:Angiosperm Classification Systems:

Phenetic versus Phylogenetic systems; Cladistics in Taxonomy

Phylogenetic Characters in Taxonomy:

Phyletic or Evolutionary or Phylogenetic characters are used primarily in
phylogenetic classification. The term Phylogenetic is used in many ways. Haeckel (1886)
used the term in evolutionary history of a group. Recently a particular approach to
classification (cladism) have used phylogentic to refer to reconstructing only the
branching sequence of phylogeny. This approach is called cladistics with phylogenetics
retained in its original and broader usage. The character distinction is in between
homologous versus analogous. Homologue means the same organ in different animals
under every variety of form and function. Analogue means a part or organ in one animal
which has the same function as another part or organ in a different animal. After
Darwin’s theory of evolution, the homologous organs were viewed as structural
modification of the same organ, inherited from a common ancestor. Analogues are those
features developed by different organs to the same selection pressure. The detection of
homologus character is a difficult task for a phylogenetic reconstruction. Because of
complexities, botanists have tended to deal with the problem obliquely. Phylogentic and
Ontagenetic characters are simply features which are presumed to reflect information
about the phylogeny of the group; and developmental features. A regressive character is
one in which loss of appendages occur. In context of evolution these are also known as
adaptive or non-adaptive characters. Phylogenetic classification attempt to reflect the
geneology or evolutionary history of a particular group of plants.

Cladistic Characters:
The characters have developed from the cladistic approach to classification attempting to
determine branching sequence of evolution and base a classification upon them. Only
derived character states are regarded as significant cladistically. Characters are Primitive
vs. derived character states; or as synonyms, general vs. unique; generalized vs.
Specialised, Primitive vs. advanced, Plesiomorphic vs. apomorphic and Pfesiotypic vs.
apotypic etc. Plesiotypic and Apotypic terms were used by cladists like Wiley (1981),
Wagner (1983) etc. Shared derived character states between and among the taxa are
called synapomorphies (or synapotypies) and shared primitive states are
symplesiomorphies (or symplesiotypies).

Automorphy:
Derived character state occurring only in one evolutionary line and has no direct use in
constructing branching sequences.
Compatible character: Useful cladistical characters are called compatible characters
where evolutionary directionality of the states within each character is the same.
Problems with Phylogenetic Classifications:
i. Convergent Evolution: Species with similar selection pressures look alike i.e., appear
alike, and hence, can ‘trick’ a taxonomist.
ii. Lack of Fossils: Fossilization is a sporadic process. Some events happened so quickly
that fossils may not adequately document the changes i.e., the angiosperms appeared very
rapidly in fossil records.

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iii. Strict Evolutionary Classification: It assumes a monophyletic origin of groups i.e., the
ancestors can only came from one group; they cannot be polyphyletic e.g., the members
of your immediate family i.e., grandparents, parents, siblings make up a group with a
single origin (monophyletic). Neighbours are not included in this group since they have a
different origin. Logical Conclusion: All angiosperms arose from a single common
ancestor.

Phenetic Characters in Taxonomy:

An approach to biological classification which uses overall similarity to assess
relationships is called Phenetics or numerical taxonomy. Phenetic classification makes no
attempt to reflect evolution; taxa are related based on similarity and difference of
character states regardless of the evolutionary content of the characters and states
reflected. The character of choice in phenetics is the Unit character or Single character.
In phenetics each unit character is given the same or no weight while in phyletic
and cladistic classification the weighting of character is believed to be evolutionarily
important. Phenetic classification are based on overall similarities of characters.
Dendrogram: It is branching diagram in the form of tree and depicts degree of
relationship.
Phenogram: It is representation of phenetic relationship.
Cladogram: It is the depiction of cladistic relationship.
Seven main advantages of numerical taxonomy over conventional taxonomy:
(1) The greater the content of information in the taxa of a classification and the more
characters on which it is based, the better a given classification will be.
(2) A Priori every character is of equal weight in creating natural taxa.
(3) Overall similarity between any two entities is a function of their individual
similarities in each of the many characters in which they are being compared.
(4) Distinct taxa can be recognized because correlation of characters differs in the group
of organisms under study.
(5) Phylogenetic inferences can be drawn from the taxonomic structure of a group and
from character correlations, given certain assumptions about evolutionary pathways and
mechanism.
(6) Taxonomy is practiced as an empirical sciences as opposed to interpretative or
intrutive science.
(7) Classification are based on phenetic similarity.
Most of these principles bear resemblance to the aims and methods of Adanson and are
therefore, known as Neo-Adansonian principles.
Numerical Taxonomy is based on Phenetic evidences, i.e., on similarities by observed
and recorded characters of taxa, and not on phytogenetic probabilities. Since, numerical
taxonomy is operational in the sense, it is divided into a series of repeated steps, allowing
its results to be checked back step by step.

