Gymnosperm

The gymnosperms, also known as

Acrogymnospermae, are a group of
seed-producing plants that includes
conifers, cycads, Ginkgo, and
gnetophytes. The term "gymnosperm"
comes from the composite word in
Greek: γυμνόσπερμος (γυμνός, gymnos,
'naked' and σπέρμα, sperma, 'seed'),
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.[1]
 Gymnospermae
Temporal range: Carboniferous – Present
PreꞒ Ꞓ O S D C P T J K PN
g

Various gymnosperms.

Scientific classification

Kingdom: Plantae

Subkingdom: Embryophyta

(unranked): Spermatophyta

(unranked): Gymnospermae
(inc. †; paraphyletic)
Acrogymnospermae
(extant only)

Divisions
Pinophyta (or Coniferophyta) – Conifers
Ginkgophyta – Ginkgo
Cycadophyta – Cycads
Gnetophyta – Gnetum, Ephedra,
Welwitschia
Encephalartos sclavoi cone, about 30 cm long

The gymnosperms and angiosperms

together compose the spermatophytes
or seed plants. The gymnosperms are
divided into six phyla. Organisms that
belong to the Cycadophyta, Ginkgophyta,
Gnetophyta, and Pinophyta (also known
as Coniferophyta) phyla are still in
existence while those in the
Pteridospermales and Cordaitales phyla
are now extinct.[2]

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).

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.

Classification

An example Phylogenetic tree

A formal classification of the living
gymnosperms is the
"Acrogymnospermae", which form a
monophyletic group within the
spermatophytes.[3][4] 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-like
vegetative morphology (the so-called
"seed ferns" or pteridosperms).[5] 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.[1][4][6]

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 …

Order Bennettitales
Family Cycadeoidaceae
Family Williamsoniaceae
Order Erdtmanithecales
Order Pentoxylales
Order Czekanowskiales

Diversity and origin

Over 1000 living species of gymnosperm
exist.[1] It is widely accepted that the
gymnosperms originated in the late
Carboniferous period, replacing the
lycopsid rainforests of the tropical
region.[7][8] This development appears to
have resulted from a whole genome
duplication event around
319 million years ago.[9] 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.[10][11] 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.[12]
Zamia integrifolia, a cycad native to Florida

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).[13] Conifers are woody
plants and most are evergreens.[14] 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.[2]

  Pteridospermatophyta
 †
Spermatophyta   Acrogymnospermae
 
   
    Angiospermae
 
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.

Life cycle

Example of gymnosperm lifecycle

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. Two spore types,
microspores(male) and
megaspores(female), are typically
produced in pollen cones or ovulate
cones, respectively. Gametophytes, as
with all heterosporous plants, 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
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.[15]

Genetics
The first published sequenced genome
for any gymnospermae was the genome
of Picea abies in 2013.[16]

References
1. "Gymnosperms on The Plant List" .
 Theplantlist.org. Retrieved
2013-07-24.
2. Raven, P.H. (2013). Biology of Plants.
New York: W.H. Freeman and Co.
3. Cantino 2007.
4. Christenhusz, M.J.M.; Reveal, J.L.;
Farjon, A.; Gardner, M.F.; Mill, R.R.;
Chase, M.W. (2011). "A new
classification and linear sequence of
extant gymnosperms" (PDF).
Phytotaxa. 19: 55–70.
doi:10.11646/phytotaxa.19.1.3 .
5. Hilton, Jason, and Richard M.
Bateman. 2006. Pteridosperms are
the backbone of seed-plant
phylogeny. Journal of the Torrey
 Botanical Society 133: 119–168
(abstract )
6. Christenhusz, M. J. M.; Byng, J. W.
(2016). "The number of known
plants species in the world and its
annual increase" . Phytotaxa. 261
(3): 201–217.
doi:10.11646/phytotaxa.261.3.1 .
7. Sahney, S.; Benton, M.J. & Falcon-
Lang, H.J. (2010). "Rainforest
collapse triggered Pennsylvanian
tetrapod diversification in
Euramerica". Geology. 38 (12):
1079–1082.
Bibcode:2010Geo....38.1079S .
doi:10.1130/G31182.1 .
 8. Campbell and Reece; Biology, Eighth
edition
9. Jiao Y, Wickett NJ, Ayyampalayam S,
Chanderbali AS, Landherr L, Ralph
PE, Tomsho LP, Hu Y, Liang H, Soltis
PS, Soltis DE, Clifton SW,
Schlarbaum SE, Schuster SC, Ma H,
Leebens-Mack J, Depamphilis CW
(2011) Ancestral polyploidy in seed
plants and angiosperms. Nature
10. Ollerton, J.; Coulthard, E. (2009).
"Evolution of Animal Pollination".
Science. 326 (5954): 808–809.
Bibcode:2009Sci...326..808O .
doi:10.1126/science.1181154 .
PMID 19892970 . S2CID 856038 .
11. Ren, D; Labandeira, CC; Santiago-
Blay, JA; Rasnitsyn, A; et al. (2009).
"A Probable Pollination Mode Before
Angiosperms: Eurasian, Long-
Proboscid Scorpionflies" . Science.
326 (5954): 840–847.
Bibcode:2009Sci...326..840R .
doi:10.1126/science.1178338 .
PMC 2944650 . PMID 19892981 .
12. Labandeira, Conrad C.; Yang, Qiang;
Santiago-Blay, Jorge A.; Hotton,
Carol L.; Monteiro, Antónia; Wang,
Yong-Jie; Goreva, Yulia; Shih,
ChungKun; Siljeström, Sandra; Rose,
Tim R.; Dilcher, David L.; Ren, Dong
(2016). "The evolutionary
convergence of mid-Mesozoic
 lacewings and Cenozoic
butterflies" . Proceedings of the
Royal Society B: Biological Sciences.
283 (1824): 20152893.
doi:10.1098/rspb.2015.2893 .
PMC 4760178 . PMID 26842570 .
13. Catalogue of Life: 2007 Annual
checklist – Conifer database
Archived January 15, 2009, at the
Wayback Machine
14. Campbell, Reece, "Phylum
Coniferophyta."Biology. 7th. 2005.
Print. P.595
15. Walters, Dirk R Walters Bonnie By
(1996). Vascular plant taxonomy .
Dubuque, Iowa: Kendall/Hunt Pub.
 Co. p. 124 . ISBN 978-0-7872-2108-
9. "Gymnosperm seeds."
16. Nystedt, B; Street, NR; Wetterbom, A;
et al. (May 2013). "The Norway
spruce genome sequence and
conifer genome evolution" . Nature.
497 (7451): 579–584.
Bibcode:2013Natur.497..579N .
doi:10.1038/nature12211 .
PMID 23698360 .

Bibliography
Cantino, Philip D.; Doyle, James A.; Graham,
Sean W.; Judd, Walter S.; Olmstead, Richard
G.; Soltis, Douglas E.; Soltis, Pamela S.;
Donoghue, Michael J. (August 2007).
"Towards a phylogenetic nomenclature of
 Tracheophyta". Taxon. 56 (3): 822–846.
doi:10.2307/25065864 .
JSTOR 25065864 .

External links

Wikimedia Commons has media

related to Gymnosperms.

Gymnosperm Database
Gymnosperms on the Tree of Life
Albert Seward (1911).
"Gymnosperms"  . Encyclopædia
Britannica (11th ed.).

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