Aglaophyton

Aglaophyton major (or more correctly Aglaophyton majus[2]) was the

Aglaophyton
sporophyte generation of a diplohaplontic, pre-vascular, axial, free-sporing
land plant of the Lower Devonian (Pragian stage, around Temporal range: Early Devonian
410 million years ago). It had anatomical features intermediate between those PreЄ Є O S D C P T J K PgN
of the bryophytes and vascular plants or tracheophytes.

A. major was first described by Kidston and Lang in 1920 as the new species
Rhynia major.[3] The species is known only from the Rhynie chert in
Aberdeenshire, Scotland, where it grew in the vicinity of a silica-rich hot
spring, together with a number of associated vascular plants such as a smaller
species Rhynia gwynne-vaughanii which may be interpreted as a
representative of the ancestors of modern vascular plants and Asteroxylon
mackei, which was an ancestor of modern clubmosses (Lycopsida).

Contents
Description
Taxonomy
Phylogeny Reconstruction of the sporophyte of
References Aglaophyton, illustrating bifurcating
External links axes with terminal sporangia, and
rhizoids. Insets show a cross-section
of a sporangium and the probable
Description
spores.
The stems of Aglaophyton were round in cross-section, smooth,
Scientific classification
unornamented, and up to about 6mm in diameter. Kidston and Lang[3]
interpreted the plant as growing upright, to about 50 cm in height, but Clade: Polysporangiophyta
Edwards[1] has re-interpreted it as having prostrate habit, with shorter aerial
Genus: †Aglaophyton
axes of about 15 cm height. The axes branched dichotomously, the aerial axes
D.S.Edwards 1986[1]
branching at a comparatively wide angle of up to 90°, and were terminated
with elliptical, thick-walled sporangia, which when mature, opened by spiral Species
slits, so that the sporangia appear to be spiral in form.[4] Sporangia contained
many identical spores (isospores) bearing trilete marks. The spores may †A. major (Kidston & Lang 1920)
therefore be interpreted as meiospores, the product of meiotic divisions, and Edwards 1986[1]
thus the plants described by Edwards and by Kidston and Lang were diploid,
Synonyms
sporophytes. The plant was originally interpreted as a tracheophyte, because
the stem has a simple central vascular cylinder or protostele,[3] but more
Lyonophyton Remy & Remy
recent interpretations in the light of additional data indicated that Rhynia
major had water-conducting tissue lacking the secondary thickening bars seen 1980
in the xylem of Rhynia gwynne-vaughanii, more like the water-conducting
system (hydrome) of moss sporophytes. Edwards[1] reinterpreted the species as non-vascular plant and renamed it Aglaophyton
major.

Aglaophyton is among the first plants known to have had a mycorrhizal

relationship with fungi,[5] which formed arbuscules in a well-defined zone in the
cortex of its stems. Aglaophyton lacked roots, and like other rootless land plants
of the Silurian and early Devonian may have relied on mycorrhizal fungi for
acquisition of water and nutrients from the soil.

The male gametophyte of the species has been formally described,[6] which was
assigned to a new form taxon Lyonophyton rhyniensis, but is now properly Aglaophyton major
referred to as an Aglaophyton gametophyte. The Rhynie chert bears many
examples of male and female gametophytes, which are loosely similar in their
construction to the sporophyte phase, down to bearing rhizoids.[7]

Taxonomy
Aglaophyton major was first described as Rhynia major by Kidston and Lang in 1920.[3] In 1986 D.S. Edwards re-examined
fossil specimens and reported that they did not contain true vascular tissue, but rather conducting tissue more similar to that of
bryophytes. As the diagnosis of Rhynia was that it was a vascular plant, he created a new genus, Aglaophyton, for this species.
(The other species of Rhynia, R. gwynne-vaughanii, was not affected.) As Rhynia major the species had been placed in the
rhyniophytes, but no alternative higher level classification was proposed for the new genus.[1]

Phylogeny
In 2004, Crane et al. published a cladogram for the polysporangiophytes which places Aglaophyton as a sister of the vascular
plants (tracheophytes), with the Horneophytopsida being sister to both.[8] The basis of the cladogram is that Aglaophyton has
more developed conducting tissue than the Horneophytopsida, but does not have true vascular tissue.

