Nothia aphylla

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Nothia aphylla
Temporal range: Pragian
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Nothia.png
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Lycophytes
Plesion: Zosterophylls (?)
Genus: Nothia
A.G.Lyon ex El-Saadawy & Lacy [1]
Species:
N. aphylla
Binomial name
Nothia aphylla
Lyon ex El-Saadawy & Lacy [1]

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. [1]

Contents

History of discovery

Fossilized remains, including bare stems (axes) and detached spore-forming organs (sporangia), were first described by Kidston and Lang in 1920 from the Rhynie chert of Aberdeenshire, Scotland – rocks which are of Pragian age ( 411 to 408 million years ago). The fragments were considered to be parts of Asteroxylon mackiei. In 1964 Lyon described sporangia belonging to Asteroxylon mackiei and suggested that Kidston and Lang's specimens were a new species which he called Nothia aphylla. [2] The first full description based on further specimens was published in 1979. [3] A very detailed description was published in 2001, making this species one of the best-known early polysporangiophytes. [1]

Description

Reconstruction of the rhizomes of Nothia aphylla, based on Kerp, Hass & Mosbrugger 2001 Nothia aphylla rhizomes reconstruction.svg
Reconstruction of the rhizomes of Nothia aphylla, based on Kerp, Hass & Mosbrugger 2001
Stylized reconstruction of part of the fertile region of the aerial stem of Nothia aphylla, based on Kerp, Hass & Mosbrugger 2001 Nothia aphylla fertile stem reconstruction.svg
Stylized reconstruction of part of the fertile region of the aerial stem of Nothia aphylla, based on Kerp, Hass & Mosbrugger 2001

The sporophyte of Nothia aphylla consisted of thin underground and aerial stems (axes). The underground stems or rhizomes were up to 2 mm in diameter and branched laterally. The underside of the rhizomes had a longitudinal ridge from which unicellular rhizoids emerged. There were no true roots. At intervals the rhizomes turned upwards to emerge as upright stems. Around the region of the upwards bend, horizontal branches appeared at right angles to continue the growth of the rhizomes. The upright stems were generally less than 2.5 mm in diameter; a reconstruction suggests a height of around 20 cm. Aerial stems branched dichotomously in a three-dimensional pattern, with the last two sets of branches bearing sporangia. The sporangia were attached by short stalks at the end and along the sides of the stem, in a more-or-less spiral fashion. The stalks curved upwards so that the sporangia were roughly upright. Spores were released through a longitudinal slit which appeared at the apex of the sporangia. The spores were trilete and around 65 μm in diameter. [4] Both the horizontal rhizomes and the vertical stems had vascular tissue which formed a central core (protostele), and is described as having centrarch development. The precise structure of the vascular core varied between the rhizomes and the aerial stems. [1] [3]

Although Nothia aphylla was leafless, its aerial stems were covered with 'emergences': bumps on the stems which were oval, around 0.3 mm high and 0.7 to 1.2 mm long by up to 0.5 mm wide. The emergences were formed by existing cells expanding, not by extra cells being produced as in other Early Devonian zosterophylls and trimerophytes. Each emergence had a single stoma. The density of the emergences varied so that in regions which had them there were between 3 and 5 or more stomata per mm2. The sporangia did not have emergences, although there were a few stomata. [1]

Kerp et al. suggest that Nothia aphylla was a geophyte which inhabited sandy soils and had a clonal life-style. The underground rhizomes persisted from year to year, continually spreading via lateral branches. The aerial stems appeared annually. They base this analysis on a number features, including evidence that the rhizomes were subterranean, and that rhizomes were still living when erect stems had decayed. [5]

The plant described as Kidstonophyton discoides is possibly the male gametophyte of Nothia aphylla. [6]

Taxonomy

The genus and species were first named by Lyon in 1964. [2] However, Nothia aphylla has been regarded as a nomen nudum since no description was published along with the name. Validation of the name has been variously considered to have been by Høeg in 1967, making the botanical authority "Lyon ex Høeg", [3] or by El-Saadawy and Lacy in 1979, making the authority "Lyon ex El-Saadawy & Lacy". [1]

Nothia has been placed in the group initially established by Banks as the subdivision Zosterophyllophytina or the class Zosterophyllopsida (zosterophylls). El-Saadawy and Lacy regard the plant as having affinities with both the rhyniophytes and the zosterophylls. [3] As discussed further below, Kerp et al. regard its taxonomic placement as unclear.

Phylogeny

A cladogram published in 2004 by Crane et al. places Nothia in a paraphyletic stem group of broadly defined "zosterophylls", basal to the lycopsids (living and extinct clubmosses and relatives). [7]

lycophytes
   

  Hicklingia

†basal groups

Adoketophyton , Discalis , Distichophytum (=Rebuchia), Gumuia , Huia , Zosterophyllum  myretonianum, Z. llanoveranum, Z. fertile

†'core'  zosterophylls

Zosterophyllum divaricatum, Tarella , Oricilla , Gosslingia , Hsua, Thrinkophyton , Protobarinophyton , Barinophyton  obscurum, B. citrulliforme, Sawdonia , Deheubarthia , Konioria , Anisophyton , Serrulacaulis , Crenaticaulis

†basal groups

Nothia , Zosterophyllum  deciduum

lycopsids

extant and extinct members

A detailed study of Nothia aphylla questions this positioning of the genus, concluding that its taxonomic placement remains unclear, and that the cladistic analyses of Kenrick and Crane (on which the above cladogram is based) have ignored "fundamental differences" between different kinds of emergences (protrusions from stems). Features of the vascular tissue of typical zosterophylls, such as characteristic thickenings of the cells which conduct water, are also absent in Nothia. [8] Earlier, El Saadawy and Lacey had concluded that Nothia was in some ways intermediate between the rhyniophytes and the zosterophylls. [3] Hao and Xue in 2013 listed the genus as a zosterophyll. [9]

Related Research Articles

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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.

