EVOLUTION OF ANGIOSPERMS

Angiosperms or Flowering Plants:

Angiosperms or flowering plants form the largest group of plant kingdom, including about 300 families (411
families, Hutchinson), 8,000 genera and 300,000 species. They are considered to be highest evolved plants on the surface
of the earth. From Cretaceous age, the angiosperms eclipsed all other vegetation and now they are dominant. They are
found almost everywhere in each possible type of habitat and climate.
They occur in deep lakes, deserts, in beds of seas and even on high peaks of mountains. The species of Opuntia
(Cactaceae) can survive without water in acute desert conditions, whereas on the other hand the species of Hydrilla
(aquatic plant) are extremely sensitive to drought conditions.
Some species are found on rocks, some in waterfalls and also some are marine. The species of Rhizophora,
popularly known as ‘mangrove vegetation’ are found near the water of the sea. The epiphytes, parasites, saprophytes,
symbionts and even insectivorous plants are also not uncommon.
They may be annual, biennial or perennial herbs, shrubs, trees, climbers, twiners and lianes. On one hand the
angiosperms may be as minute in size as a pin head, e.g., Wolffia microscopica, on the other extremity like eucleptiles of
Australia may reach upto 300 feet in height.

Evolution of Angiosperms:
Angiosperms are considered to be the most evolved group of plants today on the basis of the following characteristics:
a) Diversified habit and vegetative forms;
b) Higher degree of perfection of the vascular system; the xylem in addition to the tracheids, contains wood vessels,
and the phloem possesses companion cells;
c) Successful invasion of all habitats;
d) Adaptation of flower to insect pollination;
e) Bisexual flower, the bisexuality ensures self-pollination if cross-pollination fails, hence the seeds are formed in
any case;
f) Development of ovules within the ovary which ensures proper protection to the developing ovules and seeds;
g) Efficient and effective dispersal by insects, birds, other animals, wind, water and by several other methods;
h) Efficient and varied means of vegetative propagation, which result in rapid multiplication;
i) Their utmost economic importance, and
j) Besides the above listed reasons, there must be some hereditary causes such as better equipment of gene potential
and useful gene mutations, which enable them to encounter variations in temperature and other environmental
changes and led to their conquest over other plant groups.

Origin of Angiosperms:
The angiosperms appeared suddenly in Cretaceous age about 65 million years back. Charles Darwin described this
sudden appearance of angiosperms in lower or upper Cretaceous as an ‘abominable mystery. When angiosperms appeared
for the first time in lower or upper Cretaceous, they were full fledged like the trees and the herbs of today. In support of
this view Prof. Knowlton advocates in his ‘Plant of the past’, “from the time of their appearance they did not
progress at all due to their full-fledged appearance in the Cretaceous”.
The fossil records of the angiosperms also support their appearance full-fledged in lower or upper Cretaceous. The
fossils of that age are so characteristic and modem in appearance that most of them can be referred unmistakably to living
families, general and even to some species. Prof. Knowlton stated in his book, ‘Plant of the past’ “if a student of present
day trees and shrubs, could have wandered over the hills and vales in those days, he would have found himself quite at
home among the trees and shrubs growing there”. The forms of cycads and conifers, which long dominated the universe
were already pushed background and the earth had become infact the earth of flowering plants. Charles Darwin has called
this sudden appearance of angiosperms as an “abominable mystery”.
However, some workers do not agree with the doctrine of ‘abominable mystery’. According to H.H. Thomas
(1936), the angiosperms of the past replaced many of older gymnosperms in asturine and marshy waters. Graud Eury
(1906) believes that the angiosperms came into existence through mutation. Guppy (1919) however, supported the view of
mutation. Prof. Bertrand is of opinion that all the great groups of vascular plants (Pteridophyta, Gymnospermae and
Angiosperms) not only arose quite independently of each other but also they originated simultaneously as far back in the
Archian period (2000 million years old-oldest). There is a very considerable but scattered literature on the origin and
phylogeny of angiosperms. The palaeobotanical evidence shows that there seems three possibilities as regards the origin of
angiosperms.

