Pan-Coniferae P. D. Cantino, M. J. Donoghue, and J. A. Doyle in P.D. Cantino et al. (2007): E20 [J. A. Doyle, P. D. Cantino, and M. J. Donoghue], converted clade name Registration Number: 77 Diagnostic Apomorphies: Based on a phy- logenetic tree that included Gnetophyta Definition: The total clade of the crown clade within Coniferae, Doyle (2006: Fig. 11) listed Coniferae. This is a crown-based total-clade the following synapomorphies for the clade definition. Abbreviated definition: otal V of labeled "Conifers" (including Emporia but not Coniferae. Cordaitales and corresponding roughly to Pan- Coniferae as defined here): leaves with a single Etymology: From the Greek pan- or pantos (all, vein (modified to several-veined within some the whole), indicating that the name refers to groups such as Agathis and Nageia), compound a total clade, and Coniferae (see entry in this female strobili but simple male strobili, bilateral volume for erymology), the name of the corre- (dorsiventral) symmetry in ovuliferous short sponding crown clade. shoots (modified into cone scales in extant taxa), inverted ovules, and sarcotesta absent or Reference Phylogeny: The reference phylogeny uniseriate. Of these, the sarcotesta character is Doyle (2006: Fig. 8). is questionable because it is unclear whether a sarcotesta is present or absent in the immature Composition: Coniferae (this volume) and all ovules of Emporia (Mapes and Rothwell, 1984). extinct plants that share more recent ancestry with Coniferae than with any other extant seed Synonyms: The names Pinopsida, Coniferophyta, plants. Morphological homologies between Coniferopsida, and Coniferales, which are listed extant and Palacozoic conifers and Cordaitales as approximate synonyms of Coniferae (this vol- proposed by Florin (1951) could be evidence ume), as well as Coniferae itself, could equally that cordaites in but well be to belong Pan-Coniferae, considered be approximate synonyms this isthe uncertain, because Florin (1949) adduced ofPan-Coniferae, since authors have rarely made similar evidence that Ginkgo and related fossils it clear whether these names apply to the crown, are also related to Cordaitales. In phylogenetic the total clade, or an intermediate clade analyses that include fossils, depending in part on the position of Ginkgo, Pan-Coniferae may Comments: We prefer the name. Pan-Coniferae include Cordaitales, Palaeozoic conifers such as rather than its approximate synonyms because Emporia and Lebachia, both, or neither (Crane, the name Coniferae (as defined in this volume) 1985; Doyle and Donoghue, 1986; Rothwell is unambiguous and neither Coniferae nor and Serbet, 1994; Doyle, 1996, 2006, 2008; Pan-Coniferae has any implication of rank. As Hilton and Bateman, 2006). Because of uncer discussed in the entry on Coniferae (this vol- tainty whether the clade Gnetophyta is part of ume), the names Pinopsida, Coniferophyta, and Coniferae see Coniferae in this volume), Pan- Coniferopsida have been used in broader senses. Coniferae may, similarly, include or exclude Moreover, the endings -opsida and -ales imply Gnetophyta. class and ordinal rank, respectively. Pan-Coniferae Literature Cited Hilton,J., and R. M. Bateman. 2006. Pteridosperms are the backbone of seed-plant phylogeny. ]. Cantino, P D., J. A. Doyle, S. W. Graham, W. S. Torey Bot. Soc.133:119-168. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, Mapes, G., and G. W. Rothwell. 1984. and M. J. Donoghue. 2007. Towards a phy- Permineralized ovulate cones of Lebachia logenetic nomenclature of Tracheophyta. Taxon from late Palaeozoic limestones of Kansas. 56:822-846 and E1-E44. Palaeontology 27:69-94. Crane, P. R. 1985. Phylogenetic analysis of seed Rothwell, G. W, and R. Serbet. 