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and M. . in P. D. Cantino et al. Pan-TracheophytaP D. Cantino Donoghue . J. converted clade name (2007): 830 .P. D. Cantino ana Donognuej,

inclusive clade if Registration Number: 83 a synapomorphy of a more Anthocerotae is the extant sister group of tra- some a that has received The total clade of the crown clade cheophytes, hypothesis Definition: et 2004; Wolf crown-based total-clade molecular support (Kelch al., Tracheophyta. This is a et 2006, 2007; however, Abbreviated definition: total V of et al., 2005; Qiu al., defnition. Another see Wickett et al., 2014). possible syn- Tracheophyta. apomorphy of Pan-7racheophyta is the produc- Kato and tion of a stem, if Akiyama From the or pantos (all, Greek pan- not homolo- Etymology: correct that the stem is the name refers to a (2005) are the whole), indicating that in this with the seta. total clade, and Tracheophyta (see entry gous for the name of the corre- volume etymology), defined names sponding crown clade. Synonyms: The phylogenetically Polysporangiomorpha (Kenrick and Crane, 1997) and Kenrick, Reference Phylogeny: Crane et al. (2004: Fig. and Polysporangiophyta (Crane as base of the 1997) have the same known composition 1; the clade stemming from the but potentially refer to differ- branch labeled "Polysporangiophytes"). See also Pan-Tracheophyta Kenrick and Crane (1997: Fig. 4.31). t clades (see Comments). Comments: The definition of Pan-Tracheophyta Composition: Tracheophyta (this volume) one we used when we first and all extinct (e-g , is identical to the the name (Cantino et al., 2007). Horneophytopsida, and Rhyniopsida sensu published The name Polysporangiomorpha (polysporan- Kenrick and Crane 1997) that share more and Crane than with sensu Kenrick (1997: recent ancestry with Tracheophyta giophytes) Table 7.2, 4.31) has an apomorphy-based extant Musci (), Hepaticae (liverworts), Fig. definition and thus is unlikely to be fully and Anthocerotae (). Its cur- synonymous with Pan-Tracheophyta. known is the same as Diagnostic Apomorphies: An independent rently composition that of Pan-Tracheophyta, but there may have 9porophyte, multiple sporangia, and sunken been that preceded the are of P'an- pan-tracheophytes archegonia possible synapomorphies evolution of multiple sporangia. Similarly, racheophryta, All three were listed by Kenrick sensu Crane and Kenrick Table 7.2) as Polysporangiophyta ane(997: synapomorphies as both of Polysporangiomorpha ("polysporangiophytes (1997) has the same known composition of and Polysporangiomorpha, Crane et al., 2004; a less inclusive Pan-Tracheophyta slightly definition and ch but its name has a node-based Cade than Pan-Tracheophyta; see The order in which the three featuresCommen5evolved thus the clade Polysporangiophyta may be less is not known. inclusive than either of other two. The three Wn. Moreover, sunken archegonia the aln be if a also names would no longer synonyms l00Cur in Anthocerotae (Kenrick and Crane, 7: Fig. 3.33, pp. 63-64) and thus may be were to be found in the future that possesses an intermediate combination of ancestral and D. W. Taylor, C. M. Testa, M. Ambros, B. derived states of the three characters mentioned Crandall-Stotler, R.J. Duff, M. Stech, W. Frey, C. Davis. above as possible synapomorphies (for example, D. Quandt, and C. 2006. The deep. est divergences in land plants inferred from a sporophyte that is dependent on the gameto- phylogenomic evidence. Proc. Natl. Acad. Sci phyte throughout its life, as is the case in bryo- USA 103:15511-15516. phytes, but has more than one ). Wickett, N. J., S. Mirarab, N. Nguyen, T. Warnow, E. Carpenter, N. Matasci, S Literature Cited Ayyampalayam, M. S. Barker, J. G. Burleigh, M. A. Gitzendanner, B. R. Ruhfel, E. Wafula, Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. J. P. Der, S. W. Graham, S. Mathews, M. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, Melkonian, D. E. Soltis, P. S. Soltis, N. W and M. J. Donoghue. 2007. Towards a phylo- Miles, C. J. Rochfels, L. Pokorny, A. J. Shaw, genetic nomenclature of Tracheophyta. Taxon L. DeGironimo, D. W. Stevenson, B. Surek, 56:822-846 and E1-E44. J.C. Villarreal, B. Roure, H. Philippe, C. W. Crane, P. R., P. Herendeen, and E. M. Friis. 2004. dePamphilis, T. Chen, M. K. Deyholos, R. S. and phylogeny. Am. J. Bot. Baucom, T. M. Kutchan, M. M. Augustin, J. 91:1683-1699. Wang, Y. Zhang, Z. Tian, Z. Yan, X. Wu, X. Crane, P. R., and P. Kenrick. 1997. Problems in cla- Sun, G. K. Wong, and J.Leebens-Mack. 014. distic classification: higher-level relationships Phylotranscriptomic analysis of the origin and in land plants. Aliso 15:87-104. early diversification of land plants. Proc. Narl. Kato, M., and H. Akiyama. 2005. Interpolation Acad. Sci. USA 111:E4859-E4868. hypothesis for origin of the vegetative sporo- Wolf, P.G., K. G. Karol, D. F. Mandoli, J. Kuehl, K. phyte of land plants. Taxon 54:443-450. Arumuganathan, M. W. Ellis, B. D. Mishler, Kelch, D. G., A. Driskell, and B. D. Mishler. 2004. D. G. Kelch, R. G. Olmstead, and J. L. Boore. Inferring phylogeny using genomic characters: 2005. The first complete chloroplast genome a case study using land plant plastomes. Pp. sequence of a , hucidula 3-11 in Molecular Systematics of (B. (). Gene (Amst.) 350:117-128. Gofinet, V. Hollowell, and R. Magill, eds.). Missouri Botanical Garden Press, St. Louis, MO. Authors Kenrick, P., and P. R. Crane. 1997. The Origin D. and Early Land PlantsA Philip Cantino; Department of Environmental Diversification of and Plant Cladistic Study. Smithsonian Institution Press, ; Ohio University; Athens, Washington, DC. OH 45701, USA. Email: Michael [email protected] Qiu, Y-L., L. Li, B. Wang, Z. Chen, O. Dombrovska, J. Donoghue; Department of Ecology and J. Lee, L. Kent, R. Li, R. W. Jobson, T. A. Evolutionary Biology; Yale University; P.O. Hendry, D. W. Taylor, C. M. Testa, and M. Box 208106; New Haven, CT 06520, USA. Ambros. Email: [email protected]. 2007. A nonflowering land plant phy- inferred logeny from nucleotide sequences of seven chloroplast, mitochondrial, and nuclear Date Accepted: 1 April 2011; updated 9 genes. Int. ]. Plant Sci. 168:691-708. February 2018 Qiu, Y.-L., L. B. Li, \Wang, Z. Chen, V. Knoop, M. Grorh-Malonek, O. Dombrovska, J. Lee, L. Kent, Primary Editor: Kevin de J. Rest, G. F. Estabrook, T. A. Hendry, Queiroz

