Gnetophyta

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 Tertiary,[2] only three families, each containing a
single genus, are still alive today. The primary dierence 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 owering plants. Because of this, gnetophytes were
once thought to be the closest gymnosperm relatives to Ephedra distachya (male cones)
owering plants, but more recent molecular studies have
largely disproven this hypothesis.
Though it is clear they are all closely related, the exact evolutionary inter-relationships between gnetophytes
are unclear. Some classications hold that all three genera should be placed in a single order (Gnetales), while
other classications say they should be distributed among
three separate orders, each containing a single family
and genus. Most morphological and molecular studies conrm that the genera Gnetum and Welwitschia diverged from each other more recently than they did from
Ephedra.[3][4][5][6][7]

Ephedra distachya (female plant in bloom)

Welwitschia mirabilis bearing male cones

Ecology and morphology

Gnetum gnemon male strobili

Unlike most biological groupings, it is dicult to nd

many common characteristics between all of the mem- istics most commonly used are the presence of envelopbers of the gnetophytes.[2] The two common character- ing bracts around both the ovules and microsporangia as
1

CLASSIFICATION

from the base throughout the plants life. Ephedra

species, known as jointrs 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.

2 Fossil Gnetophyta

Gnetum gnemon female strobilus

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]

3 Classication

Female Ephedra californica cone

With just three well-dened genera within an entire division, there still is understandable diculty in establishing
an unambiguous interrelationship among them; in earlier times matters were even more dicult and we nd
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 conicted 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.

well as a micropylar projection of the outer membrane

of the ovule that produces a pollination droplet,[8] though
these are highly specic 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 Recent research by Lee EK, Cibrian-Jaramillo A, et al.
Namibia and Angola, is a ground-hugging species with (2011) suggests that the Gnetophyta are a sister group to
only two large strap-like leaves that grow continuously the rest of the gymnosperms,[14] contradicting the antho-

3.4

Gnetophyte-sister hypothesis

phyte hypothesis, which held that gnetophytes were sister lar studies within the last decade.[4][5][19][21][23][24][26][29]
to the owering plants.
However, the morphological evidence remains dicult to
reconcile with the gnepine hypothesis. If the gnetophytes
are nested within conifers, they must have lost several
3.1 Anthophyte hypothesis
shared derived characters of the conifers (or these characters must have evolved in parallel in the other conifer
From the early twentieth century, the anthophyte hy- lineages): narrowly triangular leaves (gnetophytes have
pothesis was the prevailing explanation for seed plant diverse leaf shapes), resin canals, a tiered proembryo, and
evolution, based on shared morphological characters at woody ovuliferous cone scales.[23] These kinds of mabetween the gnetophytes and angiosperms. In this jor morphological changes are not without precedent in
hypothesis, the gnetophytes, along with the extinct the Pinaceae, however: the Taxaceae, for example, have
order Bennettitales, are sister to the angiosperms, lost the classical cone of the conifers in favor of a singleforming the anthophytes.[8] Some morphological terminal ovule surrounded by a eshy aril.[19]
characters that were suggested to unite the anthophytes include vessels in wood, net-veined leaves (in
Gnetum only), lignin chemistry, the layering of cells 3.4 Gnetophyte-sister hypothesis
in the apical meristem, pollen and megaspore features
(including thin megaspore wall), short cambial ini- Some partitions of the genetic data suggest that the gnetials, and lignin syringal groups.[8][15][16][17] However, tophytes are sister to all of the other extant seed plant
most genetic studies have rejected the anthophyte groups.[6][8][23][26][27] However, there is no morphological
hypothesis.[4][18][19][20][21][22][23][24][25][26] Several of evidence nor examples from the fossil record to support
these studies have suggested that the gnetophytes and the gnetophyte-sister hypotheses.[29]
angiosperms have independently derived characters, including ower-like reproductive structures and tracheid
vessel elements, that appear shared but are actually the 4 References
result of parallel evolution.[4][8][19]

