Corystosperms are a group of extinct (often referred to as "seed ferns") belonging to the family Corystospermaceae (also called Umkomasiaceae) assigned to the order Corystospermales[
] Corystosperms are typified by a group of plants that bore forked Dicroidium leaves, Umkomasia cupulate ovulate structures and Pteruchus pollen organs, which grew as trees that were widespread over Gondwana during the Middle and Late Triassic. Other fossil Mesozoic seed plants with similar leaf and/or reproductive structures have also sometimes been included within the "corystosperm" concept sensu lato, such as the "doyleoids" from the Early Cretaceous of North America and Asia.
Description
Classic Umkomasiaceae have helmet-like cupules around
ovules born in complex large branching structures (
Umkomasia). The pollen organ (
Pteruchus) has numerous cigar-shaped pollen sacs hanging from epaulette-like blades, again in complex branching structures.
The architecture and size of Dicroidium leaves is highly variable, going from simple to tripinnate, to needle-like to lobed or dissected leaflets, however, all leaves are united by being bifurcated at the base.
The Dicroidium plant (which bore Dicroidium leaves, Umkomasia seed bearing structures and Pteruchus pollen organs) is thought to have grown as large , with trunks at least tall and over wide.
Evolutionary history
The oldest fossils of corystosperms, belonging to
Dicroidium, Pteruchus and
Umkomasia date to the Late Permian in the low-latitudes of eastern
Gondwana, including the Umm Irna Formation of Jordan, as well as the Indian subcontinent,
though possible pollen belonging to the group is also known from the Late Permian of South Africa.
During the Middle-Late Triassic,
Dicroidium bearing corystosperms were widespread and dominant trees over temperate areas of southern Gondwana (including the Indian subcontinent, South America, Southern Africa, Antarctica, Australia and New Zealand) including wetland and forest environments, where they formed part of the canopy vegetation.
bearing corystosperm ecosystems collapsed as a result of the end-Triassic extinction event. Late surviving
Dicroidium-bearing corystosperms are known from the
Early Jurassic (
Sinemurian) of East Antarctica.
During the Jurassic, the possible corystosperms
Pachypteris and
Komlopteris were widespread over both hemispheres,
with
Komlopteris surviving in the Southern Hemisphere through the Cretaceous and into the early
Eocene in Tasmania, making it the last surviving "seed fern" in the fossil record.
A group of plants with corystosperm-like reproductive structures, either considered corystosperms or as belonging to the separate order Doyleales, were present in North America and Asia during the Early Cretaceous,
with records of the group possibly extending back as far as the Early Jurassic.
List of genera
Relationships to other seed plants
The relationships of corystosperms to other seed plants are contentious.
It has been suggested that some "
Peltaspermales" may be closely related to corystosperms.
A 2016 paper suggested that they may be closely related to
Ginkgoales,
while a 2021 study suggested that based on the structure of their ovulate organs, corystosperms might form part of the "
Anthophyta" clade more closely related to
than to living
, alongside the
Petriellales,
Caytoniales, and
Glossopteridales.
Nomenclature
Corystospermaceae is a family name which was not derived from a specific genus, an arrangement which is discouraged by the ICN.
In light of this issue, Petriella (1981) proposed two solutions: either retain Corystospermaceae as a nomen conservandum, or replace it with a new name, Umkomasiaceae.
Both names have been used frequently since then. Petriella additionally named a new order to encompass the family, Corystospermales.
Doweld (2001) preferred a different name for the order, Umkomasiales. He also added a second family, Angaropeltidaceae (previously known as Cardiolepidaceae).
Later sources generally classify Angaropeltidaceae (or its updated spelling,
Angaropeltaceae)
within Peltaspermales instead.
Gallery
File:Umkomasia macleani.jpg| Umkomasia macleanii ovulate structure from Umkomaas
File:Pteruchus africanus.jpg| Pteruchus africanus pollen structure from Umkomaas
File:Dicroidium odontopteroides.jpg| Dicroidium odontopteroides leaf from Birds River
See also
External links
