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Gigantopterids (Gigantopteridales) is an extinct, possibly
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
group of plants known from the
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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. Gigantopterids were among the most advanced
land plant 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 sist ...
s of the
Paleozoic Era 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 ''z ...
and disappeared around the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
around 252 million years ago. Though some lineages of these plants managed to persist initially, they either disappeared entirely or adapted radically,
evolving Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
into undetermined descendants, as surviving life prospered again in much-altered ecosystems. One hypothesis proposes that at least some "gigantopterids" became the ancestors of angiosperms and/or
Bennettitales Bennettitales (also known as cycadeoids) is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales are among the most common Mesozoic seed plants ...
and/or Caytoniales.Miller (2007) Gigantopterid fossils were documented as early as 1883, but only investigated more thoroughly in the early 20th century. Some of their most significant evidence was initially found in
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
, but they might have been present worldwide. Another key region for gigantopterid fossils is in China, and the consolidation of all major
continent A continent is any of several large landmasses. Generally identified by convention rather than any strict criteria, up to seven geographical regions In geography, regions, otherwise referred to as zones, lands or territories, are areas t ...
s into
Pangea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
would have allowed for easy global dispersal. They were among the most striking and important plants of the Cathaysian flora of Sino- Malaya, also called ''Gigantopteris'' flora to reflect this.Wang (1999)


Description

They bore many of the traits of
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 th ...
s, but are not known to have flowered themselves. Gigantopterid plants had simple, bilaterally symmetrical leaf structures, woody stems and spines. They grew new parts by producing shoots, like flowering plants. Judging from the fossil remains, they were probably robust plants with
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
s that resembled
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
fronds when young. When mature they were more like flowering plant leaves with
reticulate venation A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
arranged in a frond. ''Gigantopteris nicotianaefolia'' for example is named thus because each of its leaflets resembles a
tobacco Tobacco is the common name of several plants in the genus '' Nicotiana'' of the family Solanaceae, and the general term for any product prepared from the cured leaves of these plants. More than 70 species of tobacco are known, but the ...
leaf in shape. They grew at least over 20 cm (around 10 in), probably over 50 cm (20 in) tall, depending on whether it grew as a scrambling vine (the initial assumption) or erect (nowadays considered more likely). Some apparently preferred wetlands, while others throve in rather
arid A region is arid when it severely lacks available water, to the extent of hindering or preventing the growth and development of plant and animal life. Regions with arid climates tend to lack vegetation and are called xeric or desertic. Most ...
habitats.Wang (1999), Glasspool ''et al.'' (2004) The
vascular tissue Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. The ...
contains
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 and in at least some
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
resembles that of the
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 ...
. They had a cuticle similar to that of other seed plants. Some male
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cyc ...
(''Gigantotheca'') and
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 angiospe ...
s (''Gigantonomia'', ''Cornucarpus'') have been tentatively referred to this group. While the sporangia are certainly not flowers, they differ from other sporangia of Permian plants. Chemical analysis of
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
remains indicates that gigantopterids produced
oleanane Oleanane is a natural triterpenoid. It is commonly found in woody angiosperms and as a result is often used as an indicator of these plants in the fossil record. It is a member of the oleanoid series, which consists of pentacyclic triterpenoids ( ...
s, chemical compounds otherwise known to be used by flowering plants, for which they function as a suppressant of insect pests. They are apparently also found in certain modern ferns however.