3.2

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 rst 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 strapshaped leaves of Ephedra and Welwitschia; and reduced
sporophylls.[23][26][29]

3.3

Gnepine hypothesis

The gnepine hypothesis is a modication 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
dened are not a monophyletic group, in contrast with
molecular ndings that support its monophyly.[27] All existing evidence for this hypothesis comes from molecu-

[1] Morphology and anities of an Early Cretaceous

Ephedra.
[2] Arber, E.A.N.; Parkin, J. (1908). Studies on the evolution of the angiosperms: the relationship of the angiosperms to the Gnetales. Annals of Botany. 22: 489
515.
[3] Peter R. Crane; Patrick Herendeen; Else Marie Friis
(2004).
Fossils and plant phylogeny.
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doi:10.3732/ajb.91.10.1683. PMID 21652317.
[4] Bowe, L.M.; Coat, G.; dePamphilis, C.W. (2000).
Phylogeny of seed plants based on all three genomic
compartments: Extant gymnosperms are monophyletic
and Gnetales closest relatives are conifers. Proceedings
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doi:10.1073/pnas.97.8.4092. PMC 18159 . PMID
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[5] Gugerli, F.; Sperisen, C.; Buchler, U.; Brunner, L.; Brodbeck, S.; Palmer, J.D.; Qiu, Y.L. (2001). The evolutionary split of Pinaceae from other conifers: evidence
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[6] Rai, H.S.; Reeves, P.A.; Peakall, R.; Olmstead, R.G.;
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[7] Ickert-Bond, S. M., C. Rydin, and S. S. Renner (2009).
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456. doi:10.1111/j.1759-6831.2009.00053.x.
[8] Judd, W.S.; Campbell, C.S.; Kellogg, E.A.; Stevens, P.F.;
and Donoghue, M.J. (2008) Plant Systematics: A Phylogenetics Approach. 3rd ed. Sunderland, Massachusetts,
USA: Sinauer Associates, Inc.
[9] Zi-Qiang Wang (2004). A New Permian Gnetalean
Cone as Fossil Evidence for Supporting Current Molecular Phylogeny. Annals of Botany. 94 (2): 281288.
doi:10.1093/aob/mch138. PMID 15229124.
[10] Catarina Rydin; Kaj Raunsgaard Pedersen; Peter R.
Crane; Else Marie Friis (2006). Former Diversity of
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[11] Pearson, H. H. W. Gnetales. Cambridge University Press
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[12] Lawrence, George Hill Mathewson. Taxonomy of vascular plants. Macmillan, 1951
[13] Foster, Adriance S., Giord, Ernest M. Jr. Comparative
Morphology of Vascular Plants Freeman 1974. ISBN 07167-0712-8
[14] Lee EK, Cibrian-Jaramillo A, Kolokotronis SO, Katari
MS, Stamatakis A, et al. (2011). A Functional Phylogenomic View of the Seed Plants. PLoS Genet. 7 (12):
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3240601 . PMID 22194700.
[15] Donoghue, M.J.; Doyle, J.A. (2000). Seed plant phylogeny: demise of the anthophyte hypothesis?". Current Biology. 10 (3): R106R109. doi:10.1016/S09609822(00)00304-3. PMID 10679315.
[16] Loconte, H.; Stevenson, D.W. (1990). Cladistics of
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[17] Nixon, K.C.; Crepet, W.L.; Stevenson, D.; Friis, E.M.
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[18] Chaw, S.M.; Aharkikh, A.; Sung, H.M.; Lau,
T.C.; Li, W.H. (1997).
Molecular phylogeny
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[22] Hansen, A.; Hansmann, S.; Samigullin, T.; Antonov,
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[23] Magallon, S.; Sanderson, M.J. (2002). Relationships
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commons/0/0c/Female_Ephedra_californica_cone_-_journal.[Link] License: CC BY 2.5 Contributors: Yang Y,
Wang Q (2013) The Earliest Fleshy Cone of Ephedra from the Early Cretaceous Yixian Formation of Northeast China. PLoS ONE 8(1):
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