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TAXONOMIC
STRUCTURE 4
Subordination of organisms in groups under groups is a fact of nature, and it results from evolutionary
divergence. When an ancestral form splits into subgroups which undergo divergence during course
of time, the resultant descendants are modified forms but resemble each other in decreasing degrees.
Members of a species resemble each other because they are genetically more close to each other.
This resemblance, however, decreases progressively in different species of the same genus, different
genera of the same family, different families of the same order, different orders of the same class
and so on. And, to establish such relationships is one of the main concerns of plant taxonomy. In
any biological classification, all this is accomplished by taxonomic hierarchy. Different hierarchical
categories of plant taxonomy have been mentioned under Article 1.5 of Chapter 1 (Table 1.1).

4.1 CONCEPT OF TAXA

Taxon (pl. taxa) may be defined as “a named taxonomic group of any rank”. Thus at family level, taxa
may be represented by the Ranunculaceae and Rosaceae, while Ranunculus and Rosa are examples
of generic taxa. The term “taxon” was actually coined to replace clumsy phrases such as taxonomic
entity and taxonomic unit. Furthermore, the organisms contained within a rank (e.g., species, genus,
or order) can also be referred to as taxa.

4.1.1 Taxonomic Characteristics and Taxonomic Characters

Characters by which an organism or group of organisms can be recognised is called a characteris-
tic. For example, flowers are characteristic of angiosperms and wood is characteristic of trees. On
the other hand, any part or shape of an organism that makes it possible to classify the organism is
called a character. For example, characters used in classification include the shape of the leaves,
arrangement of the reproductive organs, etc.
Taxonomic characters are actually features, such as form, structure, behaviour, and physiology,
that are assessed in isolation from the rest of the plant by taxonomists, for making comparisons and
interpretations. It is important to differentiate between characters and character states. For example,
width of the leaf may be a character, while leaves 6 mm wide are an expression of that character,
i.e., its character state. Often, characters are referred to as “good” or “bad” but this is strictly relative.
 Taxonomic Structure 47

Thus, a good diagnostic character, such as compound leaves in a group of plants that mainly possess
entire leaves, would be a bad character for separating taxa in a group in which leaf divisions were
either variable or often compound.

4.1.2 Problems of Hierarchy in Taxonomic Structures

It has been established that taxonomic structure is a hierarchical system. For the effective working
of this system, there should exist clear concepts regarding various taxa in plant taxonomy. Differing
from non-living things, a biologist faces peculiar type of problems in circumscribing the different
categories or taxa in biology. It is so because they show dynamic, evolving and variable popula-
tions. To solve this problem, any concept of taxa or taxonomic category has to be flexible to some
extent. It should be able to meet the special requirements of biology. Variability within and also in
between the groups has caused many subjective judgements. At higher levels of hierarchy, this has
also resulted in various types of controversies. For example, traditionally, there were two kingdoms,
viz., Animalia and Plantae. Such a classification has been challenged by several modern workers.
Copeland (1947) suggested four kingdoms, viz. Monera, Protista, Metaphyta and Metazoa. Whittaker
(1969) gave a five-kingdom classification of organisms, namely Monera, Protista, Plantae, Fungi and
Animalia. Edwards (1976) recognized seven kingdoms under two superkingdoms (Procaryota and
Eucaryota).
It has, however, been observed that the degree of arbitrariness in classification decreases as we
proceed down in the hierarchy, and it reaches to its minimum at a particular level i.e., the species.

4.2 CONCEPT OF SPECIES

A species is usually the smallest unit of classification. It includes individuals which are alike and
can breed with each other. Species are sometimes divided into subspecies and varieties on the basis
of small differences between populations. Species are given Latin binomial names.
Species is, thus, the fundamental unit of study in taxonomy, comprising all the populations of one
breeding group that normally are permanently separated from other such groups by marked disconti-
nuities. In case, the crossing between species does not occur then the resulting hybrids are normally
sterile, thus maintaining the reproductive barrier between species. This broadly genetic definition of
a species does not hold good for species that reproduce by self-fertilisation or by asexual means or
to extinct species. In all these cases, a species is delimited by observation of the similarities between
its members and dissimilarities between it and the other species.
With reference to botanical nomenclature, the specific epithet forms the second part of the binomial
(the first part being the generic name) and it is always written in lower case. In plants, the ending
of the specific epithet always agrees with the gender of the generic name.
There exist several infraspecific categories in the taxonomic hierarchy, but subspecies and then
the varieties are probably the most widely used. Groups of similar species are placed in genera.

