bearing this
number of legs. The largest species of millipede is the African giant millipede
(Archispirostreptus sp and can attain lengths of up to 22.8cm
Class Chilopoda (centipedes)

These are like the millipedes in having many legs, but they do not exhibit
diplosegmentation as in the millipedes and thus each segment typically bears one pair of
appendages. Unlike the millipedes their bodies are not cylindrical but typically dorso-
ventrally flattened (Figure 94). These terrestrial predators, like the millipedes have bodies
divided into a head and trunk. The head bears one pair of antennae and simple eyes and the
first pair of trunk appendages is modified into poison claws used in subduing prey (Figure
93). They can attain sizes larger than millipedes. The Giant South American Centipede –
Scolopendra gigantea is the largest centipede known with individuals attaining sizes up to
30cm.

Figure 93 Anterior view of Chilopoda
Figure 94:Photograph of a Chilopoda (Source:

http://t2.gstatic.com/images?q=tbn:ANd9GcQQcLcsPy-
KszagAZIl8sPHkDh8ALk_ze4OZMSjgVeg1ke_MYj_)
Class Insecta = Hexapoda (insects)

The insects are the only invertebrates with wings, and thus enjoy, together with birds and
bats, a life in air. The evolution of flight in insects brought with it many advantages. First the
wings, unlike in their colleagues (birds and bats) are not derived from the forelimbs and
hence flight, in this group did not evolve on the sacrifice of walking legs. The wings allow the
insects to move fast in search for new habitats, to escape predators, in search for food and
to evade pending unfavourable conditions. The latter form of behaviour is exhibited quite
well by the African migratory locust (Locusta migratoria) which is known to fly all the
distance from North-east Africa to Central and Southern Africa.
The insect body is typically divided into three regions; head, thorax and abdomen (Figure
94). Head with 5 pairs of appendages: a pair of antennae for chemoreception (and in few
species sound perception -mosquitoes) protrude in the anterior dorsal region of the head; a
fused labrum, a pair of mandibles, a pair of maxillae, and a fused labium form the feeding
mouthparts. Many orders have modified mouthparts with some groups having lost or
reduced mouthparts. The insects thus exhibit a numerous feeding strategies.

Figure 94 The distinct body regions of insects -Head bearing antennea, eyes and mouthparts,
Thorax-bearing legs, and a terminal abdomen
The thorax bears three pairs of legs and typically, two pairs of wings. Wings are absent in
primitive forms or secondarily lost in the more derived orders. The cerci, a pair of forked
structures at the terminal end of the body is all that the usually large abdomen has to show
for an appendage in the typical insect

Mostly terrestrial, but some secondarily aquatic. Figure 94.1 below shows an aquatic
insect.
Figure 94.1 An aquatic insect, a freshwater coleopteran (Beetle) (Photo by Dube T)

.

PART 5
CHAPTER 6
The Deuterostome Phyla
Phylum Echinodermata
This is a phylum of marine organism showing radial symmetry in the adult, in particular the
pent radial symmetry. The larval stages, however exhibit bilateral symmetry, and this
feature removes them from any kinships with the cnidarians that also show a radial
symmetry in the adult stage. The name Echinodermata means spiny skin, giving reference to
some of the echinoderms, the Echinoidea with spiny body coverings. The phylum contains
more than 6000 species which include the starfish, brittle stars, sea urchins, sea cucumbers
etc. Typically the mouth is ventral, an internal skeleton of calcium carbonate plates and a
water vascular system consisting of a ring canal, radial canals and tube feet are the major
distinguishing features of this group.
The internal calcareous skeleton is covered by spines and skin and it varies from group to
group and from one species to another. In sea cucumbers the skeleton is made of
degenerated calcareous plates which are buried in the fleshy body, while in starfish the
skeleton is made from movable calcareous plates thus forming flexible joints.
Their water vascular system (Figure 95) is a very vital component because almost all the
functions of these organisms, including locomotion, respiration and feeding, are facilitated
by this system. The water vascular system is basically a hydraulic network of canals - which
runs through the entire body of these marine animals. These animals carry out various life
functions by varying the internal water pressure in this hydraulic network. It is composed of
a ring canal which is connected to radial canals. The latter run within each arm of the
animal. Within the arms the radial canals associate with tube feet that project to the ventral
surface of each arm in depressed grooves (ambulacral areas) and are used to get a purchase
on the substratum as the animal moves. By varying the amount of water in the tube through
regulation in the ampulae the tube feet can be retracted or extended. The ring canal is also
connected to the outside through a stone canal which opens though a pore called a
madreporite, making the contents of the water vascular canals the same as sea water.

Fig
ure 95: The water vascular canal system of echinoderms (Source:
http://www.esu.edu/~milewski/intro_biol_two/lab__13_echinoderm/images/aster_diagr_w
ater_vasc.jpg)
Even though most of the echinoderms are benthic in nature - i.e. they live on ocean floor,
excerption are abundant. There do exist some species which have the tendency of floating
(such as sea cucumbers) and swimming (sea lilies). Similarly, there also exist some
echinoderms who take help of some floating debris (crinoids) or fish (sea cucumbers) to
move from one place to another.
This phylum of marine animals is believed to be existing on the planet since the Cambrian
period.
Class Crinoidea (the water lilies)

Figure 96: A water lily
(Source:http://t2.gstatic.com/images?q=tbn:ANd9GcQYnaUykx5dJWJi3hR1CUZ2wuYvB8SHS
PC4ahk5iaVQ2MYOoYlELQ)
These (figures 96) are thought to be most primitive of all echinoderms. They are
characterized by typical pentaradial plan with five arms that are often highly branched so
that as many as 200 arms are not uncommon. Being sessile, their bodies are attached to the
substratum by a stalk of variable length through the aboral surface. The dorsal surface bears
both the mouth and annual openings giving the gut a characteristic U-shape. A porous
calcareous plate, the madreporite, also opens in the dorsal surface. A water vascular ring
canal surrounds the oesophagus and radiate into each arm as umbulacral canals. Paired
rows of tube feet emerge dorsally from each arm along the umbulacral groove. The podia
are thus organs of feeding and not for locomotion as in the other echinoderms. Nerve tissue
surrounds the mouth as the circumoral ring which issue out radiating nerves into the arms.
Below the skin is a calcareous shell made of plates or ossicles.
Class Asteroidea (the sea stars and related forms).

These are typically pentaradial with five arms and some with as many as 40 arms that
gradually grades off the main body so that while some have the typical radiate body plan
others have almost pentagonal shapes (Fig 97). The through gut has openings on the dorsal
(mouth) and ventral surface (anus) suggesting a complete rotation of the latter through
180o as compared to the Crinoidea just described above. The water vascular ring canal is
thus moved ventrally and surrounds the oesophagus which leads into a mouth that opens
midventrally. The dorsally positioned madreporite is connected to the vascular ring through
a canal (the stone canal). Umbulacral canals radiate from the vascular ring into each
radiating arm. Paired tube feet linked to the umbulacral canals are provided with water
reservoirs (the ampula) and exit to the outside ventrally along each arm. Closely knit test
plates are covered by a skin which bears spines. A haemal system issues out haemal canals
into each arm. Nerves from the circumoral nerve ring also enervate each individual arm.
The Asteroidea, unlike the Crinoidea, are motile Echinodermata that employ the ventrally
projecting podia to gain traction and move in the ocean bottoms.

Class Ophiuroidea
Typically pentaradial with five long arms leaving the main body disc as distinct thin whip like
structures (Fig 98). In these forms the mouth opens ventrally and the anus has disappeared,
but the madreporite remains dorsally. Tube feet project ventrally along each arm and are
used together with
Figure 97: An Asteroid (Source:

http://animaldiversity.ummz.umich.edu/site/resources/jo_okeefe/Forbes_sea_star_100_54
79.jpg/medium.jpg)
the whole arm in locomotion. To this end the arms are well equipped with massive ossicles
which occlude much of the arm volume. The ossicles are fused in chains much like the
vertebrae bones of vertebrates and intervertebral muscles serve to bend the arms as they
gain purchase on the substratum and move the animal.

