The gymnosperms (
lit. revealed seeds) are a group of
seed-producing plants that includes
conifers,
cycads, ''
Ginkgo'', and
gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνόσπερμος ( el, γυμνός, translit=gymnos, lit=naked, label=none and el, σπέρμα, translit=sperma, lit=seed, label=none), literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds (called
ovules in their unfertilized state). The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (
angiosperms), which are enclosed within an
ovary. Gymnosperm seeds develop either on the surface of scales or
leaves, which are often modified to form
cones, or solitary as in
yew, ''
Torreya'', ''
Ginkgo''.
Gymnosperm lifecycles involve
alternation of generations. They have a dominant
diploid sporophyte phase and a reduced
haploid gametophyte phase which is dependent on the sporophytic phase. The term "gymnosperm" is often used in
paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern
monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.
The gymnosperms and
angiosperms together comprise the
spermatophytes or seed plants. The gymnosperms are subdivided into five
Divisions, four of which, the
Cycadophyta,
Ginkgophyta,
Gnetophyta, and
Pinophyta (also known as Coniferophyta) are still in existence while the
Pteridospermatophyta are now extinct.
Newer classification place the gnetophytes among the conifers.
By far the largest group of living gymnosperms are the conifers (pines, cypresses, and relatives), followed by cycads, gnetophytes (''
Gnetum'', ''
Ephedra'' and ''
Welwitschia''), and ''
Ginkgo biloba'' (a single living species). About 65% of gymnosperms are
dioecious, but conifers are almost all
monoecious.
Some genera have
mycorrhiza, fungal associations with roots (''Pinus''), while in some others (''Cycas'') small specialised roots called coralloid roots are associated with nitrogen-fixing
cyanobacteria.
Diversity and origin

Over 1000 living species of gymnosperm exist.
It was previously widely accepted that the gymnosperms originated in the
Late Carboniferous period, replacing the
lycopsid rainforests of the tropical region, but more recent phylogenetic evidence indicates that they diverged from the ancestors of
angiosperms during the
Early Carboniferous. The radiation of gymnosperms during the late Carboniferous appears to have resulted from a whole
genome duplication event around .
Early characteristics of seed plants are evident in fossil
progymnosperms of the late
Devonian period around 383 million years ago. It has been suggested that during the mid-Mesozoic era, pollination of some extinct groups of gymnosperms was by extinct species of
scorpionflies that had specialized
proboscis for feeding on pollination drops. The scorpionflies likely engaged in pollination mutualisms with gymnosperms, long before the similar and independent coevolution of nectar-feeding insects on angiosperms.
Evidence has also been found that mid-Mesozoic gymnosperms were pollinated by
Kalligrammatid lacewings, a now-extinct family with members which (in an example of
convergent evolution) resembled the modern butterflies that arose far later.

