Cooksonia

Last updated

Cooksonia
Temporal range: WenlockEarly Devonian [1] [2]
Cooksonia pertoni.png
A cartoon of Cooksonia, reconstructed with non-photosynthetic axes, dependent on its gametophyte, as per Boyce (2008)
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Stem group: Rhyniophytes
Form taxon: Cooksonioidea
Genus: Cooksonia
Lang 1937 emend. Gonez & Gerrienne 2010 [3] non Druce 1905
Type species
Cooksonia pertoni
Lang 1937
Species
  • C. paranensisGerrienne et al. 2001
  • C. pertoniLang 1937 [Note 1]
  •  ?C. acuminataMussa et al. 2002
  •  ?C. barrandeiLibertín et al. 2018
  •  ?C. cambrensisEdwards 1979
  •  ?C. degrezensisSenkevich
  •  ?C. downtonensisHeard 1939
  •  ?C. rusanoviiAnaniev 1960
  •  ?C. zhanyiensisLi & Cai 1978

Cooksonia is an extinct group of primitive land plants, treated as a genus, although probably not monophyletic. The earliest Cooksonia date from the middle of the Silurian (the Wenlock epoch); [1] the group continued to be an important component of the flora until the end of the Early Devonian, a total time span of 433 to 393 million years ago. While Cooksonia fossils are distributed globally, most type specimens come from Britain, where they were first discovered in 1937. [4] Cooksonia includes the oldest known plant to have a stem with vascular tissue and is thus a transitional form between the primitive non-vascular bryophytes and the vascular plants. [5]

Contents

Description

Only the sporophyte phase of Cooksonia is currently known (i.e. the phase which produces spores rather than gametes). Individuals were small, a few centimetres tall, and had a simple structure. They lacked leaves, flowers and roots—although it has been speculated that they grew from a rhizome that has not been preserved. [2] They had a simple stalk that branched dichotomously a few times. Each branch ended in a sporangium or spore-bearing capsule. In his original description of the genus, Lang described the sporangia as flattened, "with terminal sporangia that are short and wide", and in the species Cooksonia pertoni "considerably wider than high". [4] A 2010 review of the genus by Gonez and Gerrienne produced a tighter definition, which requires the sporangia to be more-or-less trumpet-shaped (as in the illustration), with a 'lid' or operculum which disintegrates to release the spores. [3]

Specimens of one species of Cooksonia have a dark stripe in the centre of their stalks, which has been interpreted as the earliest remains of water-carrying tissue. [6] Other Cooksonia species lacked such conducting tissue.

Cooksonia specimens occur in a range of sizes, and vary in stem width from about 0.3 mm to 3 mm. Specimens of different sizes were probably different species, not fragments of larger organisms: fossils occur in consistent size groupings, and sporangia and spore details are different in organisms of different sizes. The organisms probably exhibited determinate growth (i.e. stems did not grow further after producing sporangia). [2]

Some Cooksonia species bore stomata, which had a role in gas exchange; this was probably to assist in transpiration-driven transport of dissolved materials in the xylem, rather than primarily in photosynthesis, as suggested by their concentration at the tips of the axes. These clusterings of stomata are typically associated with a bulging in the axis at the neck of the sporangium, which may have contained photosynthetic tissue, reminiscent of some mosses. [2]

As the genus is circumscribed by Gonez and Gerrienne, there are six possible species. C. pertoni, [Note 1] C. paranensis and C. banksii are all relatively similar with flat-topped, trumpet-shaped sporangia; stems are somewhat narrower in C. paranensis than in C. pertoni. Only one specimen of C. bohemica is known. It has stouter, more branched stems; the original shape of the sporangia is unclear because of poor preservation. C. hemisphaerica, described from the same locality as C. pertoni, differs in having sporangia of which the tops, at least as preserved, are hemispherical rather than flat. C. cambrensis also has spherical sporangia, but without the gradual widening at the base characteristic of the other species. Preservation of the sporangia is again poor. [3] C. barrandei was described in 2018. [7]

