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Spermatophyte

The spermatophytes, also known as


phanerogams (taxon Phanerogamae) or
phaenogams (taxon Phaenogamae),
comprise those plants that produce
seeds, hence the alternative name seed
plants. They are a subset of the
embryophytes or land plants.
Seed plants
Temporal range: Famennian–Present
PreꞒ Ꞓ O S D C P T J K PN
g

Scots pine, Pinus sylvestris, a member of the


Pinophyta
Scientific classification
Kingdom: Plantae

Clade: Tracheophytes

Clade: Spermatophytes

Divisions

Gymnospermae
Cycadophyta
Ginkgophyta
Pinophyta
Gnetophyta
Bennettitales†
Angiospermae
Pteridospermatophyta†

Synonyms
Phanerogamae
Phaenogamae

The term phanerogams or phanerogamae


is derived from the Greek φανερός,
phanerós meaning "visible", in contrast to
the cryptogamae from Greek κρυπτός
kryptós = "hidden" together with the suffix
γαμέω, gameo, "to marry". These terms
distinguished those plants with hidden
sexual organs (cryptogamae) from those
with visible sexual organs
(phanerogamae).

Description
The extant spermatophytes form five
divisions, the first four of which are
traditionally grouped as gymnosperms,
plants that have unenclosed, "naked
seeds":

Cycadophyta, the cycads, a subtropical


and tropical group of plants,
Ginkgophyta, which includes a single
living species of tree in the genus
Ginkgo,
Pinophyta, the conifers, which are
cone-bearing trees and shrubs, and
Gnetophyta, the gnetophytes, various
woody plants in the relict genera
Ephedra, Gnetum, and Welwitschia.
The fifth extant division is the flowering
plants, also known as angiosperms or
magnoliophytes, the largest and most
diverse group of spermatophytes:

Angiosperms, the flowering plants,


possess seeds enclosed in a fruit,
unlike gymnosperms.

In addition to the five living taxa listed


above, the fossil record contains
evidence of many extinct taxa of seed
plants, among those:

Pteridospermae, the so-called "seed


ferns", were one of the earliest
successful groups of land plants, and
forests dominated by seed ferns were
prevalent in the late Paleozoic.
Glossopteris was the most prominent
tree genus in the ancient southern
supercontinent of Gondwana during
the Permian period.

By the Triassic period, seed ferns had


declined in ecological importance, and
representatives of modern gymnosperm
groups were abundant and dominant
through the end of the Cretaceous, when
the angiosperms radiated.

Evolution
A whole genome duplication event in the
ancestor of seed plants occurred about
319 million years ago.[1] This gave rise to
a series of evolutionary changes that
resulted in the origin of seed plants.

A middle Devonian (385-million-year-old)


precursor to seed plants from Belgium
has been identified predating the earliest
seed plants by about 20 million years.
Runcaria, small and radially symmetrical,
is an integumented megasporangium
surrounded by a cupule. The
megasporangium bears an unopened
distal extension protruding above the
mutlilobed integument. It is suspected
that the extension was involved in
anemophilous (wind) pollination.
Runcaria sheds new light on the
sequence of character acquisition
leading to the seed. Runcaria has all of
the qualities of seed plants except for a
solid seed coat and a system to guide
the pollen to the seed.[2]

Relationships and
nomenclature
Seed-bearing plants are a subclade of the
vascular plants (tracheophytes) and were
traditionally divided into angiosperms, or
flowering plants, and gymnosperms,
which includes the gnetophytes, cycads,
ginkgo, and conifers. Older
morphological studies believed in a close
relationship between the gnetophytes
and the angiosperms,[3] in particular
based on vessel elements. However,
molecular studies (and some more
recent morphological[4][5] and fossil[6]
papers) have generally shown a clade of
gymnosperms, with the gnetophytes in or
near the conifers. For example, one
common proposed set of relationships is
known as the gne-pine hypothesis and
looks like:[7][8][9]
  angiosperms (flowering
  plants)

  cycads [10]

  Ginkgo

gymnosperms   Pinaceae (the pine
    family)

      gnetophytes
   
  other conifers

However, the relationships between


these groups should not be considered
settled.[3][11]

Other classifications group all the seed


plants in a single division, with classes
for the five groups:

Division Spermatophyta
Cycadopsida, the cycads
Ginkgoopsida, the ginkgo
Pinopsida, the conifers,
("Coniferopsida")
Gnetopsida, the gnetophytes
Magnoliopsida, the flowering
plants, or Angiospermopsida

A more modern classification ranks


these groups as separate divisions
(sometimes under the Superdivision
Spermatophyta):

Cycadophyta, the cycads


Ginkgophyta, the ginkgo
Pinophyta, the conifers
Gnetophyta, the gnetophytes
Magnoliophyta, the flowering plants

An alternative phylogeny of
spermatophytes based on the work by
Novíkov & Barabaš-Krasni 2015[12] with
plant taxon authors from Anderson,
Anderson & Cleal 2007[13] showing the
relationship of extinct clades.
Spermatophytina  
  †Moresnetiopsida Doweld 2

  †Lyginopteridopsida Nová
    Anderson & Cleal 2007

    †Pachytestopsida Dowe
  
  †Callistophytales Rothw
    Anderson, Anderson &

    †Peltaspermopsida D
  
    †Umkomasiales Dow
  
   
   
Acrogymnospermae

Unassigned spermatophytes:

