M.

Sc Zoology: SEMESTER-1; PAPER-I

UNIT-I: Biosystematics and Applications
1.1 BASIC PRINCIPLES OF CLASSIFICATION
1.2 SPECIES CONCEPT, MECHANISM OF SPECIATION, ALLOPATRIC
SYMPATRIC SPECIATION
1.3 APOMICTIC SPECIES AND PANMICTIC SPECIES
1.4 TYPIFICATION AND DIFFERENT TYPES OF ZOOLOGICAL TYPES

AND

1.1 BASIC PRINCIPLES OF CLASSIFICATION

1.1.1 Introduction

The living world consists of millions of species of organisms. These present

enormous diversity ranging from micro-organisms to the highest evolved plants and
animals. The knowledge about all these organisms will be highly confusing,
meaningless and useless if they are not properly identified and arranged
systematically.
The systematic arrangement of properly identified and named organisms is called
classification, systematics or taxonomy. (taxis = arrangement, nomos = order or
law)
Thus, taxonomy is a branch of biology which deals with collection of organisms,
their identification, nomenclature and systematic grouping or classification into
various categories. This is done on the basis of similarities and differences of their
morphological, anatomical, cytological, genetical, physiological, biochemical,
developmental and other characteristics.
The similarities of characteristics between species or groups of species indicate
their relationship. This is also gives us some idea about their phylogeny (i.e. their
evolutionary history).
The classification of plants into various groups is called plant taxonomy or
systematic botany. Similarly, classification of animals is called animal taxonomy or
systematic zoology.

1.1.2 Basic Principles of Classification

While developing a system of classification of organisms, certain basic principles are
observed. Some of these are as follows:
(i) Morphological criteria:

Morphology forms the primary basis for classifying organisms into various
taxonomic groups or taxa. In earlier artificial systems, only one or a few
morphological characters were taken into consideration (e.g. plants were classified
into herbs, shrubs, trees, climbers, etc. on the basis of their habit). The sexual
system proposed by Linnaeus was based mainly on the characteristics of stamens
and carpels.

Later on, in the natural systems of classification (e.g. Bentham and Hooker's
system of classification of plants), a large number of morphological characters were
taken into consideration. As a result, classification of plant groups was more
satisfactory and their arrangement was showing natural relationships with each
other.

The similarities in the morphological characters are used for grouping the plants
together. Because, these similarities indicate their relationships. On the other hand,
differences or dissimilarities of characters are used for separating the plant groups

from each other. Plant groups with greater differences are considered to be
unrelated or distantly related.
For example, all flowering plants with ovules enclosed in an ovary cavity are
grouped together as Division - Angiosperms whereas, the angiosperms are further
classified into two classes: Dicotyledons and Monocotyledons, on the basis of
differences of the characters of root system, leaf venation, flower symmetry and
number of cotyledons in the embryo.

(ii) Phylogenetic considerations :

In the more recent systems of classification of plants, a greater emphasis is given

on the phylogenetic arrangement of plant groups, an arrangement which is based
on the evolutionary sequence of the plant groups. These systems also reflect on the
genetic similarities of the plants. Some of the phylogenetic systems of classification
of plants are the ones proposed by Engler and Prantle (1887-1899), Bessey (1915),
Hutchinson (1926 and 1934), etc.

However, none of these or any other systems is a perfect phylogenetic system. This
is because, our present knowledge of the evolutionary history of plant groups is
very fragmentary and incomplete. At best, the present day systems can be
described as the judicious combination of both natural and phylogenetic systems.

Modern taxonomy takes into consideration data available from all disciplines of
botany for classification of plants. This helps immensely in establishing interrelationships of various plant groups. As a result, taxonomic arrangement becomes
more authentic and convincing.

(iii) Chemical taxonomy or chemotaxonomy:

It is a comparatively recent discipline. Chemotaxonomy is the application of phytochemical data to the problems of systematic botany.