Figure 98: An Ophiuroidea echinoderm (Source:
http://www.oceaninn.com/wp-content/uploads/2010/06/5.gif
Class Concentricycloidea.
Of all the Echinodermata, these are relatively new to biology as the first specimens well
drenched from ocean bottoms as recent as 1986. These are disc shaped, and unlike all the
other groups, lack arms (Fig?). Ventrally, are two concentric rings of water vascular rings,
and dorsally are a series of covering plates. Two species have been described; one in the
waters of Bahamas and the other of the Coast of New Zealand. The Bahamas type has a
ventral mouth provided with a blind ending stomach and the New Zealand form has no gut
at all.
Class Echinoidea
Figure 9:
Structure of the naked echinoid test of Goniocidaris parasol. (a) Naked test. (b) Test with
radioles. (Source:
http://content.answcdn.com/main/content/img/McGrawHill/Encyclopedia/images/CE21
0900FG0010.gif)

These are spherical to flattened echinoderms whose arms have become incorporated into
the body of the animal (Figure 99). With a mouth typically ventral and the anus dorsal
(aboral), the umbulacral and interumbulacral are composed by two columns of ossicles that
curve upwards from the mouth to the dorsal anus. The calcareous skeleton is rigidly cast
and robust in most members and the skin is provided with spines or pedicelariae. The
ventral mouth is provided with a grazing organ used to graze and chew prey food. The
grazing organ, Aristotle's latern is composed of five plates plus numerous other smaller
plates and can be protracted from the mouth to nibble food which is mainly in the form of
algae and other sessile animals. Two major categories are realized; the regular and irregular
forms. In the regular forms the mouth is typically midventral and the body is provided with
numerous tube feet. In the extensive body cavity are five gonads that open through
gonopores in the aboral surface. In these forms the brooding usually occurs in the body
cavity. In the irregular forms the mouth and anus are moved to either end so that they have
an 'anterior' and 'posterior'. In these forms the mouth is surrounded by peristomial gills and
the grazing apparatus are greatly reduced and these are mainly deposit or suspension
feeders. Four gonads project into the large body cavity discharging their gametes through
aboral gonopores. Embryonic brooding doesn’t occur inside and development is indirect in
this forms.
The 950 species of Echinoidea are categorized into six orders.
Class Holothuroidea.
Figure 100: A Holothuroidea-

Pseudocolochirus
violaceushttp://animaldiversity.ummz.umich.edu/site/resources/Grzimek_inverts/Holoth
uroidea/Pseudocolochirus_violaceus.jpg/badge.jpg)
These are normally elongate echinoderms (Figure 100), with bodies elongated along the oral
aboral axis and the arms have become incorporated into the body of the animal. The result
is that they lie on their sides, thereby acquiring the equivalence of ventral and dorsal sides.
The ventral has side three umbulacral grooves and dorsally are two umbulacral grooves. A
through gut runs from the mouth anteriorly to the anus posteriorly. The mouth is usually
provided with tentacles which are highly branched or shield like. These serve for filter
feeding and are supported internally by the water vascular system. The gut terminates in as
expanded rectum, cloaca and finally the anus. Opening into the cloaca are tubes connecting

to the respiratory trees that lie inside the body. The respiratory trees serve for gaseous
exchange as water is pumped into them from the cloaca.
Tube feet scattered throughout the body wall are mainly for locomotion. In some forms
they are used as organs for firm attachment to the substratum and in others they are quite
elongated so that they are used for locomotion, wholly suspending the whole organism
above ground. Body peristaltic contraction aided by the skeletal ossicles that projects from
the body are also used for locomotion in some groups. However in all forms locomotion is
very slow. The skin is leathery and below it is a loosely organized cancerous skeleton of
minute icicles. The skin is provided with circular and five longitudinal muscles. Unlike in
other groups where the madreporite links with the surrounding sea water, in the
holothurians it lies free in the body cavity. The holothurians are gonochoristic and a single
gonopore discharge near the oral tentacles. Development may be direct or indirect. Direct
development occurs in species that brood either in the body cavity or the ovary and indirect
development occurs in those that do not brood.
The class contains 1150 extant species categorized into six orders on the basis of nature of
oral tentacles, tube feet and body shape.

Phylum Chordata
Figure 101: Some Chordates Representatives- Tunicate, fish, reptile, bird and mammalia
(Source: http://tolweb.org/tree/ToLimages/dein.gif)
Chordates (Figure 101), the Phylum to which humans belong, though the most conspicuous
group of animals, constitute a very small proportion (about 5%) of the metazoans. They are
united by their having a number of shared characteristics (Figure 102) some of which are
very transient in some groups. Collectively they have a firm flexible rod running dorsally and
longitudinally inside the animal’s body. This is called a notochord and serves as a supporting
structure at some stage in the life cycle of the animal. It is composed of vacuolated elastic
cells that provide a supporting frame to the animal. This is transient or last the whole life of
the animal. Segmental muscle blocks or myotomes are attached to the notochord and serve
as structures to bring about torques required for locomotion.

Figure 102: Unifying Chordate features
(Source:
http://t2.gstatic.com/images?q=tbn:ANd9GcSlEcEl94vt1kYR7zhhLqvvAODNXmc8KNgYz17w
U-zYSLzHw-YC)
A hollow nerve chord runs longitudinally above the notochord. A coelom, ventrally located
in the trunk region houses the heart, excretory, and reproductive organs. The coelom is
restricted to the trunk region of the animal and extends to the head region or tail region. A
series of openings from the gut to the outside occur in the pharyngeal region to allow for
water to be extruded to the outside during feeding. These are called pharyngeal slits or
visceral clefts. In the aquatic chordates they serve to house the gills. In the terminal region
behind the anus, typically, extends a tail of varying length depending on the group.
The chordates are divided into three subphyla, two of these, the Urochordates and
Cephalochordates are referred to as Protochordates or invertebrate chordates on account
of their lacking vertebrae bones and hard skeleton, distinct head, brain and cranium. The
notochord may or may not persist into adulthood and serves for locomotion in their aquatic
environments. They are usually mucofilter feeders with expanded pharyngeal regions for
filter feeding. The third subphylum the Vertebrata to which the fish, amphibians, reptiles,
birds and mammals belong are characterized by a notochord which is quite transient in its
existence; lasting only the embryonic stages, to be soon replaced by hard skeleton in the
form of bone or cartilage as adulthood is attained.
Subphylum Urochordata

Figure 103: Photograph of a Tunicate (Source:
http://www.ucmp.berkeley.edu/chordata/tunicates1.jpg)
The urochordates are free-living, sessile and solitary or colonial invertebrate chordates.
They exhibit the chordate features only in their embryonic forms; in their adult form, they
hardly resemble the chordate body plan. Included here are organism commonly called
tunicates. The notochord, nerve chord and post anal tail only appear in the larval stages (if a
larval stage is present) and disappear altogether in the adult stage excerpt in a few genera.
They secrete a hard outer covering housing the animal inside, called a tunica. This is
composed of either cellulose or protein.
A through, U-shaped gut runs from mouth to anus. The mouth, typically tentaculate leads
into a very wide, highly perforated pharynx, where filter feeding is effected. The mouth is
bordered by a branchial siphon through which water currents are directed into the animal's
body. An artrial siphon channels the water current to the outside, carrying with it the egesta
released from the anus which terminates in the atrium surrounding the pharyngeal area. A
partially closed blood circulatory system serves to move blood in either direction from the
heart which is ventrally situated. A single ganglia, in between the branchial siphon and the
artrial siphon is all there is for the tunicate to show for a brain.
Tunicates are typically hermaphroditic, with a single ovary and testes that open into the
atrium. They bear no excretory system and lack a coelom.
There are about 2 000 tunicates classified into three classes; the Ascidicea, Larvacea and
Thaliacea

Figure 104: Tunicate structure (a) whole adult specimen (b) adult section (c) larva (Source:
http://www.clarku.edu/departments/biology/biol201/2006/mtrivette/images/34-03-
Tunicate-L.jpg)
Class Ascidicea
The Ascidicea form the largest group of all urochordates with about 1 850 members. Its
members, in adult stage, are typically benthic, sessile, free-living, solitary or colonial (Figure
105). Once the larval stages settles to the bottom substratum using the anterior end, they
lose the post anal tail, notochord and nerve chord and the mouth and artrial pores develop
in their positions. The mouth and artrial pores are extended into tubular siphons by
extensions of the tunica. The mouth, which is bordered by tentacles, serves as incurrent
siphon allowing water and food particles to be drawn into the animal. From the mouth, an
expanded pharyngeal area almost fills the whole body of the animal. The Pharyngeal area is
highly perforated by slits which allow water to move out into the surrounding cavity
(atrium) from where it is channelled to the outside through the atrial siphon. Food particles
trapped in the pharynx are channelled into the stomach, which leads to the anal opening in
the atrium in the pathway of the excurrent to the exterior through the atrial siphon. A basal
heart which alternately pumps blood in either direction is situated below the stomach area.
The circulatory system which is partially closed is provided with an unusual type of blood
rich in heavy metals like vanadium, niobium or iron in association with sulphuric acid.
In the colonial form two organisms may share the same test or tunica as well as a single
atrial siphon.

F
igure 105: Solitary (top) and colonial (right) Ascidicians.
(Sourcehttp://www.getahugetank.com/images/Tunicates.jpg)
Class Thaliacea.
The Thaliacea are either solitary or colonial (Fig 106)). Each animal is elongate and in the
colonial form all the individuals share a single continuous shell and internal canal into which
all the atrial siphons open. All individuals (zooids) thus have their branchial siphons open to
the outside and opposite the atrial siphon which open to the inside releasing their contents
into the communal central tube. Many zooids make up a single colony which appears like a
single large cylindrical organism. No wonder these animals have been mistaken for large
serpents in the marine environment.

Figure 106: A salutary Thaliacea species
Some forms appear as either solitary or aggregate animals (Fig 106). The salutary animal
reproduces asexually and the aggregate form reproduces sexually.
Class Larvacea
This class bears all the typical chordate features; a persistent post anal tail, a notochord and
a dorsal hollow nerve cord and open pharyngeal slits (Fig 107). These animals usually very
small (not more 15mm), has an anterior mouth opening which leads into a pharyngeal
region perforated by slits. It secretes a test or house of chitinous material plus protein into
which it lives and is at liberty to abandon when clogged by detritus or other materials. The
house has a screen through which water and microscopic food particles can enter. A current
driven by the beating of cilia in the pharyngeal area leads the material into the buccal cavity
and into the pharyngeal area where microscopic plankton is trapped and channelled into
the gut. Water from the pharynx is drawn out and passed out through pores in the
pharyngeal area. An escape door, which is usually closed by an operculum-like structure,
allows the animals to escape and inflate another readymade house when it is clogged.

