Gnetophyta is a division of plants, grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three relict genera: Gnetum (family Gnetaceae), Welwitschia (family Welwitschiaceae), and Ephedra (family Ephedraceae). Fossilized pollen attributed to a close relative of Ephedra has been dated as far back as the Early Cretaceous.[1] Though diverse and dominant in the Paleogene and the Neogene,[2] only three families, each containing a single genus, are still alive today. The primary difference between gnetophytes and other gymnosperms is the presence of vessel elements, a system of conduits that transport water within the plant, similar to those found in flowering plants. Because of this, gnetophytes were once thought to be the closest gymnosperm relatives to flowering plants, but more recent molecular studies have brought this hypothesis into question.

Though it is clear they are all closely related, the exact evolutionary inter-relationships between gnetophytes are unclear. Some classifications hold that all three genera should be placed in a single order (Gnetales), while other classifications say they should be distributed among three separate orders, each containing a single family and genus. Most morphological and molecular studies confirm that the genera Gnetum and Welwitschia diverged from each other more recently than they did from Ephedra.[3][4][5][6][7]

Welwitschia mirabilis S&J6
Welwitschia mirabilis bearing male cones
Ephedra distachya (male flowers) 1
Ephedra distachya (male cones)
Ephedra distachya (female plant in bloom)
Ephedra distachya (female plant in bloom)
Gnetum gnemon male
Gnetum gnemon male strobili
Gnetum gnemon BotGardBln1105C
Gnetum gnemon female strobilus
Temporal range: Cretaceous–recent
Welwitschia at Ugab River basin
Welwitschia mirabilis female plant with cones
Scientific classification
Kingdom: Plantae
Division: Gnetophyta
Class: Gnetopsida
Families & Genera


A distribution map of Gnetophyta colour-coded by genus
Distribution, separated by genus:
Green – Welwitschia
Blue – Gnetum
Red – Ephedra
Purple – Gnetum and Ephedra

Ecology and morphology

Unlike most biological groupings, it is difficult to find many common characteristics between all of the members of the gnetophytes.[2] The two common characteristics most commonly used are the presence of enveloping bracts around both the ovules and microsporangia as well as a micropylar projection of the outer membrane of the ovule that produces a pollination droplet,[8] though these are highly specific compared to the similarities between most other plant divisions. L. M. Bowe refers to the gnetophyte genera as a "bizarre and enigmatic" trio[4] because, the gnetophytes' specialization to their respective environments is so complete that they hardly resemble each other at all. Gnetum species are mostly woody vines in tropical forests, though the best-known member of this group, Gnetum gnemon, is a tree native to western Malesia. The one remaining species of Welwitschia, Welwitschia mirabilis, native only to the dry deserts of Namibia and Angola, is a ground-hugging species with only two large strap-like leaves that grow continuously from the base throughout the plant's life. Ephedra species, known as "jointfirs" in the United States, have long slender branches which bear tiny scale-like leaves at their nodes. Infusions from these plants have been traditionally used as a stimulant, but ephedrine is a controlled substance today in many places because of the risk of harmful or even fatal overdosing.

Fossil Gnetophyta

Knowledge of gnetophyte history through fossil discovery has increased greatly since the 1980s.[3] Gnetophyte fossils have been found that date from the Permian[9] and the Triassic. Fossils dating back to the Jurassic have been found, though whether or not they belong to the gnetophytes is uncertain.[10] Overall, the fossil record is richest in the early Cretaceous, with fossils of plants, seeds, and pollen have been found that can clearly be assigned to the gnetophytes.[10]


With just three well-defined genera within an entire division, there still is understandable difficulty in establishing an unambiguous interrelationship among them; in earlier times matters were even more difficult and we find for example Pearson in the early 20th century speaking of the class Gnetales, rather than the order.[11] G. H. M. Lawrence referred to them as an order, but remarked that the three families were distinct enough to deserve recognition as separate orders.[12] Foster & Gifford accepted this principle, and placed the three orders together in a common class for convenience, which they called Gnetopsida.[13] In general the evolutionary relationships among the seed plants still are unresolved, and the Gnetophyta have played an important role in the formation of phylogenetic hypotheses. Molecular phylogenies of extant gymnosperms have conflicted with morphological characters with regard to whether the gymnosperms as a whole (including gnetophytes) comprise a monophyletic group or a paraphyletic one that gave rise to angiosperms. At issue is whether the Gnetophyta are the sister group of angiosperms, or whether they are sister to, or nested within, other extant gymnosperms. Numerous fossil gymnosperm clades once existed that are morphologically at least as distinctive as the four living gymnosperm groups, such as Bennettitales, Caytonia and the glossopterids. When these gymnosperm fossils are considered, the question of gnetophyte relationships to other seed plants becomes even more complicated. Several hypotheses, illustrated below, have been presented to explain seed plant evolution.