The presence and distribution of various chemical compounds in plants serve as

taxonomic evidences. Nearly 33 different groups of chemical compounds have
been found to be of taxonomic significance.

(iv) Numerical taxonomy :

Application of numerical methods (data) in the classification of taxonomic units is

called numerical taxonomy.

Edgar Anderson (1949) was the first to make use of numerical taxonomy in the
classification of flowering plants. It involves exhaustive quantitative estimation of
taxonomic characters from all parts of the plant as well as from all stages in the life
cycle. The numerical data thus collected for various plant groups is tabulated
systematically. Computers are used for this purpose.

The main objective of numerical taxonomy is to clarify and illustrate degrees of

relationship or similarity in an objective manner. This branch is becoming an
indispensable aid in modern systematics.

1.1.3 Conclusion
Classification is essential for the proper study and easy reference to the immense variety
of life forms. Systematics deals with identification, nomenclature and taxonomic
classification of organisms. Species has a great significance as a taxonomic unit. Recent
taxonomy gives more importance to sub-species and populations. In the systematic
classification of organisms, various taxa are arranged in the descending order of their
taxonomic categories as per the taxonomic hierarchy. Modern taxonomy makes use of the
data from all branches of botany, including genetics, cytology, ecology, chemotaxonomy,
numerical taxonomy, etc. in order to develop a phylogenetic system of classification of
plants.

1.2 SPECIES CONCEPT

1.2.1 Introduction:

A species (plural: species) is one of the basic units of biological classification and a
taxonomic rank. A species is often defined as the largest group of organisms
capable of interbreeding and producing fertile offspring. While in many cases this
definition is adequate, the difficulty of defining species is known as the species
problem. Differing measures are often used, such as similarity of DNA, morphology,
or ecological niche. Presence of specific locally adapted traits may further subdivide
species into "infraspecific taxa" such as subspecies (and in botany other taxa are
used, such as varieties, subvarieties, and formae).

1.2.2 Species Concept:

The original concept of species has undergone a considerable change during the
progress of taxonomy. John Ray (1627-1705) was the first to distinguish genus and
species. However, the clear morphological concept of species was first given by
Linnaeus (1707-1778). Later on, Darwin proposed the biological concept of species.
The concept was further modified by Ernst Meyr.

The following species concepts are developed by the different taxonomists:

1) Biological species concept:

Species are groups of actually or potentially interbreeding natural populations,

which are reproductively isolated from other such groups (Mayr, 1940).

Biological species concept: A species is a reproductive community of

populations (reproductively isolated from others) that occupies a specific niche
in nature (Mayr, 1982).

Biological species concept: Species are the members in aggregate of a group of

populations that breed or potentially interbreed with each other under natural
conditions (Futuyma, 1986)

2) Cladistic species concept:

A species is a set of organisms (an evolutionary lineage) between two branch

points or between one branch point and an extinction event or a modern
population (Ridley 1993).

Fig: Cladistic concept: every time a speciation event occur, two new species are created
and the ancestral species becomes extinct
3) Cohesion species concept:
A species is the most inclusive group of organisms having the potential for genetic
and/or demographic exchangeability. (Templeton, 1989).
4) Competition species concept:
Species are the most extensive units in the natural economy such that reproductive
competition occurs among their parts (Ghiselin, 1974).
5) Ecological species concept:

A species is a set of organisms exploiting (or adapted to) a single niche (Ridley
1993).

Ecological species concept: A species is either 1) a lineage which occupies an

adaptive zone minimally different from that of any other lineage in its range, and
which evolves separately from all lineages outside its range, or 2) a closely-related
set of lineages which occupy an adaptive zone minimally different from that of any
other lineage in their range and which evolve separately from all other lineages
outside their range (translation of Van Valen, 1975).

Ecological species concept: A species is a lineage or a closely related set of

lineages, which occupies an adaptive zone minimally different from that of any other
lineage in its range and which evolves separately from all lineages outside its range
(Van Valen, 1976).

