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A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
that produces
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s, hence the alternative name seed plant. Spermatophytes are a subset of the
embryophyte The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sis ...
s or land plants. They include most familiar types of plants, including all
flowers A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
and most
trees In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are u ...
, but exclude some other types of plants such as ferns,
mosses Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hornw ...
,
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
. The term ''phanerogams'' or ''phanerogamae'' is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
(), meaning "visible", in contrast to the cryptogamae (), together with the
suffix In linguistics, a suffix is an affix which is placed after the stem of a word. Common examples are case endings, which indicate the grammatical case of nouns, adjectives, and verb endings, which form the conjugation of verbs. Suffixes can carr ...
(), meaning "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
gymnosperm 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, γυμν ...
s, plants that have unenclosed, "naked seeds": *
Cycadophyta Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
, the cycads, a subtropical and tropical group of plants, * Ginkgophyta, which includes a single living species of tree in the genus '' Ginkgo'', *
Pinophyta Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
, the conifers, which are cone-bearing trees and shrubs, and *
Gnetophyta Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three ...
, the gnetophytes, various woody plants in the relict genera '' Ephedra'', ''
Gnetum ''Gnetum'' is a genus of gymnosperms, the sole genus in the family Gnetaceae within the Gnetophyta. They are tropical evergreen trees, shrubs and lianas. Unlike other gymnosperms, they possess vessel elements in the xylem. Some species have been ...
'', and ''
Welwitschia ''Welwitschia'' is a monotypic gymnosperm genus, comprising solely the distinctive ''Welwitschia mirabilis'', endemic to the Namib desert within Namibia and Angola. ''Welwitschia'' is the only living genus of the family Welwitschiaceae and orde ...
''. The fifth extant division is the
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s, also known as angiosperms or magnoliophytes, the largest and most diverse group of spermatophytes: *
Angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
, the flowering plants, possess seeds enclosed in a
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
, unlike gymnosperms. In addition to the five living taxa listed above, the fossil record contains evidence of many
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
taxa of seed plants, among those: *
Pteridospermae The term Pteridospermatophyta (or "seed ferns" or "Pteridospermatopsida") is a polyphyletic group of extinct seed-bearing plants (spermatophytes). The earliest fossil evidence for plants of this type is the genus ''Elkinsia'' of the late Devonian ...
, 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 The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ...
. * '' Glossopteris'' was the most prominent tree genus in the ancient southern supercontinent of
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
during the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
period. By the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
period, seed ferns had declined in ecological importance, and representatives of modern gymnosperm groups were abundant and dominant through the end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
, when the
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
radiated.


Evolutionary history

A whole genome duplication event in the ancestor of seed plants occurred about .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 This gave rise to a series of evolutionary changes that resulted in the origin of seed plants. A middle
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
(385-million-year-old)
precursor Precursor or Precursors may refer to: * Precursor (religion), a forerunner, predecessor ** The Precursor, John the Baptist Science and technology * Precursor (bird), a hypothesized genus of fossil birds that was composed of fossilized parts of u ...
to seed plants from
Belgium Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to ...
has been identified predating the earliest seed plants by about 20 million years. ''
Runcaria ''Runcaria heinzelinii'' is an extinct species of plant from the Middle Devonian of Belgium. It is the only species classified under the genus ''Runcaria'', named after Jean de Heinzelin de Braucourt. It was a precursor to seed plants, predating ...
'', 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 Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
. ''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 are a subclade of the
vascular plants Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
(tracheophytes) and were traditionally divided into
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
s, or flowering plants, and
gymnosperm 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, γυμν ...
s, 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 element A vessel element or vessel member (also called trachea or xylem vessel) is one of the cell types found in xylem, the water conducting tissue of plants. Vessel elements are typically found in angiosperms (flowering plants) but absent from most g ...
s. However, molecular studies (and some more recent morphological and fossil papers) have generally shown a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
of
gymnosperm 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, γυμν ...
s, 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: However, the relationships between these groups should not be considered settled. Other classifications group all the seed plants in a single division, with classes for the five groups: *Division Spermatophyta **
Cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
opsida, the cycads ** Ginkgoopsida, the ginkgo **
Pinopsida Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...
, the conifers, ("Coniferopsida") **
Gnetopsida Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three rel ...
, the gnetophytes **
Magnoliopsida Magnoliopsida is a valid botanical name for a class of flowering plants. By definition the class will include the family Magnoliaceae, but its circumscription can otherwise vary, being more inclusive or less inclusive depending upon the classif ...
, the
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s, or Angiospermopsida A more modern classification ranks these groups as separate divisions (sometimes under the Superdivision Spermatophyta): *
Cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
ophyta, the cycads * Ginkgophyta, the ginkgo *
Pinophyta Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
, the conifers *
Gnetophyta Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three ...
, the gnetophytes * Magnoliophyta, the
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s 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 Czekanowskiales, also known as Leptostrobales, are an extinct group of seed plants. Members of the family are distinguished by persistent leaves borne on deciduous short shoots, subtended by scale-like leaves. The leaves are highly dissected (div ...
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


Bibliography

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