Gnetophyta

Gnetophyta is a division of plants, grouped within the gymnosperms (which also

Gnetophyta
includes conifers, cycads, and ginkgos), that consists of some 70 species across
the three relict genera: Gnetum (family Gnetaceae), Welwitschia (family Temporal range: Cretaceous–recent
Welwitschiaceae), and Ephedra (family Ephedraceae). Fossilized pollen attributed PreЄ Є OS D C P T J K Pg N
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 inflowering plants. Because of this, gnetophytes were once
thought to be the closest gymnosperm relatives to flowering plants, but more
recent molecular studies have largely disproven this hypothesis.

Though it is clear they are all closely related, the exact evolutionary inter- Welwitschia mirabilis female plant with
relationships between gnetophytes are unclear. Some classifications hold that all cones
three genera should be placed in a single order (Gnetales), while other
Scientific classification
classifications say they should be distributed among three separate orders, each
containing a single family and genus. Most morphological and molecular studies Kingdom: Plantae
confirm that the genera Gnetum and Welwitschia diverged from each other more Division: Gnetophyta
recently than they did fromEphedra.[3][4][5][6][7]
Class: Gnetopsida
Families & Genera
Gnetaceae
Contents Gnetum
Ecology and morphology Welwitschiaceae

Fossil Gnetophyta Welwitschia

Ephedraceae
Classification
Anthophyte hypothesis Ephedra
Gnetifer hypothesis
Gnepine hypothesis
Gnetophyte-sister hypothesis
References

Ecology and morphology Distribution, separated by genus:

Green – Welwitschia
Unlike most biological groupings, it is difficult to find many common
Blue – Gnetum
characteristics between all of the members of the gnetophytes.[2] The two
common characteristics most commonly used are the presence of enveloping
Red – Ephedra
bracts around both the ovules and microsporangia as well as a micropylar Purple – Gnetum and Ephedra
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.

Welwitschia mirabilis bearing male

Fossil Gnetophyta cones

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 theJurassic 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
[10]
found that can clearly be assigned to the gnetophytes.

Classification
With just three well-defined genera within an entire division, there still is
Ephedra distachya (male cones)
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
Ephedra distachya (female plant in
(including gnetophytes) comprise a monophyletic group or a paraphyletic one that
bloom)
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 Gnetum gnemon male strobili
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 have rejected the anthophyte
Gnetum gnemon female strobilus
hypothesis.[4][18][19][20][21][22][23][24][25][26] Several of these studies have
suggested that the gnetophytes and angiosperms have independently derived
characters, including flower-like reproductive structures and tracheid vessel
[4][8][19]
elements, that appear shared but are actually the result of parallel evolution.

Ginkgo

cycads

conifers

angiosperms (flowering plants)

anthophytes
gnetophytes

Gnetifer hypothesis
In the gnetifer hypothesis, the gnetophytes are sister to the conifers, and the Female Ephedra californica cone
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.[18][24][27][28] 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.[23][26][29]

angiosperms (flowering plants)

cycads

Ginkgo
gymnosperms
conifers

gnetophytes

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.[27] All existing evidence for this hypothesis comes from molecular
studies since 1999.[4][5][19][21][23][24][26][29] 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.[23] 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
[19]
favor of a single-terminal ovule surrounded by a fleshy aril.

angiosperms (flowering plants)

cycad

Ginkgo

gymnosperms
Pinaceae (the pine family)

conifers gnetophytes

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][23][26][27]
etophyte-sister hypotheses.[29]
However, there is no morphological evidence nor examples from the fossil record to support the gn

gnetophytes

angiosperms (flowering plants)

cycads

Ginkgo

conifers

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 Hilton, Jason, and Richard M. Bateman. 2006. Pteridosperms are the backbone of seed-plant phylogeny
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