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And PD Cantino], Converted Cladename Gnetophyta C. E. Bessey 1907: 323 (as "Gnetales" Hoogland and Reveal, 2005) [M. J. Donoghue, J. A. Doyle, and P. D. Cantino], converted clade name Registration Number: 47 and Donoghue, 1986; Doyle, 1996), compound microsporangiate strobili, striate pollen (modi- The smallest crown clade contain- fied to echinate in Gnetum; Yao et al., 2004), Definition: with Linnaeus 1767, Ephedra di tube, and meristem Gnetum gnemon micropylar apical ing Welwitschia mirabilis one tunica layer (Doyle and Donoghue, 1986, Linnaeus 1753, and tachya Serbet, 1994; 1862. This is a minimum-crown- 1992; Crane, 1988; Rothwell and 1. D. Hooker definition: min dade definition. Abbreviated Doyle, 2006). Linnaeus 1767 & crown V (Gnetum gnemon Welwitschia and Gnetales (see Ephedra distachya Linnaeus 1753 & Synonyms: Gnetopsida mirabilis J. D. Hooker 1862). Comments); also Gnetidae (Chase and Reveal, four 2009) and Gnetatae (Kubitzki, 1990). All with the same Etymology: Derived from Gnetum (the name are approximate synonyms of an included genus) and the Greek phyto, composition. meaning plant. The name Gnetum is appar- Comments: The of Gnetophyta is derived from the same word as the specific monophyly ently both molecular and the vernacu- strongly supported by epithet gnemon (i.e., genemo, very Molucca analyses (Crane, 1985, 1988; lar name for Gnetum gnemon in the morphological and 1986, 1992; Doyle et Islands; Markgraf, 1951). Doyle Donoghue, al., 1994; Rothwell and Serbet, 1994; Doyle, 1996, 2006; Rydin et al., 2002; Soltis et al., Reference Phylogeny: The primary reference also2002; and Mathews, 2004; Hilton is et al. (2002: Fig. 1). See Burleigh phylogeny Rydin 2010, 2011; Lu Hou et al. (2015: Fig. 2). and Bateman, 2006; Zhong, et The names et al., 2014; Wickett al., 2014). and Gnetales are m Gnetopsida, widely Composition: Ephedra (40 species), Gnetophyta, to this clade (Gnetidae and Gnetatae and Welwitschia applied 50species), (1 species) (species much less frequently). Here and previously numbers from 2008). For Mabberley, descrip we have for the tions of these three taxa, see Kubitzki (1990). (Cantino et al., 2007), opted name Gnetales is -phyta ending even though the used, because, although -phyta Synapomorphies more frequently Diagnostic Apomorphies: the rank of phylum (divi- cative to other crown clades include mu is used to designate nomenclature, it tiple axillary buds, opposite phyllotaxy, er- sion) in botanical rank-based whereas and -ales nal ovules with one or two outer envelopes also means "plants," -opsida other than rank. ELVd from opposite primordia, basally fused have no meaning indicating microsporophylls with terminal microspo dngla, vessels in xylem (assuming these are Literature Cited homologous with angiosperm vessels, as of Univ E. 1907. A synopsis plant phyla. inferred in Bessey, C. even in: analyses which giosperms 7:275-373. and Stud., University of Nebraska, gnetophytes are exta sister groups: Doyle Gnetophyta Burleigh, J. G., and S. Mathews. 2004. Phylogenetic Lu, Y., J.-H. Ran, D.-M. Guo, Z.-Y. Yang, and signal in nucleotide data from seed plants: X.-Q. Wang. 2014. Phylogeny and diver- implications for resolving the seed plant tree of gence times of gymnosperms inferred from life. Am. J. Bot. 91:1599-1613. single-copy nuclear genes. PLOS ONE Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. 