Figure 107; A Larvacea species
(Source:http://t3.gstatic.com/images?q=tbn:ANd9GcSxDXJ3hC0cbU0rUVUlQagb6yOaysSQm
rsUaUzFcB_UrNhFAsqT)

Subphylum Cephalochordata.
The cephalochordates are the fish-like chordates that bear nearly all the chordate features
in their adult stages. They all fall in the genus Amphioxious (formally Branchiostoma). A
flexible notochord aids in bending the body as the animal propels itself in water. Segmental
muscle blocks or myotomes serve to bend the tail into series of S- shapes providing
propulsion to the animal. A dorsal nerve cord runs above the notochord, and a straight,
through gut run from the mouth to a sub terminal anus. The mouth, which is bound by
tentacles and velum leads into a long expanded pharynx which runs half the length of the
body, leading into the stomach, intestines, rectum and anus. Filtered water from the
pharynx enters the surrounding atrium space from where it is led to the exterior through a
preanal pore, the atriopore.
The Cephalochordates’ intermediate features between the Urochordates and Vertebrates

may serve as the bridging link between the two groups. The possession of notochord, nerve
chord and post anal tail undoubtedly links them to the Urochordate larval stage. On the
other hand their lack of true jaw connects them nicely to the simplest of vertebrates, the
Agnatha.
Figure 108: Amphioxus (Top -external), bottom -section through body. (Source:
http;//www.biosci.org/V02/p0149/ijbsV02/p0149g01.jpg

Subphylum Vertebrata
Vertebrates constitute the largest subphylum in the phylum Chordata. They are bilateral,
coelomate, typically horizontal animals, elongated along the posterior anterior axis. Their
bodies are typically divided into three regions; head, trunk and a tail region. Their bodies are
supported by an endoskeleton made of either bone or cartilage. The skeleton is divided into
an axial and an appendicular skeleton. The axial skeleton comprises the serially segmented
vertebrae bones that make up the back bone, an anteriorly placed skull and serially
arranged lateral bone, the ribs that articulate with the vertebrae. Posteriorly the tail (region
posterior to the anus), terminates in series of vertebra bones.
The appendicular skeleton consists of the pectoral and pelvis girdles together with the
associated paired pectoral and pelvis fins and the median fins (dorsal, caudal and ventral
fins) in the aquatic vertebrates. In the land vertebrates (Tetrapods) the paired fins are
replaced by some paired limbs, the fore and hind limbs, both terminating in five fingers or
toes (digits) -the typical pentadactyl limb of land Vertebrates. The median fins have
disappeared completely.
Vertebrates are highly cephalised animals, with a brain situated anteriorly and protected
within the skull and major sensory organs also in the head region. The head bears the eyes,
olfactory and auditory organs. Positioned anteriorly is the mouth, a transverse slit which
opens into the buccal cavity. Primitively there are pharyngeal slits posteriorly placed in the
lateral position in the head towards the trunk region. These are present in aquatic
vertebrates as gill slits but only present in the embryonic forms in the advanced (land)
vertebrate. Connecting the head to the trunk region is usually a constricted region- the
neck.
The trunk bears the coelomic cavity, the reproductive and excretory organs. Ventrally in the
region connecting the trunk to the tail, is the anus. Anterior to this opening are the urinary
and genital openings, sometimes united to form the urinogenital aperture and in some
groups this urinogenital aperture is united posteriorly with the gut to form the cloaca which
opens to the outside as the cloacal aperture.
Extant vertebrates are classified into two super classes; The Pisces and Tetrapods. The
Pisces in turn are grouped into three classes: the Agnatha, the Chondrichthyes and
Actinoptergii. The Tetrapods, which are largely a terrestrial fauna, are grouped into four
classes: Amphibia, Reptilia, Aves and Mammalia.
Super class Pisces. (The fishes)

Fishes are the dominant vertebrate group today with up to 19 000 species. As a group they
are characterized by bodies which tapper anteriorly and posteriorly, giving the streamlined
condition which allow for ease movement in water as frictional drag is greatly reduced. The
streamlined body allows the fish to pierce through the water and at the same time
permitting the water to flow past the body with very little hindrance. Movement in water
has been made possible mainly due to the presence of fins which come in various forms and
sizes. These allow the fishes to propel themselves in the viscous water, at the same
controlling direction of movement, preventing rolling and maintaining buoyancy. Mucous

glands in the epidermis secrete slimy mucoploysaccharides (mucus) which makes the body
slippery and further aids in movement through water. The slime material also allows the
fishes to evade predators. Scales which are present in many fish groups, overlap in an
anterior-posterior direction and hence also serve to reduce frictional. The scales also serve
to protect underlying tissues from predator attacks as well as limit loss of salts from the
body.
Gills are characteristic of fishes and they also come in various forms but they are generally
modelled along two lines; in some fishes the walls of the each gill slit is highly folded and in
others the partition between the individual gill slits have disappeared and each gill is
supported by a branchial arch. A single bony cover (operculum) overlies all the gills and
helps in the control of water flow over the gills by alternately opening and closing.
Blood circulation is primitive with a heart situated ventrally in the coelom, composed
basically of an expanded vessel differentiating into a atriosus, a ventricle and an
atriovenosus all connected to form a pulsating organ which pumps blood anteriorly to the
gills. From the gills, the blood, which is now oxygenated, is straight away channelled to all
parts of the body (systemic circulation) before finding its way back to heart where the cycle
is repeated. Movement of blood to the gills results in a marked reduction in velocity and
hence less efficient supply of oxygen to the tissues
Class Agnatha.
These are the jaw-less fish and the first vertebrate group to evolve and flourish at the end of
the Cambrian to early Ordovician (about 505 million years ago). Today they are represented
by the hagfish and lampreys among others, forming a very inconspicuous group of fish.
Being jawless, they use their sucker like mouths to attach onto other fish from which they
feed parasitically.
Class Chondrichthyes.
These are typically cartilaginous fish whose skeletons are basically composed of cartilage, a
relatively hard material composed of mainly fibrous protein called collagen forming a matrix
around collagen secreting cells. Their pharyngeal slits open to the outside as gill slits. The
gills are housed in the pharyngeal area and there are no opercular covers. This group
includes the sharks, rays and skates. The Sharks, the most conspicuous group of the
Chondrichthyes are characterized by rasp bodies covered by cosmoid scales of similar
composition to mammalian teeth. They lack paired fins and the caudal fin is heterocercal,
i.e. the upper and lower lobes are asymmetrical. They maintain buoyancy by continual
swimming because they lack swim bladders. If they stop swimming they sink. Most of them
are active swimmers and predatory, being provided with robust teeth to crush the hard
shell of molluscs which constitute their major diet. The skates are bottom dwellers.
Class Oesteichthyes
These are bone fishes that populate both the marine and fresh water environments. They
typically possess a bony skeleton. Bearing paired fins, a stream-lined body and a homocercal
caudal fin, they are active swimmers with high manoeuvrability. Their gill openings are
typically covered with a bony operculum which, in addition to the buccal pump serves to

actively move water though the gills during ventilation. This group is divided into two: the
Actinopterygii and the Crossopterygii. The former are characterized by fin rays- a condition
characterized by bony rays supporting a flap of skin. These form the vast majority of the
fishes (The Holostei and Teleost fish). The latter group, the Crossopterygii has lobed fins.
The fins are fleshy and supported by a bone axis inside. It will be of interest to note, at this
juncture, that the rest of the land vertebrates are thought to have evolved from this group
of fish. In their evolution the Crossopterygii gave rise to two stocks; the Dipnoi and the
Coelacanths. The Dipnoi (double breathers) are also called the lung fishes on account of
their swim bladders having been modified into lungs through extensive infolding. Only three
genera of these lung fish are extant; the Australian lungfish, the African lungfish and the
South-American lungfish. The Coelacanths are thought to be the progenitors of the land
vertebrates (the Tetrapods).
Class Amphibia.
The amphibians are believed to be the earliest land tetrapods, being descendent from the
Coelacanths through the genus Rhipidistia. Amphibians as a group are not quite
independent of water. Their soft bodies and shell-less eggs are prone to desiccation in the
terrestrial environment. They thus go back to water for reproduction; laying their eggs in
water or very dump places and relying on water and water films for sperm transfer to eggs.
Fertilization is thus external and prone to the vagaries of the outside environment. Many of
us have seen, first hand, the vast numbers of eggs laid by the common garden frog Rana
temporaria in pools or stagnant water, all in the vain attempt to maximize survival rates.
Apart from having lungs for respiration, many amphibians are also dependent on cutineous
respiration. To this end, the skin is highly vascularised and is kept moist by secretions from
epidermal glands which serve to dissolve atmospheric oxygen. Secretion from these
epidermal glands, in some cases act as deterrent secretions with offensive odours against
predators. The dependency of the amphibians on water has also meant that they had to
retain some of the apparati for gaseous exchange in water. Thus most amphibian larvae are
hatched with gills which will later degenerate during metamorphoses as adult stage is
attained. In some cases the gills are even retained even in the adult form. A good example is
the Mexican salamander, Axolotl.
Amphibians have significantly improved circulatory system over that of Pisces. The heart is
typically three chambered, there being a second auricle in addition to the single one in
fishes. Oxygenated blood thus goes back to the heart for pumping before being released
into the systemic circulation. Like fishes, amphibians are ectothermic.
Amphibians used to constitute a very large group of vertebrates but their diversity is
currently highly demished, and, to date a viral epidemic is on the rampage in the amphibian
world, threatening to wipe out many species of amphibians, especially the frogs and toads.
Extant amphibians are grouped into three major Orders; the wormlike caecilians or Apoda,
the tailed Salamanders (Urodela) and the Frogs and toads (Anura) (Fig? To?).
Class Reptilia
These are generally creeping animals from which they get their name (repitilios- to creep).
Their vertebrae column has a single point of articulation to the head (occipital condyle). The
lower jaw characteristically bears many bones. The teeth are rarely differentiated