General Description, Merits & Demerits of Major Systems of

Angiosperm Classification: Hutchinson, Takhtajan & Bassey

John Hutchinson: Angiosperm Classification System:

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John Hutchinson was a British botanist associated with Royal Botanic Gardens, Kew,
England. He developed and proposed his system based on Bentham and Hooker and also
on Bessey. His phylogenetic system first appeared as “The Families of Flowering Plants”
in two volumes. The first volume contains Dicotyledons (published in 1926) and second
volume contains Monocotyledons (published in 1934). He made several revisions in
different years. The final revision of “The Families of Flowering Plants” was made just
before his death on 2nd September 1972 and the 3rd i.e., the final edition, was published
in 1973.
The following principles were adopted by Hutchinson to classify the flowering
plants:
1. Evolution takes place in both upward and downward direction.
2. During evolution all organs do not evolve at the same time.
3. Generally, evolution has been consistent.
4. Trees and shrubs are more primitive than herbs in a group like
genus or family.
5. Trees and shrubs are primitive than climbers.
6. Perennials are older than annuals and biennials.
7. Terrestrial angiosperms are primitive than aquatic angiosperms.
8. Dicotyledonous plants are primitive than monocotyledonous
plants.
9. Spiral arrangement of vegetative and floral members are primitive than cyclic arrange-
ments.
10. Normally, simple leaves are more primitive than compound leaves.
11. Bisexual plants are primitive than unisexual plants and monoecious plants are
primitive than dioecious plants.
12. Solitary flowers are primitive than flowers on inflorescence.
13. Types of aestivation gradually evolved from contorted to imbricate to valvate.
14. Polymerous flowers precede oligomerous flowers.
15. Polypetalous flowers are more primitive than gamopetalous flowers.
16. Flowers with petals are more primitive than apetalous flowers.
17. Actinomorphic flowers are more primitive than zygomorphic flowers.
18. Hypogyny is considered as more primitive from which perigyny and epigyny
gradually evolved.
19. Apocarpous pistil is more primitive than syncarpous pistil.
20. Polycarpy is more primitive than gynoecium with few carpels.
21. Flowers with many stamens are primitive than flowers with few stamens.
22. Flowers with separate anthers are primitive than flowers with fused anthers and/fila-
ments.
23. Endospermic seeds with small embryo is primitive than non-endospermic one with a
large embryo.
24. Single fruits are primitive than aggregate fruits.

He divided the Phylum Angiospermae into two Subphyla Dicotyledones and

Monocotyledones. The Dicotyledones are further divided into two divisions: Lignosae
(arboreal) and Herbaceae (herbaceous).
The Lignosae includes, fundamentally, the woody representatives derived from
Magnoliales and Herbaceae includes most of the predominantly herbaceous families

