Aneurophytales

Earliest true wood-producing plants

The aneurophytes are a group of progymnosperms, which are the basal-most members of the lignophyte clade. This means that they were some of the first plants on Earth to produce robust wood from a cambium, similar to modern-day trees. These plants differed from modern woody plants in the following manner. The aneurophytes were spore-bearing plants, not seed-bearing (modern wood-producing plants are seed-bearing). Also, aneurophytes lacked leaves; they used small photosynthetic stems for photosynthesis. Evidence of a preserved forest from Gilboa, NY from the middle Devonian indicates that some aneurophytes were large woody vines that grew in-between the Earth's earliest tree-like plants, Eospermatopteris. Other aneurophytes, like Tetraxylopteris, Triloboxylon, and Rellimia may have been woody shrubs.

Above: Ancient Gilboa forest of cladoxylopsid trees with aneurophyte vines interspersed

Ecology & Form

Stem

Morphology

  • Three-dimensional branching systems

  • Lateral axes were helical or decussate

  • These plants were probably small shrubby or vine-like

Anatomy

Leaves

  • No laminate leaves

  • Ultimate appendages dichotomize were assumed to be photosynthetic

Roots

  • Adventitious from stems

Reproduction

  • Sporangia born on ultimate fertile appendages

  • Homosporous

  • Sporangia were elongated and pointed with lateral to tip & base dehiscence

  • Spores are radial, trilete, pseudosaccate

Diversity

Aneurophyton

  • Middle-Late Devonian of Europe, Russia, United States

  • Aneurophyton is similar to Tetraxylopteris in overall form, but its branches are arranged helically rather than decussately.

  • Dichotomous ultimate branches in Aneurophyton were photosynthetic but unwebbed, and sometimes interpreted as leaves

  • In the fertile axes, ultimate branches are replaced by dichotomies with multiple homosporous sporangia arising along the length of each fork of the branch.

  • Sporangia appear to recurve toward the center of the dichotomy, but this may be an artifact of preservation.

  • The main axis of Aneurophyton has a lobed protostele and produced a small amount of secondary xylem from meristematic regions between the lobes.

  • Axes are relatively slender suggesting vine growth form, similar to aneuophytes found at Gilboa, NY

A. doui

  • Jiang et al. 2013

  • At least four orders of axes and ultimate units (vegetative appendages/fertile organs)

  • Spines, 0.5–1.5 mm long, occur sparsely on the surface of all orders of axes and ultimate units.

  • First-order axes, up to 6 mm wide and at least 90 mm long, demonstrate only a slight distal taper.

  • Second-order axes, up to 4 mm wide, diverge helically from the first-order axis in closely inserted pairs.

  • From the second order, axes of subsequent orders are produced by bifurcation; third-order axes are 2.0–2.5 mm wide, and fourth-order axes are 1.5–2.0 mm wide.

  • Vegetative appendages, 7.0–13.0 mm long as a whole, diverge from the third- or fourth-order axes in helical pairs, as a one to three times dichotomizing system

  • of three-dimensionally-extended flattened branchlets.

  • Fertile organs, 3.7–8.0 mm long as a whole and borne on the third or fourth-order axes, are up to three times opposite/subopposite pinnate systems.

  • All three orders of fertile organ axes, in most cases recurved and rarely straight, bear sporangia or organ axes.

  • Sporangia, oppositely, suboppositely, or alternately, are inserted distichously to the usually adaxial side of the fertile organ axes; they are short-stalked, elliptical, 0.8–1.3 mm wide, and 2.2–3.5 mm long, and they sometimes have longitudinal dehiscence or a twisting configuration.

A. germanicum

  • Krausel & Weyland 1941

A. olnense

Above: Evidence of vine-like habit of an aneurophytalean (Gilboa, NY)

Above: Aneurophyton doui (Fig.6 from Jiang et al. 2013)

Above: Evidence of vine-like habit of an aneurophytalean (Gilboa, NY)

Above: Branches of Aneurophyton

Cairoa lamanekii

  • Matten 1973

  • Givetian (Middle Devonian) of New York

  • Fossils are preserved as petrifaction showing three orders of branching, penultimate axis, ultimate axis, leaves

  • The penultimate axis bearing the ultimate axes in an alternate manner (1/3 phyllotaxy)

  • The leaves probably dichotomizing.