4.2.1 Subspecies
A subspecies is a taxon within a species. All subspecies of a species differ in small ways. Although
they can breed with each other, they are usually found in different places or in different populations.
 48 Plant Taxonomy

While naming a subspecies, a third Latin name is put after the binomial. Binomial is the Latin name
of a species consisting of two words, of which first one is the name of the genus to which the spe-
cies belongs and the second one is the name which distinguishes the species from other species in
the same genus.
Subspecies is, thus, the rank subordinate to species in the taxonomic hierarchy. The term ‘sub-
species’ is used when two or more populations are separated in some way (e.g. morphologically or
ecologically) throughout their range. However, they are not usually genetically isolated. Generally, if
90% or more of a group of infraspecific individuals are recognizably distinct from another similar
group, then each may be ranked as subspecies. Often, this is referred to as “90% rule”.
The abbreviation “ssp.” or “subsp.” is used to indicate a subspecies, e.g. Portulaca pilosa ssp.
pilosa and Daucus carota subsp. gummifer.

4.2.2 Variety
The variety is a rank subordinate to species but above the category “form” in the taxonomic hierarchy
(Table 1.1, Chapter 1). Varieties are actually morphological variants, which may or may not have
a clear geographical distribution. Sometimes they represent only habit phase or colour. A variety
designated by one author may be designated as a subspecies or form by another author. A variety
produced by agricultural or horticultural techniques and not normally found in natural populations
is called a cultivated variety or cultivar. In a variety, there may also be recognised subvarieties.

4.2.3 Form
“Form” is the lowest rank (see Table 1.1, Chapter 1) normally used by taxonomists for sporadic
distinct variants that sometimes occur in populations. Forms may be relatively minor genetic vari-
ants of a variety or subvariety but their effects can be conspicuous. Commonly observed forms are
those in which flower colour is modified, e.g., occurrence of albino individuals in a population of
purple-flowered plants. Although, a form is more commonly used to distinguish variants of subspe-
cies and varieties, it may also be related directly to a species. Sometimes, “subforms” may also be
recognised within a form.
Above-mentioned details show that a genus is a group of species and a subspecies or a variety
or a form are parts of a species. Workers like Levin (1969), Cock (1977) and Wiley (1980) have
defined “species” in their own ways but still the definition of the term ”species” is a matter of great
discussion and controversy. Various views have been put forward by the biologists during last few
centuries. For the sake of convenience these can all be classified into four major concepts, viz.,
(i) nominalistic concept, (ii) taxonomic concept, (iii) biological concept, and (iv) alternative concept
of species. All these are discussed below, in brief.
(a) Nominalistic Concept of Species
According to Slobodchikoff (1976), the nominalistic concept of species suggests that the “nature
produces the individuals and nothing more”. Species is nothing but the brainchild of man and are not
objectively real. Different species can be explained only in terms of formal relationship and not on
the characteristics of organisms. According to workers like Burma (1954) and Spurway (1955), species
have been invented as a device to refer to large number of individuals collectively. Evolutionists like
 Taxonomic Structure 49

Haldane (1956) believe that species do not exist as taxa and that it is simply a unit of convenience.
About “species”, nominalists think that individuals are related and hence included in a species due to
similarity whereas the evolutionary point of view of workers like Mayr (1969, 1976) is that similarity
is due to evolutionary relationship, i.e., it is a common descent.
(b) Taxonomic or Typological Concept of Species
The name ‘species’ comes from the Latin root “specere” which means “to look at”. It, therefore, refers
to appearance. Aristotle, the great philosopher believed that biological species are highly variable and
also opined that hybridization between different species resulted into new species. Such examples
have also been given that different breeds of dogs have come into existence from breeding of bitches
with tigers, lions, wolves, foxes and even goats. Theophrastus, the Father of Modern Botany, wrote a
book “Enquiry into Plants” on transmutation of species. He opined that species changed when they
are transplanted in different soil and living in different climate. Several ancient workers reported that
orange was a new species created by grafting lemon on pomegranate. Similarly, banana originated
from the seed of date-palm (Zirkle, 1935) inserted into the corm of Colocasia.
(1) Aristotelian Concept of Essentialism Aristotle suggested that for every natural group, there is
an inner “essence” that makes them what they actually are and that this is real. This concept is
also known as the “typological concept of species”. According to this concept, as mentioned by
Simpson (1961), “every natural group of organisms, hence every natural taxon in classification has
an invariant, generalised or idealised pattern shared by all members of the group”. Species were
taken to be fixed units in nature, their number determinate and they were characterised by universal
types or “essence”. The essence is like floral diagram of a family, and all the genera are supposed
to fit in this and anything less or more is inconsequential. According to this concept, each species is
supposed to have a basic plan which is unchangeable and defined also by the universal characteristics
possessed by all members of the group. Several evolutionists refused to accept the taxonomic concept
of species because “they do not necessarily reflect actual species in nature.”
But, what exactly constitute the “essence” is a basic question. Supporters of the “concept of
essence” believe that for all practical purposes, the “essence” is deduced by observing a number of
individuals. The constant and invariable ones should constitute the “essence”.
(2) John Ray’s Concept of Species Botanists, such as John Ray (1686), put forward the theory of
“breeding relationship as the basis of species”. He also suggested that there exist variability within
species. In his Historia Plantarum published in 1686, Ray suggested “great care should be taken
in deciding what constitutes a species and what sort of characters are insufficient for species
delimitations”.
(3) View of Linnaeus Linnaeus (1707–1778) recognised two different kinds of variations among
organisms, the true differences created by the Creator and constituting the “essence” and the
intraspecific variations which were considered to be “sport of nature” or “accidents”. Linnaeus (1737)
warned that “sport” should not be neglected. “If neglected, these elusive ghosts glide away and are
gone”. Linnaeus, however, realised in the later years that species can arise by hybridisation.
(4) Darwin’s view Darwin (1859), however, provided a sound basis by explaining evolution of
species by natural selection. According to him, the organisms are characterised by variation. The
geometrical increase in the number of individuals takes place due to fertility of the organisms. But,
 50 Plant Taxonomy