All gymnosperms are
perennial woody plants, apart from the cycads. The soft and highly
parenchymatous wood in cycads is poorly lignified, and their main structural support comes from an armor of sclerenchymatous leaf bases covering the stem, with the exception of species with underground stems. There are no
herbaceous gymnosperms and compared to angiosperms they occupy fewer
ecological niches, but have evolved both parasites (
parasitaxus),
epiphytes (
Zamia pseudoparasitica) and
rheophytes (
Retrophyllum minus).
Conifers are by far the most abundant extant group of gymnosperms with six to eight families, with a total of 65–70 genera and 600–630 species (696 accepted names).
[Catalogue of Life: 2007 Annual checklist – Conifer database](_blank)
Most conifers are
evergreens. The
leaves of many conifers are long, thin and needle-like, other species, including most
Cupressaceae and some
Podocarpaceae, have flat, triangular scale-like leaves. ''
Agathis'' in Araucariaceae and ''
Nageia'' in Podocarpaceae have broad, flat strap-shaped leaves.
Cycads are the next most abundant group of gymnosperms, with two or three families, 11 genera, and approximately 338 species. A majority of cycads are native to tropical climates and are most abundantly found in regions near the equator. The other extant groups are the 95–100 species of
Gnetales and one species of
Ginkgo.
Today gymnosperms are the most threatened of all plant groups.
Classification
A formal classification of the living gymnosperms is the "Acrogymnospermae", which form a
monophyletic group within the
spermatophytes.
The wider "Gymnospermae" group includes extinct gymnosperms and is thought to be
paraphyletic. The fossil record of gymnosperms includes many distinctive
taxa that do not belong to the four modern groups, including seed-bearing trees that have a somewhat
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The Polypodiophyta, polypodiophytes include all living pteridop ...
-like vegetative morphology (the so-called "seed ferns" or
pteridosperms). When fossil gymnosperms such as these and the
Bennettitales,
glossopterids, and ''
Caytonia'' are considered, it is clear that angiosperms are nested within a larger gymnospermae clade, although which group of gymnosperms is their closest relative remains unclear.
The extant gymnosperms include 12 main families and 83 genera which contain more than 1000 known species.
Subclass
Cycadidae
*Order
Cycadales
**Family
Cycadaceae: ''
Cycas''
**Family
Zamiaceae: ''
Dioon'', ''
Bowenia'', ''
Macrozamia'', ''
Lepidozamia'', ''
Encephalartos'', ''
Stangeria'', ''
Ceratozamia'', ''
Microcycas'', ''
Zamia''
Subclass
Ginkgoidae
*Order
Ginkgoales
**Family
Ginkgoaceae: ''
Ginkgo''
Subclass
Gnetidae
*Order
Welwitschiales
**Family
Welwitschiaceae: ''
Welwitschia''
*Order
Gnetales
**Family
Gnetaceae: ''
Gnetum''
*Order
Ephedrales
**Family
Ephedraceae: ''
Ephedra''
Subclass
Pinidae
*Order
Pinales
**Family
Pinaceae: ''
Cedrus'', ''
Pinus'', ''
Cathaya'', ''
Picea'', ''
Pseudotsuga'', ''
Larix'', ''
Pseudolarix'', ''
Tsuga'', ''
Nothotsuga'', ''
Keteleeria'', ''
Abies''
*Order
Araucariales
**Family
Araucariaceae: ''
Araucaria'', ''
Wollemia'', ''
Agathis''
**Family
Podocarpaceae: ''
Phyllocladus'', ''
Lepidothamnus'', ''
Prumnopitys'', ''
Sundacarpus'', ''
Halocarpus'', ''
Parasitaxus'', ''
Lagarostrobos'', ''
Manoao'', ''
Saxegothaea'', ''
Microcachrys'', ''
Pherosphaera'', ''
Acmopyle'', ''
Dacrycarpus'', ''
Dacrydium'', ''
Falcatifolium'', ''
Retrophyllum'', ''
Nageia'', ''
Afrocarpus'', ''
Podocarpus''
*Order
Cupressales
**Family
Sciadopityaceae: ''
Sciadopitys''
**Family
Cupressaceae: ''
Cunninghamia'', ''
Taiwania'', ''
Athrotaxis'', ''
Metasequoia'', ''
Sequoia'', ''
Sequoiadendron'', ''
Cryptomeria'', ''
Glyptostrobus'', ''
Taxodium'', ''
Papuacedrus'', ''
Austrocedrus'', ''
Libocedrus'', ''
Pilgerodendron'', ''
Widdringtonia'', ''
Diselma'', ''
Fitzroya'', ''
Callitris'', ''
Actinostrobus'', ''
Neocallitropsis'', ''
Thujopsis'', ''
Thuja'', ''
Fokienia'', ''
Chamaecyparis'', ''
Cupressus'', ''
Juniperus'', ''
Calocedrus'', ''
Tetraclinis'', ''
Platycladus'', ''
Microbiota''
**Family
Taxaceae: ''
Austrotaxus'', ''
Pseudotaxus'', ''
Taxus'', ''
Cephalotaxus'', ''
Amentotaxus'', ''
Torreya''
Extinct groupings
* Division
Pteridospermatophyta
* Order
Bennettitales
** Family
Cycadeoidaceae
** Family
Williamsoniaceae
* Order
Erdtmanithecales
*Order
Pentoxylales
*Order
Czekanowskiales
Life cycle

Gymnosperms, like all
vascular plants, have a sporophyte-dominant life cycle, which means they spend most of their life cycle with diploid cells, while the
gametophyte (gamete-bearing phase) is relatively short-lived. Like all
seed plants, they are
heterosporous, having two spore types, microspores (male) and megaspores (female) that are typically produced in pollen cones or ovulate cones, respectively. The exception is the females in the cycad genus
Cycas, which form a loose structure called megasporophylls instead of cones. As with all heterosporous plants, the gametophytes develop within the spore wall. Pollen grains (microgametophytes) mature from microspores, and ultimately produce sperm cells. Megagametophytes develop from megaspores and are retained within the ovule. Gymnosperms produce multiple
archegonia, which produce the female gamete.
During pollination, pollen grains are physically transferred between plants from the pollen cone to the ovule. Pollen is usually moved by wind or insects. Whole grains enter each ovule through a microscopic gap in the ovule coat (
integument) called the micropyle. The pollen grains mature further inside the ovule and produce sperm cells. Two main modes of fertilization are found in gymnosperms. Cycads and ''
Ginkgo'' have
flagellated motile sperm
that swim directly to the egg inside the ovule, whereas conifers and
gnetophytes have sperm with no flagella that are moved along a
pollen tube to the egg. After
syngamy (joining of the sperm and egg cell), the zygote develops into an embryo (young sporophyte). More than one embryo is usually initiated in each gymnosperm seed. The mature seed comprises the embryo and the remains of the female
gametophyte, which serves as a food supply, and the
seed coat.
Genetics
The first published sequenced genome for any gymnosperm was the genome of ''
Picea abies'' in 2013.
Uses
Gymnosperms have major economic uses. Pine, fir, spruce, and cedar are all examples of conifers that are used for
lumber, paper production, and resin. Some other common uses for gymnosperms are
soap,
varnish,
nail polish, food, gum, and
perfumes.
References
General bibliography
*
External links
Gymnosperm DatabaseGymnosperms on the Tree of Life*
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Extant Late Devonian first appearances