  • Xylem in transverse section of penultimate axis three-armed

  • secondary xylem present; protoxylem is circular, mesarch, several;

  • Tip of xylem arm, just below separation of trace to penultimate branch, three-lobed with a protoxylem area in each lobe

  • Xylem in transverse section of penultimate branch commonly rounded to four-angled; traces to leaves terete and opposite

Above: Cross-section Cairoa lamanekii (From Fig. 14, Matten 1973)

Gmujij tetraxylopteroides

  • Pfeiler and Tomescu 2020

  • Early Devonian (Emsian) from Battery Point Formation of Quebec, Cananda

  • Mesarch actinostele with Psilophyton-type (P-type) tracheid wall thickening,

  • Gmujij differs from other Early Devonian euphyllophytes with a new type of anatomical organization in wood-production

  • Comparisons indicate similarity between Gmujij and aneurophytalean progymnosperms, such as Tetraxylopteris.

    • Its plesiomorphic P-type tracheids set this plant apart from younger actinostelic euphyllophytes

Above: Gmujij tetraxylopteroides xylem anatomy (Fig 1, Pfeiler and Tomescu 2020)

Proteokalon petryi

  • Scheckler & Banks 1971

  • Early Frasnian (Late Devonian) of New York

  • Decussate axes, except ultimate axes that dichotomize several times in a single plane

  • Abaxial appendages appear with paired axes

  • Morphology similar to Tetraxylopteris, and anatomy similar to Aneurophyton, Rellimia, and Tetraxylopteris

  • Some specimens show a sparganum-type outer cortex, with anastomosing strands of sclerenchyma

Rellimia thomsonii

  • Bonamo 1977; Dannenhoffer et al. 2007

  • Middle Devonian (Givetian-Eifelian) of Europe, Russia, and New York

  • Originally named Protopteridium and Milleria (Leclercq & Bonamo 1971, 1973)

  • Shrubby plant with helically-arranged branches up to 5 orders

  • Ultimate axes had either leaves or sporangia

  • Anatomy: three-lobed mesarch primary xylem (actinostele) surrounded by pycnoxylic secondary xylem with a bifacial cambium

  • Stele shape was the same throughout all axes (i.e. 3-ribbed), but the ultimate appendage trace was terete

Tetraxylopteris

  • Complex system of pseudomonopodial branches with three orders of decussate branches, probably giving the plant a bushy appearance

    • Laterals arranged in opposite pairs with successive pairs at right angles (decussate)

  • Primary, mesarch xylem strand was central in all axes, and cross-shaped in proximal axes

    • Ultimate axes exhibit terete primary xylem in cross-section.

    • Some specimens show a sparganum-type outer cortex, with anastomosing strands of sclerenchyma

  • Sporangia were born on fertile axes.

    • Two successive dichotomies, creates four fertile axes which were 3x pinnate.

    • Ultimate axes exhibit an elongated sporangium at the tip

T. schmidtii (Beck 1957),

  • Middle to Upper Devonian in age

  • T. schmidtii, all axes decreased in width, distally; final order axes bore appendages arranged oppositely and decussately

T. reposana (Hammond & Berry 2005)

  • Frasnian

  • T. reposana, 1st and 2nd order axes show swelling at base of branches; decrease in width begins with 3rd order axes; final order axes were spirally or helically arranged

Above: Tetraxylopteris reposana reconstructions

Triloboxylon

  • This plant gets its name from the three-lobed vascular cylinder, and may represent one of the earliest lignophyte members

  • Matten & Banks 1966

  • Middle Devonian - Late Devonian of New York

T. ashlandicum

T. arnoldii

  • Matten 1974; Stein & Beck 1983

  • Givetian (Middle Devonian)

  • Originally Aneurophyton hallii (Arnold)

  • Three-ribbed primary xylem system with protoxylem strands near the tips and along midplanes of the ribs

  • Small transversely elliptical traces produced in pairs from successive primary xylem ribs

  • Extensive secondary tissues

  • Heterogeneous inner cortex containing solitary fibers and clusters of sclereids

  • An outer cortex, with peripheral bundles consisting of fibers and sclereids separated by thin-walled parenchyma cells in ordered arrays

  • Simple but extensively developed periderm

Above: Cross-section of T. ashlandicum

Below: Cross-section of T. arnoldii