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CLASSIFICATION
2 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]
3 Classication
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.
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
3.3
Gnepine hypothesis
REFERENCES
for resolving the seed plant tree of life. International Journal of Plant Science. 168 (10): 125135.
doi:10.3732/ajb.91.10.1599.
Other Sources:
Giord, Ernest M., Adriance S. Foster. 1989. Morphology and Evolution of Vascular Plants. Third
edition. WH Freeman and Company, New York.
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)
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