228 P. D. Cantino and M. J. Donoghue in al. Apo-Tracheophyta(2007): E10[P. D. Cantino and M. J. Donoghue], convertedP. D. clade Cantinoname et

Registration Number: 15 Synonyms: Based on composition, Crane Tracheophyta sensu Kenrick and (1997: Definition: The clade characterized by the apo- Tables 7.1, 7.2, p. 236) is an approximate syn- Kenrick and Crane (1997: morphy tracheids (i.e., differentially thickened onym. Although as a water-conducting cells) as inherited by Pinus 236) listed Tracheidatae Bremer (1985) sylvestris Linnaeus 1753. This is an apomorphy- synonym of their "Eutracheophytes," imply- based definition. Abbreviated definition: V apo ing that that Tracheidatae referred to the crown

Bremer's comments tracheids [Pinus sylvestris Linnaeus 1753]. group, it is clear from (p. to be 382) that he considered rhyniopsids part Etymology: From the Greek apo-, indicating of Tracheidatae; thus, based on composition, that the name reters to an apomorphy-based Tracheidatae is an approximate synonym of clade, and Tracheophyta (see entry in this vol Apo-Tracheophyta. Preridophyta of some earlier ume for etymology). authors (e.g., Haupt, 1953) is a partial synonym; the originated from the same Reference Phylogeny: The primary reference ancestor as Apo-Tracheophyta but are paraphy- phylogeny is Crane et al. 2004: Fig. 1; the clade letic because they exclude Apo-Spermatophyta labeled "Tracheophytes"). See also Kenrick sensu Cantino and Donoghue (this volume). and Crane (1997: Fig 4.31; the clade labeled Tracheophyta). Comments: Although the name Tracheophbyta is often applied to this apomorphy-based Composition: Assuming that tracheids with clade, we opted here and previously (Cantino S-type and G-type cell walls (see Kenrick et al., 2007) to apply that name instead to the and Crane, 1997: Fig. 4.26) are homologous, crown clade, following PhyloCode (version 6) Apo-Tracheophyta includes Tracheophyta sensu Recommendation 10.1B, and a new name to Cantino and Donoghue (this volume) and the apomorphy-based clade. and Crane Rhyniopsida sensu Kenrick (1997) Rhyniaceae in the primary reference phylogeny). Literature Cited Under the alternative hypothesis that these tra- cheid types evolved independently, Rhyniopsida Bremer, K. 1985. Summary of green plant phylog- and eny and classification. ladistics 1:369-385. would not be of Apo-Tracheophyta, part Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. the currently known membership of Apo- Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, would be the Tracheophyta and Tracheophyta and M. J. Donoghue. 2007. Towards a phylo- same. genetic nomenclature of Tracheoplhyta. Taxon 56:822-846 and E1-E44 Diagnostic Apomorphies: Tracheids (see Crane, P. R., P. Herendeen, and E. M. Friis. 2004. Definition), and possibly lignin deposition Fossils and plant phylogeny. Am. J. Bot wall on the inner surface of the tracheid cell 91:1683-1699. Haupt, A. W. 1953. Plant Morphology. McGraw-Hil (Kenrick and Crane, 1997: Table 7.2, under New York. Tracheophryt). Apo-Tracheophyta

Kenrick, P., and P. R. Crane. 1997. 7he Origin Michael J. Donoghue; Department of Ecology and and Early Diversification of Land Plants-A Evolutionary Biology; Yale Universiry; P.O. Cladistic Study. Smithsonian Institution Press, Box 208106; New Haven, CT 06520, USA. Washington, DC. Email: [email protected].

Authors Date Accepted: 1 April 2011; updated 9 February 2018 Philip D. Cantino; Department of Environmental and Plant Biology: Ohio University; Athens, Primary Editor: Kevin de Queiroz OH 45701, USA. Email: [email protected].

