INTRODUCTION

Seeds first appeared in the Devonian (395-359 my = million years). These seeds are not enclosed
in ovaries, as in flowering plants, and are designated naked. This character is the basis for
grouping a large number of varied plants as a natural group, the Gymnospermae (gymnos =
naked, sperm = seed). Palaeobotanical evidence suggests that the seeds evolved independently in
more than one group of Palaeozoic plants and diversified rapidly during the Lower
Carboniferous (345 my). In the course of their evolution, the seed plants have tended to evolve a
variety of structures to protect the ovules. Some of these evolutionary "experiments" were
successful, and provide insight into the origin of the carpel, while others ended in extinction.
Presently, with the advanced knowledge of fossil plants, the gymnosperms are regarded as a
heterogeneous group of seed plants. These plants constituted most of the world's dominant
vegetation throughout the late Paleozoic and Mesozoic, and steadily declined thereafter. A
modern approach is to interpret the phylogeny of gymnosperms on the basis of
phytogeographical and ecological conditions in which the taxa survived.

Characteristic Features

The seed plants are usually classified into two major groups: gymonsperms and angiosperms,
based on the protection to the ovule (at the time of pollination). The gymnosperms bear naked
ovules, i.e. the ovules are borne directly on the sporophyll or an equivalent structure, and are
exposed. In some of the gymnosperms, overlapping scales and sporophylls may protect the
ovules but they are freely exposed at pollination. In the angiosperms, the ovules develop within
an ovary. This distinction between naked and enclosed ovules and later seeds is of considerable
importance.

Some of the other features typical of gymnosperms are: There are no herbaceous
gymnosperms, and the plants whether trees, shrubs or are woody and evergreen. They have a tap
root which usually persists for a long time. The xylem consists of tracheids, parenchyma and
rays. Vessels are present in Ephedra, Welwitschia and Gnetum. They have evolved from pitted
tracheids, as shown by intermediate stages between pits and perforations. In phloem only sieve
cells differentiate; sieve areas commonly occur on the radial walls as well, and are numerous
where the end of one sieve element overlaps that of another. The companion cells are absent.
Secondary growth takes place in all gymnosperms; mature metaxylem shows bordered pits of
various types; Stangeria and Zamia show scalariform thickenings. The anther has an exothecium.
There are numerous light pollen grains which land directly on the surface of the nucellus during
pollination. Prothallial cells are formed in the male gametophyte. The ovule is unitegmic and
orthotropous. There is a prolonged free-nuclear phase in the development of the female
gametophyte, a long interval between pollination and fertilization, a free-nuclear phase in the
development of the proembryo, and haploid nutritive and storage tissue (the female
gametophyte).

History

The gymnosperms are an ancient group which dates back to the Devonian. They began ca. 395
my, lasted for ca. 50 my, and gave rise to aneurophytes, progymnosperms, archaeopterids and
pteridosperms. During the Carboniferous (ca. 345-280 my), a large variety of cordaitels and seed
ferns existed. In the Permian and Triassic (ca. 280 and 225 my) the Carboniferous pteridosperms
became extinct. The early conifers (Voltziales) diversified and the cycads and cycadeoids
became evident for the first time. Glossopteridales formed a noticeable flora of the Southern
Hemisphere during the Permian. The ginkgophytes appeared some time during the Permian and
became more widely spread in the Triassic. In the Jurassic (ca. 195-141 my), the cycads,
cycadeoids, conifers and ginkgophytes reached their peak of diversification, and the
glossopterids became extinct. During the Upper Cretaceous (ca. 141-65 my), the angiosperms
appeared and diversified rapidly. They began to replace the already declining cycads,
cycadeoids, conifers and ginkgophytes. Mesozoic pteridosperms and other smaller groups
became extinct.