Systematics

It is probable that the gigantopterids are a non- monophyletic
form taxon Form classification is the classification of organisms based on their morphology, which does not necessarily reflect their biological relationships. Form classification, generally restricted to palaeontology, reflects uncertainty; the goal of s ...
. Plant life from their era is very difficult to document and organize. They have been variously allied with the "
seed fern A seed is an Plant embryogenesis, embryonic plant enclosed in a testa (botany), 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, includ ...
s" (another paraphyletic group of early seed plants) Peltaspermales and Callistophytales, ''
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 bee ...
'', and the
Magnoliophyta 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 ...
. All that can be said at present is that they were
spermatophyte A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
s. It is also highly unlikely that they were close relatives of
acrogymnosperm 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, γυμνόσ ...
s,
cycads 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 ...
or ginkgos, because these lineages were already established and distinct in the Late Permian. Vegetative leaves of ''Emplectopteris'' were at one time included in this group. However, they had ovules attached to the underside of the fronds and are now placed in their own family ( Emplectopteridaceae) within the Callistophytales. Some prefer to refer to the presumed "core"
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
of this group as ''Gigantonomiales''. However, many experts remain skeptical of the new name, because they consider such a move premature until more is known about the interrelationships of the genera of "gigantopterids" and the relationships of these to other plants. Several genera have been named. Those then known were assigned to a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
''Gigantopteridaceae'' and numerous subfamilies by
Gen-ichi Koidzumi was a Japanese botanist, author of several papers and monographs on phytogeography including work on roses and Amygdaloideae (Rosaceae), maples (Aceraceae), mulberries (the genus ''Morus''), and many other plants. His name is sometimes translit ...
. It appears as if at least some of his divisions ought to prove useful when a more definite
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
, systematic and taxonomic arrangement for this group is eventually found. On the other hand, the Gigantopteridaceae would with near certainty have to be more narrowly circumscribed even if the Gigantopteridales are by and large verified as a clade; they might even become limited to the two or so genera now placed in the Gigantopteridieae.Glasspool ''et al.'' (2004)


Proposed subfamilies and genera

Cathaysiopteridieae - may be synonym of Palaeogoniopteridieae * ''Cathaysiopteris'' * ''Cathaysiopteridium'' Gigantopteridieae (including Cardioglossieae) * ''Gigantonoclea'' - includes ''Cardioglossum'' and apparently ''Progigantopteris'' * ''Gigantopteris'' Gothanopteridieae - may be synonym of Cathaysiopteridieae/Palaeogoniopteridieae * ''Gothanopteris'' Palaeogoniopteridieae * ''Gigantopteridium'' * ''Palaeogoniopteris'' * ''Zeilleropteris'' Unresolved * ''Delnortea'' * ''Emplectopteridium'' * ''Emplectopteris'' * ''Neogigantopteridium'' - probably Cathaysiopteridieae/Palaeogoniopteridieae * ''Vasovinea''


See also

Other Permian-Triassic seed plants of unclear relationships are: *
Glossopteridales Glossopteridales is an extinct order of plants belonging to Pteridospermatophyta, or seed ferns, also known as Arberiales and Ottokariales. They arose at the beginning of the Permian () on the southern continent of Gondwana, but became extinct a ...
* Lyginopteridopsida * Pentoxyales


Footnotes


References

* (1982): Evolution and phylogeny of vascular plants based on the principles of growth retardation: Part 5. Origin of angiosperms inferred from the evolution of leaf forms. Bulletin of the National Science Museum, Tokyo, Series C (Geology) 8: 43-58. * (2004): Defining the gigantopterid concept: a reinvestigation of ''Gigantopteris (Megalopteris) nicotianaefolia'' Schenck and its taxonomic implications. '' Palaeontology'' 47(6): 1339-1361. (HTML abstract) * (1936): On the Gigantopteris flora. ''Acta Phytotaxonomica et Geobotanica'' 5: 130–144. * (1983): Fructifications of gigantopterids from South China. ''Palaeontographica Abteilung B'' 185: 11-26. * (1987): ''Fundamentals of palaeobotany''. Chapman and Hall, London and New York. * (2007)
Paleobotany of Angiosperm Origins
Version of 2007-DEC-10. Retrieved 2007-DEC-12. * (1999): ''Gigantonoclea'': an enigmatic Permian plant from North China. ''Palaeontology'' 42(2): 329–373. (HTML abstract)


External links


Image
from Li & Yao (1983), showing fossils and life reconstructions of ''Gigantonomia'' (denoted F) and ''Gigantotheca'' (denoted V). Retrieved 2007-DEC-12.
Photo of ''Delnortea abbottiae'' leaflet
specimen USNM 387473. Retrieved 2007-DEC-12.
Reconstruction
of ''Gigantopteris'' leaves. Retrieved 2007-DEC-12. * Wikiadugi
Version of this article
in Cherokee language. Retrieved 2007-DEC-12. {{Taxonbar, from=Q5560234 Permian plants Prehistoric plant families Cisuralian life Cisuralian first appearances Cisuralian extinctions Pteridospermatophyta