6) Evolutionary species concept:

A species is a lineage (an ancestral-descendant sequence of populations) evolving

separately from others and with its own unitary evolutionary roles and tendencies
(Simpson, 1961).

Evolutionary species concept: A species is a single lineage of ancestor-descendant

populations which maintain its identity from other such lineages and which has it
own evolutionary tendencies and historical fate (Wiley, 1981).

Evolutionary species concept: A species is a population or group of populations that

shares a common evolutionary fate through time (Templeton, 1989).

Figure: Evolutionary concept: a species does not necessarily become extinct during a
speciation event. Species 1 is paraphyletic after split from species 2.
7) Isolation species concept:

Species are systems of populations: the gene exchange between these systems is
limited or prevented by a reproductive isolating mechanism or perhaps by a
combination of several such mechanisms. (as defined by Dobzhansky 1970; in
Templeton, 1989).

8) Phenetic species concept:

A species is a set of organisms that look similar to each other and distinct from
other sets (Ridley, 1993).

9) Phylogenetic species concept:

A species is the smallest diagnosable cluster of individual organisms within which

there is a parental pattern of ancestry and descent (Cracraft 1983).

Phylogenetic species concept: A species is an irreducible (basal) cluster of

organisms, diagnosably distinct from other such clusters, and within which there is
a parental pattern of ancestry and descent (Cracraft 1989).

Fig: Example of Phylogenetic Species Concept: Ensatina salamander leneages

In the above example (Figure), Ensatina salamander lineages A and B are separate
species. Each has a common ancestor that individuals of other species do not.
Even though it has diversified a lot, Lineage C is a single species, according to the
phylogenetic species concept. None of the subspecies of Lineage C has a single
common ancestor separate from the other subspecies.

10) Recognition species concept:

A species is the most inclusive population of individual biparental organisms which

share a common fertilization system. (as defined by Paterson, 1985; in Templeton,
1989).

11) Typological species concept:

A species is a group of organisms conforming to a common morphological plan,

emphasizing the species as an essentially static, non-variable assemblage.
According to this concept the observed diversity of the universe reflects the
existence of a limited number of underlying "universals" or types (eidos of Plato).
Individuals do not stand in any special relation to each other, being merely
expressions of the same type. Variation is the result of imperfect manifestations of
the idea implicit in each species (Mayr 1969; Lincoln et al. 1982).

1.2 MECHANISM OF SPECIATION: ALLOPATRIC AND SYMPATRIC SPECIATION

1.2.1 Introduction

If species arent special creations, where do new species come from? Darwin found
the answer by concluding that lineages change over time and also multiply they
split in two. For Darwin, and all who followed, speciation is this process of
multiplication, occurring when one population splits into two reproductively isolated
populations.

Of major importance to Darwins thinking about speciation were the mockingbirds

and finches of the Galapagos Island which Darwin correctly believed had each
descended from one Central American species and multiplied on the islands. New
species were created as populations became adapted to filling different roles on the
island. This observation led Darwin to conclude that new species form by gradual
evolution and isolation from an ancestral population; eventually this leads to two
distinct species. If you catch it in the act, half way through this process you observe
what are called varieties, populations of one species which differ, but not enough
to justify being labelled as separate species. For Darwin therefore varieties and
species were just different degrees of the same thing: there is no fundamental
distinction between species and varieties.