However, the majority of the conifer families have continued up to the present. In the
Tertiary (65 my) the angiosperms evolved steadily while the conifers declined in diversity.
Thereafter, the angiosperms became dominant and the gymnosperms occupied a secondary
position, although they still dominated landscapes that angiosperms have yet to conquer. The
long evolutionary history of gymnosperms provides many examples of taxa which flourished,
and finally became completely or nearly extinct. Presently, there are ca. 13000 genera and
240000 spp. of angiosperms, while the gymnosperms comprise only 69 genera and 750-760 spp.
These are grouped in seven orders: Cycadales, Ginkgoales, Coniferales, Taxales, Ephedrales,
Welwitschiales, and Gnetales.
Geographical Distribution

The Cycadales and Ginkgoales represent the surviving members of extremely ancient groups.
The cycads were once a large and dominant group with widest distribution in the Mesozoic,
which is sometimes referred to as the age of cycads. Now there are only 11 living taxa (including
Chigua, a new genus) confined to limited areas of the tropics and subtropics. They form neither
extensive nor noticeable features of the vegetation. Six taxa occur in the Eastern Hemisphere,
and five in the Western Hemisphere; no single genus is represented in both hemispheres. Among
the eastern cycads, Macrozamia, Lepidozamia and Bowenia are confined to Australia, and
Encephalartos and Stangeria exclusively to South Africa. The genus Cycas occurs from
Australia to Japan, touching India and China. Of the western genera, Dioon and Ceratozamia are
confined entirely to Mexico, Microcycas to western Cuba, Zamia to both areas, and Chigua in
Colombia (the only endemic cycad in South America). Stangeria resembles the ferns so much
that it was placed, for a long time, in the family Polypodiaceae. These isolated genera are
"leftovers" from the widely distributed cycads of the Mesozoic. The order Ginkgoales included
many genera and species, reported from the Permian; from the Triassic onwards and during the
Mesozoic they had worldwide distribution, but, before the end of the Jurassic, they began to
disappear, and all the members of this order, except Ginkgo, became extinct by the Cretaceous.
The sole surviving member, G. biloba, is restricted in geographical distribution. At present its
natural occurrence is confined to a small, inaccessible region in southeastern China. There is,
however, some uncertainty whether or not such specimens are escapes from gardens. It is
doubtful whether it actually exists in the wild/ natural state today. One of the reasons for its
survival from extinction in that for centuries priests in China and Japan cultivated and
worshipped it.

The modern Ginkgo is remarkably resistant to attacks of insects and fungi. The reason probably
is its immense vigour, which enabled it to survive for millions of years. Ginkgo is to be regarded
as one of the wonders of the world because it has persisted with very little change through a long
succession of ages inhabited by plants and animals quite different from those present in the
modern age. The cycads and Ginkgo are designated living fossils because they still exhibit many
of their ancestral features without much change. One such character is the presence of motile or
swimming sperms, displayed only in these plants among the existing seed plants. The
Coniferales form 75% of the modem gymnosperms, and are an important constituent of the flora
today. There are ca. 52 genera and 560 spp., usually grouped into six families. They have a
markedly disjunct geographical distribution and several taxa are endemic. Each family is widely
distributed, although individual taxa often show extremely limited distribution. Such a
distribution pattern indicates antiquity. This is evident in the evolutionary history of each of the
six families, which extend back to the Mesozoic. Most modem taxa of the Coniferales were
already present by the onset of the Tertiary. The conifers are plants of the more temperate
regions of the world, and only a few taxa are strictly tropical. Western North America, Eastern
and Central China, and parts of Australia and New Zealand have abundant conifers, which show
exceptional diversity.

Pinaceae: The Pinaceae have ten genera, which form predominant coniferous forests of the
Northern Hemisphere. This family is totally unrepresented in the Southern Hemisphere.

Taxodiaceae: There are ten genera, seven monotypic. In the past, these taxa were an important
constituent of the forests of the Northern Hemisphere. Of the ten taxa, only three occur in the
USA; Sequoia sempervirens (Californian redwood) restricted to a narrow coastal belt in
California, Sequoiadendron giganteum (big tree) in central California, and Taxodium spp. which
grow in lowland swamps ofthe Southeastern USA and Mexico. Of the remaining seven taxa,
Cryptomeria japonica (Japanese cedar) and Cunninghamia spp. are distributed in Japan and parts
of China, Sciadopitys verticillata (Japanese pine) in Japan, Glyptostrobus (Chinese deciduous
cypress) in China, and Taiwania in Formosa. Metasequoia glyptostroboides (Dawn redwood)
was known only as a fossil from Pliocene deposits until 1948, when living specimens were
discovered from Szechuan Province, China. Athrotaxis spp. (Tasmanian cedars) is the only
southern taxon confined to Tasmania.

