Asteroxylon

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Asteroxylon
Temporal range: Early Devonian
Asteroxylon life restoration.jpg
Life restoration
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Lycophytes
Order: Drepanophycales
Family: Asteroxylaceae
Kidston & Lang
Genus: Asteroxylon
Kidston & Lang 1920
Type species
Asteroxylon mackiei
Kidston & Lang

Asteroxylon ("star-shaped xylem") is an extinct genus of vascular plants of the Division Lycopodiophyta known from anatomically preserved specimens described from the famous Early Devonian Rhynie chert and Windyfield chert in Aberdeenshire, Scotland. [1] [2] Asteroxylon is considered a basal member of the Lycopsida. [3]

Contents

Description

Asteroxylon is a genus of terrestrial vascular plant which flourished in the Early Devonian period. This plant consisted of aerial, isotomously and anisotomously branching stems that reached 12 mm in diameter and 40 cm in length. [4] The possibly procumbent aerial stems arose from a leaf-less rhizome which bore smaller-diameter, positively geotropic root-like branches. [4] The rhizomes, which represent an independent origin of roots, [5] reached a depth of up to 20 cm below the surface. [6] A 407 million-year-old fossil from the Rhynie chert shows the roots formed through a modified version of a mechanism called “dichotomous branching”, where one of the branches that formed from a shoot-like axis buried into the soil. This method of root formation no longer exists. [7] The xylem or conducting tissue at the center of the aerial stems is distinctly star-shaped in cross-section and has been considered an early actinostele or an "Asteroxylon-type" protostele. [8] The tracheids are of the primitive annular or helical type (so-called G-type). [9] "Leaves" – not true leaves, but protrusions – were of the form of unbranched strap-shaped enations up to 5 mm long; a single vascular trace branched from the main bundle in the centre of the stem to terminate at the base of each enation. [3] [8] Enations and axes bore stomata, indicating that their tissues were capable of photosynthesis. [10]

"Sporangia, consisting of two kidney-shaped valves, are interspersed among the nonvascularized leaflike appendages and attached to the axis with a short pedicel. The sporangia are curved and lie close to the axis." [8] [11] Fertile regions of the axes alternate with sterile regions, suggesting periodic episodes of fertility. [8]

Asteroxylon differs from other similar Early Devonian lycopsids such as Drepanophycus and Baragwanathia in that the singular vascular leaf trace in these latter plants extends into the leaf. [3] The leaves of Drepanophycus and Baragwanathia are therefore considered to be true microphylls or, alternatively, small leaves. [12]

Species

The type species is Asteroxylon mackiei. One other species has been described, Asteroxylon elberfeldense, [13] but this is now considered to be generically distinct and assigned to Thursophyton . [9] A fossil originally named as Asteroxylon setchellii is now considered to be a fern, and is known as Stenokoleos setchellii.

Taxonomy

A 2021 reanalysis found that Asteroxylon was more closely related to crown lycophytes than the more basal zosterophylls and Nothia . [14]

See also

Related Research Articles

<span class="mw-page-title-main">Lycophyte</span> Broadly circumscribed group of spore bearing plants

The lycophytes, when broadly circumscribed, are a group of vascular plants that include the clubmosses. They are sometimes placed in a division Lycopodiophyta or Lycophyta or in a subdivision Lycopodiophytina. They are one of the oldest lineages of extant (living) vascular plants; the group contains extinct plants that have been dated from the Silurian. Lycophytes were some of the dominating plant species of the Carboniferous period, and included the tree-like Lepidodendrales, some of which grew over 40 metres (130 ft) in height, although extant lycophytes are relatively small plants.

In plant anatomy and evolution a microphyll is a type of plant leaf with one single, unbranched leaf vein. Plants with microphyll leaves occur early in the fossil record, and few such plants exist today. In the classical concept of a microphyll, the leaf vein emerges from the protostele without leaving a leaf gap. Leaf gaps are small areas above the node of some leaves where there is no vascular tissue, as it has all been diverted to the leaf. Megaphylls, in contrast, have multiple veins within the leaf and leaf gaps above them in the stem.