1.2.2 Mechanism of Speciation

There are four geographic modes of speciation in nature, based on the extent to
which speciating populations are isolated from one another: allopatric, peripatric,
parapatric, and sympatric.

a) Allopatric Speciation
During allopatric (from the ancient Greek allos, "other" + Greek patr, "fatherland")
speciation, a population splits into two geographically isolated populations (for
example, by habitat fragmentation due to geographical change such as mountain
building). The isolated populations then undergo genotypic and/or phenotypic
divergence as: (a) they become subjected to dissimilar selective pressures; (b) they
independently undergo genetic drift; (c) different mutations arise in the two
populations. When the populations come back into contact, they have evolved such
that they are reproductively isolated and are no longer capable of exchanging
genes.
Examples:
Examples include insular dwarfism and the radical changes among certain famous
island chains, for example on Komodo. The Galpagos islands are particularly
famous for their influence on Charles Darwin. During his five weeks there he heard
that Galpagos tortoises could be identified by island, and noticed that Finches
differed from one island to another, but it was only nine months later that he
reflected that such facts could show that species were changeable. When he
returned to England, his speculation on evolution deepened after experts informed
him that these were separate species, not just varieties, and famously that other
differing Galpagos birds were all species of finches. Though the finches were less
important for Darwin, more recent research has shown the birds now known as
Darwin's finches to be a classic case of adaptive evolutionary radiation.

b) Peripatric Speciation:

Peripatric and peripatry are terms from biogeography, referring to organisms

whose ranges are closely adjacent but do not overlap, being separated where these
organisms do not occur for example on an oceanic island compared to the
mainland. Such organisms are usually closely related (e.g. sister species), their
distribution being the result of peripatric speciation.

Peripatric speciation is a form of speciation, the formation of new species through

evolution. In this form, new species are formed in isolated peripheral populations;
this is similar to allopatric speciation in that populations are isolated and prevented
from exchanging genes. However, peripatric speciation, unlike allopatric speciation,
proposes that one of the populations is much smaller than the other. One possible
consequence of peripatric speciation is that a geographically widespread ancestral
species becomes paraphyletic, thereby becoming a paraspecies. The concept of a
paraspecies is therefore a logical consequence of the Evolutionary Species
Concept, by which one species give rise to a daughter species. The evolution of the
polar bear from the brown bear is a well-documented example of a living species
that gave rise to another living species through the evolution of a population located
at the margin of the ancestral species' range.

Peripatric speciation was originally proposed by Ernst Mayr, and is related to the
founder effect, because small living populations may undergo selection bottlenecks.
Genetic drift is often proposed to play a significant role in peripatric speciation.
Examples:

The Australian bird Petroica multicolor and

Reproductive isolation occurs in populations of Drosophila subject to population
bottlenecking

Fig: Comparison of allopatric, peripatric, parapatric and sympatric speciation

c) Parapatric Speciation

In parapatric speciation, there is only partial separation of the zones of two

diverging populations afforded by geography; individuals of each species may come
in contact or cross habitats from time to time, but reduced fitness of the
heterozygote leads to selection for behaviours or mechanisms that prevent their
inter-breeding. Parapatric speciation is modelled on continuous variation within a
"single", connected habitat acting as a source of natural selection rather than the
effects of isolation of habitats produced in peripatric and allopatric speciation.

Parapatric speciation may be associated with differential landscape-dependent

selection. Even if there is a gene flow between two populations, strong differential
selection may impede assimilation and different species may eventually develop.[8]
Habitat differences may be more important in the development of reproductive
isolation than the isolation time. Caucasian rock lizards Darevskia rudis, D. valentini
and D. portschinskii all hybridize with each other in their hybrid zone; however,
hybridization is stronger between D. portschinskii and D. rudis, which separated
earlier but live in similar habitats than between D. valentini and two other species,
which separated later but live in climatically different habitats.

Ecologists refer to parapatric and peripatric speciation in terms of ecological niches.

A niche must be available in order for a new species to be successful.

Examples:
Ring species
o The Larus gulls form a ring species around the North Pole.
o The Ensatina salamanders, which form a ring round the Central Valley in
California.
o The Greenish Warbler (Phylloscopus trochiloides), around the Himalayas.
the grass Anthoxanthum has been known to undergo parapatric speciation in such
cases as mine contamination of an area.
d) Sympatric Speciation

Sympatric speciation refers to the formation of two or more descendant species

from a single ancestral species all occupying the same geographic location.

