Contents 1 Description 2 Taxonomy 2.1 Phylogeny 2.2 Subdivision 3 Ecology 4 See also 5 References 6 Bibliography 7 External links

Description[edit] Ferns and lycophytes (pteridophytes) are free-sporing vascular plants that share a unique life cycle with independent gametophyte and sporophyte phases that are frequently discussed together due to common characteristics, including vascular plant apomorphies (e.g., vascular tissue) and land plant plesiomorphies (e.g., spore dispersal and the absence of seeds).[1] [2]

Taxonomy[edit] Phylogeny[edit] Of the pteridophytes, ferns account for nearly 90% of the extant diversity.[2] Smith et al. (2006), the first higher-level pteridophyte classification published in the molecular phylogenetic era, considered the ferns as monilophytes, as follows:[3] Division Tracheophyta (tracheophytes) - vascular plants Sub division Euphyllophytina (euphyllophytes) Infradivision Moniliformopses (monilophytes) Infradivision Spermatophyta - seed plants, ~260,000 species Subdivision Lycopodiophyta (lycophytes) - less than 1% of extant vascular plants where the monilophytes comprise about 9,000 species, including horsetails (Equisetaceae), whisk ferns (Psilotaceae), and all eusporangiate and all leptosporangiate ferns. Historically both lycophytes and monilophytes were grouped together as pteridophytes (ferns and fern allies) on the basis of being spore-bearing ("seed-free"). In Smith's molecular phylogenetic study the ferns are characterised by lateral root origin in the endodermis, usually mesarch protoxylem in shoots, a pseudoendospore, plasmodial tapetum, and sperm cells with 30-1000 flagella.[3] The term "moniliform" as in Moniliformopses and monilophytes means "bead-shaped" and was introduced by Kenrick and Crane (1997)[4] as a scientific replacement for "fern" (including Equisetaceae) and became established by Pryer et al. (2004).[5] Christenhusz and Chase (2014) in their review of classification schemes provide a critique of this usage, which they discouraged as irrational. In fact the alternative name Filicopsida was already in use.[6] By comparison "lycopod" or lycophyte (club moss) means wolf-plant. The term "fern ally" included under Pteridophyta generally refers to vascular spore-bearing plants that are not ferns, including lycopods, horsetails, whisk ferns and water ferns (Marsileaceae, Salviniaceae and Ceratopteris), and even a much wider range of taxa. This is not a natural grouping but rather a convenient term for non-fern, and is also discouraged, as is eusporangiate for non-leptosporangiate ferns.[7] However both Infradivision and Moniliformopses are also invalid names under the International Code of Botanical Nomenclature. Ferns, despite forming a monophyletic clade, are formally only considered as four classes (Psilotopsida; Equisetopsida; Marattiopsida; Polypodiopsida), 11 orders and 37 families, without assigning a higher taxonomic rank.[3] Furthermore, within the Polypodiopsida, the largest grouping, a number of informal clades were recognised, including leptosporangiates, core leptosporangiates, polypods (Polypodiales), and eupolypods (including Eupolypods I and Eupolypods II).[3] In 2014 Christenhusz and Chase, summarising the known knowledge at that time, treated this group as two separate unrelated taxa in a consensus classification;[7] Lycopodiophyta (lycopods) 1 subclass, 3 orders, each with one family, 5 genera, approx. 1,300 species Polypodiophyta (ferns) 4 sublasses, 11 orders, 21 families, approx. 212 genera, approx. 10,535 species Subclass Equisetidae Warm. Subclass Ophioglossidae Klinge Subclass Marattiidae Klinge Subclass Polypodiidae Cronquist, Takht. & Zimmerm. These subclasses correspond to Smith's four classes, with Ophioglossidae corresponding to Psilotopsida. The two major groups previously included in Pteridophyta are phylogenetically related as follows:[7][8][9] Tracheophyta – vascular plants Lycopodiophyta Euphyllophyta Polypodiophyta – ferns Spermatophyta – seed plants Gymnospermae Angiospermae – flowering plants Subdivision[edit] Pteridophytes consist of two separate but related classes, whose nomenclature has varied.[3][10] The terminology used by the Pteridophyte Phylogeny Group (2016)[2] (with some synonyms) is used here: Classes, subclasses and orders Lycopodiopsida (lycophytes) Lycopodiidae (clubmosses) Selaginellidae (spikemosses and quillworts) Polypodiopsida (ferns) Equisetidae (horsetails, single genus Equisetum) Ophioglossidae (Psilotidae) Psilotales (One family, whisk ferns) Ophioglossales (One family, grape ferns) Marattiidae (six genera, marattioid ferns) Polypodiidae (leptosporangiate ferns, largest subclass, seven orders) In addition to these living groups, several groups of pteridophytes are now extinct and known only from fossils. These groups include the Rhyniopsida, Zosterophyllopsida, Trimerophytopsida, the Lepidodendrales and the Progymnospermopsida. Modern studies of the land plants agree that all pteridophytes share a common ancestor with seed plants. Therefore, pteridophytes do not form a clade but constitute a paraphyletic group.