<span class="mw-page-title-main">Equisetidae</span> Subclass of ferns

Equisetidae is one of the four subclasses of Polypodiopsida (ferns), a group of vascular plants with a fossil record going back to the Devonian. They are commonly known as horsetails. They typically grow in wet areas, with whorls of needle-like branches radiating at regular intervals from a single vertical stem.

<span class="mw-page-title-main">Rhynie chert</span> Early Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness

The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness. It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located some 700 m away. The Rhynie chert contains exceptionally preserved plant, fungus, lichen and animal material preserved in place by an overlying volcanic deposit. The bulk of the Devonian fossil bed consists of primitive plants, along with arthropods, lichens, algae and fungi.

<span class="mw-page-title-main">Zosterophyll</span> Group of extinct land plants that first appeared in the Silurian period

The zosterophylls are a group of extinct land plants that first appeared in the Silurian period. The taxon was first established by Banks in 1968 as the subdivision Zosterophyllophytina; they have since also been treated as the division Zosterophyllophyta or Zosterophyta and the class or plesion Zosterophyllopsida or Zosteropsida. They were among the first vascular plants in the fossil record, and had a world-wide distribution. They were probably stem-group lycophytes, forming a sister group to the ancestors of the living lycophytes. By the late Silurian a diverse assemblage of species existed, examples of which have been found fossilised in what is now Bathurst Island in Arctic Canada.

<i>Baragwanathia</i> Extinct genus of spore-bearing plants

Baragwanathia is a genus of extinct lycopsid plants of Late Silurian to Early Devonian age, fossils of which have been found in Australia, Canada, China and Czechia. The name derives from William Baragwanath who discovered the first specimens of the type species, Baragwanathia longifolia, at Thomson River.

<i>Drepanophycus</i> Extinct genus of spore-bearing plants

Drepanophycus is a genus of extinct plants of the division Lycopodiophyta of Early to Late Devonian age, found in Eastern Canada and Northeast US, China, Russia, Egypt and various parts of Northern Europe and Britain.

Drepanophycaceae is a family of extinct lycophytes of Late Silurian to Late Devonian age, found in North America, China, Russia, Europe, and Australia.

<span class="mw-page-title-main">Drepanophycales</span> Extinct order of spore-bearing plants

Drepanophycales is an order of extinct lycophyte plants of Late Silurian to Late Devonian age, found in North America, China, Russia, Europe, and Australia. Sometimes known as the Asteroxylales or Baragwanathiales.

<span class="mw-page-title-main">Rhyniophyte</span> Extinct group of plants

The rhyniophytes are a group of extinct early vascular plants that are considered to be similar to the genus Rhynia, found in the Early Devonian. Sources vary in the name and rank used for this group, some treating it as the class Rhyniopsida, others as the subdivision Rhyniophytina or the division Rhyniophyta. The first definition of the group, under the name Rhyniophytina, was by Banks, since when there have been many redefinitions, including by Banks himself. "As a result, the Rhyniophytina have slowly dissolved into a heterogeneous collection of plants ... the group contains only one species on which all authors agree: the type species Rhynia gwynne-vaughanii". When defined very broadly, the group consists of plants with dichotomously branched, naked aerial axes ("stems") with terminal spore-bearing structures (sporangia). The rhyniophytes are considered to be stem group tracheophytes.

<span class="mw-page-title-main">Evolutionary history of plants</span> History of plants

The evolution of plants has resulted in a wide range of complexity, from the earliest algal mats of unicellular archaeplastids evolved through endosymbiosis, through multicellular marine and freshwater green algae, to spore-bearing terrestrial bryophytes, lycopods and ferns, and eventually to the complex seed-bearing gymnosperms and angiosperms of today. While many of the earliest groups continue to thrive, as exemplified by red and green algae in marine environments, more recently derived groups have displaced previously ecologically dominant ones; for example, the ascendance of flowering plants over gymnosperms in terrestrial environments.