230 Tracheophyta E. W. Sinnott 1935: 441 [P. D. Cantino and M. J. Donoghue], converted clade name

Registration Number: 107 in the tracheid wall are listed by Kenrick and Crane (1997: Table 7.2, pp. 114, 120) as synapo- Definition: The smallest crown clade contain- morphies of "eutracheophytes" (= Tracheophyta data ing Magnolia tripetala (Linnaeus) Linnaeus as defined here), but the extent of missing 1759 (Euphyllophyta) and clava- for fossils combined with the apparent loss of um Linnaeus 1753 (Lycopodiophyta). This is a these traits in all extant tracheophytes reduces minimum-crown-clade definition. Abbreviated confidence in their inferred originations. definition: min crown V (Magnolia ripetala (Linnaeus) Linnaeus 1759 & Lycopodium clava- Synonyms: "Eutracheophytes" sensu Kenrick tum Linnacus 1753). and Crane (1997: 236) was described as "the tracheophyte and is thus an Etymology: From the Latin tracheida, refering unambiguous synonym. Crane and Kenrick to the presence of tracheids (differentially thick (1997) formalized this name as Eutracheophyta ened water conducting cells in the ), and and defined it (node-based) using of the Greek pbyton (plant). Huperzia and Nymphaea as specifiers; this is an unambiguous synonym. Cormatae Jeffrey Reference Phylogeny: The primary refer- (1982) is an approximate synonym; all listed ence phylogeny is Qiu et al. (2007: Fig. 1). See subordinate taxa are extant. also Kenrick and Crane (1997: Fig. 4.31; as The or Eutracheophytes"), Pryer et al. (2001: Fig. 1), Comments: vascular plants, tracheo- have been as a clade Wolf et al. (2005: Fig. 3), Qiu et al. (2006: Fig phytes, long recognized ), and Ruhfel et al. (2014: Figs. 5-7). based on their possession of tracheids as a syn- apomorphy, and the crown cdade is strongly Composition: All extant vascular plants and supported by DNA analyses (Nickrent et al., their extinct relatives that fall within the crown 2000; Pryer et al., 2001; Qiu et al., 2006, 2007; clade. The crown clade is composed of two pri- Ruhfel et al., 2014; Wickett et al., 2014). Here mary subclades, Pan-Lycopodiophyta and Pan- and previously (Cantino et al., 2007: 829) Euphyllophyta (see entries in this volume). we have defined the name Tracheophyta to refer to the crown clade, tollowing PhyloCode Diagnostic Apomorphies: The walls of the Recommendation 10.1B. This applicationis Water-conducting cells in the xylem have a thick, somewhat unconventional in that this name decay-resistant layer. free-living has more often been applied to slightly lgnified,Tophyte and multiple sporangiaA per sporophytespo- more inclusive clade originating withthe the evo- lution of tracheids (i.., Apo-Tracheophyta in synapomorphies relative to other crown introduced the ades;s however, when fossils are considered, this volume). Sinnotr (1935) C traits are synapomorphies at a more inclu- name Tracheophyta for the vascular plants but SIve level (see Pan-Tracheophyta),. Sterome (a well- the Latin diagnosis required by the botanical aeveloped peripheral zone of the stem consisting code (Turland et al., 2018) was first provided by Or thick-walled, decay-resistant cells) and pitlets Cavalier-Smith (1998: 251). We earlier Ambros. 2007. A nonfowering land planr. (Cantino et al., 2007) used a logeny in ferred from nucleotide sequencesphy- of maximum-crown-clade definition with exter- nal seven chloroplast, mitochondrial, and nuel. specifiers representing liverworts, mosses, and Int. J. Plant Sci. 168:691-708. hornworts. because we thought this defi- genes. nition would Qiu, Y-L., L. Li, B. Wang, Z. Chen, V. Knoop. NA provide greater compositional Groth-Malonek, O. mbrovska, than a J. Lee, L. stability simple node-based definition. Kent, J. Rest, G. F. Estabrook, T. A. Hendry, However, in view of the very strong D. W. C. M. Testa, M. for the basal support Taylor, Ambros, R dichotomy within Crandall-Stotler, R. J. Duff, M. Stech, W. in recent molecular Tracheophyta Frey, analyses (Qiu et al., 2006, D. Quandt, and C. C. Davis. 2006. The deen 2007; Wickett et ep- al., 2014), we are here est in land plants inferred for a opting divergences from simpler minimum-crown-clade definition phylogenomic evidence. Proc. Natl. Acad, se with the two internal specifiers the USA 103:15511-15516. two clades representing B. M. A. originating from the basal split: Pan- Ruhfel, R., Gitzendanner, P. S. Soltis, D. Lycopodiophyta and Pan-Euphyllophyta. E. Soltis, and J. G. Burleigh. 2014. From algae to angiosperms-inferring the phylogeny of Literature Cited green plants () from 360 plastid genomes. BMC Evol. Biol. 14:23. E. W. Cantino, P. D., A. Sinnot, 1935. : J. Doyle, S. W. Graham, W. S. Principles and Problems. Judd, R. G. 3rd edition. McGraw-Hill, New York. Olmstead, D. E. Soltis, P. S. Soltis, and M. J. Turland, N. J.. J. H. Wiersema, F. R. 2007. Towards a Barrie, W. Donoghue. phylo- D. genetic nomenclature of Greuter, L. Hawksworth, P. S. Tracheophyta. 1Taxon S. Herendeen, 56:822-846 and El-E44. Knapp, W-H. Kusber, D.-Z. Li, K. T. W. Marhold, Cavalier-Smith, T. 1998. A revised May, J. McNeill, A. M. Monro,J. of life. Biol. Rev. six- system Prado, M. J. Price, and Camb. Philos. Soc. 73:203-266. G. F. Smith, eds. 2018. Crane, P. R., and P. Kenrick. International Code 1997. Problems in cla- of Nomenclature for Algae, distic classification: Fungi and Plants (Shenzhen in higher-level relationships Code). Adopred land plants. Aliso by the Nineteenth International C. 15:87-104. Botanical Jeffrey, 1982. Kingdoms, codes and classifica- Congress, Shenzhen, , tion. Kew July 2017. Bull. 37:403-416. Regnum Vegetabile 159. Koeltz Kenrick, P., and P. R. Books, Glashütten. Botanical Crane. 1997. The Origin and Early Land Wickett, N. J., S. Diversification of Plants--A Mirarab, N. 1. Cladistic Study. Smithsonian Institution Warnow, E. Nguyen, Press, Carpenter, N. Matasci, S. Washington, DC. Ayyampalayam, M. S. Nickrent, D. L, C. L. M. A. Barker, J. G. Parkinson, J. D. Palmer, and B. R. Burleigh R. J. Duff. 2000. J. P. Gitzendanner, Ruhfel, E. Wafula, Multigene of land Der, S. W. with phylogeny Graham, S. Mathews, plants special reference to and E. M. the earliest land bryophytes Melkonian, D. Soltis, P. S. Soltis, N. plants. Mol. Evol. Miles, C. J. 17:1885-1895. Phylogenet. L. Rothfels, L. A. J. DeGironimo, D. W. Pokorny, Sha Pryer, K. M., H. A. J. C. Stevenson, B. Schneider, R. Smith, R. Villarreal, B. Surek, P.G. S. Cranfill, Roure, H. C. Wolf, J. Hunt, and S. D. Sipes. 2001. Philippe, W. Horsetails and dePamphilis, T. Chen, M. K. R. are a Baucom, T. M. Deyholos, and the closest monophyletic group Kutchan, M. M. living relatives to seed Wang, Y. Nature 409:618-622. plants. Zhang, Z. Augustin, Sun, G. K. Tian, Z. Yan, X. Wu, L. Li, B. Wong, and A Qiu, Y-L Wang. Z. Chen, O. Dombrovska, J. Leebens-Mack. 201 J. Lee, L. Kent, R. Li, R. W. earlyPhylotranscriptomic analysis of the Jobson, T. A. of origin and Hendry, D. W. Taylor, C. M. Acad. diversification land Testa, and M. Sci. USA 111:E4859-E4868.plants. Proc. a 232 Tracheophyta

K. G. Karol, D. F. Mandoli, J. Kuehl, Michael J. Donoghue; Department of Ecology and Wolf P. G. M. W. Ellis, B. D. Yale P.O. K. Arumuganathan, Evolutionary Biology; University; R. G. Olmstead, and Box 208106; New Haven, CT 06520, USA. Mishler. D. G. Kelch, 2005. The first complete chlo- Email: [email protected]. 1. L. Boore. of a roplast genome sequence lycophyte, Huperzia ucidula (Lycopodiaceae). Gene Date Accepted: 23 March 2011; updated 9 (Amst.) 350:117-128. February 2018, 11 July 2018

Authors Primary Editor: Kevin de Queiroz

Philip D. Cantino; Department of Environmental and Plant Biology; Ohio University; Athens, OH 45701, USA. Email: [email protected]. Pan-Lycopodiophyta P. D. Cantino and M. J. Donoghue in P. D. Cantino et al. (2007): E11 [P. D. Cantino and M. J. Donoghue), converted clade name