Ecology[edit] Pteridophyte life cycle Just as with seed plants and mosses, the life cycle of pteridophytes involves alternation of generations. This means that a diploid generation (the sporophyte, which produces spores) is followed by a haploid generation (the gametophyte or prothallus, which produces gametes). Pteridophytes differ from mosses and seed plants in that both generations are independent and free-living, although the sporophyte is generally much larger and more conspicuous. The sexuality of pteridophyte gametophytes can be classified as follows: Dioicous: each individual gametophyte is either male (producing antheridia and hence sperm) or female (producing archegonia and hence egg cells). Monoicous: each individual gametophyte produces both antheridia and archegonia and can function both as a male and as a female. Protandrous: the antheridia mature before the archegonia (male first, then female). Protogynous: the archegonia mature before the antheridia (female first, then male). These terms are not the same as monoecious and dioecious, which refer to whether a seed plant's sporophyte bears both male and female gametophytes, i. e., produces both pollen and seeds, or just one of the sexes.

See also[edit] Embryophyte Fern ally Plant sexuality

References[edit] ^ Schneider & Schuettpelz 2016. ^ a b c Pteridophyte Phylogeny Group 2016. ^ a b c d e Smith et al.2006. ^ Kenrick & Crane 1997. ^ Pryer et al. 2004. ^ Kenrick & Crane 1997a. ^ a b c Christenhusz & Chase 2014. ^ Cantino et al. 2007. ^ Chase & Reveal 2009. ^ Kenrick & Crane 1996.

Bibliography[edit] Cantino, Philip D.; Doyle, James A.; Graham, Sean W.; Judd, Walter S.; Olmstead, Richard G.; Soltis, Douglas E.; Soltis, Pamela S.; Donoghue, Michael J. (1 August 2007). "Towards a Phylogenetic Nomenclature of Tracheophyta". Taxon. 56 (3): 822. doi:10.2307/25065865.  Christenhusz, M. J. M.; Zhang, X. C.; Schneider, H. (18 February 2011). "A linear sequence of extant families and genera of lycophytes and ferns" (PDF). Phytotaxa. 19 (1): 7. doi:10.11646/phytotaxa.19.1.2.  Christenhusz, Maarten J.M. & Chase, Mark W. (2014). "Trends and concepts in fern classification". Annals of Botany. 113 (9): 571–594. doi:10.1093/aob/mct299. PMC 3936591 . PMID 24532607.  Clark, James; Hidalgo, Oriane; Pellicer, Jaume; Liu, Hongmei; Marquardt, Jeannine; Robert, Yannis; Christenhusz, Maarten; Zhang, Shouzhou; Gibby, Mary; Leitch, Ilia J.; Schneider, Harald (May 2016). "Genome evolution of ferns: evidence for relative stasis of genome size across the fern phylogeny". New Phytologist. 210 (3): 1072–1082. doi:10.1111/nph.13833.  Chase, Mark W. & Reveal, James L. (2009). "A phylogenetic classification of the land plants to accompany APG III". Botanical Journal of the Linnean Society. 161: 122–127. doi:10.1111/j.1095-8339.2009.01002.x.  Gifford, Ernest M.; Foster, Adriance S. (1996). Morphology and evolution of vascular plants (3rd ed.). New York: Freeman. ISBN 0-7167-1946-0.  Kenrick, Paul; Crane, Peter (1996). "Embryophytes: Land plants". Tree of Life Web Project. Retrieved 19 April 2017.  Kenrick, Paul; Crane, Peter R. (4 September 1997). "The origin and early evolution of plants on land" (PDF). Nature. 389: 33–39.  Kenrick, Paul; Crane, Peter (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D.C.: Smithsonian Institution Press. ISBN 9781560987291.  Pryer, K. M.; Schuettpelz, E.; Wolf, P. G.; Schneider, H.; Smith, A. R.; Cranfill, R. (1 October 2004). "Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences". American Journal of Botany. 91 (10): 1582–1598. doi:10.3732/ajb.91.10.1582.  Pteridophyte Phylogeny Group (November 2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi:10.1111/jse.12229.  Ranker, Tom A.; Haufler, Christopher H. (2008). Biology and Evolution of Ferns and Lycophytes. Cambridge University Press. ISBN 978-0-521-87411-3.  Raven, Peter H.; Evert, Ray F.; Eichhorn, Susan E. (2005). Biology of plants (7th ed.). New York, NY: Freeman and Company. ISBN 0-7167-1007-2.  Schneider, Harald; Schuettpelz, Eric (November 2016). "Systematics and evolution of lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 561–562. doi:10.1111/jse.12231.  Smith, Alan R.; Kathleen M. Pryer; Eric Schuettpelz; Petra Korall; Harald Schneider; Paul G. Wolf (2006). "A classification for extant ferns" (PDF). Taxon. 55 (3): 705–731. doi:10.2307/25065646. JSTOR 25065646.  Pteridophyte Phylogeny Group (November 2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi:10.1111/jse.12229. 