<span class="mw-page-title-main">Polysporangiophyte</span> Spore-bearing plants with branched sporophytes

Polysporangiophytes, also called polysporangiates or formally Polysporangiophyta, are plants in which the spore-bearing generation (sporophyte) has branching stems (axes) that bear sporangia. The name literally means 'many sporangia plant'. The clade includes all land plants (embryophytes) except for the bryophytes whose sporophytes are normally unbranched, even if a few exceptional cases occur. While the definition is independent of the presence of vascular tissue, all living polysporangiophytes also have vascular tissue, i.e., are vascular plants or tracheophytes. Extinct polysporangiophytes are known that have no vascular tissue and so are not tracheophytes.

<i>Aglaophyton</i> Extinct (Devonian) prevascular land plant

Aglaophyton major was the sporophyte generation of a diplohaplontic, pre-vascular, axial, free-sporing land plant of the Lower Devonian. It had anatomical features intermediate between those of the bryophytes and vascular plants or tracheophytes.

<i>Rhynia</i> Extinct species of vascular plant

Rhynia is a single-species genus of Devonian vascular plants. Rhynia gwynne-vaughanii was the sporophyte generation of a vascular, axial, free-sporing diplohaplontic embryophytic land plant of the Early Devonian that had anatomical features more advanced than those of the bryophytes. Rhynia gwynne-vaughanii was a member of a sister group to all other eutracheophytes, including modern vascular plants.

Sawdonia is an extinct genus of early vascular plants, known from the Upper Silurian to the Lower Carboniferous. Sawdonia is best recognized by the large number of spikes (enations) covering the plant. These are vascular plants that do not have vascular systems in their enations. The first species of this genus was described in 1859 by Sir J. William Dawson and, was originally attributed to the genus Psilophyton. He named this plant Psilophyton princeps. In 1971 Francis Hueber proposed a new genus for this species due to its "Divergent technical characters from the generic description for Psilophyton." The holotype used for description is Dawson Collection Number 48, pro parte, Museum Specimen Number 3243. Sir J. William Dawson Collection, Peter Redpath Museum, McGill University, Montreal, Quebec, Canada.

<i>Horneophyton</i> Extinct genus of early plants

Horneophyton is an extinct early plant which may form a "missing link" between the hornworts and the Rhyniopsida. It is a member of the class Horneophytopsida. Horneophyton is among the most abundant fossil organisms found in the Rhynie chert, a Devonian Lagerstätte in Aberdeenshire, UK. A single species, Horneophyton lignieri, is known. Its probable female gametophyte is the form taxon Langiophyton mackiei.

Hsua is a genus of extinct vascular plants, known from the Devonian. The name of the genus honours the Chinese palaeobotanist, Jen Hsü (徐仁).

<i>Nothia aphylla</i> Extinct species of spore-bearing plant

Nothia was a genus of Early Devonian vascular plants whose fossils were found in the Rhynie chert in Scotland. It had branching horizontal underground stems (rhizomes) and leafless aerial stems (axes) bearing lateral and terminal spore-forming organs (sporangia). Its aerial stems were covered with small 'bumps' (emergences), each bearing a stoma. It is one of the best described early land plants. Its classification remains uncertain, although it has been treated as a zosterophyll. There is one species, Nothia aphylla.

<i>Ventarura</i> Extinct genus of spore-bearing plants

Ventarura is a genus of extinct vascular plants of the Early Devonian. Fossils were found in the Windyfield chert, Rhynie, Scotland. Some features, such as bivalved sporangia borne laterally and the anatomy of the xylem, relate this genus to the zosterophylls. Other features are unclear due to poor preservation.

Trichopherophyton is a genus of extinct vascular plants of the Early Devonian. Fossils were found in the Rhynie chert, Scotland. The remains are very fragmentary, but the plant appears to be related to the zosterophylls.