Often-cited examples of sympatric speciation are found in insects that become

dependent on different host plants in the same area.[10][11] However, the existence
of sympatric speciation as a mechanism of speciation is still hotly contested.
Scientists have argued that the evidences of sympatric speciation are in fact
examples of micro-allopatric, or heteropatric speciation.

The best illustrated example of sympatric speciation is that of the cichlids of East
Africa inhabiting the Rift Valley lakes, particularly Lake Victoria, Lake Malawi and
Lake Tanganyika. There are over 800 described species, and according to
estimate, there could be well over 1,600 species in the region. All the species have
diversified from a common ancestral fish (Oryzias latipes) about 113 million years
ago. Their evolution is cited as an example of both natural and sexual selection.

Until recently, there has been a dearth of strong evidence that supports this form of
speciation, with a general feeling that interbreeding would soon eliminate any
genetic differences that might appear. But there has been at least one study, in
2008, that suggests that sympatric speciation has occurred in Tennessee cave
salamanders.

Sympatric speciation driven by ecological factors may also account for the
extraordinary diversity of crustaceans living in the depths of Siberia's Lake Baikal.

Examples: three-spined sticklebacks

Fig: Difference between Allopatric and Sympatric speciation

1.3 APOMICTIC SPECIES AND PANMICTIC SPECIES
Apomictic Species

The species formed by without involving the fusion of male and female gametes in
reproduction is called as Apomictic Species.

Because apomictic specis are genetically identical from one generation to the next,
each lineage has some of the characters of a true species, maintaining distinctions
from other apomictic lineages within the same genus, while having much smaller
differences than is normal between species of most genera. They are therefore
often called microspecies.

In some genera, it is possible to identify and name hundreds or even thousands of

microspecies, which may be grouped together as aggregate species, typically
listed in floras with the convention "Genus species agg." (e.g., the bramble, Rubus
fruticosus agg.). In some plant families, genera with apomixis are quite common,
e.g. in Asteraceae, Poaceae, and Rosaceae. Examples of apomixis can be found in
the genera Crataegus (hawthorns), Amelanchier (shadbush), Sorbus (rowans and
whitebeams), Rubus (brambles or blackberries), Poa (meadow grasses), Hieracium
(hawkweeds) and Taraxacum (dandelions).

Although the evolutionary advantages of sexual reproduction are lost, apomixis can
pass along traits fortuitous for evolutionary fitness.

Panmictic Species

A panmictic population is one where all individuals are potential partners. This
assumes that there are no mating restrictions, neither genetic nor behavioural, upon
the population, and that therefore all recombination is possible. The Wahlund effect
assumes that the overall population is panmictic.

In genetics, random mating involves the mating of individuals regardless of any

physical, genetic, or social preference. In other words, the mating between two
organisms is not influenced by any environmental, hereditary, or social interaction.
Hence, potential mates have an equal chance of being selected. Random mating is
a factor assumed in the Hardy-Weinberg principle and is distinct from lack of natural
selection: in viability selection for instance, selection occurs before mating.

In simpler terms, it is the ability of individuals in a population to move about freely

within their habitat, possibly over a range of hundreds to thousands of miles, and
thus breed with other members of the population that defines panmixia (or
panmicticism).

To signify the importance of this, imagine several different finite populations of the
same species (for example: a grazing herbivore), isolated from each other by some
physical characteristic of the environment (dense forest areas separating grazing
lands). As time progresses, natural selection and genetic drift will slowly move each
population toward genetic differentiation that would make each population
genetically unique (that could eventually lead to speciation events or extirpation).

However, if the separating factor is removed before this happens (ex. a road is cut
through the forest), and the individuals are allowed to move about freely, the
individual populations will still be able to interbreed. As the species's populations
interbreed over time, they become more genetically uniform, functioning again as a
single panmictic population.