(homodont), and capable of being replaced throughout life. They are ectothermic and hence
poikliothermic, i.e. their body temperatures are dependent on external or ambient
temperatures and thus cannot maintain a constant internal body temperature. This puts
them at the mercy of the vagaries of weather and to a large extend this determines their
spatial distribution in their terrestrial environment.
A single occipital condyle enjoins the vertebrae column to the head. The vertebrae column
is variable; being very long in some groups like the Serpentes (snakes) and somewhat very
reduced in others like the Chelonia (tortoises). Typically four limbed, but vestigial or
completely lacking in the Serpentes.
Reptiles are characterized by a dry scaly skin which is impervious to water. This feature,
together with the shelled leathery eggs they lay, has enabled them to be quite independent
of water although some have secondarily gone back to the aquatic environment. The
reptiles are thus typically a terrestrial group. The shelled reptilian egg has a special
membrane (the amnion) which houses the developing embryo during development. The
presence of this membrane (the amnion) in reptiles as well as in the birds and mammals as
earned these vertebrates the name Amniotes.
The reptilian scales are modified epidermal cells deposited with keratin. These scales can be
quite enlarged in some reptiles to form plate-like structures called scutes as found in the
dorsal tail regions of the crocodiles.
Blood circulation in the reptiles is more efficient than in the Amphibians owing to the
evolution of some septum partially separating the right and left ventricles. The heart is thus
typically three chambered as in Amphibians but the partial separation of the ventricle and
the inherent physiological mechanisms to limit the mixing of oxygenated and deoxygenated
blood makes the reptilian heart more efficient than the amphibian heart. In some reptilian
groups like the Crocodilia, the heart is almost completely separated into four chambers, a
feature which is close to the aves, their very close relatives. Fertilization is essentially
internal and in some groups there are elaborate courtship behaviours as well as guarding
and provisioning for the young. Eggs are usually laid and buried in soil or under rocks and
the hatchling is an exact replica of the adult.
Reptiles evolved from amphibians in the Carboniferous era of the Palaeozoic and become
the dominant group in the Mesozoic for over 200 million years. They diversified to give a
number of lineages amongst them the now extinct Dinosaurs which were the largest land
animals that ever roamed the earth. To date only four Orders of reptiles remain: the
Chelonian (tortoises, porpoises and related forms); the Crocodilia (crocodiles, gavials and
relatives); the Squamata (lizards and snakes) and the Rhychocephalia (the tautara or
sphenodons of New Zealand) (Figure? to?).
Class Aves (Birds).

Aves or birds are typically endothermic. Endothermism is made possible by the very high
metabolic rates associated with diets very rich in carbohydrates and oils such as seeds,
grains and fruits, as well as other animals. Their high metabolic rates allow them to power
their fight as well as maintaining a constant body temperature (homeothermy). They have a
single occipital condyle and a skull roof mainly composed of fused bones.

Homeothermy (mantainance of constant body temperature) is made possible by the high
metabolic rates characteristic of birds, which has been made possible by an efficient blood
circulatory system and gaseous exchange system. The heart is composed of four chambers,
all completely separated so that there is no mixing of oxygenated and deoxygenated blood.
The bird's lung is constructed in such a way that there is unidirectional flow of air and hence
no mixing (and the inadvertent dilution) of gases between inhaled and exhaled air.
Birds are amongst the most colourful of the vertebrates and their plumage makes them
undoubtedly the most easily distinguished animals even to the laymen. This, they owe to
their feathers which are very often highly coloured. Birds are the only vertebrates with
feathers and this feature has distinguished birds ever since their evolution in the Triassic to
Jurassic era. Bird's feathers are believed to be modified reptilian scales which become
insunk into the dermis for nourishment, and, extending outwards through a shaft equipped
with barbs and barbules that hook to each other to provide a close-knit structure (vane) that
can be used as an aerofoil to push down on air to provide a lifting effect. Feathers thus give
birds their gift of flight. The forelimbs are typically constructed for flight as are many
features of birds. This forelimb, which is generously provided with feathers, has been made
more rigid and firm by way of a shortened humerus bone and fusion of the carpus and
metacarpus to give the carpometacarpus bone. Linked to the humerus are flight muscles
whose origins are in the breast bone. The latter is extended ventrally into a keel (or carina)
to provide sufficient attachment for the insertion of the flight muscles.
Typical of the bird's anatomy is that the hind limbs are brought forward so that the leg only
begins to separate from the body offer the knee joint as the femur is tightly held close to
the body. The drumstick that some cultures reserve for the head of the household on the
table is in fact not a bird’s thigh but the shank. The hind legs serve for landing and take-off
and are thus modified into strong structures that are held vertically below the body. The
tibia is quite massive and robust although the fibula is reduced into a rudimentary structure
held close to the latter; the tarsus and metatarsus have fused to form a firm structures; the
tarsometatarsus. Together, these modifications serve to ensure that the hind limbs can
absorb the shock that comes with landing and flight take-off. Terminating at the tip of the
hind leg are four fingers that are all clawed, three facing forwards and one facing
backwards. The fifth finger is lost in many birds but occur in some groups like the Numididae
(e.g. guinea fowls) and chicken-like birds as a short horny structure called a hallux used in
offense and defence. Covering the distal portion of the legs are scales, which, point to a
reptilian ancestry.
To further increase the rigidity of the landing device, the femur is linked proximally to a rigid
structure, the synsacrum, which is composed of central sacrum linked anteriorly to the
lumbar and a few thoracic vertebrae, and posteriorly too few caudal vertebrae all fused to
form a rigid girdle.
The birds mouth is modified into a horny structure (the rhombotheca) bearing no teeth. The
nature of bird’s beak is very often species specific and forms a very important taxonomic
feature. The beak comes in various forms; short and pointed as in the weavers like the
quelia, long and spear like in the hammer kops, long and spoon shaped as in the spoonbills,
short and hooked like in the Aquidae (the eagles) etc. The loss of teeth has been
compensated by the evolution of the gizzard in the place of the stomach which takes over

the role of grinding food. The gizzard is strong muscular bag lodged with sand grains
consciously ingested by the bird to act as a grinding mechanism for the food which is
swallowed whole. It is important to point out that the loss of teeth in birds was a further
adaptation to a life in the air where lightness counts above everything.
Pneumatic bones (hollow bones) connected to the air sacs in the respiratory system serve to
make a bird's body less dense and light for locomotion in air. Life in air has also meant that
the major apparati for flight (the wings) tends to be greatly enlarged in comparison to the
rest of the body. Albatrosses highlight this phenomenon well with their wing spans attaining
lengths of up to 3.6metres.
To further aid in weight reduction birds have lost one ovary and only the left one remains.
Fertilization is essentially internal although very few are equipped with a penis for sperm
transfer. For most birds, just like their reptilian ancestors, the male deposits his sperm into
the common urinogenital aperture, the cloaca, which also serves for egestion. After
fertilization a shell is deposited around the egg. Eggs are laid in clutches either in well
prepared nest in a tree or on the ground, incubated by heat from the parent that sit over
the eggs for the greater part of the incubation period or from the heat from the surrounding
soil. At hatching the nestling is an exact replica of the adult into which it develops.
Provisioning of food and protection to young is nearly universal in aves.
Extant birds are generally grouped into two major groups; the ratites and the carinates. The
ratites are those that have secondarily lost the ability to fly and are thus flightless. These
include the Kiwis, the rheas and the ostriches. These, unlike the carinates have degenerated
most of the features important for flight. The carina and flight muscles are quite demished.
The Carinates on the other hand, are birds of flight, and, are characterized by enlarged keels
(carinas), large wing spans and generally light bodies. These form the largest group of birds
that populate the air and the seas.
Wing and feet form reflect functions performed. Bird diversification has seen some capable
of souring and gliding high up in the air with long narrow wings. High speed or migratory
birds have narrow, often swept back wings. Those living in dense forested areas where
maneuverability is crucial have broad elliptical wings and slotted high lift wings are found in
birds that sour on warm air currents over inland areas. Raptors like eagles, falcons and owls
which rely on seizing their prey by surprise have talons which are used to stun and grasping.
Those that rely on scratching like chickens and chicken-like birds have long claws.
Extant birds are grouped into 26 orders and many families. The families include the
Struthionididae (the Ostriches), Sphenecidae (the penguins), the Numididae (Guinea fowls),
the Peliconidae (pellicons), the Caprimulgidae (nightjars), the Sciconidae (stocks), the
Aquidae (the eagles), the Columbidae (the doves and related forms), the Ostrigidae (owls)
(Figures ? to ?).
Class Mammalia.
Mammals are the present day dominant vertebrate group on land. They are typically
terrestrial although some groups have secondarily gone back to water. As a group,
mammals are unified by a number of characteristics: possession of mammary glands from
which they suckle their young; presence of hairs or furs, anucleate red blood cells;
endothermic metabolism; presence of diaphragm separating the thoracic cavity from the