Registration Number: 81 the (unnamed) lycopodiophyte total clade. Kenrick and Crane (1997: Table 4.6) cited it as a Definition: The total clade of the crown clade possible synapomorphy of their node 52, which Lycopodiophyta. This is a crown-based total- is near the base of the total clade. Crane and clade definition. Abbreviated definition: total V Kenrick (1997) listed the following additional of Lycopodiophyta. potential synapomorphies for their Lycophytina (which is somewhat less inclusive than the Etymology: From the crown clade name, total clade but much more inclusive than the Lycopodiophyta (see entry in this volume for its crown; see Comments): reniform sporangia; etymology), and the Greek pan- or pantos (all, marked sporangial dorsiventrality; inconspicu- the whole), indicating reference to a total clade. ous cellular thickening of the dehiscence line; sporangia on short, laterally inserted stalks; Reference Phylogeny: Doyle (2013: Fig. 1.1). and exarch xylem differentiation. However, the See also Crane et al. (2004: Fig. 1). In both of more taxonomically comprehensive analysis of these phylogenies, the clade here named Pan- Kenrick and Crane (1997: Fig. 4.32 and Table Lycopodiophyta is not labeled as such but is the 4.6) suggests that some of these characters may most inclusive clade containing the be synapomorphic for more inclusive clades but not the . than Lycophytina within the total clade (also see Doyle, 2013). Composition: Lycopodiophyta (this volume) and all extinct plants that share more recent ances Synonyms: Based on its composition, the name Doweld (2001) is an try with Lycopodiophyta than with Euphyllophyta 1ycopodiobiotina approxi- mate (Kenrick and Crane, (this volume). According to the phylogenies synonym. Lycophytina of Kenrick and Crane (1997), Crane et al. 1997; Crane and Kenrick, 1997; Bateman et al.,

to a clade that is less inclusive than (2004), and Doyle (2013), extinct taxa outside 1998) applies of the crown include (not an exhaustive list): Pan-Lycopodiophyta (see Comments). the relatively apical stem groups Asteroxrylon, zostero- Comments: Our definition of Pan- Drepanophbycus, ; the to that used phylls; and the relatively basal stem group Lycopodiophyta here is identical we the name (Cantino et al. , , Uskiella, Sartilmania, and when published the name rather cambrensis. The inclusion of some 2007). We chose panclade Other species of Cooksonia is uncertain because than its approximate synonym Lycopodiobiotina Code of their unresolved position on these trees. (Doweld, 2001), as permitted by Phylo to a total Article 10.6, so that its application Diagnostic Apomorphies: A possible synapo clade would be evident. The name Lycophytina and a trans- sensu Kenrick and Crane (1997: Fig. 4.31 morphy is sporangium dehiscence by showed Table has a "synapomorphy-based defini- verse, apical slit. Doyle (1998, 2013) 7.2) this character as arising at or near the base of tion (not a formal definition; perhaps better Pan-Lycopodiophyta

described as an P. R., P. Herendeen, and E. M. Friis, 2004 apomorphy-based conceptual- Crane, 4. ization of the taxon, with six Fossils and plant phylogeny. Am. J. Bot synapomorphies 91:1683-1699. cited) and is somewhat less inclusive than Pan- Lycopodiophyta. The name Lycophytina sensu Crane, P. R., and P. Kenrick. 1997. Problems in ca. DiMichele and distic classification: higher-level relationshins Bateman (1996) and Bateman et in land plants. Aliso 15:87-104. al. (1998) to appears be applied to a clade that, DiMichele, W. A., and R. M. Bateman. 1996. The based on its synapomorphies and composition, rhizomorphic lycopsids: a case-study in paleo is circumscribed similarly to Lycophytina sensu botanical classification. Syst. Bot. 21:535-552, Kenrick and Crane (1997). Lycophytina sensu Doweld, A. 2001. Prosyllabus Tracheophytorum Crane and Tentamen Systematis Plantarum Vascularum Kenrick (1997) has a minimum-clade definition and approximates (Tracheophyta). Institutum National in Pan-Lycopodiophyta Carpologiae, Moscow. composition if only the taxa in their clado- gram are considered, but it is less inclusive than Doyle, J. A. 1998. Phylogeny of vascular plants. Annu. Rev. Ecol. Syst. 29:567-5999. the total clade in the context of trees that include Doyle, J. A. 2013. Phylogenetic analyses and mor- more stem fossils (e.g.. Crane et al., 2004). For phological innovations in land plants. Ann. example, Yunia and Renalia, which were not Plant Rev. 45:1-50. included in Crane and Kenrick's (1997) tree, Kenrick, P., and P. R. Crane. 1997. The Origin are part of Pan-Lycopodiophyta but outside of and Early Diversificarion of Land PlantsA Lyoophytina when Crane and Kenrick's (1997) Cladistic Study. Smithsonian Institution Press, definition of Washington, DC. Lycophytina is applied in the con- text of the tree of Crane et al. (2004). Authors Literature Cited Philip D. Cantino; Department of Environmental Bateman, R. M., P. R. Crane, W. A. DiMichele, and Plant Biology; Ohio University; Athens, P. R. Kenrick, N. P. Rowe, T. Speck, and W. OH 45701, USA. Email: [email protected]. E. Stein. 1998. Early evolution of land plants: Michael J. Donoghue; Department of Ecology and phylogeny, physiology, and ecology of the pri- Evolutionary Biology: Yale University; P.O. mary terrestrial radiation. Annu. Rev. Ecol. Box 208106; New Haven, CT 06520, USA. Syst. 29:263-292. Email: [email protected]. Cantino, P. D., J. A. Doyle, S. W. Graham, W.s. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, Date Accepted: 1 April 2011; updated 9 and M. J. Donoghue. 2007. Towards a phylo- February 2018 genetic nomenclature of Tracheophyta. Taxon 56:822-846 and E1-E44. Primary Editor: Kevin de Queiroz

236 Lycopodiophyta A. Cronquist, A. Takhtajan, and W. Zimmermann 1966: 133 [P. D. Cantino and M. J. Donoghue], converted clade name

