The cladoxylalean cladoxylosids are the crown group of taxa within the Cladoxylopsids, which include the genus Cladoxylon. They possess complex anatomical patterns with multiple radiating arms of xylem within their stems.
Mid‐ to late Emsian
Axes, 1–1.5mm thick
Primary xylem is an actinostele with five or six lobes that protrude from the central area
Phloem layer thin around xylem, with thin‐walled fusiform cells
Epidermal cells with robust primary walls and secondary metabolite content fossilized inside as solid rods of dark, compact material
Xylem traces to ultimate appendages hourglass‐shaped, tangentially elongated
Ultimate appendages isotomously branched at least twice, some with recurved tips.
Sporangia erect, attached at tips of ultimate appendage segments, fusiform,
Unger, 1854
Mississippian from Germany
Anatomically preserved stem axis with many stelar segments
Lateral axes are spirally arranged
Primary xylem has scalariform tracheids
Secondary xylem abundant in some species, but nearly absent in others (e.g., Cladoxylon radiatum)
Periderm has been observed in some specimens of Cladoxylon (Decombeix & Galtier, 2021)
C. centrale
Unger, 1854
Tournasian of Germany
C. mirabile
Unger, 1854
Tournasian of Germany
C. radiatum
C. taeniatum
Solms-Laubach, 1897
Tournasian of Germany
Originally named Schizoxylon taeniatum
Cladoxylon taeniatum might have had a creeping or climbing habit
Wang & Geng, 1997
M. tetraxylum †
Wang & Geng, 1997
Middle Devonian of China
M. ziguinum †
Wang & Geng, 1997; Berry & Stein, 2000; Wang & Lin, 2007
Frasnian of China
Eight stelar ribs arranged in four U-shaped strands
Each rib shows one (rarely, two) mesarch strands of protoxylem in peripheral position and produces a C-shaped trace at nodes.
Distribution of the traces within the two to seven branches described from compressions is unclear.
Middle Devonian (Givetian) Haikou Formation, Yunnan Province, southwestern China
Stout stem, two orders of branching, and recurved ultimate units forming more or less planate sterile branching structures laterally
The fertile unit is complex, with many stalked discoidal sporangia aggregated around the second order fertile branch
This plant shares few morphological characters with the only well known and well defined order of cladoxylopsid plants
The primary xylem in main axes consists of numerous primary xylem segments, most of which are radially directed and elongate.
Some xylem segments are connected toward the center to form V, Y, W, or more complex-shaped configurations
One permanent protoxylem strand occurs near the outer tip of each xylem segment of the main axes
A dissected primary xylem column occurs in first-order branches, but the xylem segments are fewer in number and smaller in size than those in main axes
Most xylem segments in first-order branches are unidirectional and parallel in orientation, some are connected to form Y, twice-dichotomous, or more complex-shaped configurations, and one protoxylem strand occurs near each of the two ends of individual xylem segment or a segment group
Metaxylem tracheids are characterized by multiseriate circular bordered pits
Vascular traces depart from the tip of primary xylem segments, produced by elongation and division of protoxylem strands
These traces are of various sizes and shapes, from circular, elliptical to elongate-elliptical in transverse outline, with one or two protoxylem strands
Cortex tissues are mainly composed of thin-walled, approximately isodiametric cells
Above/Below: Anatomy of Panxia
Emsian of Quebec, CA
Paracladophyton appears to be closely aligned with Cladophyton, and is a cladoxylalean cladoxylopsid
Axes with primary xylem forming multiple long radially oriented ribs, sometimes converging and fusing with adjacent ribs centripetally, and some of the xylem segments converging at the center of the axis.
Primary xylem maturation mesarch, with multiple protoxylem strands located along each xylem rib midplane.
Lateral appendages with whorled or helical taxis, supplied by single terete centrarch xylem traces that diverge radially from the xylem ribs of the axis.
Lateral appendages three-dimensionally dissected, with appendage segments supplied by small centrarch xylem traces.
Above: Paracladophyton from Chu et al., 2024, Figures 16-21 showing anatomy
Gothan, 1927; Cross & Hoskins, 1951
Late Devonian (Famennian) of Morocco
Similar anatomically to Pseudosporochnus
The stems have a diameter of about 2.5 cm, and have a cylinder of radially arranged xylem plates
P. schulleri †
Gothan, 1927
Famennian of Morocco
Primary tissue only; the trunk diameter is often more than 15 cm, the tribe has both central as well as peripheral xylem strands
P. polyupsilon †
Read & Campbell, 1939
From the New Albany Shale of Kentucky, the number of xylem segments is 54
The main axis had determinate growth; the lateral axes were arranged in whorls
P. timanica †
Lepechina, 1968
P. levis †
Soria & Meyer-Berthaud, 2004
Lower Famennian (Upper Devonian) of Morocco
Possesses aerenchyma, suggesting that it lived in wetlands
Late Devonian of China
Branches were produced in an irregular helix, but ultimate appendages were borne in a pinnate arrangement on a frondlike system
Vegetative "pinnules" were dichotomous and planar (Sphenopteridium-type), whereas fertile "pinnules" were decussately branched and bore fusiform sporangia terminally
Anatomically, Polypetalophyton is unusual because it has a stele that consists of several primary vascular segments, each of which consists of tracheids and parenchyma surrounded by wood comprising radially aligned tracheids with ray-like slits
Similar in anatomy to Cladoxylon taeniatum
Probably aligned with the Pseudosporochnales.
Polypetalophyton differs from the handful of well-known members of this group in possessing radially aligned xylem and in lacking sclereid nests in the stem cortex.
It is also unique in its plagiotropic pinnate frond-like branching system, which we interpret to be the functional equivalent of a leaf or frond.
Polypetalophyton represents the first evidence of Sphenopteridium-type foliage occurring within Cladoxylopsids and demonstrates that this kind of foliage has evolved independently within at least three distinct plant clades: zygopterids, cladoxylopsids, and seed plants.
Above: Foliage of Polypetalophyton
Read & Campbell, 1939
P. elegans
Read & Campbell, 1939
Tournasian of Kentucky & Ohio
P. australe
Late Devonian of Australia
Small stem axis with a diameter of 2.7 cm.
Anatomy has 19 elongated arms with mesarch primary xylem
Each of the arms branches at the end of dichotomous and forms at least six traces running in the whorled standing side axes.
Above: Polyxylon australe line drawing of axis (Fig 2, Meyer-Berthaud et al. 2007)
Late Devonian of New York, USA
The primary xylem consists of 18 radially oriented and elongate primary xylem ribs.
Toward the center of the system, some xylem ribs are laterally continuous with adjacent ribs, forming elongate xylem segments, whereas others appear discrete.
Order of xylem maturation is mesarch, with one to three protoxylem strands located near the tip of each primary xylem rib.
Primary phloem immediately surrounds each primary xylem rib and consists of sieve cells.
To the outside of the primary phloem, a conspicuous ground tissue is comprised of roughly isodiametric parenchyma with dark contents in their lumina as well as parenchyma cells that are longitudinally elongate with clear lumina.
Beyond the vascular column, a fragmentary cortex contains numerous vascular traces embedded in parenchyma.
These latter cells are surrounded by conspicuous masses of longitudinally elongate cortical thick-walled parenchyma.
Vascular traces depart from the tips of the primary xylem ribs in a whorled phyllotaxis.
Most traces emerge along the midplane of the primary xylem ribs. However, some depart at an oblique angle.
Above: Fig 1 (left) and Fig 6 (right) of Cordi & Stein (2005) of the distal stem of Rotoxylon dawsonii