it is subject to nature check i.e. the natural selection. Amongst these individuals only those survive
which possess an inherent advantage over the others in the population. These inherent advantages
are inheritable. For thousands of generations, the selection continues and new variants take the place
of the original ones in a rapidly changing environment. Mechanism of heredity was, however, not
known much during the times of Darwin.
(5) Neo-Darwinism Rediscovery of Mendel’s work in 1900 brought forward a new theory of evolution
in the 20th century, after the death of Darwin. It is called neo-Darwinism. It includes Darwin’s
theory of natural selection and the more recent knowledge of genetics and inheritance through
chromosomes. Due to this theory, phylogenetic relationship came to be accepted as the basis of
organismal similarities. It has been suggested that groups of organisms are not related because they
are similar, but they are similar because they have a common descent.
(c) Biological Concept of Species
Concept of species has altogether changed during last century due to considerable amount of work
done on various aspects, such as (i) genetic basis of variations, (ii) reproductive mechanisms, (iii)
population structure, (iv) breeding behaviour, etc. All these have led to the formulation of biologi-
cal concept of species by Mayr (1969), which has been supported by workers like Stebbins (1970),
Heywood (1974), and others. Mayr (1969) defined the biological concept of species by genetic kin-
ship as evidenced by breeding behaviour. Grant (1959) defined such a species as “a community of
cross-fertilising individuals linked together by bonds of mating and isolated reproductively from
other species”. This shows that biological species is formed by “groups of interbreeding natural
populations that are reproductively isolated from such other groups”. Two points mainly considered
in biological concept of species are (i) interbreeding between the members of the same species, and
(ii) reproductive isolation between the members of different species. The biological concept of spe-
cies has received wide acceptance among the modern biologists and it has inspired several studies
on various aspects of biology, especially genetics and breeding behaviour.
One of the greatest shortcomings of biological species concept is that it is not applicable to the
non-sexual organisms.
(d) Alternative Concepts of Species
(i) Concept of Evolutionary Species Grant (1971) proposed the concept of evolutionary species which
is equally applicable both sexual and asexual organisms. This concept as defined by Grant represents
“a spatio-temporal lineage of populations that evolve separately from other lineages and has its
own ecological niche”. In its present form, however, this concept is too vague to be applied in all
forms.
(ii) Concept of Sibling Species In several plant and animal species, there exist many instances where
“morphologically similar or identical populations are reproductively isolated”. Such species have
been named as sibling species by Mayr (1963). According to Amadon and Short (1976), the term
“sibling species” is used as a “purely descriptive term having no genetic or taxonomic implication”.
Stayskal (1972) criticized the use of the term “sibling species” because it exactly means of “same
parentage”. Instead, he used the alternative term “aphanic species” for “sibling species”.
(iii) Concept of Ecological Species According to Van Velen (1976), “species are maintained for the
most part ecologically and not reproductively”, and this is called concept of ecological species. This
 Taxonomic Structure 51