and Crane Registration Number: 160 Diagnostic Apomorphies: Kenrick (1997: Table 6.3 and Fig. 6.19; node 35), Crane Doyle Definition: The smallest crown clade con- and Kenrick (1997: Table 9; "Lycopsida"), the follow- ra ining Linnaeus 1753, (1998), and Gensel (1992) listed dose Httperzia selago (Linnaeus) Schrank & Martius ing synapomorphies for the crown dade: axillary or 1829 (originally described as Lycopodium sel- developmental association of a single absence ago Linnaeus 1753), lacustris Linnaeus adaxial sporangium with a ; metaxylem 1753 , and apoda (Linnaeus) Spring of vasculature in the sporangium; symmet- 1840 (originally described as Lycopodium tracheids pitted; stele bilaterally one side of apodum Linnaeus 1753). This is a minimum- rical, with phloem located on only of missing data for crown-clade definition. Abbreviated defini- the stele (but there are a lot , so this trait may be tion: min crown V (Lycopodium clavatum fossils outside the crown a more inclusive clade); cres- Linnaeus 1753 & (Linnaeus) synapomorphic for (but there are a lot of Schrank & Martius 1829 & cent-shaped root xylem fossils outside the crown). The Linnaeus 1753 & (Linnaeus) missing data for synapomorphies of this crown Spring 1840). following are clade relative to other crowns but are known to at a more inclusive level when Etymology: Based on the name Lycopodium, be apomorphic considered (Kenrick and Crane, which is derived from Greek lycos and podus, stem fossils are and Table 7.2; Doyle, 2013: meaning "wolf's foot", from a fancied resem- 1997: Fig. 6.18 ("lycophylls"; Schneider blance (Fernald, 1970), and Greek phyton Fig. 1.1): microphylls Pryer et al., 2004a); exarch xylem (plane) (Stearn, 1973). et al., 2002; differentiation in stem (Kenrick and Crane, 1998; Schneider et al., 2002); stel- Reference Phylogeny: The primary reference 1997; Doyle, in stem; reniform sporangia phylogeny is Korall et al. (1999: Fig. 2). See also late xylem strand (Doyle, 1998). This Rydin and Wikstrom (2002: Fig. 2), Pryer et al. with transverse dehiscence see Kenrick and Crane (2004b: Fig. 3; labeled "lycophytes"), Qiu et al. list is not exhaustive; and Bateman (2007: Fig. 1), Ruhfel et al. (2014: Fig. 6), and (1997: Table 7.2) and DiMichele listed under Wickett et al. (2014: Fig. 2). (1996) for other synapomorphies Lycophytina and Lycopsida. ~omposition: The total clades of Lycopodiaceae and (including Huperziaceae) and Isoetopsida (Isoetes Synonyms: Lycophyta, Lycopsida, synonyms (see + Selaginella; Camino et al., 2007). According are approximate to current understanding of phylogeny (Doyle, Comments below). Lycopsida sensu Crane and l998; Pryer et al., 2004a; Judd et al., 2016), the Kenrick (1997) is an unambiguous synonym in total clade of lsoetes includes Palaeozoic arbores- that it was phylogenetically defined as apply- cent forms such as . ing to the same crown clade, but the definition Lycopodiophyta - conflicts with the labeling of the reference phy- l6.l). We are narrowing our choice to the lat- logeny (see Comments). ter pair of names in the interest of promoting consistency between rank-based and phyloge- Comments: The monophyly of this clade is netic nomenclature. We prefer the -phyta end- strongly supported by molecular data (Korall ing because it means "plant" while -opsida is et al., 1999; Pryer et al., 2001, 20046; Qiu et solely an indicator of (class) rank, and we have al., 2006, 2007; Ruhfel et al., 2014; Wickett therefore chosen Lycopodiophyta, as in our pre- et al., 2014) as well as by numerous morpho- vious paper (Cancino et al., 2007). We also fol- logical synapomorphies (detailed above). One low that paper (p. El 1) in applying the name study (Carbary et al., 1993) based solely on explicitly to the lycopodiophyte crown clade. male gametogenesis characters found this The clade Lycopsida sensu Kenrick and Crane group to be polyphyletic, with Selaginella shar- (1997) is somewhat larger in that it includes fos- ing closer ancestry with bryophytes than with sils such as and Baragwanathia that Lycopodium, but this hypothesis has not been are shown (p. 239; see also Pryer et al., 2004a: supported by any other analysis. Fig. 10.3) as being outside the crown clade. While the monophyly of the morphologi- The same is true of Microphyllophyta sensu cally distinctive taxa Selaginella and Isoi!tes is not Bold (1957) and Bold et al. (1980), Lepidophyta in question, the monophyly of Lycopodiaceae sensu Smith (1955), and Lycopodiopsida sensu (including Huperziaceae) is supported by only Bierhorst (1971). Crane and Kenrick (1997) one possible morphological synapomorphy gave Lycopsida a minimum-clade definition (foveolate-fossulate wall; Kenrick with extant species of Huperzia and Isoetes as and Crane, 1997). Although there is molecular the specifiers, thereby effectively (but not explic- support for its monophyly, those studies either itly) applying the name to the crown clade. were based solely on rbcL (Korall et al., 1999) However, in the accompanying cladogram and or included too small a sample of Lycopodiaceae classification, they applied this same name to a to be convincing (Qiu et al., 2006, 2007; larger clade by including an extinct stem group Wickett et al. 2014). We remain concerned (Drepanophycales); thus, their conceptualization that Huperzia could end up being sister to the of the name was ambiguous. rest of Lycopodiophyta. Consequently, the clade Following the PhyloCode (Version 6, Rec. Lycopodiaceae is represented by two specifiers 9.15A), we attribute the preexisting name (Lycopodium clavatum and Huperzia selago) in Lycopodiophyta to the definition of Lycopodiophyta. Cronquist et al. (1966), but under the botanical code The names Lycophyta, Lycopodiophyta, (Turland et al., 2018; Art. l6.3), the name is attributed Lycopsida, and Lycopodiopsida have histori- to Scott (l909), who spelled cally applied to the same set it Lycopsida (Hoogland and of clades (from Reveal, 2005). crown to total), but most phylogenetic stud- ies have instead used the informal equivalents, "lycophytes" and "lycopsids". Since Lycophyta Literature Cited and Lycopsida are apparently based on the name Lycopodium, they should be corrected Bierhorst, D. W. 1971. Morphology of Vascular to Lycopodiophyta and Lycopodiopsida Plants. Macmillan C N v under Bold H C o., k the botanical ew 1-or . code (Turland et al. , 2018; Art. ' · · l957. Morphology of Plants. Harper & Row, New York.