<i>Psilotum</i> Genus of ferns in the family Psilotaceae

Psilotum is a genus of fern-like vascular plants. It is one of two genera in the family Psilotaceae commonly known as whisk ferns, the other being Tmesipteris. Plants in these two genera were once thought to be descended from the earliest surviving vascular plants, but more recent phylogenies place them as basal ferns, as a sister group to Ophioglossales. They lack true roots and leaves are very reduced, the stems being the organs containing photosynthetic and conducting tissue. There are only two species in Psilotum and a hybrid between the two. They differ from those in Tmesipteris in having stems with many branches and a synangium with three lobes rather than two.

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

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<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.

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

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

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<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

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

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

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Hicklingia is a genus of extinct plants of the Middle Devonian. Compressed specimens were first described in 1923 from the Old Red Sandstone of Scotland. Initially the genus was placed in the "rhyniophytes", but this group is defined as having terminal sporangia, and later work showed that the sporangia of Hicklingia were lateral rather than strictly terminal, so that it is now regarded as having affinities with the zosterophylls.

Huia is a genus of extinct vascular plants of the Early Devonian. The genus was first described in 1985 based on fossil specimens from the Posongchong Formation, Wenshan district, Yunnan, China.

Discalis is a genus of extinct vascular plants of the Early Devonian. The name is derived from the Greek δίσκος, referring to the disc-shaped sporangia. The genus was first described by Hao in 1989 based on fossil specimens from the Posongchong Formation, Wenshan district, Yunnan, China.

Dutoitia is a genus of Devonian rhyniophyte, named after the renowned South African geologist Alex du Toit. It is one of the earliest plants from Gondwana to colonize land. Its fossils were preserved in fine mudstones of the 400-million-year-old Bokkeveld and Witteberg Groups of South Africa. This erect, gracile plant is less than 10 cm high and very simple in structure. Its diminutive stems, which are devoid of leaflike appendages, branch in two and end in club- or cup-shaped sporangia, occasionally containing its reproductive spores. Stomata are present in the cuticle of their stems for gas exchange, and primitive cells inside the stems transported water from the roots to the aerial parts of the plant. Three species are recognized, D. pulchra Hoeg 1930, D. alfreda Plumstead 1967 and D. maraisia Plumstead 1967.

<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.

Wenshania is a genus of extinct vascular plants found in the Posongchong Formation, Yunnan, China, which is of Early Devonian age. Plants consisted of leafless stems with simple dichotomous branching, and bore spore-forming organs or sporangia all around the sides of stems. Wenshania is part of the broadly defined group of zosterophylls.

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

Macivera is a genus of extinct vascular plants. Fossils were found in sediments in Bathust Island, Nunavut, Canada, from the upper Silurian. The leafless stems (axes) branched dichotomously and were relatively thin, being between 0.7 and 1.0 mm wide. Spore-forming organs or sporangia, which were elliptical, being longer than wide, were borne on the end regions of stems. Macivera is considered to be a zosterophyll.

References

  1. 1 2 3 4 5 6 7 Kerp, H.; Hass, M.H. & Mosbrugger, V. (2001), "New Data on Nothia aphylla Lyon 1964 ex El-Saadawy et Lacey 1979, a Poorly Known Plant from the Lower Devonian Rhynie Chert", in Gensel, P.G. & Edwards, D. (eds.), Plants invade the Land : Evolutionary & Environmental Perspectives, New York: Columbia University Press, pp. 52–82, ISBN   978-0-231-11161-4
  2. 1 2 Lyon, A.G. (1964), "The probable fertile region of Asteroxylon mackiei K. and L.", Nature, 203 (4949): 1082–1083, Bibcode:1964Natur.203.1082L, doi:10.1038/2031082b0, S2CID   4292879
  3. 1 2 3 4 5 El-Saadawy, W.El-S. & Lacey, W.S. (1979), "Observations on Nothia aphylla Lyon ex Høeg", Review of Palaeobotany and Palynology, 27 (2): 119–147, doi:10.1016/0034-6667(79)90037-X
  4. Kerp, Hass & Mosbrugger 2001, particularly pp. 54–55
  5. Kerp, Hass & Mosbrugger 2001 , pp. 80–82
  6. Kerp, Hass & Mosbrugger 2001 , p. 55
  7. Crane, P.R.; Herendeen, P.; Friis, E.M. (2004). "Fossils and plant phylogeny". American Journal of Botany. 91 (10): 1683–99. doi: 10.3732/ajb.91.10.1683 . PMID   21652317.
  8. Kerp, Hass & Mosbrugger 2001 , pp. 78–79
  9. 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, p. 329, ISBN   978-7-03-036616-0 , retrieved 2019-10-25