Archaeopteridales †
Wood-bearing, spore-producing plants
The Archaeopteridales were a group of progymnosperms, meaning that they were spore-bearing plants that possessed a cambium with robust woody growth. This combination of traits is not found in any living plants, since all extant spore-bearing plants lack robust woody growth. The group is named after the taxon Archaeopteris, which is the earliest known woody tree found in the fossil record (Middle Devonian). This plant had fern-like foliage that bore sporangia on specialized fertile leaves, which is the form genus for Archaeopteris. For many years it was assumed that Archaeopteris was a fern, until the 1960s when Charles Beck was able to demonstrate that these fern-like leaves were actually connected to fossil wood known as Callixylon. From a distance, these plant probably appeared quite conifer-like, but upon closer inspection, the leaves would appear fern-like. These trees were a major component of the Late Devonian and Mississippian ecosystems, and may have played a role in the Late Devonian extinction event.
Ecology & Form
Probably appeared quite conifer-like
Major component of ecosystems
Originally thought to be ferns
First true woody trees
Stem
Protostele-Eustele with large amounts of pycnoxylic wood (secondary xylem)
Callixylon is form genus for stem w/ wood
Trunk diameters in excess of 1 m
Estimated heights of up to 30 m have been reported
Pseudomonopodial and monopodial branching with axillary buds
Leaves
Laminate leaves that were roughly fan-shaped, but highly variable (see Archaeopteris below)
Archaeopteris is form genus for branching systems with sporangia
Reproduction
Sporangia adaxial on special, helically-arranged leaves (see Archaeopteris below)
Sporangia are elongated and pointed with lateral dehiscence
Heterosporous with spores that are radial, trilete, pseudosaccate
Microspores are 70 micrometers, and align with the form taxon Aneurospora
16-32 megaspores (110-400 micrometers) are in a megasporangium; assigned to Contagisporites
Classification
└Archaeopteridales †
Geologic Age
Middle Devonian (Eifelian)-Carboniferous (Mississippian)
Above: Archaeopteris root system at the Cairo fossil forest site (Credit: Charles Ver Straeten)
Above: Archaeopteris root system at the Cairo fossil forest site (Image credit: Bill Stein)
Diversity
Actinopodium nathorstii †
Schweitzer 1999
Middle Devonian of Spitsbergen
Permineralized axes with medullated actinostele with wedges of mesarch xylem, and some secondary xylem
Maybe the main axis of Archeopteris (Gensel & Andrews 1984)
Actinoxylon banksii †
Middle Devonian
Several orders of branching, with ultimate axes bearing sub-opposite leaves in a decussate arrangement
Stele contains actinostele with 6 ribs; each rib contains one protoxylem point at the peripheral tip, and another along the radius of the rib
Secondary xylem is present
Archaeopteris †
Dawson 1871, Lesquereux 1880, Fairon-Demaret et al. 2001
Devonian - Carboniferous taxon for vegetative and fertile branch systems
Known from North America, Russia, China, Morocco, Europe and Australia
Lateral branch systems were pseudomonopodial in nature, developing from primordia and not axillary buds
Leaves are helically-arranged, but appear opposite and decussate in pentultimate axes
Lamina was high-variable (i.e. dissected, entire, obovate, spatulate) and anisophylly is occurs in some species
Fertile ultimate axes are usually occur as the basalmost units of branching system
Fertile leaves are helically-arranged on fertile axes, with two rows of fusiform sporangia
Heterospory is present
A. halliana
A. hibernica
A. macilenta
A. obtusa
Callixylon †
Zalessky 1911
Twenty species of Callixylon described from North America, Morocco, Europe, China and several parts of Russia
Late Givetian to the Mississippian
Pycnoxylic woody stems up to 150 cm in diameter, and 10 m in length
Eustelic with pith and mesarch xylem of thick-walled tracheids and narrow rays
Growth rings present
Eddya sullivanensis †
Beck 1967
Late Devonian (Frasnian) from New York
Possible immature plant, or sapling, of Archaeopteris
30cm long, with shoots, leaves, and well-developed roots
Leaves are fan-shaped, with dichotomous veins, and helically-arranged
Stem is an eustele with mesarch development and small amount of wood
? Fuellingia gilkinetii
Schweitzer & Geisen 2002
Axes up to 4 cm in diameter and 1 m in length
Fan-shaped, helically-arranged leaves are attached
Sporangiophores are attached to midrib of wing-shaped leaves, with numerous helically -arranged lateral branches
? Langoxylon asterochlaenoideum †
Middle Devonian of Belgium
Fern-like, permineralized stem with possible archaeopterid affinities
"Langoxylon compares with Middle Devonian archaeopterid progymnosperms like Actinopodium and Actinoxylon, but has a mixture of anatomical characters that were previously used to subdivide Euphyllophytina into Moniliformopses and Radiatopses types. Langoxylon is thus a ‘fern-like’ Middle Devonian euphyllophyte of uncertain phylogenetic relationship" (Scheckler et al. 2006)
Siderella †
Read 1936
Lower Mississippian (Tournaisian) from the New Albany Shale of North America
Permineralized axes with 7-10 primary xylem (px) lobes, with patches of px that is reminiscent of peripheral loops at the tips of the lobes
Anatomically, this taxon is larger, but otherwise identical to the penultimate axes of Archaeopteris
Siderella leaves may have been distichous, unlike the decussate leaves in Archaeopteris (Beck & Wight 1988)
Svalbardia †
Hoeg 1942, Schweitzer 1999
Middle Devonian
Planated axes that may represent a portion of Archeopteris trees