|Scots pine, Pinus sylvestris, a member of the Pinophyta|
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. 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).
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 possess seeds enclosed in a fruit, unlike gymnosperms.
In addition to the taxa listed above, the fossil record contains evidence of many extinct taxa of seed plants. The so-called "seed ferns" (Pteridospermae) 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 angiosperms radiated.
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.
Relationships and nomenclature
Seed-bearing plants 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, in particular based on vessel elements. However, molecular studies (and some more recent morphological and fossil 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:
angiosperms (flowering plants)
- Division Spermatophyta
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 with plant taxon authors from Anderson, Anderson & Cleal 2007 showing the relationship of extinct clades.
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
- 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
- "Science Magazine". Runcaria, a Middle Devonian Seed Plant Precursor. American Association for the Advancement of Science. 2011. Retrieved March 22, 2011.
- 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.
- 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.
- 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.
- 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.
- 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. doi:10.1073/pnas.97.8.4086. PMC 18157. PMID 10760277.
- 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. doi:10.1073/pnas.97.8.4092. PMC 18159. PMID 10760278.
- 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. Cite uses deprecated parameter
- 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.CS1 maint: multiple names: authors list (link)
- 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.
- 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.
- 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.
- 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.