
George Bentham and Joseph Dalton
Hooker – Two English taxonomists who were closely associated with the Royal
Botanical Garden at Kew, England have given a detailed classification of plant
kingdom, particularly the angiosperms.They gave an outstanding system of classification of phanerogams in their
Genera Plantarum which was published in three volumes between the years 1862 to
1883. It is a natural system of classification. However, it does not show the
evolutionary relationship between different groups of plants, in the strict
sense. Nevertheless, it is the most popular system of classification
particularly for angiosperms. The popularity comes from the face that very
clear key characters have been listed for each of the families. These key
characters enable the students of taxonomy to easily identify and assign any
angiosperm plant to its family.Bentham and Hooker have grouped advanced, seed bearing plants into a major
division called Phanerogamia. This division has been divided into three classes
namely:1. Dicotyledonae 2.
Gymnospermae and 3.
Monocotyledoneae
Class Dicotyledonae
This group includes angiosperms in which the seed bears two cotyledons and
leaves exhibit reticulate venation. It is divided into three subclasses –
Polypetalae, Gamopetalae and Metachlamydae.
Sub-class
Polypetalae
The flowers contain distinct
non-essential whorls calyx and corolla. In the corolla petals are free. This
sub-class includes 8 series Thalamiflorae, Disciflorae and Calyciflorae.
- Series Thalamiflorae: Many stamens in the androecium.
Flower is hypogynous.
- Series Disciflorae: Hypogynous flowers with a
cushion-like disc around or below the ovary.
- Series Calyciflorae: Flowers epigynous or perigynous.
Thalamus is in the form of a cup.
Sub-class
GamopetalaeFlowers with distinct calyx and
corolla. In the corolla petals are fused. This sub-class includes 3 series.
- Series Inferae: Flowers with inferior ovary.
- Series Heteromerae: Flowers with superior ovary. Number
of carpels – more than two.
- Series Bicarpellatae: Flowers with superior ovary.
Number of carpels – two.Sub-class
MonochlamydaeThe flowers are with only one
non-essential whorl (perianth) or absence of non-essential whorls. It includes
8 series.
- Curvembryae: Usually single ovule, embryo coiled around
the endosperm.
- Multiovulate Aquaticae: Aquatic plants with syncarpous
ovary and many ovules.
- Multiovulate Terrestris: Terrestrial plants with
syncarpous ovary and many ovules.
- Microembryae: Only one ovule, small, tiny embryo
endospermic seed.
- Daphnales: Only one carpel and single ovule.
- Achlamydosporae: Ovary inferior, 1 to 3 ovules –
unilocular.
- Unisexuales: Flower unisexual, perianth usually absent.
- Ordines Anomali: (Anomolous families) Plants with
uncertain systematic position but closer to unisexualesClass GymnospermaeThis group includes the gymnosperms in which seeds are not enclosed in
fruits. This class is divided into three families Gnetaceae, Confiraceae and
Cycadaceae.
Class Monocotyledonae
This group includes angiosperms in
which the seed bears only one cotyledon. The leaves exhibit parallel venation.
It is divided into the following seven series.
- Microspermae: Ovary is inferior, seeds are minute and
non-endospermic.
- Epigynae: Ovary inferior, seeds are large and
endospermic.
- Coronarieae: Ovary superior, perianth petalloid.
- Calycinae: Ovary superior, perianth sepalloid.
- Nudiflorae: Perianth reduced or absent. Seeds are
endospermic.
- Apocarpae: Carpels more than one, free, seeds are
endospermic.
- Glumaceae: Perianth reduced or absent, scaly bracts
present.
Each of the series mentioned under
dicotyledonae and monocotyledonae have been further divided into orders and
families. Bentham and Hooker classified the angiosperms into 202 families. They
were able to provide distinct diagnostic key characters to each of these
families. This is the reason for the popularity of Bentham and Hooker’s
classification particularly amongst the taxonomists.
Engler and Prantl regarded
Angiosperms as having arisen from a hypothetical extinct group of gymnosperms
called Protoangiosperms related to unknown taxa of gymnosperms. They considered
gymnosperms to be primitive and hence placed them before angiosperms.
Similarly, monocots were initially considered more primitive and hence placed
before dicots. However, in the last edition of the “Syllabus” (Melchior,
1964), the dicots were placed before monocots. They distinguished between
primitive and advanced conditions.Engler believed in the Amentiferous
origin of the angiosperm flower. According to this concept, the unisexual
flowers without perianth (achlamydeous) or with one whorl of perianth
(monochlamydeous; and borne in catkin inflorescence with wind pollination) were
primitive whereas those with two whorls of perianth (dichlamydeous –
distinguished into sepals and petals) were advanced. The bisexual flowers were
believed to have evolved from a cluster of male and female flowers (held
together in the same inflorescence and appearing like a single flower (called a
pseudoanthium), while the surrounding perianth evolved later.