abdominal cavity; a single dentary bone in the lower jaw and a four chambered heart
divided into two pumps. In addition the quadrate and articular bones which are used as
components of the lower jaw in the lower vertebrates have become incorporated into the
hearing apparatus of the ear as sound transmitting ossicles (the maleus and incus).
Mammals are also characterised by presence of sebaceous and sweat glands. However,
mammals share with birds characteristics like endothermic metabolism and a four
chambered heart. Both mammals and birds owe their high activity to endothermic
metabolism, and a four chambered heart which completely occludes the mixing of
oxygenated and deoxygenated blood, thereby promoting high metabolic rates.
Mammary glands are thought to be modified sweat glands. Possession of mammary glands
enables the mammals to nurse their young ones with milk secretions and enhance the
caring for the young ones. Mammary glands occur ventrally along two milk lines which occur
on either side of the ventral side. Milk secretions are led to the exterior through either teats
or nipples. Teats are characterised by single duct which leads to the exterior and nipples
have numerous such ducts. Hair is a dermal structure which is thought to have evolved to
enhance the endothermic condition of mammals by keeping the internally generated heat
from escaping from the body surface. It is typically a column of heavily keratinized dead
epidermal cells whose live roots are insunk into the dermis for nourishment. In section, hair
is typically cylindrical in all mammals and other humans except for the Negroid hair which is
flattened. Sebaceous and sweat glands are usually associated with hairs. Their secretions
condition the skin and allow evaporative cooling of the body.
The diaphragm, separating the thoracic cavity from the abdominal cavity is functionally an
important structure for efficient gaseous exchange. It aids in the ventilation mechanism,
and, together with the efficient four chambered double pump heart, make oxygen supply to
body tissues very efficient- no wonder mammals are amongst the most active group of
animals. The endothermic nature of mammals arises from their ability to efficiently produce
enough energy for bodily requirements as well as for keeping the body warm and hence
constant regardless of fluctuations in ambient temperatures. This, as we have seen in birds,
allow them to be independent of temperature changes in the external environment and
hence enable them to inhabit wide ranging habitats. The presence of hair serves to enhance
this endothermic nature. By keeping warmed air close to the body the hairs or furs help to
keep the internal body temperature constant.
Extant mammals are grouped into three subclasses, the Prototheria (Monotremes),
Metatheria (Marsupials) and Eutheria (Placentals) (Figure? to?). The Monotremes are
thought to be the most primitive mammalian forms and are believed to be the closest to the
reptilian mammalian ancestors. These, like reptiles, they lay eggs and their legs are covered
with scales. Upon hatching the young ones suck milk oozing from mammary glands below
the skin in between hairs on the mother's belly. Although they suckle their young like all the
other mammals they lack nipples or teats and do not have external ears. Adult monotremes
lack teeth which are only present in the young ones. In the adult the teeth are replaced by a
horny beaklike structure. They lack a vagina but insteady possess a cloaca. Their testicles are
abdominal and not suspended in scrotum outide the body like in the other mammalian

subclasses. In addition their penis is not used for passage of urine. The echidnas and the
platypus are present day common examples of this group.
The Marsupials are more advanced than the Monotremes. They are viviparous, bearing the
young at a very early stage in a very immature form. Although born very immature, the
young can, within hours of birth, craw into a pouch on the mother's inguinal region, attach
onto one of the teats protruding from mammary glands enclosed in the pouch and start
feeding to complete embryonic development. Unlike the monotremes whose testis are
abdominal the marsipuials testis are contained in a scrotum. They have double uteri and
vaginas. The marsupial bag is characteristic of female marsupials although some females of
some species do not possess this structure. The kangaroos of Australia are examples of
marsipials that quickly come into the minds of many people.
The Placentals are the most advanced of all Mammalia. They are viviparous, carrying the
one in their uteri and nourishing it through food supplies via the placenta. The young is
bone fully mature, and can, within hours after birth follow its mother as in the whale. The
young is nursed with milk by the mother until it can switch onto the adult diet. Protection of
the young is quite extensive in this group and very often prolonged, reaching its zenith in
humans.

PART 6
CHAPTER 7
The insects: the champions of the metazoan world
The arthropods are the largest group of animals constituting about 80% of all animals. The
majority of these animals are the insects (comprising about 70% of all animals). The other
arthropods (Crustacea, Chelicerates, Chilopoda, Diplopoda, Symphyla and Pauropoda) put
together constituting only 7%. Insects are thus as Dhaliwal (1999) puts it -the lords of this
planet. They are found in every conceivable habitat; marine water, freshwater, brackish and
the terrestrial environment; under varying environmental conditions, in the ice of the Arctic
and Antarctica, in the hot and cold deserts of the Kalahari and Siberia and even in hot
springs (Dhaliwal 1999); occurring as either free-living and grading from imperceptible to
perceptible levels of parasitism; subsisting on nearly anything organic- algae, fungi animal
and detritus and even hydrocarbon in oil spills. Their food acquisition methods are
unparalleled; there being true hunters and parasites, browsers and grazers, suspension and
deposit feeders and still others utilizing chemoautotrophic and photosynthetic symbionts.
Many are solitary and still many others are quite organized and live in colonies exhibiting
varying levels of division of labour and specialization, reaching its zenith in the bees where
some members, for the wellbeing of the whole colony, have sacrificed their reproductive
capabilities. They have evolved to be amongst the best of architects; burrowing and building
galleries to consistent geometric specification and orientations. Amongst them are some of
the most proficient foragers and hunters that leave man green with envy. Still others have
ventured into agriculture and slave making, areas hitherto thought to be a preserve for
mankind. In fact insects learnt the ropes of these trades well before men evolved.
The highly cryptic nature and camouflage exhibited by some members of this group like the
praying mantis, the stick insects and melanic moths have no doubt served as great lessons
for mankind during times of war but he has not mimicked them to any level comparable
with the models (the insects themselves).

The vast repertoire of devices and mechanisms employed by insects to maintain their
dominant position in the animal world has long been appreciated by insect’s scientists
(entomologists), not to mention those in the business of insect pest control in agriculture
and veterinary sciences. Man and insect are heavy competitors for food and the farmers in
their attempts to outdo the latter have been left convinced beyond any reasonable doubt
that any victory over insects is short lived.
In this chapter we start by considering insect evolutionary history then move on to look at
the features that make insects a success story then wind up by giving a survey of the major
insect orders.
Arthropod evolutionary history

It is not possible to exhaust the evolution of insects in a module of this nature. Interested
readers should consider textbooks on entomology some of which are cited in the references
for this chapter. Here we look at the origins of arthropods to which the insects are a
member then traverse to look at the origins of insects themselves. We restrict ourselves to
concentrating on the general lineage that led to the insects and make no issue with dead
ends in the numerous evolutionary experiments that never saw the light of day.
The search for an arthropod ancestor, just like the search for the ancestor of any other
animal group, centres on opening up archives of more primitive animal forms, bearing some
similarities, which sometimes, though less developed or primitive would most probably
have formed a basis or frame for modification to give rise to the descendent whose ancestry
is being sought. This search has led biologist to zero down the progenitor of the arthropod
to an annelid like ancestor, most probably related to present day Onychophorans. Insects,
apart from having features that are annelid-like and hence can be traced directly to the
latter also possess characteristics which are not present in the annelid but present in the
Onychophorans. The Onychophorans on the other hand, apart from resembling arthropods
also have some features common with annelid and not present in the arthropods. This has
been taken to be suggestive of an intermediate group linking the two.
In a previous chapter we dwelt on the distinguishing features of both annelids and