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derived from Ranales. The subphylum Monocotyledones are divided into three divisions:
Calyciferae, Corolliferae and Glumiflorae.
1. The division Lignosae was further divided into 54 orders beginning with Magnoliales
and ending in Verbenales.
2. The division Herbaceae was divided into 28 orders beginning with Ranales and ending
in Lamiales.
3. The division Calyciferae was divided into 12 orders beginning with Butamales and
ending in Zingiberales.
4. The division Corolliferae was divided into 14 orders beginning with Liliales and
ending in Orchidales.
5. The division Glumiflorae was divided into 3 orders beginning with Juncales and
ending in Graminales.
So in the latest system of Hutchinson, the Dicotyledones consists of 83 orders and 349
families and Monocotyledones consists of 29 orders and 69 families.
Merits:
1. Hutchinson proposed the monophyletic origin of angiosperms from some hypothetical
Proangiosperms having Bennettitalean characteristics.
2. He made a valuable contribution in phylogenetic classification by his careful and
critical studies.
3. Monocots have been derived from Dicots.
4. According to him, the definitions of orders and families are mostly precise, particularly
in case of subphylum Monocotyledones.
Demerits:
1. There is undue fragmentation of families.
2. Too much emphasis is laid on habit and habitat. Thus, creation of Lignosae and
Herbaceae is thought to be a defect reflecting the Aristotelean view.
3. The origin of angiosperms from Bennettitalean-like ancestor is criticised by many,
because the anatomical structures of the early dicotyledons are not tenable with such
ancestry.

Armen Takhtajan: Angiosperm Classification System:

Takhtajan was a reputed palaeobotanist of Komarov Botanical

Institute of Leningrad, U.S.S.R. (now in Russia). He also made great
contributions in the field of angiosperm taxonomy. In 1942, he
proposed preliminary phylogenetic arrangement of the orders of
higher plants, based on the structural types of gynoecium and
placentation. After 12 years i.e., in 1954, the actual system of
classification was published in “The Origin of Angiospermous
Plants” in Russian language. It was translated in English in 1958. Later on, in 1964, he
proposed a new system in Russian language. To trace the evolution of angiosperm, he
was particularly inspired by Hallier’s attempt to develop a synthetic evolutionary
classification of flowering plants based on Darwinian philosophy.
The classification was published in ‘Flowering Plants: Origin and Dispersal’ (1969) in
English language. Later on, in 1980, a new revision of his system was published in
“Botanical Review”.
Takhtajan (1980) included the angiospermic plants under the Division Magnoliophyta.
The Magnoliophyta is divided into two classes Magnoliopsida (Dicotyledons) and
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Liliopsida (Monocotyledons). The class Magnoliopsida consists of 7 subclasses, 20
superorders, 71 orders and 333 families.
On the other hand, Liliopsida comprises of 3 subclasses, 8 super- orders, 21 orders and
77 families. The class Magnoliopsida starts with the order Magnoliales and ends in
Asterales and the class Liliopsida begins with Alismatales and ends in Arales.

The class Magnoliopsida is considered to be monophyletic in origin, probably derived

from Bennettitales-like ancestors or stocks ancestral to them. On the other hand, the
Liliopsida have been considered to be originated from the stocks ancestral to
Nymphaeales. He considered Magnoliopsida more primitive than the Liliopsida. The
principles as adopted by Takhtajan (1980) for interpreting the evolutionary lineages in
higher plants are mentioned in Table 4.2.

Merits:
1. The classification of Takhtajan is more phylogenetic than that of earlier systems.

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2. This classification is in a general agreement with the major contemporary systems of
Cronquist, Dahlgren, Thorne, and others. Both phylogenetic and phenetic informations
were adopted for delimination of orders and families.
3. Due to the abolition of several artificial groups like Polypetalae, Gamopetalae,
Lignosae, Herbaceae, many natural taxa came close together, viz. Lamiaceae (earlier
placed under Herbaceae) and Verbenaceae (placed under Lignosae) are brought together
under the order Lamiales.
4. Nomenclature adopted in this system is in accordance with the ICBN, even at the level
of division.
5. The treatment of Magnoliidae as a primitive group and the placement of Dicotyledons
before Monocotyledons are in agreement with the other contemporary systems.
6. The derivation of monocots from the extinct terrestrial hypothetical group of
Magnoliidae is found to be logical.
Demerits:
1. In this system, more weightage is given to cladistic information in comparison to
phenetic information.
2. This system provides classification only up to the family level, thus it is not suitable
for identification and for adoption in Herbaria. In addition, no key has been provided for
identification of taxa.
3. Takhtajan recognised angiosperms as division which actually deserve a class rank like
that of the systems of Dahlgren (1983) and Throne (2003).
4. Numerous monotypic families have been created in 1997 due to the further splitting
and increase in the number of families to 592 (533 in 1987), resulting into a very narrow
circumscription.
5. Takhtajan incorrectly suggested that smaller families are more “natural”.
6. Although the families Winteraceae and Canellaceae showed their 99-100%
relationship by multigene analyses, Takhtajan placed these two families in two separate
orders.

