- Cooksonia
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Cooksonia
Temporal range: Wenlock to Early Devonian[1][2]
A cartoon of Cooksonia, reconstructed with non-photosynthetic axes, dependent on its gametophyte, as per Boyce (2008) Scientific classification 
Kingdom: Plantae Subkingdom: Embryophyta clade: Polysporangiophyta Division: Tracheophyta Genus: †Cooksonia
W.H.Lang (1937) emend. Gonez & Gerrienne (2010)[3]Species - C. pertoni W.H.Lang (1937)
- C. hemisphaerica W.H.Lang (1937)
- C. cambrensis D.Edwards (1979)
- C. bohemica Schweitzer (1980)
- C. paranensis Gerrienne et al. (2001)
- C. banksii Habgood et al. (2002)
Cooksonia is an extinct grouping of primitive land plants. The earliest Cooksonia date from the middle of the Silurian (the Wenlock epoch);[1] the group continues to be an important component of the flora until the Early Devonian, a total time span of 428 to 398 million years ago. For historical reasons, while Cooksonia fossils are distributed globally, most type specimens come from Britain.
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 an unpreserved rhizome.[2] They had a simple stalk, that branched dichotomously a few times. Each branch ended in a sporangium, a rounded, spore-bearing structure. The original description of the genus by Lang described the sporangia as terminal on the stems and wider than high. 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.[4] Other Cooksonia species lacked such conducting tissue.
Cooksonia specimens occur in a range of sizes, and vary in stem width from about 0.03 mm to 3 mm.[2] 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.[2] The organisms probably exhibited determinate growth (i.e. stems did not grow further after producing sporangia).[2]
Some Cooksonia species can be shown to bear 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.[2] 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. pertonii, 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. pertonii. 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. pertonii, differs in having sporangia whose 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]
Physiology
While reconstructions traditionally depict Cooksonia as a green and red, photosynthesising, self-sufficient stem, it is likely that at least some fossils instead preserve a sporophyte generation which was dependent on a gametophyte for its nutrition – a relationship that occurs in modern mosses and liverworts.[2]
However, no fossil evidence of a gametophyte of Cooksonia has been discovered to date. 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.[2] Further, the axis thickness is what would be expected if its sole role was to support a sporangium.[2] 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.[2]
The widths of Cooksonia fossils span an order of magnitude; while the smaller ones could not possibly be self-sufficient, the larger axes could, which provides a possible illustration of the evolution of an independent sporophyte generation.[2]
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.[5] There were originally two species, Cooksonia pertonii and C. hemisphaerica.[5] The genus was defined as having narrow leafless stems (axes), which branched dichotomously, with terminal sporangia that were "short and wide". There was a 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 whose sporangia were not more-or-less trumpet-shaped. As emended by Gonez and Gerrienne, Cooksonia has the following species:
- Cooksonia pertoni – the type species designated by Gonez & Gerrienne
- Cooksonia paranensis
- Cooksonia banksii
Three further species are considered doubtful because of the poor preservation of the specimens, but are left in the genus:
- Cooksonia hemisphaerica
- Cooksonia cambrensis
- Cooksonia bohemica
Two species are excluded from the genus: C. caledonica (which was later moved to a new genus Aberlemnia[6]) and the less well-preserved C. crassiparietilis. These 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.[7]
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[8]), supplemented by further information on Cooksonia species resulting from the authors' own research.[3]
polysporangiophytes tracheophytes Cooksonia hemisphaerica
Paratracheophytes
Cooksonia banksii, C. bohemica, C. cambrensis, C. paranensis, C. pertonii
Sartilmania, Yunia, Uskiella
Renalia, Cooksonia crassiparietilis, C. caledonica (=Aberlemnia caledonica)
lycophytes
euphyllophytes
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[6]). 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]
See also
References
- ^ a b 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
- ^ a b c d e f g h i j k l 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, http://paleobiol.geoscienceworld.org/cgi/content/full/34/2/179
- ^ a b c d e f g 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
- ^ 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
- ^ a b Lang, W.H. (1937), "On the plant-remains from the Downtonian of England and Wales", Phil. Trans. Royal Soc. B 227 (544): 245–291, Bibcode 1937RSPTB.227..245L, doi:10.1098/rstb.1937.0004
- ^ a b 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, doi:10.1016/j.revpalbo.2010.09.005
- ^ 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: Biological Sciences 353 (1378): 1983–2004, doi:10.1098/rstb.1998.0349, http://journals.royalsociety.org/index/PWH1Y6JA3EVM83CD.pdf
- ^ 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
External links
Categories:- Prehistoric plants
- Silurian life
- Devonian life
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