arthropods and it is important here to point out only those features linking the two and thus
pointing out to the annelid as possible progenitor of the arthropods. Like the Annelids, the
arthropods are segmented, so is the construction of the nervous system of both groups. In
both groups an anterior brain issue out circumoesophageal commissures that link to a
suboesophageal ganglion which in turn give off a ventral nerve cord with segmental
ganglion in each segment to enervate the body wall and viscera. However some of the
arthropod features cannot be traced directly to the annelid ancestors. The insect tracheal
system has no equivalence in the annelid ancestor so is the heamocoel and the open
circulatory system. These can be traced to the Onychophora, which, being more typically
anellid like in being elongated in the anterior posterior axis, and possessing a pair of
nephridia in each segment that open at the base of each leg, also possess tracheal tubes and
a heamocoel, tracheal ventilation, an open circulatory system and appendages associated
with the mouth. The intermediate position of the Onychophorans can also be deduced from
the fact that they are typically segmented in the embryonic stages, a feature present in both
the adult annelid and arthropod.
Apart from having features common in both annelids and arthropods and thus serving as a
probable link between the two the onychophorans also have features peculiar to
themselves which warrant them being placed in a separate class. Snodgrass (1952) has thus
envisioned a classification which ties the three groups as phyla in the super phylum
Annulata (Romoser 1973). By the Cambrian each group was firmly fixed and it is thus
possible that they all emerged in the Precambrian.
That the progenitor of these three groups is an annelid or annelid-like ancestor is now
widely accepted, and the polycheate annelids, being the oldest of the annelids have been
touted as the most probable ancestors or the closest to the real ancestor (Romoser, 1973).
Opinion differs on the exact transition from the annelid-like ancestors to the arthropods.
Whilst Meglisth (1967) envisioned a direct evolution from polycheate to arthropods
Snodgrass (1982) and Sharov (1966) are of the opinion that from an ancestral wormlike
organism evolved two lines; one that gave rise to the annelid polycheate and the other one
giving rise to a lobopod, the latter subsequently branching to give the Onychophorans and
arthropod groups. (Romoser 1973) (Fig. 2.3)
Figure 23? Schematic representation of the proposed evolutionary tree of the Arthropoda
The establishment of the arthropod line then saw the branching to give three groups which
are often given subphylum status; the Trilobita (trilobites), Chelicerata (chelicerates) and the
Mandibulata (mandibulates). The trilobites, having had their heydays in much of the
Palaeozoic failed to witness the end of this era, they become extinct in the Devonian period
and are known to us only through their fossil remains. The chelicerates as we have seen in a
previous chapter are still extant and are represented today by the horseshoe crabs,
scorpions, spiders, ticks and mites. The Mandibulata are divided into a number of classes:
the Insecta or Hexapoda (beetles, butterflies, bees, locust, bugs, flies etc) the Crustacea
(crabs, shrimps, barnacles, copepods, and sow bugs) and the Myriapoda which is a
composite group composed of four classes; the Chilopoda (centipedes), the Diplopoda
(millipedes), the Symphyla (symphylans), the Pauropoda (pauropopds).
The kinships between the three subphyla is clouded in controversy with some investigators
envisioning a monophyletic origins and others a diphyletic one but it is clear that by the
Cambrian all three groups had established themselves quite well as shown by the fair
representation of each group in the rock strata dating to this age. Fossil evidence is also
conclusive on the fact that the Tribilobita are the oldest of the three groups.
Menton (1973) is of the view that these groups are polyphyletic. Thus the Mandibulata
(Crustacea, Chilopoda, Diplopoda, Symphyla, Pauropoda and Insecta) are a polyphyletic
group. She envisaged the development of the mandibles in Crustacea as being totally

different from that in other groups, having developed from the coxal endites (bases) of
limbs in the former and from the whole limb in the latter.
Insect origins
Now we narrow down to our target group, the insects. After the branching of the Trilobita,
the next to branch off were the Crustacea in the Silurian period leaving the Mryiapoda and
insect line (Sharov 1966, cited in Romser 1973). During the Devonian the Insecta branched
off, evolving from a Myriapoda or protomyriapoda-like progenitor. Menton is of the view
that the protomyriapoda from which the insects evolved from was more like a Symphylan as
evidenced by the similarities in the symphylan head appendages and mouthparts. This
landmark transition is thought to have involved the loss of locomotory organs from the rest
of elongate body and their retention in the region just distal to the head region. This region,
the thorax thus became a specialized area for locomotion, which as we will see shortly, also
developed wings.
The first insects to emerge in the Devonian were apterous (wingless). Today they are
represented by the Thysanura which are soft bodied. From these primitive insects some
insect groups were to develop which were better able to stand to the challenges of the
terrestrial environment. Thus we see in the Lower Carboniferous the evolution of insects
that were better able to cope with the ferocious predators (amphibians) that have just
emerged from the water onto land. These insects evolved wings. This allowed them to
evade the predatory amphibians and the reptiles that later evolved from the amphibians
during this time. These insects populated the terrestrial environment with very little
challenge from competitors and predators. The wing was however primitive in that although
capable of efficient flight could not be folded over the body when the insect was at rest
because of the lack of a wing flexion mechanism. The bearers of this primitive wing
(Paleopterous insects as they are called) thus could not fully utilize refuge provided by
microhabitats away from predators. The paleopterous insects had their heydays in the
Carboniferous and Permian and today are represented by the dragonflies (Odonata) and
mayflies (Ephemeroptera).
The Upper Carboniferous saw the emergence of insects with the ability to flex their wings
(Neopterous insects). The evolution of the wing flexion meant that they could fold their
wing over their abdomens when at rest. This had unprecedendent effects on the bearers.
These insects had the dual advantage of being capable of flight to escape predators and at
the same time occupy microhabitats like crevicies to evade the same when at rest. They
became the dominant insect group and have maintained this pole position to this day,
constituting 97% of all insects and 90% of the insect orders (Romoser (1973). The Neoptera
include all present day insects serve for the Apterygota and Paleoptera.
The last major or milestone achievement in the evolution of insects was the evolution of
metamorphoses in the neopterous insects in the Upper Carboniferous. Metamorphoses
involve a gradual change in body form from larval form to adulthood. This can be simple or
complex. In the simple or incomplete form, also referred to as hemimetabolous
metamorphosis, the young insect is a small wingless replica of the adult which develops into
the winged adult over a period of time, growing in spurts during periods of moulting or
ecdysis. Complete metamorphosis involves the development of a young one which is totally

different in form from the adult into the adult form through a number of larval stages.
Hemimetabolous metamorphosis was the first to evolve followed by holometabolous
metamorphoses in the late Upper Carboniferous. The evolution of metamorphoses had
marked survival advantages on the neopterous insects. The larval stages and the adult
stages are usually separated in space, nutrition and role with the result that there is no
competition for food. The young ones are usually ferocious feeders separated from the
adult whose role, in many species is reproduction,
Extant insect are categorised into as many as 29 orders basing on relationships between
them. We will come back to look at some of the characteristic of the major insect orders but
for now let us make a detour and look at the some of the characteristics that make insects
the champions in the metazoan world.
Biological success- what do we mean by

this?
Whilst the term success has an obvious meaning in the world of examinations and business
and related trades, in the biological world it assumes a slightly different meaning, which
however becomes quite obvious and apparent even to the uninitiated upon slight unveiling.
In the biological world the main yardstick for measuring success in a group of organisms is
species diversity. The more speciose a group is the more successful it is said to be. Using
this criteria the insects come up position one without any rival contender. If you doubt it
consider the following fact: to date the insects constitute the largest class not only of the
animal kingdom but also of the whole living world. The number of known insect species vary
from 700 000 to 1 500 000 which constitute about 70 to 90% of known species (Dhaliwal,
1999).
Organisms owe their success to inherent characteristics (inherited traits which thus can be
passed from parent to offspring) that confer a survival advantages on the bearers. This
means that the bearer/s can survive to attain reproductive age, reproduce offspring that will
in turn pass on the button stick as they in turn reproduce. We are thus carriers of the 'good'
genes that we should pass onto to the next generation! With the environment constantly
changing (as it has always been the case) it follows that the larger the repertoire of
advantageous traits the better off are the bearers; many of the traits offering survival
advantages under contemporary conditions and still many others to serve for any
eventualities. The latter constitute preadaptions. The present day adaptions, the
preadaptations and the novel adaptations which may crop up in the gene pool due to
mutations all serve to ensure that the bearer's survival is not limited to the present , but
also to the future so that the group continues to survive and diversify today and in future (in
the next epochs, eras and eons).
The aspect we have just alluded to brings us to the second dimension to success in
biological entities. This is survival of group through time, and, here we talk of deep time,
time into the past and into the future. This, as it were, has largely been availed through the

study of past life forms (Palaeontology) as presented in sedimentary rock strata, a branch of
great concern not only to the biologist but also to the geologist. Insects as a group are fairly
represented as fossils in rock strata from the Devonian (some 480million years ago). If the
insect have been in existence as early as this and are still extant, we can, with all due
respect hail them for having been able to stand the question of time.
Ever since these early times insects have occupied the habitats they occupy today serve for
the terrestrial environment. But even though, they were the first to occupy the terrestrial
environment, with the result that when the land vertebrates moved onto land in the
Carboniferous times, some 280 million years ago, they were ‘surprised’ to find out that the
new territory already had inhibitors in the form of insects populating the soil in burrows,
climbing and hoping from grass to grass, tree to tree and still others, in the air, flying. If
these second invaders were not surprised, then, part of the reason for their moving onto
the land was in pursuit of the insects as food items. The insects provided abundant food for
the first vertebrate invaders, the amphibians and later the reptiles that evolved from them
in the late Carboniferous to Permian. To deny the insects their top position, even on this
criterion alone, would be unjust to one group immensely favoured by nature; but denial
may not surprise me because the judge (humankind) is an interested part in the contest and
his strong bias is pervasive: consider this fact: dinosaurs roamed the earth in their diversity
for close to 200million years, but mankind, who has only been around for less than a million
years and is already under threat of climate change caused by his carelessness, want to
judge the dinosaurs as being one group 'least adapted' to cope up with the troubles of this
world.
We have already alluded to the cosmopolitan nature of insects in our opening remarks in
this chapter. Yes insects inhabit every conceivable habitat in the earth's biosphere. This
brings us to the third dimension on our measure of success- extent of geographical
distribution. It is only the widest variety of antics that can guarantee the bearer to survive in
any game in the repertoire of demanding games for survival in the different environments.
Except in the marine environments, insects are not found wanting in this aspect and they
have their representatives in the very cold of the cold environments, the very hot of the
hottest deserts. By geographical extent, insects occupy undoubtedly the most extensive of
territories and this makes them once again the most successful group of animals. Perhaps it
is because of their cosmopolitan nature that every human culture practices entomophagy!
As you pose to reflect on what you are reading about and gaze into the air and the
surroundings, perhaps the first animal that comes into your sight is an insect of some sort.
This brings us to our fourth dimension of biological success- large population numbers. We
have already noted that insects are the most speciose of not only animals but the whole
living world. In addition, insects are also good contenders for the pole position for
champions in the animal world in terms of population numbers. If the unicellular animals
are disregarded and only the metazoans are considered, then no doubt insects come out
tops. Many of us have seen, as we take a casual walk in the home garden, a single vegetable
leaf being home to several hundreds aphids, not to mention the thousands of ants that visit
our homes on foraging engagements, the hundreds of thousands of elate termites that
decorate the urban tower lights at dusk after rains in early summer and the tens of millions