99):e107679. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, Mabberley, D. J. 2008. Mabberley's Plant-Book. 3rd and M. J. Donoghue. 2007. Towards a phylo- edition. Cambridge University Press, Cambridge, genetic nomenclature of Tracheophyta. Taxon UK. 56:822-846 and E1-E44. Markgraf, F. 1951. Gnetaceae. Pp. 336-347 in Flora Chase, M. W., and J. L. Reveal. 2009. A phyloge Malesiana, Series 1, Vol. 4(3). (C. G. G. J. van netic classification of the land plants to accom- Steenis, ed.). Noordhof-Kolf, Djakarta. pany APG III. Bot. J. Linn. Soc. 161:122-127. Rothwell, G. W., and R. Serbet. 1994. Lignophyte Crane, P. R. 1985. Phylogenetic analysis of seed phylogeny and the evolution of spermato- plants and the origin of angiosperms. Ann. phytes: a numerical cladistic analysis. Syst. Bot Mo. Bot. Gard. 72:716-793. 19:443-482. Crane, PR 1988. Major clades and relationships in the Rydin, C., M. Källersjö, and E. Friis. 2002. Seed "higher gymnosperms. Pp. 218-272 in Origin plant relationships and the systematic position and Evolution of Gymnosperms (C. B. Beck, ed.). of Gnetales based on nuclear and chloroplast Columbia University Press, New York. DNA: conflicting data, rooting problems, and Doyle, J. A. 1996. Seed plant phylogeny and the the monophyly of conifers. Int. J. Plant Sci. relationships of Gnetales. Int. J. Plant Sci. 157 163:197-214. (suppl.):$3-s39. oltis, D. E., P. S. Soltis, and M. J. Zanis. 2002. Doyle, J. A. 2006. Seed ferns and the origin of Phylogeny of seed plants based on evidence angiosperms. J. Torrey Bot. Soc. 133:169-209. from eight genes. Am. J. Bot. 89:1670-1681. Doyle, J. A., and M. J. Donoghue. 1986. Seed plant Wickett, N. J., S. Mirarab, N. Nguyen, T. phylogeny and the origin of angiosperms: an Warnow, E. Carpenter, N. experimental cladistic approach. Bot. Reu Matasci, S. Ayyampalayam, M. S. Barker, J. G. 52:321-431. Burleigh, M. A. Gitzendanner, B. R. Ruhfel, Doyle, J. A., and M. J. Donoghue, 1992. Fossils E. Wafula, J. P. Der, S. W. Graham, S. and seed plant phylogeny reanalyzed. Brittonia Mathews, M. Melkonian, D. E. Soltis, P. 44:89-106. S. Soltis, N. W. Miles, C. J. Rothfels, L. Doyle, J. A., M. J. Donoghue, and E. A. Zimmer. Pokorny, A. J. Shaw, L. DeGironimo, D. 1994. Integration of and ribo- morphological W. Stevenson, B. C. B. somal RNA data on Surek, J. Villarreal, the origin of angiosperms. Roure, H. Philippe, C. W. dePamphilis, T. Ann. Mo. Bot. Gard. 81:419-450. Chen, M. K. Deyholos, R. S. Baucom, T. Hilton, J., and R. M. Bateman. 2006. Pteridosperms M. M. M. are the Kutchan, J. Y. backbone of seed-plant J. Augustin, Wang, Torrey Bot. Soc. 133:119-168. phylogeny. Zhang, Z. Tian, Z. Yan, X. Wu, X. Sun, G. K. Wong, and J. Leebens-Mack. 2014. Hou, C., A. M. Humphreys, O. Thureborn, and C. 2015. New Phylotranscriptomic analysis of the origin Rydin. insights into the and lutionary history of Gnetum (Gnetales). Taxonevo early diversification of land plants. Proc 64:239-252. Natl. Acad. Sci. USA 111:E4859-E4868. Yao, Y., Y. B. Kubitzki, K. 1990. Xi, Geng, and C. Li. 2004. The exine Gnetatae. Pp. 378-391 in Families and ultrastructure of pollen in Gnetum Genera of Vascular Plants, Vol. grains U. (Gnetaceae) from China and its bearing on the 1 (K. Kramer and P. S. vol. Green, eds.). with relationship the ANITA group. Springer-Verlag, Berlin. Linn. Soc. 146:415-425. Bot. 266 Gnetophyta V. V. D. 7hong, B., 0. Deusch, Goremykin, Penny, James A. Doyle; Department of Evolution and S. V. P. J. Biggs, R. A. Atherton, Nikiforova, Ecology; University of California; Davis, CA and P. J. Lockhart. 