These possibilities are:

1. That the angiosperms are monophyletic in their origin but have had a very much longer history than at present
known, perhaps stretching back into Palaeozoic times and with a whole series of missing links;
2. That the angiosperms are monophyletic but that the first and at present unknown group diverged quickly in terms
of geological time, into a considerable number of different groups;
3. That the angiosperms are polyphyletic.
 According to Campbell, “both comparative morphology and the geological record indicate that the existing
angiosperms represent a number of distinct phyla which cannot be traced back to a single ancestral type”. This statement
shows that he does not believe in monophyletic origin of angiosperms. According to Thomas, the evolutionary tendencies
detected in the three groups, i.e., Caytoniales, Bennettitales and Pteriodsperms furnish reasonable grounds for the idea
that the angiosperms were derived from various pteridosperms early in the Mesozoic period. Parkin also argues for the
monophyletic origin of the angiosperms.
In conclusion we can say, that the history of the angiosperms is still almost as great a mystery as it was in the time
of Darwin. We do not know, when, where or from what the presumably most recent and now dominant large group of
existing terrestrial plants arose. Lotsy says, that hybridization is the key to evolution of angiosperms. This view has been
supported by Anderson. He has suggested on the basis of cytological investigations that the angiosperms may have arisen
as a result of hybridization between two gymnosperms. According to Hagerup, the origin of some angiosperms took place
from the Coniferales through Gnetales. According to Eames, Sinnott and Bailey the more primitive angiosperms were
arboreal in habit and the herbaceous angiosperms have been evolved from them. According to Hutchinson, in certain
groups, trees and shrubs are probably more primitive than herbs. Evidently views in this respect are divergent and
speculative, the available data being meagre, fragmentary and isolated.

Some of the theories proposed from time to time in this connection, are as follows:

1. Bennettitales-Ranales Theory:
a) Hallier’s view (1906) regarding the origin of angiosperms is that Ranales, (e.g., Magnolia) seems to be related to
Bennettitales and may have been derived from Cycadeoidea, and that the monocotyledons are an offshoot of the
dicotyledons. The elongated floral axis of Ranales with spirally arranged male and female sporophylls and the
cone of Cycadeoidea are definitely alike. Both the groups were also abundant in the Cretaceous.

Ranales is regarded as the earliest stock from which the polycarpic families of dicotyledons have arisen, and also
the monocotyledons as an offshoot. But the differences in the anatomical structure of the wood, types of
sporophylls, nature and position of ovules, etc., in the two groups, i.e., Bennettitales and Ranales, have led to
difficulties in accepting this view. It is more probable that both the groups have evolved from a common ancestry
and developed in unrelated parallel lines.

b) Arber and Parkin (1907), proposed the existence of an imaginary group (Hemiangiospermae) having cycadeoid
type of flowers linking the above two groups. From this imaginary group the Ranalian type of flowers might have
originated, and from this Ranalian type all other angiosperms had sprung up. They were of the opinion that the
earliest monocotyledons were more primitive than the dicotyledons. They also supported the Hallier’s view.

c) Hutchinson (1925) considered the origin of angiosperms as monophyletic, and supported the views of Hallier
(1906) and Arber and Parkin (1907). He believed in the Bennettitalean origin of angiosperms and stressed on two
parallel evolutionary lines for the primitive dicotyledons a woody (arborescent) line, called Lignosae, starting from
Magnoliales, and a herbaceous line, called Herbaceae starting from Ranales. He further held the view that the
monocotyledons were derived from a primitive dicotyledonous order. The Ranales.