concept suggests that (i) genes are of minor importance in evolution, (ii) the control of evolution is
mainly by ecology and the constraints of individual development, and (iii) selection works primarily
on phenotypes which are the main building blocks of community. Van-Velen defined species as
“a lineage which occupies an adaptive zone minimally different from that of any other lineages
outside its range”.
(iv) Concept of Selection Species Slobodchikoff (1976) viewed species as a unit of selection and this
formed the basis of the concept of selection species. According to him, “species is a system of
genetically similar individuals and populations maintained as a cohesive unit by a set of selection
pressures that balance the disruptive forces imposed by environmental factor, mutations or genetic
recombinations”. The entire concept of Slobodchikoff turns round selection pressures.
Conclusion: All the above-mentioned concepts clearly indicate that there is no universally appli-
cable definition for species. In biology, the rank of species covers a variety of situations, and therefore,
the concept or definition of species should be excercised by considerable flexibility. To conclude, it
may be mentioned that species is a group of individuals that (1) actually or potentially interbreed
with each other but not with other such groups, (2) show continuous morphological variation within
the group but which is distinct from other such groups. Taxonomically, species are grouped into
genera and divided into subspecies and varieties or, horticulturally, into cultivars.

4.3 CONCEPT OF GENUS

Most widely accepted definition of the word “genus” (pl. genera) is that “it is a group of related spe-
cies”. In the taxonomic hierarchy, “genus” is an important rank which is subordinate to family, but
above the rank of species. It is a group of obviously homogeneous species. The generic name forms
the first part of the binomial (the second part being the specific epithet), e.g. Cicer arietinum.
Genus is usually a singular noun and is written in Latin with a capital initial letter. Collections
of similar genera are grouped into families. Davis and Heywood (1963) have suggested three main
parameters that should provide guidelines for the determination of the generic status, viz. (i) natu-
ralness, (ii) delimitation of closely related genera, and (iii) practicability of keeping them distinct or
including them in other genera.
The concept of monophyletic taxa has proved quite satisfactory in determining the naturalness
and demarkation between different genera. Monophyletic group means that its members must have
descended from a common ancestor and it must include all the descendants of that ancestor. Legendre
and Vaillancourt (1969) defined genus as a monophyletic group of species which occupies a definite
adaptive zone. According to them, all the species descending from a common ancestor have to be
included in the same genus. All non-monophyletic genera have to be abandoned because they do not
represent natural groups.
In the light of recent developments the circumscription and delimitation of the genera may also be
redefined. For example, the “Eugenia complex” of Myrtaceae may include all the new world species
in the genus Eugenia Linn. and all the old world species in the genus Syzygium Gaertn.
However, while revising the generic limits, it should be done mainly on the basis of a worldwide
study because it needs in-depth knowledge of the range and patterns of variations. Instead of taking
a single arbitrarily selected character for revising a genus, taxonomists should better be guided by
what is called a synthetic approach. More emphasis should be given on natural grouping.
 52 Plant Taxonomy

4.4 CONCEPT OF FAMILY

A taxon consisting of related genera is called a family. The Latin names of families usually end
with “–aceae” e.g. Ranunculaceae, Malvaceae, etc. Eight exceptions to such an ending, however, are
Compositae, Cruciferae, Gramineae, Guttiferae, Labiatae, Leguminosae, Palmae and Umbelliferae.
International Code of Botanical Nomenclature has, however, proposed alternative names for these fam-
ilies as Asteraceae for Compositae, Brassicaceae for Cruciferae, Poaceae for Gramineae, Hypericaceae
for Guttiferae, Lamiaceae for Labiatae, Fabaceae for Leguminosae, Arecaceae for Palmae, and
Apiaceae for Umbelliferae. Many workers, however, still use older names.
A family is a major category in taxonomic hierarchy comprising groups of similar genera. Some
taxonomists believe that “families” represent the highest natural grouping. Groups of similar families
are placed in orders.
Larger families may be split into tribes. The tribe is, therefore, a rank subordinate to family but
superior to genus in the taxonomic hierarchy (see Table 1.1, Chapter 1). The term ‘tribe’ is applied
to assemblages of similar genera within large families. The Latin names of tribes have the ending
“-eae”, e.g. tribe Saniculeae in the family Umbelliferae or Apiaceae. Similar tribes may be grouped
together in subfamilies. Tribes may also be split into subtribes. The Latin names of subtribes have
the ending “-inae”.
Ideally, families have to be natural and monophyletic. The characters, by which families are
delimited vary with the groups, but they are of more obvious nature than that of genera and species.
Both vegetative and reproductive characters are used in delimiting families. For example, Cactaceae
is recognised by the cactoid habit of the members while insectivorous families (e.g. Nepenthaceae
and Droseraceae) are characterised by their distinct insect-trapping mechanisms. Compositae and
Umbelliferae are recognised by their characteristic inflorescences while Poaceae is recognised by
their fruits.
Two different types of families are definable and indefinable (Walters, 1961). The families which
are very homogenous and natural groups are called definable, e.g. Brassicaceae and Apiaceae. These
can be easily recognised from such other familier because they are homogenous natural groups
and their component genera are ill-defined. On the other hand, indefinable families include great
diversity of structure and are not as distinctive as the definable families. Most of the present day
plant families are indefinable. Their member genera differ among themselves so much that they can
be easily separated, as in Ranunculaceae. Genera of indefinable families are easily definable due to
clear morphological discontinuities.
Sometimes, the question of inclusion of a particular genus in a given family becomes largely a
matter of choice of taxonomist. For example, Nyctanthes is included by some taxonomists in Oleaceae,
by others in Verbenaceae and by still others in Nyctanthaceae.