Cupressaceae: The largest family of the conifers includes ca. 19 genera, 8 monotypic. Of these
19 genera 9 are distributed in the Northern and 10 in the Southern Hemisphere. Of the northern
taxa, Cupressus (cypress), Chamaecyparis (false cypress) and Thuja (arbor vitae) have a disjunct
distribution. Juniperus (Juniper) is distributed in a broad belt round the Northern Hemisphere.
This may be due to its female cones which are berry-like and distributed by birds. Some of the
southern taxa are Callitris (Cypress pines) confined to Australia, Tasmania, and New Caledonia,
Libocedrus to New Zealand and New Caledonia, and Papuacedrus to New Guinea, extending
somewhat across the equator into the Northern Hemisphere.
Podocarpaceae: Podocarpaceae is the most important family of the Southern Hemisphere,
where it originated. It comprises seven taxa, two monotypic. Some of the taxa are represented in
the Northern Hemisphere also, and are assumed to be recent arrivals. The largest genus,
Podocarpus (yellow woods), occurs in the mountain forests of warm temperate and subtropical
regions of the Southern Hemisphere. Some species occur in Japan, China, India, Malaya and The
Philippines. Dacrydium and Phyllocladus are distributed chiefly in New Zealand and Tasmania,
Dacrydium also occurs in Malaysia, Burma, Southern China, Patagonia and Chile, while
Phyllocladus is found in New Guinea, North Borneo and the Philippines. They have
representatives north of the Equator. Saxegothaea (Prince Alberts' yew) and Microstrobus (=
Pherosphera) are monotypic. The former is confined to Chile and the latter to Tasmania.
Acmopyle is confined to New Caledonia and Fiji.

Araucariaceae: An extremely old family with fossil record extending back to the Triassic of
both North and South Hemispheres. Both the present day taxa are restricted to the Southern
Hemisphere. Araucaria occurs in South America, Australia, New Guinea and New Caledonia,
and Agathis (kauri pines) is exclusively eastern, extending from The Philippines to New Zealand
and Malaya to Fiji.

Cephalotaxacae: A single taxon, Cephalotaxus, is restricted to eastern Asia. It is distributed

Taxales: The order Taxales includes a single family, Taxaceae, with five genera. Taxus (Yew)
occurs in North America, Europe and Asia, and extends up to Malaya. This wide geographical
distribution is partly due to birds. Pseudotaxus (=Nothotaxus) grows in a small part of east
China. Amentotaxus presently occurs in East Asia, but fossil remains have been reported from
Europe and western America. Austrotaxus is restricted to New Caledonia. Torreya occurs only in
California, Florida and eastern Asia.

Ephedrales: The Ephedrales comprise a single monogeneric family, Ephedraceae, and Ephedra
with ca. 42 spp. has a worldwide distribution. None of the species is common to the Old and the
New Worlds or Eastern and Western Hemispheres. Most of the species inhabit arid or desert
regions including saline tracts where they act as a sand-binder. The plants grow up to 5000 m
above sea level.
Welwitschiales: Welwitschia is a monotypic genus of limited distribution. It is restricted to a
strip ca. 1200 km long along the west coast of southern Africa from Angola (Nicolau River) to
South-West Africa, Namibia (Kuiseb River). In the Welwitschia Flats 5000-6000 specimens
grow. Welwitschia never reaches the coast. It occupies only the northern and central parts of the
Namib Desert in its south-north extension and stretches over subtropical grassland, entering the
Mopane Savanna in its west-east extension. Thus, it occupies an area with a wide-ecological
range. It grows mainly where the annual rainfall is between 0.0 and 100 mm and precipitation
(except dew and fog) is only in the summer months, January to March.

Gnetales: The single family, Gnetaceae, includes only one genus, Gnetum. It inhabits moist
tropical forests in parts of Asia, Africa, northern South America, and certain islands between
Asia and Australia. Most species, ca 30, are endemic within the areas of their distribution. There
is no species common to both the hemisphere, and none of the Asian species occurs in Africa or
America. The center of the present diversification appears to be Eastern Malayasia.