In attempting to describe the mathematical properties of structured populations,

Sewall Wright proposed a "factor of Panmixia" (P) to include in the equations
describing the gene frequencies in a population, and accounting for a population's
tendency towards panmixia, while a "factor of Fixation" (F) would account for a
population's departure from the Hardy-Weinberg expectation, due to less than
panmictic mating. In this formulation, the two quantities are complementary, i.e. P =
1 - F. From this factor of fixation, he later developed the F statistics.

1.4 TYPIFICATION AND DIFFERENT TYPES OF ZOOLOGICAL TYPES

1.4.1 TYPIFICATION

The designation of a nomenclatural type is called typification. It is the means by

whgicgh ames are allocated to taxa. The type method is the only way to determine
objectively and unequivocally the correct application of names to various taxa.
A type is a zoological object on which the original published description of a name
is based. It is the objective basis to which a given zoological name is permanently
linked. In other words it is the nucleus of a taxon and foundation of its name. Once
designated the type cannot be changed.
The type of a nominal species is a specimen, that of a nominal genus isa nominal
species and that of a nominal family is a nominal genus. According to this concept
all specimens conformed to the taxonomists concept of the type of a taxon were
considered typical. There were as many types as there were typical specimens
which formed the basis of species description. The concept was followed by
Linnaeus and his contemporaries.
The typification of the higher taxa was done by the process of elimination of a
typical element and even the
1001 Code did ot ratify this method for old cases but ruled for the designation of
types for the future works.

1.4.2 Different Types of Zoological types

The Process of inventing names relating with the term type continued. This resulted
in numerous different names which led us to believe that earlier workers were more
concerned with the invention of new names for replacing type or expanding it rather
than defining the exact role of the type. Frizzell listed as many as 233 such names.
Fernald listed 108, grouped in three categories:
1) Primary types or Proterotypes 2) Supplementary types 3) Ecotypes

The above three types are typical, specimens that have been used in published
descriptions of figures but consist of material which the authors have worked on or
such as have been collected at the original locality. Fischer gave list of principle
kinds of types. Blackwelder grouped such names of the types into the following
seven categories:

1) Primary types (the single nomenclatural types e.g. Holotype, Lectotype, Neotype).
2) Secondary types (the specimens from which the primary type must be selected e.g.
Syntypes, paralectotypes).
3) Tertiary types (other specimens originally set aside as of special taxonomic interest
to supplement the primary type; e.g. Paratype, Allotype)
4) Specimens identified as of special origin. E.g., Topotypes
5) Specimens identified as to time or person of identification. Eg. Metatype,
Homotypes or Homeotypes, etc.

6) Specimens identified as to special treatment or use, eg. Plesiotypes, Hypotypes.

7) Replicas of type specimens, e.g. Plastotypes.
All these terms together with some more are very often of occasionally used in the
zoological literature. All such terms are explained below:
1) Allotypes: It is a specimen of the opposite sex to the type. It can be designated
either at the time of typification or subsequently.
2) Apotype: A specimen, not the type upon which a subsequent or supplementary
description or figure is bases.
3) Autotype: Any specimen identified by the describer as an illustration of his species
and compared with the type.
4) Chirotype: A type specimen upon which a manuscript name is based.
5) Genotype: The species which is designated as the type species of a genus, upon
which it is based.
6) Geotype: A specimen from the type locality.
7) Haplotype: A generic type by a single reference (only species).
8) Holotype: The single specimen selected by the author of a species as its type, or
the only specimen known at the time of description.
9) Ideotype: A specimen named by the author after comparison with the type, but has
not been from the type locality.
10) Morphotype: A selected specimen of the second or later form of a dimorphic or
polymorphic species; its use is permitted only by international code of nomenclature
of bacteria but not by zoological code.
11) Neotype: A specimen designated or selected subsequently to serve as the type of a
name when all the original type specimens are destroyed or missing or believed to
be so.