Therefore the families with naked (Achlamydeous = without perianth), unisexual
and wind pollinated flowers were placed at the beginning, while those with dichlamydeous
(perianth in two whorls – calyx and corolla), bisexual and insect pollinated
flowers were placed at a later (= advanced) position in the classification.
Thus the different families were arranged in an ascending series with simple
groups at the beginning, followed by families with increasing complexity and
elaboration of flowers. Engler’s system thus did not recognize the significance
of reduction in evolution, and equated simplicity with primitiveness. However,
it is now known that in many cases simplicity may be the result of evolutionary
reduction. This has led to the suggestion that the classification was not
phylogenetic in a true sense, and may be regarded as a step towards phylogeny.Engler and Prantl considered the
angiosperms to be polyphyletic in origin, and derived from seed ferns and
gymnosperms through Amentiferae.The classification covered the
entire plant kingdom from algae to angiosperms which has been divided into 13
divisions. The first 11 divisions are Thallophytes, 12th division is
Embryophyta Asiphonogama (plants with embryos but no pollen tubes;
Bryophytes and Pteridophytes) and the 13th division is Embryophyta
Siphonogama (plants with embryos and pollen tubes) which includes seed
plants. This is divided into 2 subdivisions: 1. Gymnospermae, 2. Angiospermae.The subdivision Angiospermae is
further divided into 2 classes:Class 1. MonocotyledoneaeClass 2. DicotyledoneaeClass 1. Monocotyledoneae include 11
orders starting with order Pandanales with naked unisexual flowers and ending
with order Microspermae with family Orchidaceae. Orchidaceae is considered as
the most advanced family and evolved over grasses.Class 2. Dicotyledoneae is divided
into 2 subclasses: Subclass 1. Archichlamydeae and Sublass 2. Sympetalae or
Metachlamydeae. Archichlamydeae include 33 orders starting with order
Verticillatae and ending with order Umbelliflorae. It included achlamydeous or
monochlamydeous families at the beginning and dichlamydeous and polypetalous
families in the end. Hence, Archichlamydeae comprised of the Poylpetalae and
Monochlamydeae of Bentham and Hooker’s classification. Sympetalae included
Gamopetalous flowers with 11 orders starting with order Diapensiales and ending
with order Campanulatae.
- Merits of Engler and Prantl System of
Classification- The
classification incorporated evolution and was the beginning of
phylogenetic classifications.- The
system classified all the plant families from the algae to angiosperms.- Gymnosperms
were considered primitive and placed before angiosperms.- The
taxa were fully illustrated, described along with identification keys upto
family level in Syllabus der Pflanzenfamilien, up to genus level in
Die naturlichen Pflanzenfamilien and for many families along with
their species in Das Pflanzenreich.- The
description of families also contained information on embryology, anatomy
and geographical distribution.- The
classification was very elaborate, extensive and is still followed in many
herbaria and floras of the world.- The
polypetalous condition was considered primitive than gamopetalous
condition which is also the present phylogenetic view.- The
Polypetalae and Monochlamydeae of Bentham and Hooker’s classification were
merged into a single subclass Archichlaymdeae which resulted in proper
placement of many families. For example family Illecebraceae is combined
with family Caryophyllaceae. Families Amaranthaceae, Chenopodiaceae and
Caryophyllaceae are included in the same order Centrospermae.- Families
Compositae and Orchidaceae in dicots and monocots respectively are
considered advanced are placed at the end of each group.
- Demerits of Engler and Prantl System of Classification
- The
classification though not intentionally was more natural and less
phylogenetic.
- The
classification equated simplicity of structure with primitiveness and did
not recognize the simplicity due to evolutionary reduction. Thus the
classification was not phylogenetic in true sense.
- Monocotyledons
were placed before dicots but this placement was reversed in the 1964
edition of Syllabus der Pflanzenfamilien.
- The
naked flowers of Amentiferous families were considered primitive, but
evidences from floral anatomy, wood anatomy and palynology suggest these
are not primitive but simple due to the disappearance of perianth and
evolutionary reduction.
- Unisexual
flowers were treated as primitive, but evidence from floral anatomy has
shown that the unisexual condition is derived from bisexual condition by
suppression of either androecium or gynoecium.
- Monochlamydeous
flowers (one whorl of perianth) were treated primitive over dichlamydeous
flowers (distinct calyx and corolla) which is not accepted by modern
taxonomists.
- Angiosperms
were considered to be polyphyletic in origin, but modern taxonomists
regard angiosperm to be monophyletic in origin.
- Ranales
are considered primitive by modern taxonomists, but they have been placed
after Centrospermae at a position not accepted by modern taxonomists.
- Araceae
in Monocotyledons are derived from Liliaceae, but Araceae have been placed
before Liliaceae, a position not accepted by modern taxonomists.
- Helobieae
consisting of primitive groups have been placed between two advanced
orders Pandanales and Glumiflorae.
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