238 Lycopodiophyta

Bold, H. C., C. J. Alexopoulos, and T. Delevoryas. Korall, P., P. Kenrick, and J. P. Therrien. 1999. 1980. Morphology ofPlan ts and Fungi. 4th edi- Phylogeny of Selaginellaceae: evaluation of tion. Harper & Row, New York. generic/subgeneric relationships based on rbcL Cantino, P. D. , J. A. Doyle, S. W. Graham, W. S. gene sequences. Int. J Plant Sci. 160:585-594. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, Pryer, K. M., H. Schneider, and S. Magallon. 2004a. and M. J. Donoghue. 2007. Towards a phylo- The radiation of vascular plants. Pp. 138-153 genetic nomenclature of Tracheophyta . T axon in Assembling the Tree of Life (J. Cracraft and 56:822-846 and El-E44. M. J. Donoghue, eds.). Oxford University Crane, P. R. , and P. Kenrick. 1997. Problems in cla- Press, Oxford. distic classification: higher-level relationships Pryer, K. M., H. Schneider, A. R. Smith, R. Cranfill, in land plants. Aliso 15:87-104. P. G. Wolf, J. S. Hunt, and S. D. Sipes. 2001. Cronquist, A., A. Takhtajan, and W. Zimmermann. Horsetails and ferns are a monophyletic group 1966. On the higher taxa of Embryobionta. and the closest living relatives to seed plants. Taxon 15:129-168. Nature 409:618-622. DiMichele, W. A., and R. M. Bateman. 1996. Pryer, K. M., E. Schuettpelz, P. G. Wolf, H. The rhizomorphic lycopsids: a case-study in Schneider, A. R. Smith, and R. Cranfill. paleobotanical classification. Syst. Bot. 21: 2004b. Phylogeny and evolution of ferns 535-552. (monilophytes) with a focus on the early lep- Doyle, J. A. 1998. Phylogeny of vascular plants. tosporangiate divergences. Am. J Bot. 91: Annu. Rev. Ecol. Syst. 29:567-599. 1582-1598. Doyle, J. A. 2013. Phylogenetic analyses and mor- Qiu, Y.-L., L. Li, B. Wang, Z. Chen, 0. Dombrovska, phological innovations in land plants. Pp. J. Lee, L. Kent, R. Li, R. W. Jobson, T. A. 1-50 in The Evolution of Plant Form (B. A. Hendry, D. W. Taylor, C. M. Testa, and M. Ambrose and M. Purugganan, eds.). Annual Ambros. 2007. A nonflowering land plant phy- Plant Reviews 45. Blackwell, Oxford. logeny inferred from nucleotide sequences of Fernald, M. L. 1970. Gray's Manual ofBotany. 8th seven chloroplast, mitochondrial, and nuclear edition, corrected printing. Van Nostrand, genes. Int. J Plant Sci. 168:691-708. New York. Qiu, Y.-L., L. Li, B. Wang, Z. Chen, V. Knoop, M. Garbary, D. J., K. S. Renzaglia, and J. G. Duckett. Groth-Malonek, 0. Dombrovska, J. Lee, L. 1993. The phylogeny ofland plants: a cladistic Kent, J. Rest, G. F. Estabrook, T. A. Hendry, analysis based on male gametogenesis. Plant D. W. Taylor, C. M. Testa, M. Ambros, B. Syst. Evol. 188:237-269. Crandall-Stotler, R. J. Duff, M. Stech, W. Frey, Gensel, P. G. 1992. Phylogenetic relationships D. Quandt, and C. C. Davis. 2006. The deep- of the and lycopsids: evidence est divergences in land plants inferred from from morphology, paleoecology, and cladistic phylogenomic evidence. Proc. Natl. Acad. Sci. methods of inference. Ann. Mo. Bot. Gard. USA 103:15511-15516. 79:450-473. Ruhfel, B. R., M.A. Gitzendanner, P. S. Soltis, D. Hoogland, R. D., and J. L. Reveal. 2005. Index E. Soltis, and J. G. Burleigh. 2014. From algae nominum familiarum plantarum vascular- to angiosperms-inferring the phylogeny of ium. Bot. Rev. 71:1-291. green plants (Viridiplantae) from 360 plastid Judd, W. S., C. S. Campbell, E. A. Kellogg, P. F. genomes. BMC Evol. Biol. 14:23. Stevens, and M. J. Donoghue. 2016. Plant Rydin, C., and N. Wikstrom. 2002. Phylogeny of Systematics: A Phylogenetic Approach. 4th edi- Isoetes (Lycopsida): resolving basal relationships tion. Sinauer Associates, Sunderland, MA. using rbcl sequences. Taxon 51:83-89. Kenrick, P., and P. R. Crane. 1997. The Origin Schneider, H., K. M. Pryer, R. Cranfill, A. R. and Early Diversification of Land Plants-A Smith, and P. G. Wolf. 2002. Evolution ofvas- Cladistic Study. Smithsonian Institution Press, cular plant body plans: a phylogenetic perspec- Washington, DC. tive. Pp. 330-364 in Developmental Genetics

239 Lycopodiophyta

L. DeGironimo, D. W. Stevenson, B. Surek, and Plant Evolution (Q. C. B. Cronk, R. M. J. C. Villarreal, B. Roure, H. Philippe, C. W. Bateman, and J. A. Hawkins, eds.). Taylor & dePamphilis, T. Chen, M. K. Deyholos, R. S. Francis, London. Baucom, T. M. Kutchan, M. M. Augustin, J. 2nd edi- Scott, D. H. 1909. Studies in Fossil Botany. Wang, Y. Zhang, Z. Tian, Z. Yan, X. Wu, X. tion. Adam & Charles Black, London. Sun, G. K. Wong, andJ. Leebens-Mack. 2014. Smith, G. M. 1955. Cryptogamic Botany. 2nd edi- Phylotranscriptomic analysis of the origin and tion, Vol. 2. McGraw-Hill, New York. early diversification of land plants. Proc. Natl. Stearn, W. T. 1973. Botanical Latin. David & Acad. Sci. , USA. 1ll:E4859-E4868. Charles, Newton Abbot, Devon. Turland, N. J., J. H. Wiersema, F. R. Barrie, W. Greuter, D. L. Hawksworth, P. S. Herendeen, Authors S. Knapp, W.-H. Kusber, D.-Z. Li, K. Marhold, T. W. May, J. McNeill, A. M. Monro, J. Philip D. Cantine; Department of Environmental Prado, M. J. Price, and G. F. Smith, eds. 2018. and Plant Biology; Ohio University; Athens, International Code of Nomenclature for Algae, OH 45701, USA. Email: [email protected]. Fungi, and Plants (Shenzhen Code). Adopted Michael J. Donoghue; Department of Ecology and by the Nineteenth International Botanical Evolutionary Biology; Yale University; P.O. Congress, Shenzhen, China, July 20l7. Box 208106; New Haven, CT 06520, USA. Regnum Vegetabile 159. Koeltz Botanical Email: [email protected]. Books, Glashiitten. Wickett, N. J., S. Mirarab, N. Nguyen, T. Date Accepted: 1 April 2011; updated 21 Ma Warnow, E. Carpenter, N. Matasci S Ayyampalayam, M. S. Barker, J. G. Burl~igh. 2018, 11 July 2018 y M. A. Giczendanner, B. R. Ruhfel, E. Wafula: J. P. Der, S. W. Graham, S. Mathews, M. Primary Editor·· Kev1· n de Que1roz · Melkonian, D. E. Soltis, P. S. Soltis N W Miles, C. J. Rochfels, L. Pokorny, .A . 1' Sh. aw,.