Recent research by Lee EK, Cibrian-Jaramillo A, et al. (2011) suggests that the Gnetophyta are a sister group to the rest of the gymnosperms,[14] contradicting the anthophyte hypothesis, which held that gnetophytes were sister to the flowering plants.

Anthophyte hypothesis

From the early twentieth century, the anthophyte hypothesis was the prevailing explanation for seed plant evolution, based on shared morphological characters between the gnetophytes and angiosperms. In this hypothesis, the gnetophytes, along with the extinct order Bennettitales, are sister to the angiosperms, forming the "anthophytes".[8] Some morphological characters that were suggested to unite the anthophytes include vessels in wood, net-veined leaves (in Gnetum only), lignin chemistry, the layering of cells in the apical meristem, pollen and megaspore features (including thin megaspore wall), short cambial initials, and lignin syringal groups.[8][15][16][17] However, most genetic studies, as well as more recent morphological analyses[18], have rejected the anthophyte hypothesis.[4][19][20][21][22][23][24][25][26][27] Several of these studies have suggested that the gnetophytes and angiosperms have independently derived characters, including flower-like reproductive structures and tracheid vessel elements, that appear shared but are actually the result of parallel evolution.[4][8][20]





angiosperms (flowering plants)


Gnetifer hypothesis

In the gnetifer hypothesis, the gnetophytes are sister to the conifers, and the gymnosperms are a monophyletic group, sister to the angiosperms. The gnetifer hypothesis first emerged formally in the mid-twentieth century, when vessel elements in the gnetophytes were interpreted as being derived from tracheids with circular bordered pits, as in conifers.[8] It did not gain strong support, however, until the emergence of molecular data in the late 1990s.[19][25][28][29] Although the most salient morphological evidence still largely supports the anthophyte hypothesis, there are some more obscure morphological commonalities between the gnetophytes and conifers that lend support to the gnetifer hypothesis. These shared traits include: tracheids with scalariform pits with tori interspersed with annular thickenings, absence of scalariform pitting in primary xylem, scale-like and strap-shaped leaves of Ephedra and Welwitschia; and reduced sporophylls.[24][27][30]

angiosperms (flowering plants)






Gnepine hypothesis

The gnepine hypothesis is a modification of the gnetifer hypothesis, and suggests that the gnetophytes belong within the conifers as a sister group to the Pinaceae.[8] According to this hypothesis, the conifers as currently defined are not a monophyletic group, in contrast with molecular findings that support its monophyly.[28] All existing evidence for this hypothesis comes from molecular studies since 1999.[4][5][20][22][24][25][27][30] However, the morphological evidence remains difficult to reconcile with the gnepine hypothesis. If the gnetophytes are nested within conifers, they must have lost several shared derived characters of the conifers (or these characters must have evolved in parallel in the other two conifer lineages): narrowly triangular leaves (gnetophytes have diverse leaf shapes), resin canals, a tiered proembryo, and flat woody ovuliferous cone scales.[24] These kinds of major morphological changes are not without precedent in the Pinaceae, however: the Taxaceae, for example, have lost the classical cone of the conifers in favor of a single-terminal ovule surrounded by a fleshy aril.[20]

angiosperms (flowering plants)





Pinaceae (the pine family)


other conifers

Gnetophyte-sister hypothesis

Some partitions of the genetic data suggest that the gnetophytes are sister to all of the other extant seed plant groups.[6][8][24][27][28] However, there is no morphological evidence nor examples from the fossil record to support the gnetophyte-sister hypotheses.[30]


angiosperms (flowering plants)





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  • 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)
Division (biology)

This article discusses categorisations of organisms. For a different meaning in biology, see cell division.Division is a taxonomic rank in biological classification that is used differently in zoology and in botany.

In botany and mycology, division refers to a rank equivalent to phylum. The use of either term is allowed under the International Code of Botanical Nomenclature, and both are commonly used in scientific literature.

The main Divisions of land plants, in the order in which they probably evolved, are the Marchantiophyta (liverworts), Anthocerotophyta (hornworts), Bryophyta (mosses), Filicophyta (ferns), Sphenophyta (horsetails), Cycadophyta (cycads), Ginkgophyta (ginkgo)s, Pinophyta (conifers), Gnetophyta (gnetophytes), and the Magnoliophyta (Angiosperms, flowering plants). The flowering plants now dominate terrestrial ecosystems, comprising 80% of vascular plant species.