of African migratory locusts that cloud the skies and dim the high intensity tropical sunlight
in spring in a fateful year.
Using all the four criteria commonly used to measure biological success; species diversity,
survival through geological time, extent of geographical distribution and population
numbers, insects come up tops on all aspects. They are indeed the undoubted land lords
and man should simply learn to coexist with them - a fact long known by the entomologists
and crop protectionists after the dismal failure of chemical control measures during the
insecticide boom of the 1940s to 60s.
What attributes make insects a very

successful group
We will turn on to look at some representative orders of insects later, for now we consider
those aspects of insects that have and still contribute to their success.
Throughout their extensive evolutionary history insects have acquired many distinctive
structural, developmental, physiological and behavioural attributes, well perfected, which
have and still enable them to remain the dominant group.
Waterproof exoskeleton
In a previous chapter we mentioned that the Cambrian boom was characterised by the
evolution of novel characteristics which resulted in the complete change of the fortunes of
the bearers. The evolution of a wax covered chitinous waterproof exoskeleton was one such
evolutionary invention that changed the fortunes of the insects (and the arthropods as a
whole) on a scale that was never to be repeated again in the evolution of this group. It was
a milestone evolutionary invention because the bearers could move onto the desiccating
terrestrial environment. What else would this group want? Because they were now
immune to desiccation which otherwise would have rendered the whole exercise futile
because of their small size, they could now occupy the land environment feeding on the
numerous plant food available. The Bryophytes and Pteridophytes had already occupied the
vast terrestrial environments. With abundant food, many opportunities opened up; with
fewer predators on land, the insects multiplied to populate all the terrestrial habitats and
niches hitherto vacant. They populated, as we have already noted, most favourable and
even the least of favourable environments.
Apart from waterproofing, the insect exoskeleton also serves a variety of purposes. It is light
for easy movement both on land and in air. It is of great strength for muscle attachment and
for protection against predators and acts as the first line of defence against pathogens
(viruses, mycoplasmas, bacteria, and protozoa). It is vital in the water relations of aquatic
insect by curbing excessive loss of salts from the body. It has been modified into a tool for
digging (as in the sclerotized claws of burrowing insects), for preying (as in the spined
forelegs of mantids for example) and oviposition (as in the ovipositor of many insects).
Above all it maintains the shape of the insect.

Differentiation of the body
The insect body is segmented and different regions have become specialized for various
purposes (tagmosis). It is in the insect world that tagmosis has been exploited in an
unparalleled way in the metazoan world. This division of the body into head, thorax and
abdomen is really suitable for terrestrial life. The head region has been specialized for
feeding and stimulus perception bearing as it does the mouth and associated mouthparts,
compound eyes, osceli and antennae. Its mobility about the adjoining thorax allows the
head to be flexible and this has allowed the development of various mouthparts and thus
feeding habits. This also allows for better orientation during flight and landing. The thorax is
highly integrated and this has made it a firm region equipped with the tools for locomotion.
Its rigidity provides a firm structure for leg attachment and yet a flexible base for wing
insertion (Dhaliwal 1999). The appendageless, telescoped abdomen allows for distension to
serve a food storage and digestive function as well as reproductive function by
accommodating eggs and often times used for embryo brooding. It also houses the
excretory organs and the fat body, the latter being used as an energy store to power growth
and flight.
Adaptability of body appendages

Closely linked to the differentiation of the body is the adaptability of the body appendages.
The insect body appendages are highly adaptable. The appendages surrounding the mouth,
the mouth parts, have been modified for various modes of feeding. In the orthopteran and
coleopteran orders they are typically mandibulate and are used by phytophagous insects to
bite off chunks of vegetation and chew it before swallowing. In the Hemiptera and some
Diptera the mouthparts are structured into dagger like, piercing and sucking mouthparts
used to reach deep into the vascular tissue of their plant or animal host to extract body
fluids that constitute their diets. The butterflies have long drawn out mouth parts that form
tubes used in sucking out nectar from nectaries of plants. Some Diptera have mouthparts
produced into proboscis used to deliver digestive saliva onto food source and suck the
liquefied food, and still others have rasping mouthparts designed to wear down host tissue
and suck the oozing blood. This diversity of mouthparts allow insect to feed on a wide range
of food sources and thus limiting the levels of competition by exploiting different niches.
The locomotory appendages whose centre is the thoracic region are also highly adaptable
and have been modified to perform various functions. The primitive orthopteran legs are
basically for walking and jumping or hoping from grass to grass or from tree branch to
another. To this end the hind leg is long and thick to propel the insect during takeoff as they
jump or take to flight. In cockroaches, speed is of paramount importance and the leg just
serves this. The legs are long to allow for speed and at the same time can be folded under
the body so that the insect can move through crevices and cracks and hide from predators.
In the burrowing insects such as cicada whose larva are subterranean, the legs (especially
the forelegs), are heavily clawed and highly sclerotized to enable them to act as digging
apparati. The same adaptation is seen in the crickets. In praying mantis the foreleg is
modified into forceps-like structure equipped with spines to grab and puncture prey. The
stick legs of the stick insects are ideal for the aquatic life, having been designed such that
the bearer can walk on water without breaking the hydrostatic attractions between water
molecules caused by the hydrogen bonds.

The terminal structures in the periproct, which in most insect are ovipositing structures,
have also been modified for offence and defence in the bees. In this group, the ovipositor
has lost their reproductive functions completely in the non reproductive female bees and is
used as an organ to inflict a painful sting to whoever dares to disturb the colony. The sting is
provided with a poison gland that produces toxins that are injected into the victim. The
toxin contains a lot of histamines that cause severe inflammation of the affected area.
The adaptability of the insect appendages thus allow the bearers to utilize various food
sources, escape predators, find mates with ease and defend the bearer from intruder,
amongst other things.
Small size of insects

Insect are in general small animals. Their small size confers a lot of advantages on the
insects. It allows them to occupy small habitats or microhabitats which other animals cannot
access. This serves to utilize habitats in which there are no competitors and at the same
time evading the larger predators. It also allows them to utilize small amounts of food in
large numbers- the situation of hundreds of aphids all obtaining nourishment from a single
vegetable leaf quickly comes to mind.
High reproductive powers
Insects have prolific powers of reproduction. A single insect can lay up to several hundreds
or even thousands of eggs in very short period of time. These high reproductive powers
compensate for mortality, so that even if mortalities are high (as is often the case in the
insectan world) still many offspring will survive and reach reproductive age. Many insect can
have one reproductive cycle in a year (univoltine) and still many do have more cycles in a
single year (multivoltine).
Quick speciation
Closely linked to the issue of high reproductive powers are the inherent opportunities which
come with rapid reproduction. With so many offspring being produced, chances are also
high that there will be manifold chances of gene reshuffling through independent
assortment of chromosomes during meiosis as well as new gene combinations due to
crossing over and mutations. The effects of these are wide variations in the offspring. The
wide variation pays dividends especially when it results in slight behavioural and structural
changes which may inhibit successful courtship and copulation between siblings. In this way
new species are rapidly generated. This is ease to conceptualize when we realize that a
slight change in the male or female genitalia may hinder successful copulation given that the
insect exoskeleton is hard.
Hexapod locomotion
The insect locomotion is such that at any given moment the insect will be balanced on a
tripod. This is a rigid structure which gives stability and hence high speeds can be attained at
the same time maximizing manoeuvrability. This is unlike the case with other arthropod
groups where numerous legs tend to cause clumsy movement. Efficient stability, high
manoeuvrability and speed allow for ease escape from predators, finding mates, colonising
new habitats, etc

