Gigantopterids †
Order Gigantopteridales
The Gigantopteridales are various taxa of advanced seed ferns from the Permian and Triassic of Eastern Asia, Greenland, and North America. They are known from compression and impression fossils with a diverse array of leaf forms that are angiosperm-like in morphology and with a chemical signature of oleanane. In addition, their stems are woody with vessels which are only found in angiosperms and gnetophytes. Because reproductive structures are poorly known, the evolutionary affinities of this group is poorly known, with the Callistophytales, Caytoniales, Ginkgoales, and others being suggested as related to the Giganopterids (Meyen 1984).
Ecology and Form
Possible vine or upright plant
Grew in wetlands as well as arid conditions
Known mostly from compression and impression fossils
They were among the most striking and important plants of the Cathaysian flora of Sino-Malaya, also called Gigantopteris flora (Wang 1999)
Stem
Woody stems with spines
Wood contains vessel elements similar to gnetophytes and angiosperms
Aculeovinea yunguiensis
These stems are woody and vessel-less with a sparganum cortex and prickles
Stems were probably connected to Gigantopteris guizhouensis leaves
Vasovinea tianii
A permineralized eustelic stem for the Late Permian of China
Eustelic with 5-9 wedge-shaped mesarch segments, each with wood with vessels
Prickles, trichomes, and compound hooks are found along stems suggesting a climbing habit
Classification
└Gigantopteridales †
Geologic Age
Permian of China, Japan, Korea, southern and western Asia, Greenland, and North America
Above: The world map, around 300 million years ago, showing Cathaysia (in pink) which was dominated by Giganopterids during the Permian
Leaves
Large bilaterally symmetrical leaves in mature plants (Giganopteris, Gigantopteridium, Cathaysiopteris, Zeilleropteris, Gigantonoclea)
Dichotomous venation known is some;
Reticulate venation found in others with anastomosing higher-order veins
Some leaves are compound fronds; mature leaves may have been simple
Oleanane found in leaves (similar to angiosperms)
Earliest evidence of foliar nyctinasty has been documented in gigantopterid leaves (Feng et al. 2023)
Symmetrical insect damage indicates folding of the leaves, probably during the night
Gigantopteris nicotianaefolia (type species)
Leaves from Permian of South China (Yan 1983)
Leaves up to 50 cm long with an entire margin (Glasspoll et al. 2004)
Pinnate venation with 4 orders of veins which give rise to "large, polygonal meshes with finer veinlets anastomosing to form meshes and blind endings" (Taylor et al. 2009)
Gigantopteris guizhouensis
Permineralized simple leaves from the Late Permian of southeast China (Li & Tian 1990)
Stomata are paracytic with uniseriate trichomes on the abaxial surface
Many secretory cavities are within the mesophyll of the leaf
Prickles are found along the abaxial midrib
Gigantopteridium americana
Leaves from Permian of North America (White 1912)
Gigantopteridium marginervum
Leaves from Middle Permian of China (Yao & Liu 2004)
Amphistomic leaves (both sides of leaf) with cyclocytic stomata
Subsidiary cells on abaxial (lower) surface have papillae
Subsidiary cells on adaxial (upper) surface lack papillae and are heavily cutinized
Cathaysiopteris
Leaves from the Permian of Asia
Forked and pinnately-compound leaves up to 20 cm in length with a sinuous margin and pinnate veins with three orders of venation
The tertiary veins are closely-spaced and the meshes are long and narrow (Mamay et al. 1986)
Zeilleropteris
Leaves from the Permian of Asia
Dichotomously-divided leaves over 30 cm in length and up to 27 cm wide with four orders of venations
Gigantonoclea
Leaves from Asia and North America, which were possibly lianas (Li et al. 1994) or possibly aquatic, based on frond dimorphism (Wang 1999)
Leaves similar to Gigantopteris, but with a toothed margin and veins that are less complex
Delnortea abbottiae
Permineralized leaf from Early Permian of North America (Mamay et al. 1988)
Simple, petiolate leaves mostly (3-5 cm) are oblong-elliptical with an entire to crenulate margin and a clasping base
Sclerenchyma occurs on the enrolled leaf margin, which may aid in the structure of the leaf
Venation is pinnate with four orders of veins; veins terminate at the margin
Roots
Unknown
Reproductive organs
Reproductive parts are not well understood
Pollen-producing structures
Synangiate structures (Gigantotheca) in linear rows on the abaxial surface of Gigantonoclea are assumed to be pollen organs
They are 4 cm long and extend over a secondary vein
Linophyllum xuanweiense
Supposed pollen structures are rounded (0.5 mm) and arranged in two rows on some species of Gigantopteris and Gigantonoclea (Zhao et al. 1980; Yao & Liu 2004)
Jiaochengia
Late Permian of North China
Supposed pollen structures in 3 pairs of microsporphylls bearing synangia, each consisting of 2-8 elongate sporangia
Found in association with Gigantonoclea
Ovulate structures
Leaves of Gigantonoclea have been found with seeds on both sides of the midvein (Li & Yao 1983)
The ovules or seeds (Gigantonomia) are ellipsoidal and attached to the abaxial surface with the micropyle exposed on the adaxial surface