In zoology, the term division is applied to an optional rank subordinate to the infraclass and superordinate to the cohort. A widely used classification (e.g. Carroll 1988) recognises teleost fishes as a Division Teleostei within Class Actinopterygii (the ray-finned fishes). Less commonly (as in Milner 1988), living tetrapods are ranked as Divisions Amphibia and Amniota within the clade of vertebrates with fleshy limbs (Sarcopterygii).


Drewria potomacensis is a Cretaceous megafossil member of the Gnetales, from the Potomac Group, hence its name. It was possibly a shrub. It is the only known species in the genus Drewria.


The Embryophyta, or land plants, are the most familiar group of green plants that form vegetation on earth. Embryophyta is a clade within the Phragmoplastophyta, a larger clade that also includes several green algae groups (including the Charophyceae and Coleochaetales), and within this large clade the embryophytes are sister to the Zygnematophyceae/Mesotaeniaceae and consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, ferns, lycophytes, gymnosperms and flowering plants. The Embryophyta are informally called land plants because they live primarily in terrestrial habitats, while the related green algae are primarily aquatic. Embryophytes are complex multicellular eukaryotes with specialized reproductive organs. The name derives from their innovative characteristic of nurturing the young embryo sporophyte during the early stages of its multicellular development within the tissues of the parent gametophyte. With very few exceptions, embryophytes obtain their energy by photosynthesis, that is by using the energy of sunlight to synthesize their food from carbon dioxide and water.

Ephedra breana

Ephedra breana (frutilla de campo, pingo-pingo) is a species of Ephedra growing from northwest Argentina through to Chile and Bolivia.

Ephedra intermedia

Ephedra intermedia, with the Chinese common name of Zhong Ma Huang, is a species of Ephedra that is native to Siberia, Central Asia, Iran, Afghanistan, Pakistan, the western Himalayas, Tibet, Mongolia, and China.

Ephedra sinica

Ephedra sinica (also known as Chinese ephedra or Ma Huang) is a plant species native to Mongolia, Russia (Buryatiya, Chita, Primorye), and northeastern China (Gansu, Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shanxi).

Ephedra viridis

Ephedra viridis, known by the common names green Mormon tea, green ephedra, and Indian tea, is a species of Ephedra. It is indigenous to the Western United States, where it is a member of varied scrub, woodland, desert, and open habitats. It grows at 900–2,300 metres (3,000–7,500 ft) elevations.


Gnetidae is a subclass of Equisetopsida in the sense used by Mark W. Chase and James L. Reveal in their 2009 article "A phylogenetic classification of the land plants to accompany APG III." This subclass comprises the gnetophytes. The Gnetidae subclass is equivalent to the division Gnetophyta and class Gnetopsida of previous treatments.


Gnetum is a genus of gymnosperms, the sole genus in the family Gnetaceae and order Gnetales. They are tropical evergreen trees, shrubs and lianas. Unlike other gymnosperms, they possess vessel elements in the xylem. Some species have been proposed to have been the first plants to be insect-pollinated as their fossils occur in association with extinct pollinating scorpionflies. Molecular phylogenies based on nuclear and plastid sequences from most of the species indicate hybridization among some of the Southeast Asian species. Fossil-calibrated molecular-clocks suggest that the Gnetum lineages now found in Africa, South America and Southeast Asia are the result of ancient long-distance dispersal across seawater.

Gnetum macrostachyum

Gnetum macrostachyum is a species of vine gymnosperm, native to tropical Asia.


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 Greek composite word γυμνόσπερμος (γυμνός gymnos, "naked" and σπέρμα sperma, "seed"), 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 stands in contrast to 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.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.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).

Roots in some genera have fungal association with roots in the form of mycorrhiza (Pinus), while in some others (Cycas) small specialised roots called coralloid roots are associated with nitrogen-fixing cyanobacteria.


A liana is any of various long-stemmed, woody vines that are rooted in the soil at ground level and use trees, as well as other means of vertical support, to climb up to the canopy to get access to well-lit areas of the forest. Lianas are characteristic of tropical moist deciduous forests (especially seasonal forests), but may be found in temperate rainforests. There are also temperate lianas, for example the members of the Clematis or Vitis (wild grape) genera. Lianas can form bridges amidst the forest canopy, providing arboreal animals with paths across the forest. These bridges can protect weaker trees from strong winds. Lianas compete with forest trees for sunlight, water and nutrients from the soil. Forests without lianas grow 150% more fruit; trees with lianas have twice the probability of dying.The term "liana" is not a taxonomic grouping, but rather a description of the way the plant grows – much like "tree" or "shrub". Lianas may be found in many different plant families. One way of distinguishing lianas from trees and shrubs is based on the stiffness, specifically, the Young's modulus of various parts of the stem. Trees and shrubs have young twigs and smaller branches which are quite flexible and older growth such as trunks and large branches which are stiffer. A liana often has stiff young growths and older, more flexible growth at the base of the stem.

