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The gymnosperms are a group of seed-bearing plants that includes conifers, cycads, Ginkgo, and Gnetales. The term "gymnosperm" comes from the Greek word gymnospermos (?????spe????), meaning "naked seeds", after the unenclosed condition of their seeds (called ovules in their unfertilized state). Their naked condition stands in contrast to the seeds or ovules of flowering plants (angiosperms), which are enclosed during pollination. Gymnosperm seeds develop either on the surface of scale- or leaf-like appendages of cones, or at the end of short stalks (Ginkgo).

The gymnosperms and angiosperms together comprise the spermatophytes or seed plants. By far the largest group of living gymnosperms are the conifers (pines, cypresses, and relatives), followed by cycads, Gnetales (Gnetophyta, Ephedra and Welwitschia), and Ginkgo (a single living species).

In early classif ication schemes, the gymnosperms (Gymnospermae) were regarded as a "natural" group. There is conflicting evidence on the question of whether the living gymnosperms form a clade.^[1][2] 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-like vegetative morphology (the so-called "seed ferns" or pteridosperms.)^[3] When fossil gymnosperms such as Bennettitales, Caytonia and the glossopterids are considered, it is clear that angiosperms are nested within a larger gymnosperm clade, although which group of gymnosperms is their closest relative remains unclear.

For the most recent classification on extant gymnosperms see Christenhusz et al. (2011)^[4] and Lee EK, Cibrian-Jaramillo A, et al. (2011)^[5]

Classification:'

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 (incl. Actinostrobus), Neocallitropsis, Thujopsis, Thuja, Fokienia, Chamaecyparis, Cupressus, Juniperus, Calocedrus, Tetraclinis, Platycladus, Microbiota
  • Family Taxaceae: Austrotaxus, Pseudotaxus, Taxus, Cephalotaxus, Amentotaxus, Torreya

Diversity and origin

There are between 700 and 900 extant or currently living species of Gymnosperms.

It is widely accepted ^[6] that the gymnosperms originated i n the late Carboniferous period. This appears to have been the result of a whole genome duplication event around 319 million years ago^[7] Early characteristics of seed plants were evident in fossil progymnosperms of the late Devonian period around 380 million years ago. It has been suggested that during the mid-Mesozoic era, pollination of some extinct groups of gymnosperms were 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.^[8][9]

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).^[10] Conifers are woody plants and most are evergreens.^[11] The leaves of many conifers are long, thin and needle-like, others 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 about 130 species. The other extant groups are the 75 - 80 species of Gnetales and one species of Ginkgo.

Uses

Gymnosperms have major economic uses. Pine, fir, spruce, and cedar are all examples of conifers that are used for lumber. Some other common uses for gymnosperms are soap, varnish, nail polish, food, and perfumes.

Life cycle

Gymnosperms are spore-bearing plants (sporophytes), with a sporophyte-dominant life-cycle; as in all other vascular plants, the gametophyte (gamete-bearing phase) is relatively short-lived. Two spore types, microspores and megaspores, are, in general, produced in pollen cones or ovulate cones, respectively. A short-lived multicellular haploid, gamete-bearing phase (gametophyte) develops inside the spore wall. Pollen grains (microgametophytes) mature from microspores, and ultimately produce sperm cells; megagametophyte tissue develops in the megaspore of each ovule, and produces multiple egg cells. Thus, megaspores are enclosed in ovules (unfertilized seeds) and give rise to megagametophytes and ultimately to egg cells. During pollination, pollen grains are physically transferred between plants, from pollen cone to the ovule, being transferred 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 motile sperm that swim directly to the egg inside the ovule, whereas conifers and gnetophytes have sperm with no flagella that are conveyed to the egg along a pollen tube. After fertilization (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. Competition between the embryos for nutritional resources within polyembryonic seeds produces programmed cell death to all but one embryo. The mature seed comprises the embryo and the remains of the female gametophyte, which serves as a food supply, and the seed coat (integument).^[12]

million years ago^[7] Early characteristics of seed plants were evident in fossil progymnosperms of the late Devonian period around 380 million years ago. It has been suggested that during the mid-Mesozoic era, pollination of some extinct groups of gymnosperms were 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.^[8][9]

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).^[10] Conifers are woody plants and most are evergreens.^[11] The leaves of many conifers are long, thin and needle-like, others 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 about 130 species. The other extant groups are the 75 - 80 species of Gnetales and one species of Ginkgo.

Uses

Gymnosperms have major economic uses. Pine, fir, spruce, and cedar are all examples of conifers that are used for lumber. Some other common uses for gymnosperms are soap, varnish, nail polish, food, and perfumes.

Life cycle

Gymnosperms are spore-bearing plants (sporophytes), with a sporophyte-dominant life-cycle; as in all other vascula r plants, the gametophyte (gamete-bearing phase) is relatively short-lived. Two spore types, microspores and megaspores, are, in general, produced in pollen cones or ovulate cones, respectively. A short-lived multicellular haploid, gamete-bearing phase (gametophyte) develops inside the spore wall. Pollen grains (microgametophytes) mature from microspores, and ultimately produce sperm cells; megagametophyte tissue develops in the megaspore of each ovule, and produces multiple egg cells. Thus, megaspores are enclosed in ovules (unfertilized seeds) and give rise to megagametophytes and ultimately to egg cells. During pollination, pollen grains are physically transferred between plants, from pollen cone to the ovule, being transferred 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 motile sperm that swim directly to the egg inside the ovule, whereas conifers and gnetophytes have sperm with no flagella that are conveyed to the egg along a pollen tube. After fertilization (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. Competition between the embryos for nutritional resources within polyembryonic seeds produces programmed cell death to all but one embryo. The mature seed comprises the embryo and the remains of the female gametophyte, which serves as a food supply, and the seed coat (integument).^[12]

References

External links

• Gymnosperm Database
• Gymnosperms on the Tree of Life

The Infraphylum Gymnospermae is a member of the Subphylum Spermatophytina. Here is the complete "parentage" of Gymnospermae:

• Domain: Eukaryota Whittaker & Margulis,1978 - eukaryotes
  • Kingdom: Plantae Haeckel, 1866 - Plants
    • Subkingdom: Viridaeplantae Cavalier-Smith, 1981 - Green Plants
      • Phylum: Magnoliophyta Cronquist, Takhtajan & W. Zimmermann, 1966 - Flowering Plants
        • Subphylum: Spermatophytina (Auct.) Cavalier-Smith, 1998 - Seed Plants
          • Infraphylum: Gymnospermae Auct.