2. Coniferae-Amentiferae Theory:
Engler and Prantl (1924) rejected the Cycadeoidean origin of angiosperms, as proposed by Hallier (1906) earlier.
They held the view that dicotyledons and monocotyledons had arisen independently from a hypothetical group of extinct
gymnosperms (allied coniferae) with unisexual strobilus which developed in the Mesozoic.
Thus, according to them, the angiosperms had a polyphyletic origin, and evolution took place on several parallel
lines from the beginning. They considered the monocotyledons to be more primitive than the dicotyledons. The unisexual
naked (without perianth) condition of the angiospermic flowers, such as Pandanales (monocotyledons) and Amentiferae
(catkin-bearing dicotyledons, e.g., Casuarina. Salix, Betula, etc.) was most primitive. But according to the modem
classification these orders are regarded more advanced.

3. Gnetales-Casuarinales Theory:
Wettstein (1935) held the view that angiosperms of Casuarina type evolved from Gnetales, (particularly Ephedra),
a highly advanced group of gymnosperms, which branched off from the main gymnospermic line. However, there are
some angiospermic features in Gnetales but this group has meagre fossil records, and not gone earlier than Tertiary.
Fagerland (1947) was of the opinion that both Gnetales and Proangiosperms had a common ancestor and the modern
angiosperms evolved from the Proangiosperms in Polyphyletic lines.

4. Caytoniales-Angiosperm Theory:
Thomas (1925) suggested that Caytoniales, a small group of angiosperm-like plants, discovered in the Jurassic
rocks of Yorkshire (England), might be the ancestor of angiosperms. However, Harris (1932-33) opposed this view.
Knowlton (1927) expressed the view in his book Plants of the Past that both Caytoniales and angiosperms evolved from the
large extinct Palaeozoic group of Pteridosperms. Arnold (1948), expressed the view that Caytoniales were allied to the
Pteridosperms rather than to the angiosperms.

5. Pteridosperm-Angiosperm Theory:
Andrews (1947) was of opinion that seed ferns or Pteridosperms, an ancient group of the Palaeozoic, might be the
starting point for the angiospermic plants.
There are two large groups of angiosperms:
(1) Dicotyledons and
(2) Monocotyledons.

Origin of Dicotyledons and Monocotyledons of Angiosperms:

A. Origin of Dicotyledons:
The dicots are more important and they are supposed to have originated before the monocotyledons. It is thought that for
the first time dicotyledons appeared in the early Mesozoic era, or perhaps the late Palaeozoic. The oldest known fossils of
dicots are from the lower Cretaceous rocks.
It is generally believed and agreed that the dicotyledons have been originated from gymnosperms of a type somewhat
different from present day forms of gymnosperms. According to some similarities noted in these two groups, the
development seems to be parallel, or in some respects convergent development of the two groups.

B. Origin of Monocotyledons:
This is the subject of great controversy. For some time this was thought that the monocotyledons were more primitive
than the dicotyledons and probably they have given rise to the dicotyledons. Now, this belief has been totally given up. It is
now generally thought that the dicotyledons are more primitive and they have given rise to the monocotyledons.
This is also believed, that the monocotyledons were an offshoot of the primitive dicotyledons back in the early Mesozoic
era. They are thought to be monophyletic {i.e., of one origin) in their origin. According to another conception the
monocotyledons are not monophyletic but polyphyletic in their origin.

Principles of Taxonomy and Phylogeny of Angiosperms:

The problem of classifying the angiospermic plants in a systematic way may either be termed as, ‘taxonomy of
angiosperitis’ or ‘systematic botany’. This is a functional science and deals with identification, nomenclature and
classification of the plants found all over the world.
The angiosperms are widely distributed with so many morphological variations that sometimes it seems almost impossible
to arrange them in systematic order.
Since the prehistoric times, people were interested in placing the plants in a systematic way and for the first time a few
plants were classified according to their medicinal and food value and thus the study of taxonomy of flowering plants
began. In nineteenth century, the formation of the principles of taxonomy began and several principles were formed,
which are still very helpful in arranging the plants in a systematic order.