†Avatiaceae Anderson & Anderson


2003
†Axelrodiopsida Anderson & Anderson
†Alexiales Anderson & Anderson 2003
†Hamshawviales Anderson &
Anderson 2003
†Hexapterospermales Doweld 2001
†Hlatimbiales Anderson & Anderson
2003
†Matatiellales Anderson & Anderson
2003
†Petriellales Taylor et al. 1994
†Arberiopsida Doweld 2001
†Czekanowskiales Taylor et al. 2008
†Iraniales E. Taylor et al. 2008
†Vojnovskyales E. Taylor et al. 2008
†Hermanophytales E. Taylor et al. 2008
†Dirhopalostachyaceae E. Taylor et al.
2008

References
1. 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
2. "Science Magazine" . Runcaria, a
Middle Devonian Seed Plant
Precursor. American Association for
the Advancement of Science. 2011.
Retrieved March 22, 2011.
3. Palmer, Jeffrey D.; Soltis, Douglas E.;
Chase, Mark W. (2004). "The plant
tree of life: an overview and some
points of view" . American Journal of
Botany. 91 (10): 1437–1445.
doi:10.3732/ajb.91.10.1437 .
PMID 21652302 .
4. James A. Doyle (January 2006).
"Seed ferns and the origin of
angiosperms". The Journal of the
Torrey Botanical Society. 133 (1):
169–209. doi:10.3159/1095-
5674(2006)133[169:SFATOO]2.0.CO
;2 . ISSN 1095-5674 .
5. Coiro, Mario; Chomicki, Guillaume;
Doyle, James A. (n.d.). "Experimental
signal dissection and method
sensitivity analyses reaffirm the
potential of fossils and morphology
in the resolution of the relationship
of angiosperms and Gnetales".
Paleobiology. 44 (3): 490–510.
doi:10.1017/pab.2018.23 .
ISSN 0094-8373 . S2CID 91488394 .
6. Zi-Qiang Wang (2004). "A New
Permian Gnetalean Cone as Fossil
Evidence for Supporting Current
Molecular Phylogeny" . Annals of
Botany. 94 (2): 281–288.
doi:10.1093/aob/mch138 .
PMC 4242163 . PMID 15229124 .
7. Chaw, Shu-Miaw; Parkinson,
Christopher L.; Cheng, Yuchang;
Vincent, Thomas M.; Palmer, Jeffrey
D. (2000). "Seed plant phylogeny
inferred from all three plant
genomes: Monophyly of extant
gymnosperms and origin of Gnetales
from conifers" . Proceedings of the
National Academy of Sciences. 97
(8): 4086–4091.
Bibcode:2000PNAS...97.4086C .
doi:10.1073/pnas.97.8.4086 .
PMC 18157 . PMID 10760277 .
8. Bowe, L. M.; Michelle, L.; Coat,
Gwénaële; Claude (2000).
"Phylogeny of seed plants based on
all three genomic compartments:
Extant gymnosperms are
monophyletic and Gnetales' closest
relatives are conifers" . Proceedings
of the National Academy of
Sciences. 97 (8): 4092–4097.
Bibcode:2000PNAS...97.4092B .
doi:10.1073/pnas.97.8.4092 .
PMC 18159 . PMID 10760278 .
9. Soltis, Douglas E.; Soltis, Pamela S.;
Zanis, Michael J. (2002). "Phylogeny
of seed plants based on evidence
from eight genes" . American
Journal of Botany. 89 (10): 1670–
1681. doi:10.3732/ajb.89.10.1670 .
PMID 21665594 . Archived from the
original on 2012-07-10.
10. Chung-Shien Wu, Ya-Nan Wang, Shu-
Mei Liu and Shu-Miaw Chaw (2007).
"Chloroplast Genome (cpDNA) of
Cycas taitungensis and 56 cp
Protein-Coding Genes of Gnetum
parvifolium: Insights into cpDNA
Evolution and Phylogeny of Extant
Seed Plants" . Molecular Biology and
Evolution. 24 (6): 1366–1379.
doi:10.1093/molbev/msm059 .
PMID 17383970 .
11. Won, Hyosig; Renner, Susanne
(August 2006). "Dating Dispersal and
Radiation in the Gymnosperm
Gnetum (Gnetales)—Clock
Calibration When Outgroup
Relationships Are Uncertain" .
Systematic Biology. 55 (4): 610–622.
doi:10.1080/10635150600812619 .
PMID 16969937 .
12. Novíkov & Barabaš-Krasni (2015).
Modern plant systematics. Liga-
Pres. p. 685.
doi:10.13140/RG.2.1.4745.6164 .
ISBN 978-966-397-276-3.
13. Anderson, Anderson & Cleal (2007).
Brief history of the gymnosperms:
classification, biodiversity,
phytogeography and ecology.
Strelitzia. 20. SANBI. p. 280.
ISBN 978-1-919976-39-6.

Bibliography
Kron, Kathleen A; Chase, Mark W (2005-11-
17). Molecular systematics and seed plant
phylogeny: a summary of a parsimony
analysis of rbcL sequence data . pp. 243–
252. ISBN 9780521022897., in Gibbs et al
(1995)
Gibbs, Adrian J.; Calisher, Charles H.;
García-Arenal, Fernando, eds. (1995).
Molecular basis of virus evolution .
Cambridge: Cambridge University Press.
ISBN 9780521022897.
Soltis, D. E.; Soltis, P. S.; Zanis, M. J. (1
October 2002). "Phylogeny of seed plants
based on evidence from eight genes" .
American Journal of Botany. 89 (10): 1670–
1681. doi:10.3732/ajb.89.10.1670 .
PMID 21665594 . S2CID 2444652 .

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