240 Pan-Euphyllophyta P. D. Cantino and M J D h . Cantino a~d·MoJnoog ue tnh ] P. D. Cantino et al. (2007): E12 [P. o. . . onog ue, converted clade name

Registration Number: 78 Table 7.2, and pages listed below), most of which have been lost or modified in some or all extant Definition: The total clade of the crown clade members of the clade: pseudomonopodial or Ettphyllophyta. This is a crown-based total-clade monopodial branching (Kenrick and Crane, definition. Abbreviated definition: total V of 1997: 109, 359; Doyle, 2013) (although if the Euphyllophyta. fernlike of early seed plants were derived from pseudomonopodial branch systems of Etymology: From the Greek eu- (true, original, more basal lignophytes (Doyle, 1998, 2013), the primitive), phyllon (), and phyton (plant), axillary monopodial branching of seed plants plus pan- or pantos (all, the whole), indicating and the pseudomonopodial branching of more that this is a total clade. Euphylls (also known basal lignophytes may not be homologous; as megaphylls; i.e., leaves that usually have more Doyle, 2013); helical arrangement of branches than one vein and are thought to have evolved (Kenrick and Crane, 1997: 110, 360); dichoto- from branch systems) are characteristic of the mous appendages (Kenrick and Crane, 1997: crown clade Euphyllophyta. 113, 361); recurved branch apexes (Kenrick and Crane, 1997: 112-113, 360); paired sporangia Reference Phylogeny: Crane and Kenrick grouped into terminal trusses (Kenrick and (1997: Fig. 1, the clade labeled Euphyllophytina) Crane, 1997: 121-122, 364); sporangial dehis- (see Synonyms). cence along one side through a single longitu- dinal slit (Kenrick and Crane, 1997: 125, 366); Composition: Euphyllophyta (this volume) and Doyle, 2013; and radially aligned xylem in all extinct plants that share more recent ancestry larger axes (Crane and Kenrick, 1997). Kenrick with Euphyllophyta than with Lycopodiophyta. and Crane also cited scalariform bordered pit- The stem euphyllophytes are often classified as ting of metaxylem cells as a synapomorphy, but trimerophytes, a paraphyletic group that has it does not occur in Eophyllophyton and there- been named at various ranks (see Synonyms). fore is synapomorphic for a less inclusive group The trimerophytes include species of than the total clade (Kenrick and Crane, 1997: and Eophyllophyton, which lie outside the 120, 363, Fig. 7.10). crown (Crane and Kenrick, 1997; Kenrick and Crane, 1997; Doyle, 2013), Synonyms: Euphyllophytina Kenrick and but (which is also classified as a trim- Crane (1997: Table 7.1) and Crane and Kenrick erophyte; Stewart and Rothwell, 1993; Taylor, (1997) may be an approximate synonym (see 1981) is thought to lie within the crown Comments). Trimerophytophyta sensu Bold (Kenrick and Crane, 1997: Figs. 4.31, 7.10). et al. (1980), Trimerophytina Banks 1968, Trimerophytopsida Foster and Gifford 1974 have Diagnostic Apomorphies: Several synapomor- been applied to a paraphyletic group originat- phies were listed by Crane and Kenrick (1997: ing in approximately the same ancestor as Pan- Table 9) and Kenrick and Crane (1997: 240, Euphyllophyta according to the phylogenies of Pan-Euphyllophyta

P. D., A. Doyle, S. W. Graham, W. S. Kenrick and Crane (1997: Figs. 4.31, 4.32) and Camino, J. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, Crane and Kenrick (1997: Fig. 1) and therefore and M. J. Donoghue. 2007. Towards a phylo- are partial (and approximate) synonyms. genetic nomenclature of Tracheophyta. Taxon 56:822-846 and El-E44. and Comments: Euphyllophytina (Crane Crane, P.R., and P. Kenrick. 1997. Problems in cla- Kenrick, 1997; Kenrick and Crane, 1997) distic classification: higher-level relationships referred to a clade that is similar in composition in land plants. Aliso 15:87-104. plants. to Pan-Euphyllophyta. However, there is conflict Doyle, J. A. 1998. Phylogeny of vascular 29:567-599. within and between these two papers regard- Annu. Rev. Ecol. Syst. A. 2013. Phylogenetic analyses and mor- ing whether the name Euphyllophytina applies Doyle, J. phological innovations in land plants. Pp. to a node-based, apomorphy-based, or branch- 1-50 in 1he Evolution of Plant Form (B. A. based clade. Kenrick and Crane (1997) gave the Ambrose and M. Purugganan, eds.). Annual "synapomorphy-based definition" in name a Plant Reviews 45. Blackwell, Oxford. the clade (p. 240) Table 7.2, but they described Foster, A. S., and E. M. Gifford. 1974. Comparative as the sister group of Lycophytina, suggesting Morphology of Vascular Plants. 2nd edition. W. that both of these clades were conceptualized H. Freeman, San Francisco, CA. as originating from their point of divergence Kenrick, P., and P. R. Crane. 1997. 1he Origin rather than from the origin of a particular apo- and Early Diversification of Land Plants-A Institution Press, morphy. In contrast, the same authors (Crane Ciadistic Study. Smithsonian Washington, DC. and Kenrick, 1997) used a "node-based" (mini- Stewart, W. N., and G. W. Rothwell. 1993. mum-clade) definition for Euphyllophytina with Paleobotany and the Evolution of Plants. Eophyllophyton bellum and Nymphaea odorata 2nd edition. Cambridge University Press, the composition as the specifiers. In any case, Cambridge, UK. on the reference of Euphyllophytina, as shown Taylor, T. N. 1981. Paleobotany. McGraw-Hill, trees in these two papers by Crane and Kenrick, New York. seems to approximate that of Pan-Euphyllophyta as defined here. However, there could be undis- Authors covered fossil pan-euphyllophytes that lie out- side the node-based Euphyllophytina of Crane Philip D. Camino; Department of Environmental and Kenrick. and Plant Biology; Ohio University; Athens, OH 45701, USA. Email: [email protected]. Literature Cited Michael J. Donoghue; Department of Ecology and Evolutionary Biology; Yale University; P.O. 06520, USA. Banks, H. P. 1968. The early history of land plants. Box 208106; New Haven, CT Pp. 73-107 in Evolution and Environment (E. Email: [email protected]. T. Drake, ed.). Yale University Press, New Haven. Date Accepted: 31 January 2013; updated 9 Bold, H. C., C. J. Alexopoulos, and T. Delevoryas. February 2018 1980. Morphology ofPlants and Fungi. 4th edi- tion. Harper & Row, New York. Primary Editor: Kevin de Queiroz