Evolutionary Principles proposed by Bessey:

The following principles of taxonomy proposed by C.E. Bessey (1915) are of vital importance and are used by all
modern taxonomists to establish the evolutionary trends in plants. These are known as the Besseyan principles.
They are as follows:
1. There is progressive evolution, i.e., life advances from simplicity to complexity.
2. The existing simpler forms resemble more to their simple ancestors than to the present complex ones.
3. The evolved forms never become like ancestors.
4. The herbaceous habit of plants is more advanced than arboreal habit.
5. Annuals are more advanced than biennials and biennials are more advanced than perennials.
6. Hydrophytes, epiphytes, saprophytes and parasites are more advanced than ordinary terrestrial forms.
7. Compound leaves are more advanced than simple leaves.
8. Bisexual flowers are more primitive than unisexual ones.
9. Dioecious plants are more advanced than monoecious ones.
10. Inflorescence is more advanced than solitary flowers.
11. Gamopetalous flowers are more advanced than polypetalous flowers. The gamopetaly has been derived from
polypetaly by the fusion of the petals.
12. Flowers lacking petals (apetaly) are more advanced.
13. Actinomorphic flowers are more primitive than zygomorphic ones.
14. Hypogyny precedes epigyny.
15. Apocarpous condition is more primitive than syncarpous condition.
16. Exalbuminous seeds are more advanced than albuminous seeds.
17. Polyandrous stamens precede jointed condition of stamens.
18. Simple fruits are more primitive than compound fruits.
 Turril states, that “the angiosperms were the last of the great groups (Thallophyta, Bryophyta,
Pteridophyta, Gymnosperms, Angiosperms) to appear. Hence, changes towards characters peculiar to
the angiosperms may with some justification be read as progressive, the more so that they are now the
dominant group ecologically over much of the land surface of the globe.”
The ‘conservative’ characters have been supposed by many taxonomists to be most useful and valuable in
phylogenetic studies. For plants, the reproductive organs are supposed to be more conservative than the vegetative organs.
For example, the morphogenetic evolutionary changes in carpels have been far greater than in any vegetative organ
considered throughout Spermophyta. According to Tuzson, the monocots form an older group than the dicots, because the
smaller groups of monocots are separable into series and families which show greater gaps on the whole than those in
dicots.
As a Gray is of opinion that a natural system of classification of plants aim to arrange all the known plants of the
plant kingdom, in a series of grades according to their resemblances, in all respects, so that each species, genus, tribe,
family, order, etc., shall stand next to those which it most resembles in all respects, or rather in the whole plan of
structure. For example, two plants may be very much alike in external appearance, yet very different in their principal
structure.
According to Sprague the monophyletic groups are regarded as the only natural ones at and above the family level in
existing angiosperms. On the other hand, Hutchinson in his phylogenetic scheme and classification definitely retains
groups he considers polyphyletic, e.g., Asterales and Euphorbiales.

Oldest Angiosperms:
Caytoniales:
In Jurassic rocks have been found the oldest
known plants which were angiospermous in the true
sense of the term; that is, in having seed enclosed in a
carpel. These plants represent two closely-related genera
of the order Caytoniales. The carpels were borne on
sporophylls. Each sporophyll consisted of a central stalk
with pinnately arranged short side branches each of
which was terminated by a carpel or fruit. The carpel was
completely closed, and the tip of the portion of the
pinnule which bent over became the stigma. Pollen grains
have been found attached to the stigmas. The seed were
borne within the closed carpel (Fig. 6.3). The integument
of the seed is rather strikingly similar to that of certain
seed of seed-ferns. The fruits were fleshy, so the seeds
may have been scattered by animals that ate the fruit.
The anthers are found on sporophylls similar in
general outline to those which bore the carpels. The
branches of the sporophylls divided, and the tips of the
divisions bore groups of stamens (Fig. 6.2). The anthers
were sessile or at the ends of short filaments. They had a
four- winged form, and each wing seems to have
contained a pollen sac. Thus they had the same general
form as the stamen of a modern angiosperm.
The leaves which appear to belong to the known
Caytoniales are of a type which was very widespread
during the Jurassic period and extended from Triassic to
Cretaceous times. The venation was netted, but similar to
that of Glossopteris which is generally regarded as a seed-
fern. The general character of the leaves, sporophylls, and
seed indicate that the Caytoniales were derived from the
seed ferns.