2011. Systematic errorin 95616, USA. Email: [email protected]. seed plant phylogenomics. Genome Biol. Evol Philip D. Cantino; Department of Environmental 3:1340-1348. and Plant Biology; Ohio University; Athens, Y. and M. Zhong, B., T Yonezawa, Zhong, OH 45701, USA. Email: [email protected]. Hasegawa. 2010. The position of Gnetales among seed plants: overcoming pitfalls of Date Accepted: 29 August 2011; updated 09 Mol. Biol. chloroplast phylogenomics. Evol February 2018 27:2855-2863. Primary Editor: Kevin de Queiroz Authors Michael J. Donoghue; Department of Ecology and Evolutionary Biology; Yale University: PO. Box 208106;: New Haven, CT 06520, USA. Email: [email protected]. 267 Mesangiosperm_ae M. J. Donoghue, J. A. Doyle, and P. o. cantino in P. D. Cantmo et al. (2007): 834 [M. J. Donoghue, J. A. Doyle, and P. D. Cantine], converted clade name Registration Number: 65 ascidiate carpels sealed by secretion, compara- ble to those of Amborella trichopoda and other Definition: The largest crown clade contain- members of what Qiu et al. (1999) called the ing Platanus occidentalis Linnaeus 1753 but ''ANITA'' grade (i.e., Amborella, Nymphaeales, not Amborella trichopoda Baillon 1869 and and Austrobaileyales, later extended to include Nymphaea odorata Aiton 1789 (Nymphaeales) Hydatellaceae: Saarela et al., 2007). Most and Austrobaileya scandens C. T. White 1933 molecular analyses have indicated that (Austrobaileyales). This is a maximum-crown- Chloranthaceae and Ceratophyllum are nested clade definition. Abbreviated definition: max within Mesangiospermae, and with some topolo- crown V (Platanus occidentalis Linnaeus 1753 ~ gies (Endress and Doyle, 2009: Fig. 8B) it is Amborella trichopoda Baillon 1869 & Nymphaea equally parsimonious to assume either that pli- odorata Aiton 1789 & Austrobaileya scandens C. cate carpels and sealing by postgenital fusion are T. White 1933). apomorphies at the level of Mesangiospermae, with reversals in some lines, or that these fea- Etymology: Mesangiospermae is a rough trans- tures originated more than once within the lation of"core angiosperms," the informal name clade. However, with other topologies (Soltis et applied to this clade by Judd et al. (2002). The al., 2005: Fig. 3.17) the latter scenario is more prefix mes- (Greek) means "middle" (Stearn, parsimonious. Combined molecular and mor- 1973), and Angjospermae (this volume) is the phological analyses (Doyle and Endress, 2000; name of a larger clade. Endress and Doyle, 2009) and some molecular analyses (Qiu et al., 2005, 2010) supported the Reference Phylogeny: The primary reference placement of Chloranthaceae or a clade con- phylogeny is Qiu et al. (2005: Fig. 2). See also sisting of Chloranthaceae and Ceratophyllum Soltis et al. (2011: Figs. 1, 2) and Wickett et al. as sister to all remaining Mesangjospermae, in (2014: Figs. 2, 3). which case it could be inferred that the ascidi- ate carpels of Chloranthaceae (with or without Composition: Chloranthaceae, Ceratophyllum, Ceratophyllum) are plesiomorphic and plicate Magnoliidae, Monocotyledoneae, and Eudi- carpels sealed by postgenital fusion evolved cotyledoneae (see entries in this volume for the once within Mesangiospermae.
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