Dicroidium Zuberi 2
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''Dicroidium'' is an extinct genus of fork-leaved
seed plant A seed plant or spermatophyte (; New Latin ''spermat-'' and Greek ' (phytón), plant), also known as a phanerogam (taxon Phanerogamae) or a phaenogam (taxon Phaenogamae), is any plant that produces seeds. It is a category of embryophyte (i.e. la ...
s. It is the archetypal genus of the corystosperms, an extinct group of seed plants, often called "
seed fern In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be sown, which may include seed and husk or tuber. Seeds are the ...
s", assigned to the order Corystospermales or Umkomasiales. Species of ''Dicroidium,'' which grew as large trees, were widely distributed and dominant over
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
during the
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
(). Their fossils are known from
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, the
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,
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,
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
,
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,
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, the
Indian subcontinent The Indian subcontinent is a physiographic region of Asia below the Himalayas which projects into the Indian Ocean between the Bay of Bengal to the east and the Arabian Sea to the west. It is now divided between Bangladesh, India, and Pakista ...
and
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
.


Description

Within the form genus classification system used in paleobotany, the genus ''Dicroidium'' refers specifically to the leaves. Some authors have suggested dividing ''Dicroidium'' up into several genera, including ''Dicroidiopsis, Diplasiophyllum, Zuberia'', ''Xylopteris'', ''Johnstonia'' and ''Tetraptilon,'' but this is rejected by other authors. The leaves of ''Dicroidium'' bifurcate (fork) at their base, which is characteristic of all species. The leaves are highly variable in size and morphology, ranging from simple to
tripinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...
, with the individual leaflets having varying morphologies, including dissected,
lobed The following terms are used to describe leaf plant morphology, morphology in the description and taxonomy (biology), taxonomy of plants. Leaves may be simple (that is, the leaf blade or 'lamina' is undivided) or compound (that is, the leaf blade ...
, needle-like and
entire Entire may refer to: * Entire function, a function that is holomorphic on the whole complex plane * Entire (animal), an indication that an animal is not neutered * Entire (botany) This glossary of botanical terms is a list of definitions o ...
. Some leaf specimens have more than one type of leaflet morphology, which may have been the result of hybridisation between different species. The
venation Venation may refer to: * Venation (botany), the arrangement of veins in leaves * Wing venation, the arrangement of veins in insect wings See also * * Vernation Vernation or leafing is the formation of new leaves or fronds. In plant anatomy, it ...
of the leaves is also highly variable, encompassing taeniopteroid, odontopteroid, alethopteroid and simple morphologies.


Whole plant

''Dicroidium'' plants grew as medium-large sized trees, with some preserved trunk sections tall and over wide, with the wood assigned to the genera ''Kykloxylon'' and ''Rhexoxylon.'' The
ovulate Ovulation is an important part of the menstrual cycle in female vertebrates where the egg cells are released from the ovaries as part of the ovarian cycle. In female humans ovulation typically occurs near the midpoint in the menstrual cycle and ...
reproductive structures are usually assigned to the genus ''
Umkomasia ''Umkomasia'' is a genus of seed bearing organs produced by corystosperm seed ferns, first based on fossils collected by Hamshaw Thomas from the Burnera Waterfall locality near the Umkomaas River of South Africa. He recognized on the basis of ...
,'' while the pollen-producing organs are assigned to the genus ''
Pteruchus ''Pteruchus'' is a form genus for pollen organs of the seed fern (Pteridospermatophyta family Umkomasiaceae. It was first described by Hamshaw Thomas from the Umkomaas locality of South Africa. It is associated with the seed bearing organs ''Umk ...
,'' with pollen attributable to the genus '' Falcisporites.'' These structures are almost never found in organic connection, and their placement as part of the same plant primarily relies on their repeated co-occurrence with each other, and the similarities in the morphology of their
cuticles A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
. The leaves grew on short protuberances attached to the stem in a similar way to living ''
Ginkgo biloba ''Ginkgo biloba'', commonly known as ginkgo or gingko ( ), also known as the maidenhair tree, is a species of gymnosperm tree native to East Asia. It is the last living species in the order Ginkgoales, which first appeared over 290 million year ...
''. The leaves of ''Dicroidium'' are suggested to have been
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
, and shed along with the reproductive organs during the winter. The seeds and pollen are suggested to have been wind dispersed. At least some of the Antarctic species are suggested to have engaged in prolific bark shedding. Possible whole plant associations include: * ''
Dicroidium odontopteroides ''Dicroidium odontopteroides'' was a common and widespread species of '' Dicroidium'' known from South Africa, Australia, New Zealand, South America and Antarctica Antarctica () is Earth's southernmost and least-populated continent. S ...
'' may have been produced by the same plant as '' Umkomasia macleanii'' (ovulate structures) and ''
Pteruchus africanus ''Pteruchus africanus'' is a pollen organ of a seed fern (Pteridospermatophyta). It was first described by Hamshaw Thomas from the Umkomaas locality of South Africa. Description The pollen organs ''Pteruchus africanus'' differ from other spe ...
'' (pollen organs). * ''
Dicroidium zuberi ''Dicroidium zuberi'' is a large bipinnate species of the seed fern '' Dicroidium'' with a forked rachis. The leaves are affiliated with ''Umkomasia feistmantellii'' megasporophylls and ''Petruchus'' ''barrealensis'' microsporophylls. ''D. zu ...
'' may have been produced by the same plant as ''
Umkomasia feistmantelii ''Umkomasia feistmantelii'' is an unusually large species of ''Umkomasia'' from the Early Triassic of New South Wales, Australia. Description ''Umkomasia feistmantelii'' is found both with cupules enclosing the large seeds and with cupules ...
'' (ovulate structures) and ''
Pteruchus barrealensis ''Pteruchus barrealensis'' is an unusually large species of ''Pteruchus'' with very elongate polleniferous heads from Early Triassic of Australia and Argentina. Description ''Pteruchus barrealensis'' is one of the geologically earliest species ...
'' (pollen organs)


Evolution

The earliest ''Dicroidium'' species are known from the
Late Permian Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
aged
Umm Irna Formation The Umm Irna Formation is a geological formation in Jordan. It is found in several outcrops in Jordan in the area around the eastern shore of the Dead Sea. It is Late Permian (likely Changhsingian) in age, and is the oldest unit in the succession, ...
of
Jordan Jordan, officially the Hashemite Kingdom of Jordan, is a country in the Southern Levant region of West Asia. Jordan is bordered by Syria to the north, Iraq to the east, Saudi Arabia to the south, and Israel and the occupied Palestinian ter ...
, which inhabited equatorial humid tropical environments, as well as equivalently aged more southerly subtropical deposits on the Indian subcontinent. Following the end-Permian mass extinction, ''Dicroidium'' expanded its range southwards across
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
, including South Africa, Antarctica, Australia, New Zealand and South America, with a record also reported from the Middle Triassic of China. Later Triassic ''Dicroidium''-bearing plants were dominant large
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an a ...
forming trees in temperate wetland and forested habitats at mid-high latitudes, extending to the South Pole. ''Dicroidium''-dominated ecosystems in Gondwana collapsed during the end-Triassic mass extinction, with ''Dicroidium'' surviving in parts of East Antarctica into the Early Jurassic (
Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...
).


References

* Bomfleur, B. and Kerp, H. (2010). ''Dicroidium diversity in the Upper Triassic of north Victoria Land, East Antarctica.'' {{Taxonbar, from=Q5273916 Triassic plants Fossil taxa described in 1912 Pteridospermatophyta