Lycopodiophyta

Lycopodiophyta Temporal range: 428–0 Ma PreЄ Є O S D C P T J K Pg N Silurian[1] to recent
Lycopodiella inundata
Scientific classification
Kingdom:	Plantae
(unranked):	Tracheophyta
Division:	Lycopodiophyta Cronquist, Takht. & W.Zimm.[2] [P.D. Cantino & M.J. Donoghue][3]
Classes
Lycopodiopsida - clubmosses Isoetopsida - spikemosses, quillworts, scale trees † Zosterophyllopsida - zosterophylls

The Division Lycopodiophyta (sometimes called Lycophyta or Lycopods) is a tracheophyte subdivision of the Kingdom Plantae. It is the oldest extant (living) vascular plant division at around 410 million years old.^[4]:99 These species reproduce by shedding spores and have macroscopic alternation of generations, although some are homosporous while others are heterosporous. Most members of Lycopodiophyta bear a protostele, and the sporophyte generation is dominant.^[5] They differ from all other vascular plants in having microphylls, leaves that have only a single vascular trace (vein) rather than the much more complex megaphylls found in ferns and seed plants.

Classification

There are around 1,200^[6] living (extant) species of Lycopodiophyta which are generally divided into three orders (Lycopodiales, Isoetales, and Selaginellales); in addition there are extinct groups. There is some variation in how the extant orders are grouped into classes: they may be put into a single class; they may be put into two classes, with the Isoetales and Selaginellales combined into one class;^[7] or they may be put into three classes, one order in each.^[8]:8 The system which uses two classes for extant species is:

• Class Lycopodiopsida – clubmosses and firmosses
• Class Isoetopsida – quillworts, scale trees, and spikemosses
• Class † Zosterophyllopsida – extinct zosterophylls.

The following phylogram shows a likely relationship between Lycopodiophyta orders.

Lycopodiophyta	Lycopodiopsida Lycopodiales Drepanophycales † Isoetopsida Selaginellales Lepidodendrales † Pleuromeiales † Isoetales

Evolution

The members of this division have a long evolutionary history, and fossils are abundant worldwide, especially in coal deposits. In fact, most known genera are extinct. The Silurian species Baragwanathia longifolia represents the earliest identifable Lycopodiophyta, while some Cooksonia seem to be related. Lycopodolica is another Silurian genus which appears to be an early member of this group.^[9]

Fossils ascribed to the Lycopodiophyta first appear in the Silurian period, along with a number of other vascular plants. Phylogenetic analysis places them at the base of the vascular plants; they are distinguished by their microphylls and by transverse dehiscence of their sporangia (as contrasted with longitudinal in other vascular plants). Sporangia of living species are borne on the upper surfaces of microphylls (called sporophylls). In some groups, these sporophylls are clustered into strobili.

During the Carboniferous Period, tree-like Lycopodiophyta (such as Lepidodendron) formed huge forests that dominated the landscape. The complex ecology of these tropical rainforests collapsed during the mid Pennsylvanian due to a change in climate.^[10]

Unlike modern trees, leaves grew out of the entire surface of the trunk and branches, but would fall off as the plant grew, leaving only a small cluster of leaves at the top. Their remains formed many fossil coal deposits. In Fossil Park, Glasgow, Scotland, fossilized Lycopodiophyta trees can be found in sandstone. The trees are marked with diamond-shaped scars where they once had leaves.

The Lycopodiophyta had their maximum diversity in the Upper Carboniferous, particularly tree-like Lepidodendron and Sigillaria, that dominated tropical wetlands. In Euramerica these became apparently extinct in the Late Pennsylvanian, as a result of a transition to a much drier climate, to give way to conifers, ferns and horsetails. In Cathaysia (now South China) tree-like Lycopodiophytes survived into the Permian. Nevertheless, lycopsids are rare in the Lopingian (latest Permian), but regained dominance in the Induan (earliest Triassic), particularly Pleuromeia. After the worldwide Permian–Triassic extinction event Lycopodiophyta pioneered the repopulation of habitats as opportunistic plants. The heterogeneity of the terrestrial plant communities increased markedly during the Middle Triassic when plant groups like sphenopsids, ferns, pteridosperms, cycadophytes, ginkgophytes and conifers resurfaced and diversified quickly.^[11]

Characteristics

Club-mosses are homosporous, but spike-mosses and quillworts are heterosporous, with female spores larger than the male, and gametophytes forming entirely within the spore walls.