Evolution of wings
The evolution of the insect wing brought, as we have seen many benefit to the insects. It
brought speed to locomotion in insects, thus instead of only relying on legs the insect could
also took to flight. The insect could evade predators with ease; colonize new habitats in
search for food or escaping adverse conditions. The migration of the African migratory
locust is mainly triggered by a deterioration of food quality as well as environmental
changes which act as cues for adverse conditions to follow. The insect wing, unlike the wing
in other flying animals such as birds and bats, did not evolve from the modification of the
walking appendage, but was a result of cuticular extensions and thus did not result in the
compromise of the animals walking capabilities. This wing, driven by very efficient thoracic
flight muscle is powered by one; very high energy molecule, trehalose, and can attain wing
vibration of up to 1000Hz. The noise that you hear when a mosquito approaches is
produced solely by rapid wing vibrations!
Tracheal respiration
The insect tracheal system is, unlike that in the vertebrates, independent of a circulatory
fluid and hence oxygen is delivered directly to the body tissues or cell. This makes it a very
efficient oxygen conveying system. The tracheal system is a system of tubes that ramify the
insect body and terminate, internally by poking in or between cells without penetrating
them and open to the outside through spiracles. The spiracles are in most insects guarded
by spiracular valves that regulate entry of oxygen and at the same time limit water loss from
the insect body. Regulation of air flow is brought about by alternate opening and closing of
spiracles in different regions of the body, allowing air to enter for instance in the thoracic
spiracles and exiting in the abdominal spiracles. This provides an efficient exchange system
as the air flow becomes unidirectional. From the series of numerous spiracles in the pleural
surface of the insect, short tracheal tubes lead and terminate in longitudinal lateral tracheal
trunks that run posteriorly and anteriorly. Primitively, these lateral tracheal trunks are
linked by dorsal and ventral trachea that connect to two dorsal and two ventral longitudinal
tracheal trunks that also run anteriorly and posteriorly. A series of visceral trachea link the
lateral tracheal trunks to the visceral organs.
In aquatic insects the insect tracheal system is connected to a rectal system which extracts
air from water.
Developmental characteristics
We have seen that insects, during their development undergo through a series of changes
collectively called metamorphosis. This can be incomplete or complete metamorphosis. In
either case, for many insect, the developmental stages are quite different in form and
requirements from the adult stages. This means that there is utilization of different food
sources and occupation of different habitats. As a result there is very little, if any
competition at all between adults and young ones. For example in the Odonata the young
ones (nymph/naiads) are carnivorous and aquatic whereas the adults, although carnivorous,
are not aquatic. In the holometabolous insects, there is a non-feeding stage, the pupa and it
may serve to overcome adverse conditions.
Chemical communication
Insect get a lot of information from conspecifics, other species and food items. This
information is usually in the form of pheromones and kairomones. Pheromones are released

by other conspecifics or other species of insects. These serves to warn others of impending
danger or eminent danger (alarm pheromone) or food sources (aggregation pheromones).
Other chemicals serve in courtship to attract members of the opposite sex and, in still other
members serve to curtail certain behaviours in the colony which may bring disharmony to
the members of the colony and at the same time serve to fix the castes system as in social
termites. Kairomes are chemical produced by the insect food sources and these serve to
discourage feeding by herbivorous insects or to attract insect for feeding.
Physiological adaptations.
Many insects exhibit some developmental arrests in their life cycles. During this period of
developmental arrest the insect is inactive and no development takes place. This is called
diapause and often serves to overcome adverse condition like very cold periods or periods
of inadequate food. Diapause can occur at any stage in the development of an insect; it can
be in the egg, larvae or adult stage and is elicited by environmental cues such as long day
length which may act as an indicator to the pending adverse conditions. When it occurs in
the adult the adult insect suspends all adult behaviours like courtship and mating.
Insects, being mainly terrestrial have also evolved ways to limit water loss. Insects are
uricotellic implying that they excrete uric acid, a relatively non-toxic nitrogenous waste
which can be flushed out of the insect body with minimal amounts of water. This thus
serves to conserve water in the body as minimum amounts of water are lost from the body.
Basic classification of insects

We have already alluded to the fact that the 29 extant insect orders are grouped into two
subclasses; the Apterygota and Pteryogota (Fig?). The Apterygota comprises of primitive
wingless insects which show little or no metamorphosis, a thorax not clearly demarcated
from the abdomen and the abdomen having bristle-like appendages. Their bodies are not
heavily cutinized and thus relatively soft and they have chewing and sucking mouthparts.
Only four orders belong to this subclass: the Protura, Diplura, Collembola and Thysanura.
The most common member in the Thysanura is the silverfish so named because of its silvery
body. This makes its home our libraries and bookshelves feeding on paper and glue.
The Pteryogota are primitively winged insects although some have secondarily lost the
winged condition. They are subdivided into two super orders; Exopterygota and
Endopterygota, depending on the nature of early development. The Exopterygota are
characterised by wings which develop externally and the Endopterygota by wings which
develop internally from imaginal buds. Both groups exhibit some form of metamorphoses.
The Exopterygota exhibit incomplete metamorphoses, with the wingless young one (nymph)
resembling the adult into which it develops. Because they exhibit incomplete
metamorphoses they are said to be hemimetabolous and they constitute the Hemimetabola
group. They are divided into the Paleoptera and Neoptera orders. The Endopterygota
exhibit complete metamorphosis in which the young (larva) is completely different form the

adult, and develops into the adult through a number of stages. For exhibiting complete
metamorphoses, they are said to be holometabolous and constitute the Holometabola
group. They constitute the four largest orders (Coleoptera, Hymenoptera, Lepidoptera and
Diptera) and few other minor orders.
Some of the Orders of extant insects
APTERYGOTA (Primitively wingless insects)

Order Diplura
Order Protura
Order Collembola- springtails
Order Thysanura- silverfish
PTERYGOTA (Winged, or secondarily lost)
(A) Exopterygota (Wings developing externally, hemimetabolous))
(I) Paleoptera (Wing without flexion mechanism, held up and not folded on abdomen)
Order Odonata- Damselflies and dragonflies
Order Ephemeroptera- mayflies
(II) Neoptera (Wings with flexion mechanism, fold top of abdomen, or tent like)
Order Orthoptera -grasshoppers, locusts, katydids
Order Phasmida- stick insects and leaf insects
Order Anoplura- sucking lice
Order Mallophaga -biting lice
Order Dermiptera- lacewings
Order Hemiptera- aphids, cicadas, stink bugs
Order Plecoptera- stone flies
Order Thysanoptera- thrips
Embioptera- web spinners
Isoptera- termites
Dictyioptera-cockroaches and praying mantids
(B) Endopterygota (Wings develop internally from imarginal discs, holometabolous))
Order Coleoptera- beetles
Order Lepidoptera- butterflies and moths

Order Diptera -flies (houseflies, tsetse flies, horseflies, fruit flies, carrion flies and
mosquitoes)
Order Hymenoptera- ants, bees and wasps
Order Trichoptera- insects with hairs on wings
Order Neuropteran- ant lions and related forms
Mecoptera- scorpion flies
Siphonaptera- fleas -insects with laterally compressed bodies
References
BARNES, R.S.K. (et al) (1993). The invertebrates: a new synthesis. Publisher: Oxford:
Blackwell, 1
Diversity of animals in Sweden
Brusca RC and Brusca GJ (2003). Invertebrates. 2nd edn. Sinauer Associates Inc. Publisher,
Saunderland, Massachusetts
Galas, F. and Sinervo B. (2002). Divergence and Convergence in early embryonic stages of
Metazoans. Contributions to Zoology, 71 (1/3)
Bush, Albert (et al) (2001). Parasitism: the diversity and ecology of animal parasites.
Publisher: Cambridge: Cambridge Univ. Press,
Bush et al (2003)
Gilbert, SF, Raunio, AM. (1997). Embryology. Constructing the organism. Sinauer Associates,
Sunderland, MA.
Moore, Janet (2003). An Introduction to the Invertebrates
Romoser, W.S. (1973). The Science of entomology. New Kork: Macmillan Publishing.
Wilson, EO (2001). The diversity of life. London, Penguin
Valentine, J. W. (1997). Cleavage patterns and the topology of the metazoan tree of life.
Proc Natl Acad: 94(15): 8001–8005.
RANA, S.V.S (2003). .Essentials of Ecology and Environmental Science. New Delhi : Prentice
- Hall,

WATSON, James D. (ed.) (2005).Darwin: The Indelible Stamp: The Evolution of an Idea.
Philadelphia: Running Press Book Publishers,

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Diversity of Life: The

major Animal Phyla

Module HBio 102

1 Caston Makaka (in prep)

CHAPTER 2 Basic biological classification and criteria

PART 2- THE PARAZOA

CHAPTER 3 Phylum Porifera (Sponges)

PART 3-THE RADIATA

CHAPTER 4 The Cnidaria

PART 4-THE BILATERIA I

CHAPTER 5 The Protostome Phyla.

PART 5-THE BILATERIA II

CHAPTER 6 The Deuterostome Phyla

PART 6-THE MOST SUCCESSFUL

CHAPTER 7 The insects: the champions of the metazoan world

CHAPTER 8 Fishes dominate the Vertebrate world.

5 Caston Makaka (in prep)

What are metazoans/Animals?

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The hypothetical multicellular metazoan or protometazoan is called a blastea and is similar

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Measuring Genetic diversity

Measuring Ecosystem diversity

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There is a maximum of 17,500 species of butterflies.

Therefore, there are approximately 2,430,000 species of insects.

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Animal group Total in group Number estimated

Birds 9 040 Known species account 98% of

Alpha (α), beta (β) and gamma (γ) diversity

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Diversity Grassland Miombo Acacia woodland

Where QS is the quotient of similarity;

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S = the total number of species recorded in the second locality (community)

c= the number of species common to both communities.

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H'=-∑ (185/438In234/438) + (87/438In87/438) + (54/438In54/438) + (44/438In44/438) +

Diversity Grassland Inividuals Miombo Individuals Acacia Individuals

And dividing H' by H'max we have 1.63/2.079= 0.78

For the grassland E H'=1.63/In8 =1.63/2.079=0,78

Where is the proportion of individuals belonging to

n = the total number of organisms of a particular species

16 Caston Makaka (in prep)

For the Grassland community d is given by

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Rarity or commonness of individual species (taxon) can be evaluated by measuring the

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Fig 1.4: Biodiversity hotspots (Source: http://www.bbc.co.uk/2/science/nature)

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Endemism and biodiversity

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Basic biological classification

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Categories used in Biological Classification.

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Figure 2.1: Categories used in the hierarch of biological classification

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The hierarchical classification of organisms is characterised by two inherent features. The

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Role of the International Code on