Olaf Hagerup

Olaf Hagerup (29 September 1889 – 2 March 1961) was a Danish botanist. He studied botany at the University of Copenhagen from 1911 under the professors Eugenius Warming, Christen C. Raunkiær, L. Kolderup Rosenvinge og W. Johannsen. He took his Ph.D. from the same university in 1930. From 1934 to 1960, he was superintendent at the Botanical Museum of the University of Copenhagen.

Hagerup’s scientific works concern evolution, polyploidy and pollination, among other things. He showed that the tetraploid Empetrum hermaphroditum is a separate species from the diploid Empetrum nigrum. He thereby initiated the use of chromosome numbers in systematic botany, a field later known as cytotaxonomy. He put forward the hypothesis that the ploidy level is an important factor in the distribution and ecology of plant species. In contrast, another of his scientific ideas has been disproven by later modern research – the idea of a direct ancestry of the centrosperms (approximately equal to Caryophyllales) from the gymnospermous Gnetophyta and, hence, two separate evolutionary lineages within the flowering plants. Many of Hagerup’s studies were concerned with plant species of the Ericaceae, Empetraceae and related families, or ’’Bicornes’’ as they were known in the Wettstein system.

The cranberry Oxycoccus hagerupii (Ericaceae) was named to his honour by Á. & D. Löve (later transferred to Vaccinium by Hannu Ahokas as Vaccinium hagerupii.

Outline of botany

The following outline is provided as an overview of and topical guide to botany:

Botany – biological discipline which involves the study of plants.

Sherwin Carlquist

Sherwin John Carlquist FMLS (born July 7, 1930) is an American botanist and photographer. He received his undergraduate degree from the University of California, Berkeley in 1952 and a Ph.D. in botany in 1956, also at Berkeley. Carlquist did a postdoctoral study at Harvard University from 1955 to 1956. After his postdoctoral studies, he began his teaching career at the Claremont Graduate School. In 1977 he also began teaching at Pomona College and continued teaching at both institutions until 1992. From 1984 to 1992 Carlquist was the resident Plant Anatomist at Rancho Santa Ana Botanic Garden. His last post was as an adjunct professor at University of California at Santa Barbara from 1993 to 1998.Carlquist studied wood anatomy of the Gnetophyta and was an author of many plant taxa, including species of the carnivorous plant genus Drosera, the Western Australian genus Stylidium, and the odd Australian genus Alexgeorgea whose female flowers are almost entirely underground.The California plant genus Carlquistia is named for Carlquist.


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, gameein, "to marry". These terms distinguished those plants with hidden sexual organs (cryptogamae) from those with visible sexual organs (phanerogamae).


A strobilus (plural: strobili) is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but many botanists restrict the use of the term cone to the woody seed strobili of conifers. Strobili are characterized by a central axis (anatomically a stem) surrounded by spirally arranged or decussate structures that may be modified leaves or modified stems.

Leaves that bear sporangia are called sporophylls, while sporangia-bearing stems are called sporangiophores.

Vascular plant

Vascular plants (from Latin vasculum: duct), also known as tracheophytes (from the equivalent Greek term trachea), form a large group of plants (c. 308,312 accepted known species) that are defined as those land plants that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified tissue (the phloem) to conduct products of photosynthesis. Vascular plants include the clubmosses, horsetails, ferns, gymnosperms (including conifers) and angiosperms (flowering plants). Scientific names for the group include Tracheophyta, Tracheobionta and Equisetopsida sensu lato. The term higher plants should be avoided as a synonym for vascular plants as it is a remnant of the abandoned concept of the great chain of being.


Welwitschia is a monotypic gymnosperm genus, comprising solely the distinctive Welwitschia mirabilis. The plant is commonly known simply as welwitschia in English, but the name tree tumbo is also used. It is called kharos or khurub in Nama, tweeblaarkanniedood in Afrikaans, nyanka in Damara, and onyanga in Herero. Welwitschia is the only living genus of the family Welwitschiaceae and order Welwitschiales, in the division Gnetophyta. Informal sources commonly refer to the plant as a "living fossil". Welwitschia mirabilis is endemic to the Namib desert within Namibia and Angola.

(red algae)
(green algae,
& land plants)
Extant Life phyla/divisions by domain

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