The spores of Lycopodiophyta are highly flammable and so have been used in fireworks.^[12] Currently, huperzine, a chemical isolated from a Chinese clubmoss, is under investigation as a possible treatment for Alzheimer's disease.

Gallery

• 

  Lycopodites, an early lycopod-like fossil.

• 

  External mold of Lepidodendron from the Upper Carboniferous of Ohio.

• 

  Fossil in situ lycopsid, probably Sigillaria, with attached stigmarian roots.

• 

  Base of a fossil lycopsid showing connection with stigmarian roots.

References

1. Kenrick, Paul; Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D. C.: Smithsonian Institution Press. pp. 339–340. ISBN 1-56098-730-8.
2. Cronquist, A.; A. Takhtajan; W. Zimmermann (1966). "On the higher taxa of Embryobionta". Taxon (International Association for Plant Taxonomy (IAPT)) 15 (15): 129–134. doi:10.2307/1217531. JSTOR 1217531.
3. Cantino, Philip D.; James A. Doyle; Sean W. Graham; Walter S. Judd; Richard G. Olmstead; Douglas E. Soltis; Pamela S. Soltis; Michael J. Donoghue (2007). "Towards a phylogenetic nomenclature of Tracheophyta". Taxon 56 (3): E1–E44. doi:10.2307/25065865.
4. McElwain, Jenny C.; Willis, K. G.; Willis, Kathy; McElwain, J. C. (2002). The evolution of plants. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-850065-3.
5. Eichhorn, Evert, and Raven (2005). Biology of Plants, Seventh Edition. 381-388.
6. Callow, R. S.; Cook, Laurence Martin (1999). Genetic and evolutionary diversity: the sport of nature. Cheltenham: S. Thornes. p. 8. ISBN 0-7487-4336-7.
7. Yatsentyuk, S.P.; Valiejo-Roman, K.M.; Samigullin, T.H.; Wilkström, N.; Troitsky, A.V. (2001). "Evolution of Lycopodiaceae Inferred from Spacer Sequencing of Chloroplast rRNA Genes". Russian Journal of Genetics 37 (9): 1068–73. doi:10.1023/A:1011969716528.
8. "www.ncbi.nlm.nih.gov". Retrieved 2009-03-19.
9. Raymond, A.; Gensel, P. & Stein, W.E. (2006). "Phytogeography of Late Silurian macrofloras". Review of Palaeobotany and Palynology 142 (3-4): 165–192. doi:10.1016/j.revpalbo.2006.02.005.
10. Sahney, S., Benton, M.J. & Falcon-Lang, H.J. (2010). "Rainforest collapse triggered Pennsylvanian tetrapod diversification in Euramerica" (PDF). Geology 38 (12): 1079–1082. doi:10.1130/G31182.1.
11. Moisan, Philippe; Voigt, Sebastian (2013). "Lycopsids from the Madygen Lagerstätte (Middle to Late Triassic, Kyrgyzstan, Central Asia)". Review of Palaeobotany and Palynology 192: 42–64. doi:10.1016/j.revpalbo.2012.12.003. Retrieved 2015-03-20.
12. Cobb, B (1956) A Field Guide to Ferns and their related families: Northeastern and Central North America with a section on species also found in the British Isles and Western Europe (Peterson Field Guides), 215

External links

• Introduction to the Lycophyta from the University of California Museum of Paleontology
• Lycophytes
• Fossil Groves
• Paleo Plants