242 Euphyllophyta G. Lecointre and H. Le Guyader 2006: 181 [P. D. Cantino and M. J. Donoghue], converted clade name

Registration Number: 43 crown clade. The crown clade is composed of two primary subclades, Pan-Spermatophyta (see Definition: The largest crown clade containing entry in this volume) and the unnamed total biloba Linnaeus 1771 (Spermatophyta) clade of Monilophyta. and Pteridium aquilinum (Linnaeus) Kuhn 1879 (originally described as Pteris aquilina Diagnostic Apomorphies: Apomorphies rela- Linnaeus 1753) (Leptosporangiatae), but not tive to other crown clades include with Se/,aginella apoda (Linnaeus) Spring 1840 (origi- monopodial branching and endogenous lateral nally described as Lycopodium apodum Linnaeus roots (Schneider et al., 2002); sporangia termi- 1753) (Lycopodiophyta). This is a maximum- nal on lateral branches (Pryer et al., 2004a) and crown-clade definition. Abbreviated definition: dehiscing longitudinally (Doyle, 1998, 2013) max crown V (Ginkgo biloba Linnaeus 1771 & (these features characterize the earliest members Pteridium aquilinum (Linnaeus) Kuhn 1879 ~ of Pan-Euphyllophyta and were modified in most Se/,aginella apoda (Linnaeus) Spring 1840). extant representatives); lobed, mesarch primary xylem strand (Stein, 1993; Kenrick and Crane, Etymology: From the Greek eu (true), phyllon 1997: Fig. 7.10 and p. 241; Doyle, 1998, 2013), (leaf), and phyton (plant). Megaphylls (some- which has been modified in the stems of most times referred to as euphylls; e.g., Simpson, extant members; multiflagellate spermatozoids 2006; Schneider et al., 2009), which are leaves (apparently convergent in Isoetes) (Garbary et that usually have more than one vein and are al., 1993; Kenrick and Crane, 1997: 240, 275); thought to have evolved either from dichoto- a 30-kb inversion in the chloroplast genome mously forking lateral branches or from branch (Raubeson and Jansen, 1992). Megaphylls are systems bearing such units, are characteristic of sometimes cited as a synapomorphy of this clade chis clade though probably not a synapomorphy (Schneider et al., 2002), but analyses that include (see Diagnostic Apomorphies). fossils suggest that megaphylls of monilophytes and seed plants evolved independently (Stewart Reference Phylogeny: The primary reference and Rothwell, 1993; Kenrick and Crane, 1997; phylogeny is Pryer et al. (2001: Fig. 1, the clade Doyle, 1998, 2013; Boyce and Knoll, 2002; labeled Euphyllophytina). See also Kenrick and Friedman et al., 2004; Galtier, 2010). Even Crane (1997: Fig. 7.10), Pryer et al. (2004: Fig. within Lignophyta (i.e., Spermatophyta and their 3), Qiu et al. (2007: Fig. 1), Schneider et al. stem relatives that have a bifacial vascular cam- (2009: Figs. 1 and 2), Ruhfel et al. (2014: Figs. 6 bium; phylogenetically defined by Donoghue and 7), and Wickett et al. (2014: Figs. 2 and 3). and Doyle in Cantino et al., 2007), the small, wedge-shaped leaves of Archaeopteris may not Composition: All extant seed plants be homologous with the whole fernlike fronds (Spermatophyta), ferns (Leptosporangiatae, of "seed ferns" (a non-monophyletic assortment Ophioglossales, and Marattiales), horsetails of extinct seed plants with more or less - (), and whisk ferns (Psilotaceae), as like leaves), but rather with individual leaflets well as their extinct relatives that fall within the of such fronds (Doyle and Donoghue, 1986; Euphyllophyta

Doyle, 1998, 2013). The single-veined leaves of Euphyllophytina Kenrick and Crane (1997: Equisetum may be characterized as microphylls Table 7.1 and Fig. 7.10) referred to a more inclu- or megaphylls , depending on whether one's def- sive clade than the crown, though it is unclear inition of these terms is based on structure (spe- whether the name was applied to an apomor- cifically, presence of only one vein in the case phy-based or a total clade. The name was given of microphylls) or evolutionary origin (specifi- a "synapomorphy-based definition" in Table cally, from dichotomously forking branches or 7.2, but the clade was described (p. 240) as the branch systems in the case of megaphylls versus sister group of Lycophytina, suggesting that both from unvascularized enations [or possibly from of these clades were conceptualized as originat- sterilized sporangia; Kenrick and Crane, 1997] ing from their point of divergence rather than in the case of microphylls). Some fossil relatives from the origin of a particular apomorphy. In of Equisetum such as Sphenophyllum had multi- contrast, the same authors (Crane and Kenrick, ple-veined leaves that apparently evolved from 1997) used a "node-based" (minimum-clade) dichotomous branches (reviewed by Doyle, definition for Euphyllophytina. Although there 2013), suggesting that the one-veined leaves of is conflict within and between these two papers Equisetum would qualify as megaphylls if the as to whether the name Euphyllophytina applies definition is based on origin. to a minimum, apomorphy-based, or total clade, all three are more inclusive than the crown clade Synonyms: There are no synonyms; to which we apply the name Euphyllophyta here. Euphyllophytina (Kenrick and Crane, 1997; A maximum-crown-clade definition requires Crane and Kenrick, 1997) refers to a more only one internal specifier, but two are used inclusive clade (see Comments). here in order to disqualify the name under certain conditions. In the context of a phylog- Comments: A group composed of 'ferns' eny in which either 'ferns' or seed plants share (excluding Equisetum and Psilotophyta) and more recent ancestry with lycophytes than they seed plants has been recognized on mor- do with each other (e.g., Rothwell and Nixon phological grounds for many years (Banks, [2006: Fig. 6]; Ruhfel et al. [2014: Fig. 5]), the 1968), and the monophyly of this group, when name Euphyllophyta would not apply to any Equisetum and Psilotophyta are included, is clade. In our earlier definition (Cancino et al., strongly supported by molecular data (Pryer 2007), we included a species of Equisetum as a et al., 2001, 2004; Qiu et al., 2007; Wickett third internal specifier, but we have omitted it et al., 2014). However, there was no scientific here because it is unnecessary. To the best of our name for the crown clade until we explicitly knowledge, Euphyllophyta has the same compo- applied the name Euphyllophyta to it (Camino sition with either definition in the context of all et al., 2007). We incorrectly referred to it as a recently published phylogenies. new name, rather than converted, because we were unaware of its use by Lecointre and Le Literature Cited Guyader (2006). The latter authors were not explicit about the clade to which the name Banks, H. P. 1968. The early history of land plan rs. Euphyllophyta was intended to apply, bur their Pp. 73-107 in Evolution and Environment (E. inclusion of Psilophyton indicates that it was T. Drake, ed.). Yale University Press, New more inclusive than rhe crown (see Kenrick Haven, CT. and Crane, 1997: Fig. 4.32). Boyce, C. K., and A. H. Knoll. 2002. Evolution of developmencal potential and the multiple

244 Euphyllophyta

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dePamphilis, T. Chen, M. K. Deyholos, R. S. Michael J. Donoghue; Department of Ecology and Baucom, T. M. Kurchan, M. M. Augustin, J. Evolutionary Biology; Yale University; P.O. Wang, Y. Zhang, Z. Tian, Z. Yan, X. Wu, X. Box 208106; New Haven, CT 06520, USA. Sun, G. K. Wong, and]. Leebens-Mack. 2014. Email: [email protected]. Phylotranscripcomic analysis of the origin and early diversification of land plants. Proc. Natl. Date Accepted: 21 January 2013; updated 9 A cad. Sci. USA . 111 :E4859-E4868. February 2018

Authors Primary Editor: Kevin de Queiroz

Philip D. Camino; Department of Environmental and Plant Biology; Ohio University; Athens, OH 45701, USA. Email: [email protected].

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