References

  1. Kidston, R.; Lang, W. H. (1920). "On Old Red Sandstone Plants showing Structure, from the Rhynie Chert Bed, Aberdeenshire. Part III. Asteroxylon Mackiei, Kidston and Lang". Transactions of the Royal Society of Edinburgh. 52 (3): 643–680. doi:10.1017/S0080456800004506. S2CID   131282834.
  2. Rice, C. M.; Ashcroft, W. A.; Batten, D. J.; Boyce, A. J.; Caulfield, J. B. D.; Fallick, A. E.; Hole, M. J.; Jones, E.; Pearson, M. J.; Rogers, G.; Saxton, J. M.; Stuart, F. M.; Trewin, N. H.; Turner, G.; et al. (1995). "A Devonian auriferous hot spring system, Rhynie, Scotland". Journal of the Geological Society, London. 152 (2): 229–250. Bibcode:1995JGSoc.152..229R. doi:10.1144/gsjgs.152.2.0229. S2CID   128977213.
  3. 1 2 3 Hao, Shougang; Xue, Jinzhuang (2013). The Early Devonian Posongchong Flora of Yunnan - A Contribution to an Understanding of the Evolution and Early Diversification of Vascular Plants. Beijing: Science Press. pp. 244–245. ISBN   978-7-03-036616-0.
  4. 1 2 Strullu-Derrien, Christine; Wawrzyniak, Zuzanna; Goral, Tomasz; Kenrick, Paul (2015). "Fungal colonization of the rooting system of the early land plant Asteroxylon mackiei from the 407-Myr-old Rhynie Chert (Scotland, UK)". Botanical Journal of the Linnean Society. 179 (1): 201–213. doi: 10.1111/boj.12307 .
  5. Hetherington, Alexander J.; Dolan, Liam (2018). "Stepwise and independent origins of roots among land plants". Nature. 561 (7722): 235–238. Bibcode:2018Natur.561..235H. doi:10.1038/s41586-018-0445-z. PMC   6175059 . PMID   30135586.
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  7. 400-Million-Year-Old Fossils Reveal How the First Roots Evolved
  8. 1 2 3 4 Kerp, Hans; Wellman, Charles H.; Krings, Michael; Kearney, Patricia; Hass, Hagen (2013). "Reproductive organs and in situ spores of Asteroxylon mackiei Kidston & Lang, the most complex plant from the lower Devonian Rhynie chert". International Journal of Plant Sciences. 174 (3): 293–308. doi:10.1086/668613. S2CID   84542464.
  9. 1 2 Kenrick, Paul & Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D.C.: Smithsonian Institution Press. ISBN   978-1-56098-730-7.
  10. Wilson, Jonathon P.; Fischer, Woodward W. (2011). "Hydraulics of Asteroxylon mackei, an early Devonian vascular plant, and the early evolution of water transport tissue in terrestrial plants". Geobiology. 9 (2): 121–130. Bibcode:2011Gbio....9..121W. doi: 10.1111/j.1472-4669.2010.00269.x . PMID   21244621.
  11. In their original description, Kidston & Lang thought that some unconnected sporangial branches, similar to those of rhyniophytoids, possibly belonged to Asteroxylon, and in spite of the lack of certainty, many subsequent authors constructed suprageneric taxonomies based on this assumption. The true nature of Asteroxylon sporangia, and of these unconnected sporangia (which were found to belong to Nothia aphylla ) were later shown by: Lyon, A.G. (1964). "Probable Fertile Region of Asteroxylon mackiei K. and L.". Nature. 203 (4949): 1082–1083. Bibcode:1964Natur.203.1082L. doi:10.1038/2031082b0. S2CID   4292879..
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  14. Hetherington, Alexander J; Bridson, Siobhán L; Lee Jones, Anna; Hass, Hagen; Kerp, Hans; Dolan, Liam (2021-08-24). "An evidence-based 3D reconstruction of Asteroxylon mackiei, the most complex plant preserved from the Rhynie chert". eLife. 10: e69447. doi: 10.7554/eLife.69447 . ISSN   2050-084X. PMC   8384418 . PMID   34425940.