Rhynia Temporal range: | |
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Reconstruction of Rhynia gwynne-vaughanii, redrawn after Kenrick & Crane (1997:101) [1] | |
Scientific classification ![]() | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Subdivision: | † Rhyniophytina |
Class: | † Rhyniopsida |
Order: | † Rhyniales |
Family: | † Rhyniaceae |
Genus: | † Rhynia Kidst. & W.H.Lang (1917) |
Type species | |
R. gwynne-vaughanii Kidst. & W.H.Lang (1917) | |
Species | |
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Rhynia is a single-species genus of Devonian vascular plants. Rhynia gwynne-vaughanii was the sporophyte [2] 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.
Rhynia gwynne-vaughanii was first described as a new species by Robert Kidston and William H. Lang in 1917. [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. Rhynia was a vascular plant, and grew in association with other vascular plants such as Asteroxylon mackei , a probable ancestor of modern clubmosses (Lycopsida), and with pre-vascular plants such as Aglaophyton major , which is interpreted as basal to true vascular plants. [4]
Rhynia is thought to have had deciduous lateral branches, which it used to disperse laterally over the substrate [5] [6] and stands of the plant may therefore have been clonal populations.
Evidence of the gametophyte generation of Rhynia has been described in the form of crowded tufts of diminutive stems only a few mm in height, with the form genus name Remyophyton delicatum. [7] Like those of Aglaophyton major, [8] [9] Horneophyton lignieri [10] and Nothia aphylla [11] the gametophytes of Rhynia were dioicous, bearing male and female gametangia (antheridia and archegonia) on different axes. A significant finding is that the axes of the gametophytes were vascular, unlike almost all of the gametophytes of modern pteridophytes except for that of Psilotum . [12]
Two species of Rhynia were initially described by R. Kidston and W. H. Lang from the Rhynie chert bed: R. gwynne-vaughnii in 1917, [3] and R. major in 1920. [13] R. gwynne-vaughanii was named by Kidston and Lang in honour of their late friend and colleague, the botanist David Thomas Gwynne-Vaughan. [3]
A study of the vascular tissue of the two by David S. Edwards in 1986 lead to the conclusion that the cell walls of the water-conducting cells of R. major lacked the secondary thickening bars seen in the xylem of R. gwynne-vaughanii, and were more like the water-conducting hydroids of moss sporophytes. His conclusion was that R. gwynne-vaughanii belongs in the vascular plants, while R. major belongs among the bryophytes. Accordingly, he transferred it to a new genus Aglaophyton , leaving R. gwynne-vaughnii as the only known species of Rhynia. [14] Rhynia is the type genus for the rhyniophytes, established as the subdivision Rhyniophytina by Banks, [15] but since treated at various ranks.
In 2004, Crane et al. published a cladogram for the polysporangiophytes, in which Rhynia and the other Rhyniaceae are placed as basal vascular plants (tracheophyes). [16]
polysporangiophytes |
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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.
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.
Asteroxylon 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. Asteroxylon is considered the most basal member of the Lycopsida.
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.
Ornatifilum is an artificial form genus, which is used to categorise any small, branched filaments with external ornamentation.
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.
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.
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.
The Horneophytopsida, informally called horneophytes, are a class of extinct plants which consisted of branched stems without leaves, true roots or vascular tissue, found from the Late Silurian to the Early Devonian. They are the simplest known polysporangiophytes, i.e. plants with sporophytes bearing many spore-forming organs (sporangia) on branched stems. They were formerly classified among the rhyniophytes, but it was later found that some of the original members of the group had simple vascular tissue and others did not. The group has also been treated as the division Horneophyta.
Professor Dianne Edwards CBE, FRS, FRSE, FLS, FLSW is a palaeobotanist, who studies the colonisation of land by plants, and early land plant interactions.
Sciadophyton is a morphotaxon of lower Devonian plants known only from compression fossils. It is interpreted as the monoicous gametophyte of a vascular land plant, because its vascularised branches end in a cup-shaped structure bearing gametangia, both antheridia and archegonia, but little structural information is preserved at the cellular level. It formed rosettes of stems, which may have radiated from a basal gametophytic corm-like thallus or from a central 'stem' or even from a root system, although there is not enough evidence to discriminate between these possibilities.
Psilophytopsida is a now obsolete class containing one order, Psilophytales, which was previously used to classify a number of extinct plants which are now placed elsewhere. The class was established in 1917, under the name Psilophyta, with only three genera for a group of fossil plants from the Upper Silurian and Devonian periods which lack true roots and leaves, but have a vascular system within a branching cylindrical stem. The living Psilotaceae, the whisk-ferns, were sometimes added to the class, which was then usually called Psilopsida. This classification is no longer in use.
The lowermost Upper Devonian fossil Pallaviciniites was for a time the oldest known liverwort until Metzgeriothallus was recovered from earlier Devonian strata.
Huvenia is a genus of extinct plants of the Early Devonian, found in slate deposits of the Rhenish Massif. The sporophyte generation consisted of leafless stems (axes), which appear to be flattened, and which branch dichotomously. The strand of conducting tissue contains simple tracheids, making this a vascular plant (tracheophyte). The sporangia are borne on the ends of short branching stems (sporangiophores) rather than terminating main stems as in some other early land plants. Sporangia appear to be twisted, but it is not clear whether this feature was present in life or developed after death.
Stockmansella is a genus of extinct plants of the Middle Devonian, fossils of which have been found in north-west Germany. The sporophyte generation consists of prostrate dichotomizing stems (axes) up to 10 cm long and around 3mm wide, which at intervals produce narrower smooth upright stems. These bear sporangia on short lateral branches (sporangiophores). The prostrate stems have bulges from which rhizoids form. Both prostrate and upright stems have a central strand of conducting tissue which contains simple tracheids, so that Stockmansella is a vascular plant.
Renalia is a genus of extinct vascular plants from the Early Devonian. It was first described in 1976 from compressed fossils in the Battery Point Formation. It is difficult to reconstruct the original form of the complete plant, but it appears to have consisted of leafless branching stems whose side branches had sporangia at their tips. It is regarded as an early relative of the lycophytes.
Sartilmania is a genus of extinct vascular plants of the Early Devonian. Fossils were found on the Sart Tilman campus of the University of Liège, Belgium.
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.
Adoketophyton is a genus of extinct vascular plants of the Early Devonian. The plant was first described in 1977 based on fossil specimens from the Posongchong Formation, Wenshan district, Yunnan, China. These were originally named Zosterophyllum subverticillatum; later the species was transferred to a new genus as Adoketophyton subverticillatum. One cladistic analysis suggested that it is a lycophyte, related to the zosterophylls. Other researchers regard its placement within the vascular plants as uncertain.
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, N. aphylla.