Physiology

Reconstruction as an independent plant Cooksonia sp. - MUSE.jpg
Reconstruction as an independent plant

While reconstructions traditionally depict Cooksonia as a green and red, photosynthesising, self-sufficient stem, it is likely that at least some fossils are of a sporophyte generation that was dependent on a gametophyte for its nutrition – a relationship that occurs in modern mosses and liverworts. However, no fossil evidence of a gametophyte of Cooksonia has been discovered to date. [2]

The widths of Cooksonia fossils span an order of magnitude. Study of smaller Cooksonia fossils showed that once the tissue required to support the axes, protect them from desiccation, and transport water had been accounted for, no room remained for photosynthetic tissue, and the sporophyte may therefore have been dependent on the gametophyte. Further, the axis thickness is what would be expected if its sole role was to support a sporangium. It appears that, originally at least, the role of the axes in smaller species was solely to ensure continued spore dispersal, even if the axis desiccated. The potentially self-sufficient larger axes may represent the evolution of an independent sporophyte generation. [2]

In 2018, the sporophyte of a new species, Cooksonia barrandei, was described, from about 432 million years ago. It is the oldest-known megafossil of land plants, as of May 2018. It was sufficiently robust to pass Boyce's test for possible self-sufficiency. [2] Together with evidence that, unlike modern mosses and liverworts, hornwort sporophytes do have a degree of nutritional independence through photosynthesis, C. barrandei suggests that independent gametophyte and sporophyte generations could have been ancestral in land plants, rather than evolving later. [7]

Taxonomy

The first Cooksonia species were described by William Henry Lang in 1937 and named in honor of Isabel Cookson, with whom he had collaborated and who collected specimens of Cooksonia pertoni in Perton Quarry, Wales, in 1934. [4] There were originally two species, Cooksonia pertoni and C. hemisphaerica. [4] The genus was defined as having narrow leafless stems (axes), which branched dichotomously, with terminal sporangia that were "short and wide". There was a central vascular cylinder consisting of annular tracheids (water-conducting cells with thickened walls). Six other species were later added to the genus: C. crassiparietilis, C. caledonica, C. cambrensis, C. bohemica, C. paranensis and C. banksii. A review in 2010 concluded that the delineation of the genus was inaccurate and that some species needed to be removed; in particular those in which sporangia were not more-or-less trumpet-shaped. As amended by Gonez and Gerrienne, Cooksonia has the following species:

Seven further species are considered doubtful because of the poor preservation of the specimens, but are left in the genus:

Four species are excluded from the genus by Gonez and Gerrienne. Species that have been transferred or removed are:

C. caledonica and the less well-preserved C. crassiparietilis have sporangia which are composed of two 'valves', splitting to release their spores along a line opposite to where they are attached to the stem (i.e. distally). [3]

Phylogeny

For some years, it was suspected that Cooksonia and its species were poorly characterized. Thus four different kinds of spore, probably representing four different species, were found in sporangia originally identified as C. pertoni. [9]

A 2010 study of the genus produced the consensus cladogram shown below (some branches have been collapsed to reduce the size of the diagram). This was based on data from an earlier study (by Kenrick and Crane [10] ), supplemented by further information on Cooksonia species resulting from the authors' own research. [3]

polysporangiophytes

Horneophytopsida

Aglaophyton

tracheophytes

Cooksonia hemisphaerica

Paratracheophytes

Cooksonia banksii, C. bohemica, C. cambrensis, C. paranensis, C. pertoni

Sartilmania , Yunia , Uskiella

Renalia , Cooksonia crassiparietilis, C. caledonica (= Aberlemnia caledonica)

lycophytes

euphyllophytes (higher plants: ferns, horsetails and seed plants)

A more recent phylogeny by Hao and Xue from 2013:

Paratrachaeophytes

This confirms that the genus Cooksonia sensu Lang (1937) is polyphyletic. A core group of five species are placed together, unresolved between the euphyllophytes and the lycophytes. The poorly preserved C. hemisphaerica is placed as the most basal tracheophyte. Two other species, C. crassiparietilis and C. caledonica, are placed in the stem group of the lycophytes. These two species have been removed from Cooksonia sensu Gonez & Gerrienne (C. caledonica has since been placed in a new genus Aberlemnia [8] ). Both have sporangia which, although borne terminally rather than laterally, have a mechanism for releasing spores similar to those of the zosterophylls. [3]

A second cladistic analysis was carried out using only the three best preserved and thus best known species, C. pertoni, C. paranensis, and C. caledonica. The position of C. caledonica was confirmed, but C. pertoni and C. paranensis now formed a single clade more clearly related to the lycophytes than the euphyllophytes. [3]

Cooksonioids

Cooksonia and similar genera have been placed in a group called "cooksonioids". Originally the term was used for a group of plants fitting the general description of Cooksonia (i.e. simple plants with naked axes showing dichotomous branching and terminal sporangia), but with uncertain evidence of vascular tissue. [11] Boyce restricted the group to forms with axes usually less than 1 mm in diameter, and hence possibly not capable of independent growth. In addition to Cooksonia, he included genera such as Salopella , Tarrantia and Tortilicaulis . [2] Hue and Xao regarded cooksonioids as a group within the rhyniophytes with radially symmetrical sporangia of roughly the same height and width, and included Cooksonia pertoni, C. paranensis and C. hemisphaerica, but not C. crassiparietilis and Aberlemnia caledonica, as they had bilaterally symmetrical sporangia. [12]

See also

Notes

  1. 1 2 Two spellings are in use: the spelling used by the original author of the name, C. pertoni, and the spelling C. pertonii (e.g. in Bateman, R.M.; Crane, P.R.; Dimichele, W.A.; Kenrick, P.R.; Rowe, N.P.; Speck, T.; Stein, W.E. (1998), "Early Evolution of Land Plants: Phylogeny, Physiology, and Ecology of the Primary Terrestrial Radiation", Annual Review of Ecology and Systematics, 29 (1): 263–92, doi:10.1146/annurev.ecolsys.29.1.263 ). The second is appropriate in botanical Latin when an epithet relates to a person, according to Article 60.11 of the International Code of Nomenclature for algae, fungi, and plants.

Related Research Articles

<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">Embryophyte</span> Subclade of green plants, also known as land plants

The embryophytes are a clade of plants, also known as Embryophyta or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophytes have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of freshwater charophyte green algae as a sister taxon of Charophyceae, Coleochaetophyceae and Zygnematophyceae. Embryophytes consist of the bryophytes and the polysporangiophytes. Living embryophytes include hornworts, liverworts, mosses, lycophytes, ferns, gymnosperms and angiosperms. Embryophytes have diplobiontic life cycles.

<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.

<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">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>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.

<span class="mw-page-title-main">Horneophytopsida</span> Extinct class of leafless plants

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.

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.

<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.

<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.

Junggaria was a genus of rhyniophyte-like land plants known from fossils found in China in Upper Silurian strata. It bore leafless dichotomously or pseudomonopodially branching axes, some of which ended in spore-forming organs or sporangia of complex shape. The genus Cooksonella, found in Kazakhstan from deposits of a similar age, is considered to be an illegitimate synonym.

Danziella is a genus of extinct vascular plants of the Early Devonian. Fossils found in the Artois region of northern France were first described as Zosterophyllum artesianum, but a later review by Edwards showed that they did not fit the circumscription of that genus.

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

Aberlemnia is a genus of extinct vascular plants of the Early Devonian, which consisted of leafless stems with terminal spore-forming organs (sporangia). Fossils found in Scotland were initially described as Cooksonia caledonica. A later review, which included new and more complete fossils from Brazil, showed that the specimens did not fit the circumscription of the genus Cooksonia; accordingly a new genus Aberlemnia was proposed.