External links[edit] Wikimedia Commons has media related to Pteridophyta. Wikispecies has information related to Pteridophyta British Pteridological Society Annual Review of Pteridological Research  "Pteridophyta". Encyclopædia Britannica (11th ed.). 1911.  Taxon identifiers Wd: Q178249 EoL: 39280889 ITIS: 564826 v t e Botany History of botany Subdisciplines Plant systematics Ethnobotany Paleobotany Plant anatomy Plant ecology Phytogeography Geobotany Flora Phytochemistry Plant pathology Bryology Phycology Floristics Dendrology Plant groups Algae Archaeplastida Bryophyte Non-vascular plants Vascular plants Spermatophytes Pteridophyte Gymnosperm Angiosperm Plant morphology (glossary) Plant cells Cell wall Phragmoplast Plastid Plasmodesma Vacuole Tissues Meristem Vascular tissue Vascular bundle Ground tissue Mesophyll Cork Wood Storage organs Vegetative Root Rhizoid Bulb Rhizome Shoot Stem Leaf Petiole Cataphyll Bud Sessility Reproductive (Flower) Flower development Inflorescence Umbel Raceme Bract Pedicellate Flower Whorl Floral symmetry Floral diagram Floral formula Receptacle Hypanthium (Floral cup) Perianth Tepal Petal Sepal Sporophyll Gynoecium Ovary Ovule Stigma Archegonium Androecium Stamen Staminode Pollen Tapetum Gynandrium Gametophyte Sporophyte Plant embryo Fruit Fruit anatomy Berry Capsule Seed Seed dispersal Endosperm Surface structures Epicuticular wax Plant cuticle Epidermis Stoma Nectary Trichome Prickle Plant physiology Materials Nutrition Photosynthesis Chlorophyll Plant hormone Transpiration Turgor pressure Bulk flow Aleurone Phytomelanin Sugar Sap Starch Cellulose Plant growth and habit Secondary growth Woody plants Herbaceous plants Habit Vines Lianas Shrubs Subshrubs Trees Succulent plants Reproduction Evolution Ecology Alternation of generations Sporangium Spore Microsporangia Microspore Megasporangium Megaspore Pollination Pollinators Pollen tube Double fertilization Germination Evolutionary development Evolutionary history timeline Hardiness zone Plant taxonomy History of plant systematics Herbarium Biological classification Botanical nomenclature Botanical name Correct name Author citation International Code of Nomenclature for algae, fungi, and plants (ICN) - for Cultivated Plants (ICNCP) Taxonomic rank International Association for Plant Taxonomy (IAPT) Plant taxonomy systems Cultivated plant taxonomy Citrus taxonomy cultigen cultivar Group grex Practice Agronomy Floriculture Forestry Horticulture Lists Related topics Botanical terms Botanists by author abbreviation Botanical expedition Category Portal WikiProject Retrieved from "" Categories: PteridophytaCryptogamsHidden categories: Wikipedia articles incorporating a citation from the 1911 Encyclopaedia Britannica with Wikisource reference

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