Barinophyte

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Barinophyte
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Barinophytes
Genera

See text.

The barinophytes are a group of extinct vascular plants (tracheophytes). Their relationship with other vascular plants is unclear. They have been treated as the separate class Barinophytopsida, the order Barinophytales of uncertain class and as a family or clade Barinophytaceae within the zosterophylls. They have also been considered to be possible lycopodiopsids. [1]

Contents

Description

Barinophyton is the type genus of the group; Protobarinophyton is similar. They were vascular plants with an exarch protostele. Plants consisted of alternatively arranged branches, apparently without leaves or enations, with their sporangia arranged in two rows on one-sided spike-like structures that developed on side shoots. Each sporangium was born on a curved bract-like appendage (a "sporangiferous appendage") and contained several thousand microspores, about 30–40 µm in diameter and about 30 megaspores, 410–560 µm in diameter, so that the plants were heterosporous. A similar pattern of spore sizes and numbers has been found in the progymnosperm Archaeopteris . [2]

Dibracophyton has been tentatively assigned to this group. It does have some outgrowths on the lower parts of the stems (axes), which raises the possibility that more fossil material might show that other barinophytes did not have naked stems. [1]

Taxonomy

Phylogeny

Two very different cladograms have been produced showing the evolutionary relationships of taxa assigned to the barinophytes. Kenrick and Crane in 1997 placed two species of Barinophyton and the genus Protobarinophyton in the Barinophytaceae in their Sawdoniales, well nested within the zosterophylls. [3] A summary cladogram produced by Crane et al. in 2004, shows Barinophyton and Protobarinophyton as 'core' zosterophylls. [4]

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

Taylor et al. in 2009 did not present a cladogram, but considered the barinophytes to be possible lycopsids rather than zosterophylls. [2] By contrast, Hao and Xue in 2013 do not even place Dibracophyton , which they consider to be a possible barinophyte, within the lycopodiopsids, instead putting it between this group and the euphyllophytes. [5]

Lycopodiopsida

Yunia

Dibracophyton (possible barinophyte)

euphyllophytes

Genera

Genera that are placed, or tentatively placed, in the barinophytes (with the group treated at different ranks) include: [1] [6] [3] [7] [8]

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.

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

Zosterophyllum was a genus of Silurian-Devonian vascular land plants with naked branching axes on which usually kidney-shaped sporangia were arranged in lateral positions. It is the type genus for the group known as zosterophylls, thought to be part of the lineage from which modern lycophytes evolved. More than 20 species have been described.

<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.

Krithodeophyton is a genus of lower Devonian plant with branching axes. It is considered to be a barinophyte.

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

Yunia is a genus of extinct vascular plants from the Early Devonian. It was first described from the Posongchong Formation of Yunnan, China. The leafless plant consisted of spiny stems, some 2 to 5 cm wide, which branched dichotomously at wide angles in a cruciate arrangement. Each stem contained vascular tissue with one or two strands of protoxylem. The spore-forming organs (sporangia) were elongated and borne on short stalks. The spores had a relatively smooth sculptural pattern and were trilete.

<i>Renalia</i> Extinct genus of vascular plants

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.

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.

Gumuia is a genus of extinct vascular plants of the Early Devonian. The genus was first described in 1989 based on fossil specimens from the Posongchong Formation, Wenshan district, Yunnan, China.

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

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.

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.

Distichophytum is a genus of extinct vascular plants of the Late Silurian (Ludfordian) to Early Devonian (Emsian), around 426 to 393 million years ago. The genus has a tangled taxonomic history, also being known as Bucheria and Rebuchia.

<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>Barinophyton</i> Extinct genus of plants

Barinophyton was a genus of early land plant with branching axes. It is placed in a group of early vascular plants (tracheophytes), the barinophytes, a group that has been given various ranks and scientific names. Known fossils are of Devonian to Carboniferous age.

Protobarinophyton was a genus of Silu-Devonian land plant with branching axes. It is placed in a group of early vascular plants (tracheophytes), the barinophytes, a group that has been given various ranks and scientific names.

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.

Demersatheca is a genus of extinct vascular plants of the Early Devonian. Fossils were first found in the Posongchong Formation of eastern Yunnan, China. The plant had smooth leafless stems at least 1 mm in diameter, but only regions which bore spore-forming organs or sporangia are well-known. Sporangia were borne in 'spikes' or strobili, at least 40 mm long; one had 32 sporangia. Sporangia were arranged in four rows, two sporangia being opposite to one another on the stem with the next two being at right angles. Each sporangium consisted of two 'valves' which opened at the top to release their spores. A particular feature of Demersatheca which distinguishes it from other zosterophylls is that the stalk-less sporangia were sunken into the stem of the spike, so that the outer valve was flush with the surface.

Gosslingiales is an order of extinct zosterophylls. The zosterophylls were among the first vascular plants in the fossil record, and share an ancestor with the living lycophytes. The group has been divided up in various ways. Hao and Xue in 2013 used the presence or absence of terminal sporangia as a major dividing feature. The order Zosterophyllales was used for species with terminal as well as lateral sporangia, which were considered to have determinate growth, with their sporangia generally being arranged in spikes. The paraphyletic order Gosslingiales was used for species without terminal sporangia, which were considered to have indeterminate growth, with fertile branches generally circinate. Species assignable to the Gosslingiales made up about 9% of all confirmed species in the Early Devonian flora.

References

  1. 1 2 3 Hao & Xue (2013), p. 55.
  2. 1 2 Taylor, T.N.; Taylor, E.L. & Krings, M. (2009), Paleobotany, The Biology and Evolution of Fossil Plants (2nd ed.), Amsterdam; Boston: Academic Press, pp. 325–326, ISBN   978-0-12-373972-8
  3. 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, p. 172, ISBN   978-1-56098-730-7
  4. 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
  5. Hao & Xue (2013), p. 246.
  6. Hao & Xue (2013), p. 329.
  7. Brauer, D.F. (1981), "Heterosporous, barinophytacean plants from the Upper Devonian of North America and a discussion of the possible affinities of the Barinophytaceae", Review of Palaeobotany and Palynology, 33 (2–4): 347–362, doi:10.1016/0034-6667(81)90092-0
  8. Cichan, M.; Taylor, T. & Brauer, D. (1984), "Ultrastructural studies of in situ Devonian spores: Protobarinophyton pennsylvanicum Brauer", Review of Palaeobotany and Palynology, 41 (3–4): 167, doi:10.1016/0034-6667(84)90044-7

Bibliography