<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.

<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.

Hollandophyton is a genus of extinct plants known from fossils found in Shropshire, England, in rocks of upper Silurian age. The specimens are fragmentary, consisting of leafless stems (axes) which branched dichotomously and bore kidney-shaped spore-forming organs or sporangia, apparently at their tips. The internal structure of the stems is unknown.

<i>Asterotheca</i> Genus of plants

Asterotheca is a genus of seedless, spore-bearing, vascularized ferns dating from the Carboniferous of the Paleozoic to the Triassic of the Mesozoic.

References

  1. 1 2 Edwards, D. & Feehan, J. (1980), "Records of Cooksonia-type sporangia from late Wenlock strata in Ireland", Nature, 287 (5777): 41–42, Bibcode:1980Natur.287...41E, doi:10.1038/287041a0, S2CID   7958927
  2. 1 2 3 4 5 6 7 8 Boyce, C. Kevin (2008), "How green was Cooksonia? The importance of size in understanding the early evolution of physiology in the vascular plant lineage", Paleobiology, 34 (2): 179–194, doi:10.1666/0094-8373(2008)034[0179:HGWCTI]2.0.CO;2, S2CID   36688488
  3. 1 2 3 4 5 6 7 Gonez, P. & Gerrienne, P. (2010a), "A New Definition and a Lectotypification of the Genus Cooksonia Lang 1937", International Journal of Plant Sciences, 171 (2): 199–215, doi:10.1086/648988, S2CID   84956576
  4. 1 2 3 4 Lang, W.H. (1937), "On the plant-remains from the Downtonian of England and Wales", Philosophical Transactions of the Royal Society B , 227 (544): 245–291, Bibcode:1937RSPTB.227..245L, doi:10.1098/rstb.1937.0004
  5. Freeman, Scott; Herron, Jon C. (2004). Evolutionary analysis (3rd ed.). Upper Saddle River, NJ: Pearson Education. p. 816. ISBN   978-0-13-101859-4.
  6. Edwards, D.; Davies, K.L. & Axe, L. (1992), "A vascular conducting strand in the early land plant Cooksonia", Nature, 357 (6380): 683–685, Bibcode:1992Natur.357..683E, doi:10.1038/357683a0, S2CID   4264332
  7. 1 2 3 Libertín, Milan; Kvaček, Jiří; Bek, Jiří; Žárský, Viktor & Štorch, Petr (2018), "Sporophytes of polysporangiate land plants from the early Silurian period may have been photosynthetically autonomous", Nature Plants, 4 (5): 269–271, Bibcode:2018NatPl...4..269L, doi:10.1038/s41477-018-0140-y, PMID   29725100, S2CID   19151297
  8. 1 2 Gonez, P. & Gerrienne, P. (2010b), "Aberlemnia caledonica gen. et comb. nov., a new name for Cooksonia caledonica Edwards 1970", Review of Palaeobotany and Palynology, 163 (1–2): 64–72, Bibcode:2010RPaPa.163...64G, doi:10.1016/j.revpalbo.2010.09.005
  9. Wellman, C. H.; Edwards, D. & Axe, L. (1998), "Ultrastructure of laevigate hilate spores in sporangia and spore masses from the Upper Silurian and Lower Devonian of the Welsh Borderland", Philosophical Transactions of the Royal Society B , 353 (1378): 1983–2004, doi:10.1098/rstb.1998.0349, PMC   1692433
  10. Kenrick, Paul & Crane, Peter R. (1997a), The Origin and Early Diversification of Land Plants: A Cladistic Study, Washington, D.C.: Smithsonian Institution Press, ISBN   978-1-56098-730-7
  11. Taylor, Thomas N. (1988), "The origin of land plants: some answers, more questions", Taxon, 37 (4): 805–833, doi:10.2307/1222087, JSTOR   1222087 , cited in Boyce (2008)
  12. 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. 45, 47, ISBN   978-7-03-036616-0 , retrieved 2019-10-25