4.5 TAXA ABOVE FAMILY LEVEL

Order, Class, Division, and Kingdom are the categories above the level of family used in taxonomic
hierarchy. They are also sometimes divided into their lower categories like suborder, subclass and
subdivision. One more category “superorder” has been included by Takhtajan (1969) to accommodate
 Taxonomic Structure 53

one more evolutionary node. Ideally, all these groups should be monophyletic. Regarding evolution
of these higher taxa (Kubitzki, 1977), much information is, however, not available.
Order is a taxon consisting of families. The Latin names of orders usually end with “-ales”, e.g.
Rosales. However, some orders, which were erected prior to the compilation of the International Code
of Botanical Nomenclature, end in “-ae”, e.g. Tubiflorae, Glumiflorae. Groups of families, thought
to possess a degree of phylogenetic unity, are placed in an order.
Class is a taxon consisting of orders. In Botany, class is a taxonomic rank below division and
above order. The names of classes end in “-phyceae” in algae, -mycetes in fungi, or -opsida in other
plants.
Division is a major taxon, which is made up of classes. Three main divisions of land plants
are bryophytes, pteridophtes and spermatophytes. A division is the second highest category in the
taxonomic hierarchy, placed above the classes and below the kingdom. The Latin names of division
terminate in “-phyta”, e.g. Tracheophyta. In place of division, several botanists now use the term
phylum.
Kingdom, the largest of all the taxa, is actually the highest level in the hierarchy of taxonomic
ranks. In older systems, of classification, there are only two kingdoms, viz., plant kingdom and
animal kingdom. In some modern systems of classifications, fungi are considered in a separate
kingdom—Mycota. Similarly, some taxonomists prefer to place unicellular organisms in their own
kingdom, the Protista. Whittaker (1969) proposed a five-kingdom system of classification of living
organisms, namely, Monera, Protista, Plantae, Fungi and Animalia.

Test Your Understanding

1. Give a brief account of concept of species.
2. Explain briefly the concept of taxa.
3. Give an account of problems of hierarchy in taxonomic structures.
4. Describe briefly the biological concept of species.
5. Differentiate between: (a) subspecies and variety, and (b) variety and form.

Suggested Reading
Amadon, D. and L.L. Short, 1976, Treatment of subspecies approaching species status. Syst. Zool. 25:
161–167.
Cock, A.G. 1977, Bernard’s symposium – the species concept. Bio. J. Linn. Soc. 9: 1–30.
Edwards, P. 1976, A classification of plants into higher taxa based on cytological and biochemical criteria.
Taxon 25: 529–54.
Grant, V. 1971, Plant speciation. New York.
Haldane, J.B.S. 1956, Can a species concept be justified? Syst. Assoc. Publ. 2: 95–96.
Heywood, V.H. 1974, Principles and concepts in the classification of higher taxa. Pl. Syst. Evol. Suppl.
1: 1–12.
54 Plant Taxonomy

Legendre, P. and P. Vaillancourt 1969, A mathematical model for the entities, species and genus, Taxon
18: 234–252.
Levin, D.A. 1969, The nature of plant species, Science 204: 381–384.
Mayr, E. 1969, The biological meaning of species. Biol. J. Linn. Soc. 1: 311–320.
________ 1976, Is the species a class or an individual? Syst. Zool. 25: 19.
Slobodchikoff, C.N. 1976. Concepts of species. Vol. III. Pennsylvania.
Van Valen L. 1976. Ecological species, multispecies and oaks. Taxon 25: 233–239.
Walters, S.M. 1961, The shaping of angiosperm taxonomy, New Phytol. 60: 74–84.
Wiley, E.O. 1980, Is the evolutionary species fiction? Syst. Zool. 29: 76.