The Marattiales are a group of ancient tree ferns that date back to the Carboniferous coal age. They are uncommon tree ferns of the modern tropics with six living genera and 90 species. This group was very abundant and diverse in Paleozoic lowland swamps, even dominant at the end of the Carboniferous, before seed plants began to take over dominance. A well-known marattialean fern of the Carboniferous is Psaronius, which existed in the understory of lepidodendrid swamps.
Stems
Stump-like, unbranched rhizome that can attain tree stature
Dictyostele present in stem
Leaves
Large compound leaves (fronds)
Croziers or fiddle heads
Roots
Abundant adventitious roots that are large and fleshy
Completely encase the stem of plant
Reproduction
Spore-bearing with large sporangia
Eusporangiate: large sporangia with thick walls and 100s of spores
Sporangia fuse into a synangium borne abaxially on ultimate pinnule laminae
└Marattiales
Carboniferous - present
Above: rhizome stump of Marattia
Above: Leaves of Marattia
Above: Sporangia of Angiopteris and Marattia
6 living genera (90 species): Angiopteris, Christensenia, Danaea, Eupodium, Marattia, and Ptisana
Abundant in the Pennsylvanian
Morphotaxon for the trunk, which may have been up to eight to ten meters in length
The stem possessed a dictyostele with endarch maturation xylem
P. simplicicaulis
Dimichele & Phillips, 1977
One of the earliest marattialean ferns from the Early Pennsylvanian of North America
P. renaultii
Williamson, 1876
The ferns possessed a large crown of pinnately-compound leaves inserted in spirals or vertical rows on the trunk
The fronds were large, up to three meters in length, bilaterally symmetrical, and planate
The petioles have a U-shaped vascular trace
Devonian - Carboniferous, with the greatest diversity in the Pennsylvanian
Morphotaxon for compressions of leaves of marattialean ferns during the Carboniferous
It is also the morphotaxon for some true ferns (Polypodiidae) and at least one seed fern (Taylor et al., 2009)
There are over 250 species of Pecopteris named (Robertson & DiMichele, 1997)
Watson, 1906
Morphotaxon for permineralized leaves
Morphotaxon for compression or impression petiole scars
Fronds at least bipinnate. Pinnules taeniopteroid or neuropteroid, bases cordate, apices rounded.
Midvein distinct [andstraight]
Lateral veins curved, rising at an acute angle, perpendicular or subperpendicular to pinnule margin.
In vegetative pinnules, lateral veins commonly bifurcate twice, occasionally three times, showing first bifurcation near the midvein, with a vein density of 40–50 veins per centimeter at the pinnule margin.
In fertile pinnules, lateral veins bifurcate once near midvein, with vein density less than half of vegetative
pinnules.
Fertile pinnules identical in size and shape to vegetative pinnules.
Synangia, as pendants, abaxially attached, [arranged in one row on each side of midvein], born on every individual lateral vein branch, closely spaced, with length of nearly half-width of pinnules, and in a ‘louvered’ pattern [or an imbricate arrangement].
Each valve of bivalvate synangia composed of elongated and laterally fused sporangia.
Sporangia are long ellipsoid, exannulate, with length near synangium
Abundant roots that are large and fleshy, completely encasing the stem forming a root mantle
The root cross-section reveals an exarch actinostele
The roots that are closest to the stem have a thick, sclerenchymatous cortex, and reinforced the trunk which lacked secondary xylem
Spore-bearing
Fertile fronds bore synangia composed of eusporangia on the abaxial surface, commonly clustered near veins of ultimate pinnules
Sporangia have uniform dehiscence along the midline of the sporangia
Spore genera Punctatisporites and Laevigatosporites are associated with Pecopteris foliage
Above: Psaronius, an extinct Marattioid fern common in the Carboniferous Period
Below: a cross-section of Psaronius, showing the stem and root mantle
Giant Ferns (In Defense of Plants 2017)