In the Caytoniales from the Jurassic the pollen seems to

have been caught on a stigma, and the ovules are
enclosed in ovaries, but in a form from the upper
Triassic pollen grains are found in the micropyles of the
seed. In this form there appear to have been canals
running through the “stigma,” and through these canals
pollen grains reached the micropyles of the seed (Fig.
6.4).
This Triassic Caytonia is yet completely angiospermous.
It has been evidently not suggested that the walls of the
fruit of the Caytoniales had their origin in the fusion of
cupules which surrounded the seed of Cycadofilicales.
Unfortunately we do not know how the sporophylls of
the Caytoniales were borne, or what kind of plant bore
them, and so the relationship of the Caytoniales to
modern angiosperms is obscure.

Ancestry of Angiosperms:
The ancestry of angiosperms has long been a
moot question. There is not enough evidence to reach a
definite decision, and there is much disagreement as to
theory.
Among living gymnosperms the group which is most similar to the angiosperms is the Gnetales. While there is
great dissimilarity between the Gnetales and the angiosperms, still there is enough similarity to convince many that either
the angiosperms are descended from the Gnetales, perhaps from extinct forms, or else the two groups are closely related
and have a common ancestry.
Some regard the Gnetales as being intermediate between the conifers and angiosperms, and so are inclined to a
belief in a coniferous ancestry for angiosperms. This belief is based in part on similarity in wood structure and on the fact
that in conifers, Gnetales, and angiosperms the fertilization is by male nuclei and not by spermatozoids as in all other
living groups of land plants. A rather complete understanding of the “flower” of Cycadeoidea was followed by an extensive
discussion of a cycadophyte ancestry for angiosperms; not from Cycadeoidea with its highly specialized stem, rather from
one more nearly related to Williamsonia or Williamsoniella, but still more primitive.
It was pointed out that the bracts, stamens, and ovuliferous cone of Cycadeoidea were in the same relative
positions as the perianth, stamens, and carpels of the angiosperms. Also, the embryo had two cotyledons, while the seeds
were almost enclosed by the inter-seminal bracts. The discovery of the Caytoniales, first in the Jurrasic and later in the late
Triassic, has been taken by some as indicating that the angiosperms, through forms related to the Caytoniales, are
descended direct from the seed-ferns.
A recent theory is that the carpels of angiosperms were derived through a modification and fusion of cupules
which surrounded the seeds of Cycadofilicales, and that the ovaries of the Caytoniales represent an intermediate condition
between the seed-ferns and modern angiosperms.
One feature which is common to practically all theories as to the origin of the angiosperms is that they go far back
into early Mesozoic or latter Palaeozoic times; also that their ancestors were generalized forms and the not such
specialized ones as modem conifers or cycads or the late Mesozoic cycadeoideas. A conservative suggestion is that they
were derived from somewhere in the general gymnospermous complex, from a line in which the marked peculiarities of
more modern groups had not become so pronounced as they appear in the well-known specialized types.
Our ignorance as to the ancestry of the angiosperms is not surprising when we remember how scanty our
knowledge of plant floras is. If, as seems probable, the angiosperms evolved in Arctic regions, much of the record may be
thickly covered with an icy mantle and inaccessible to us. Also, the record of former vegetation is largely that in and
around swamps, about lakes, and in similar situations; and any trace of much of that which flourished on higher ground is
forever lost.
However, in comparatively recent years a great deal of evidence as to the history and relationships of land plants
has been discovered; so we may hope that perhaps someday we may have a fairly connected account of the development of
the angiosperms.