† Horneophytopsida (Caia, Horneophyton, Tortilicaulis)

polysporangiophytes † Aglaophyton

tracheophytes

References
1. Edwards, David S. (1986), "Aglaophyton major, a non-vascular land-plant from the Devonian Rhynie Chert",
Botanical Journal of the Linnean Society, 93 (2): 173–204, doi:10.1111/j.1095-8339.1986.tb01020.x (https://doi.or
g/10.1111%2Fj.1095-8339.1986.tb01020.x)
2. Strictly the name should have been Aglaophyton majus, as -phyton is neuter and the neuter of Latin comparative
adjectives ends in -us. Since February 2018, authors writing on the Rhynie chert have begun using the more
correct form. See Wellman, Charles H. (2018), "Palaeoecology and palaeophytogeography of the Rhynie chert
plants: Further evidence from integrated analysis of in situ and dispersed spores", Philosophical Transactions of
the Royal Society B, 373: 20160491, doi:10.1098/rstb.2016.0491 (https://doi.org/10.1098%2Frstb.2016.0491),
PMC 5745327 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5745327) and other papers in the same issue of
that journal.
3. Kidston, R. & Lang, W.H. (1920), "On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed,
Aberdeenshire. Part II. Additional notes on Rhynia gwynne-vaughani, Kidston and Lang; with descriptions of
Rhynia major, n.sp. and Hornea lignieri, n.g., n.sp." (https://zenodo.org/record/1428662/files/article.pdf) (PDF),
 Transactions of the Royal Society of Edinburgh, 52: 603–627, doi:10.1017/s0080456800004488 (https://doi.org/1
0.1017%2Fs0080456800004488)
4. Remy, W. & Hass, H. (1996), "New information on gametophytes and sporophytes of Aglaophyton major and
inferences about possible environmental adaptations", Review of Palaeobotany and Palynology, 90 (3–4): 175–
193, doi:10.1016/0034-6667(95)00082-8 (https://doi.org/10.1016%2F0034-6667%2895%2900082-8)
5. Remy W, Taylor TN, Hass H, Kerp H (1994), "4 hundred million year old vesicular-arbuscular mycorrhizae",
Proceedings of the National Academy of Sciences of the United States of America, 91 (25): 11841–11843,
Bibcode:1994PNAS...9111841R (https://ui.adsabs.harvard.edu/abs/1994PNAS...9111841R),
doi:10.1073/pnas.91.25.11841 (https://doi.org/10.1073%2Fpnas.91.25.11841), PMC 45331 (https://www.ncbi.nl
m.nih.gov/pmc/articles/PMC45331), PMID 11607500 (https://www.ncbi.nlm.nih.gov/pubmed/11607500).
6. Remy, W & Remy, R (1980) Lyonophyton rhyniensis n.gen. et nov. spec., ein Gametophyt aus dem Chert von
Rhynie (Unterdevon, Schottland). Argumenta Palaeobotanica, 6, 37-72
7. Taylor, T. N.; Kerp, H; Hass, H (2005), "Life history biology of early land plants: Deciphering the gametophyte
phase", Proceedings of the National Academy of Sciences of the United States of America, 102 (16): 5892–7,
Bibcode:2005PNAS..102.5892T (https://ui.adsabs.harvard.edu/abs/2005PNAS..102.5892T),
doi:10.1073/pnas.0501985102 (https://doi.org/10.1073%2Fpnas.0501985102), PMC 556298 (https://www.ncbi.nl
m.nih.gov/pmc/articles/PMC556298), PMID 15809414 (https://www.ncbi.nlm.nih.gov/pubmed/15809414).
8. Crane, P.R.; Herendeen, P. & Friis, E.M. (2004), "Fossils and plant phylogeny" (http://www.amjbot.org/cgi/content/
full/91/10/1683), American Journal of Botany, 91 (10): 1683–99, doi:10.3732/ajb.91.10.1683 (https://doi.org/10.37
32%2Fajb.91.10.1683), PMID 21652317 (https://www.ncbi.nlm.nih.gov/pubmed/21652317), retrieved 2011-01-27

External links
Cladogram (http://www.amjbot.org/content/vol91/issue10/images/large/abot-91-10-04-f01.jpeg) from Crane,
Herendeen & Friis 2004

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