Charles E. Bessey: Angiosperm Classification System:

Charles E. Bessey (1845-1915) proposed a modified system

of classification of Bentham and Hooker. He separated the
gymnosperms from angiosperms. He reorganized the orders of
angiosperms. His system is popularly known as Besseyan system. It
was published under the title The Phylogenetic Taxonomy of
Flowering plants. Bessey’s system was based on primitiveness and
evolutionary advancement of plant groups. According
him Ranales was a basic group. Monocots and most of the dicot
groups evolved from it. Bessey based his system on the tradition of de
Candolle, Bentham and Hooker and Hallier. He was also influenced
by Darwin and Wallace. He taught that taxonomy must be based on evolutionary
principles. Like Wettstein he placed the Ranales at the origin of Angiospermae. He
proposed the ‘Ranalian concept of primitive flower’, later modified by Hutchinson. He
proposed 28 principles for determining which characters were primitive and which were
advanced in flowering plants. These are referred as Bessey’s Dicta (guiding principles).
General dicta:

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1. Evolution is not always upward, but often it involves degradation and
degeneration.
2. In general, homogeneous structures (with many and similar parts) are lower, and
heterogeneous structures (with fewer and dissimilar parts) are higher.
3. Evolution does not necessarily incvolve all organs of the plant equally in any
particular period, and one organ may be advancing while another is retrograding.
4. Upward development is sometimes through an increase in complexity, and
sometimes by a simplification of an organ or a set of organs.
5. Evolution has generally been consistent, and when a particular progression or
retrogression has set in, it is persisted in to the end of the phylum.
6. In any phylum the holophytic (chlorophyll-green) plants precede the colourless
(hysterophytic) plants, and the latter are derived from the former.
7. Plant relationships are up and down the genetic lines, and must constitute the
framework of phylogenetic taxonomy.
Dicta having special reference to the general structure of the flowering plants:
8. The stem structure with collateral vascular bundles arranged in a cylinder is more
primitive than that with scattered bundles, and the latter are to be regarded as derived
from the former.
9. Woody stems (as of trees) are more primitive than herbaceous stems, and herbs are
held to have been derived from trees.
10. The simple, unbranched stem is an earlier type, from which branching stems have
been derived.
11. Historically the arrangement of leaves in pairs on the stem is held to have
preceded the spiral arrangement in which the leaves are solitary at the nodes.
12. Historically simple leaves preceded branched ("compound") leaves.
13. Historically leaves were first persistent ("evergreen") and later deciduous.
14. The reticulated venation of leaves is the normal structure, and the parallel venation
of some leaves is derived from it.
Dicta having reference to the flowers of flowering plants:
15. The polymerous flower structure precedes, and the oligomerous structure follows
from it, and this is accompanied by a progressive sterilization of sporophylls.
16. Petaly is the normal perianth structure, and apetaly is the result of perianth
reduction (aphanisis).
17. The apochlamydeous perianth is earlier and the gamochlamydeous perianth is
derived from it by symphysis of the members of perianth whorls.
18. Actinomorphy is an earlier structure than zygomorphy, and the latter results from
a change from similar to dissimilar growth of the members of the perianth whorls.
19. Hypogyny is the more primitive structure, and from it epigyny was derived later.
20. Apocarpy is the primitive structure, and from it syncarpy was derived later.
21. Polycarpy is the earlier condition, and oligocarpy was derived from it later.
22. The endospermous seed is primitive and lower, while the seed without endosperm
is derived and higher.
23. Consequently, the seed with a small embryo (in endosperm) is more primitive than
the seed with a large embryo (in scanty or no endosperm).
24. In earlier (primitive) flowers, there are many stamens (polystemonous), while in
later flowers there are fewer stamens (oligostemonous).
25. The stamens of primitive flowers are separate (apostemonous), while those of
derived flowers are often united (synstemonous).
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26. The condition of powdery pollen is more primitive than that with coherent or
massed pollen.
27. Flowers with both stamens and carpels (monoclinous) precede those in which
these occur in separate flowers (diclinous).
28. In diclinous plants the monoecious condition is the earlier, and the dioecious later.
He considered Spermatophyta as having a polyphyletic origin, being composed by
three different phyla, of which he only treated Anthophyta (syn.: Angiosperms). In that
he used the same names for the subclasses of both monocotyledons and dicotyledons, this
is contrary to contemporary rules on plant nomenclature that require names to be unique.
However, Bessey actually used a qualifying hyphenation (Alternifoliae-Strobiloideae and
Oppositifoliae-Strobiloideae), a distinction not always recognised in reference to this
scheme. With some modifications, most modern classifications - for example, those
of Cronquist (1981, 1983, 1988), Takhtajan (1969, 1980, 1983,
1991), Stebbins (1974), R. Dahlgren (1975, 1980, 1983; R. Dahlgren et al. 1981; R.
Dahlgren and F. N. Rasmussen 1983; R. Dahlgren and K. Bremer 1985; G. Dahlgren
1989), and Thorne (1976, 1981, 1983, 1992) - follow the Bessey tradition.
“Bessey’s Cactus,” it sounds a lot like a type of desert cactus named after a milking cow. 
However, “Bessey’s Cactus” is, in fact, not a plant at all.  It is a scientific diagram that
depicts how angiosperms, or flowering plants, evolved and are connected.  The shape of
“Bessey’s Cactus” loosely resembles a beavertail
cactus, and is named after the man who created it,
Dr. Charles E. Bessey, a renowned botanist from
the early 20th century.  
Dr. Charles E. Bessey was a botanist and educator
from Ohio.  Bessey was recognized for his well-
known diagram depicting the evolution of
angiosperms – known as “Bessey’s Cactus.”  He
was also well known for his innovative idea
regarding education.  Bessey emphasized scientific
and laboratory training in the agricultural
curriculum.   He combined the university level
agricultural program with the other university
science departments so that they shared common
core science courses.  Bessey was also committed
to the idea of learning through practice rather than
memorization.   In fact, his botany laboratory
course was the first offered to undergraduates in
the United States.  Despite his contributions to agricultural education, those were not his
most notable works in the field of botany.  
His most memorable work in the field of botany was the development of the first
explicitly phylogenetic classification of flowering plants – in other words the
classification of flowering plants using the evolutionary relationship of the different
species.  Bessey asserted that the angiosperms are the most diverse group of land plants,
as well as the most primitive of the flowering plants. 
He contended that they are the most primitive because they possess several features
considered close to gymnosperms – seed-bearing plants.  Gymnosperms are considered a
more primitive type of plant due to the fact that seeds are not enclosed in an ovule. 
Bessey argued that angiosperms derived from a single ancestral group. 
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He developed a set of 28 rules for distinguishing genetic characteristics or evolutionary
trends in plants.  Based on these characteristics and trends, he formed a classification
system, and designed a diagram to represent how the plants were connected to each other
and which of these plants were evolutionarily primitive or advanced.  This diagram was
rather intricate, and was nicknamed “Bessey’s Cactus” because the branching, bending
shape looks somewhat like a beavertail cactus.
Merits:
Besseys system has been followed by a large number of supporters in the recent years on
account of the following merits:
1. The system gains support due to a more acceptable choice of the Ranales as the
primitive order and the concept developed there upon of the primitive flower.
2. According to him the families with superior ovary precede with families with inferior
ovary.
3. He abolished Monochlarnydeae in his system and distributed the families of
Monochlarnydeae in Oppositifoliae of Dicotyledons.
4. Although initially he followed Bentham and Hooker's classification, he. however, did
not place the Cymnosperms in between Dicotyledons and Monocotyledons.
Demerits:
In light of availability of more information of phylogenetic nature, now. Besseys
classification has the following demerits:
1. A serious objection to this system has been due to two evolutionary lines considered
by him.
2. Monocots have been placed before Dicots. This is a major demerit of this system.
3. The system has overemphasised hypogyny, perigyny and epigyny.

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