1994. Lignophyte plants and the origin of angiosperms. Ann. phylogeny and the evolution of spermato- Mo. Bot. Gard. 72:716-793. phytes: a numerical cladistic analysis. Syst. Bot. Doyle, J. A. 1996. Seed plant phylogeny and the 19:443-482. relationships of Gnetales. Int. J. Plant Sci. 157 (suppl):$3-S39. Authors Doyle, J. A. 2006. Seed ferns and the origin of angiosperms. J. Torrey Bot. Soc. 133: James A. Doyle; Department of Evolution and 169-209. Ecology; Universiry of California; Davis, CA Doyle, J. A. 2008. Integrating molecular phyloge 95616, USA. Email: [email protected]. netic and paleobotanical evidence on origin of Philip D. Cantino; Department of Environmental the flower. Int. J. Plant Sci. 169:816-843. and Plant Biology: Ohio University; Athens, Doyle, J. A., and M. J. Donoghue. 1986. Seed plant OH 45701, USA. Email: [email protected]. phylogeny and the origin of angiosperms: an Michael J. Donoghue; Department of Ecology and experimental cladistic approach. Bot. Reu Evolutionary Biology: Yale University; P.O. 52:321-431. Box 208106; New Haven, CT 06520, USA. Florin, R. 1949. The morphology of Trichopitys het- Email: [email protected]. eromorpha Saporta, a seed-plant of Palaeozoic age, and the evolution of the female ffow- ers in the Ginkgoinae. Acta Horti Bergiani Date Accepted: 29 August 2011; updated 09 15:79-109. February 2018 Florin, R. 1951. Evolution in cordaites and conifers. Acta Horti Bergiani 15:285-388. Primary Editor: Kevin de Queiroz 270 Coniferae A. L. Jussieu 1789: 411 [J.A. Doyle, P. D. Cantino, and M. J. Donoghue]. Converted clade name Registration Number: 31 see Diagnostic Apomorphies) is absent in Gnetophyta (Raubeson and Jansen, 1992). Most Definition: The smallest crown clade con- analyses based on morphology (see Comments) taining Pinus strobus Linnaeus 1753, Cupressus have not supported inclusion of sempervirens Linnaeus 1753, Podocarpus mac- Gnetophyt, but the clade Gnetophyta is nested in Coniferae rophyllus (Thunberg) Sweet 1818 (originally in some most-parsimonious trees in the mor- described as Taxus macrophylla Thunberg 1783), phological analysis of Doyle (2008). Although and Taxus baccata Linnaeus 1753. This is a min- Emporia (a Palacozoic conifer) or Emporia and imum-crown-clade definition. Abbreviated def- Cordaitales are part of Coniférae, as defined here, inition: min crown V (Pinus strobus Linnaeus in some of the trees of Rothwell and Serbet 1753 & Cupressus sempervirens Linnaeus 1753 (1994: Figs. 2a,b), this is not the case in their & Podocarpus macrophylus (Thunberg) Sweet other equally parsimonious trees (Figs. 1, 2c,d), 1818 &Taxus baccata Linnaeus 1753). or in those of other authors (Miller, 1999; Doyle, 1996, 2006, 2008; Hilton and Bateman, Etymology: Derived from Latin, conus (cone) 2006). and ferre (to carry or bear). Diagnostic Apomorphies: Synapomorphies Reference Phylogeny: The reference phylogeny relative to other crown clades include one- is Rydin et al. (2002: Fig. 1). See also Rai et al. veined needlelike leaves (modiied to several- (2008: Figs. 1, 2), Leslie et al. (2012: Fig. S4), veined in some groups), resin canals, compound Lu et al. (2014: Fig. 1), and Ruhfel et al. (2014: female strobili but simple male strobili, Fig. 5). tiered proembryos, siphonogamy (Doyle and Donoghue, 1986; Rothwell and Serbet, 1994; Composition: Araucariaceae, Cephalotaxaceae, Doyle, 2006, 2008), and loss or extreme reduc- Cupressaceae including "Taxodiaceae"), tion of one copy of the inverted repeat in the Pinaceae, Podocarpaceae, Taxaceae, and chloroplast genome (Raubeson and Jansen, Sciadopitys. The clade Gnetophyta (this volume) 1992; Wakasugi et al., 1994). Siphonogamy in is included within Coniferae in some trees based Coniferae is not with that in on homologous angio- molecular data et (Bowe al., 2000; Chaw et sperms and gnetophytes if phylogenies such as al, 2000; et Sanderson al., 2000; Gugerli et al., those of Crane (1985), Doyle and 2001; Donoghue Magallón and Sanderson, 2002; (1986), and (1996) are but it and Burleigh Doyle correct, Mathews, 2004; Zhong et al., 2010, may be with in the 2011; Xi homologous siphonogamy et al., 2013; Lu et al., 2014; Ruhfel latter groups in phylogenies that link Coniferae 2014), but not in others (Sanderson et with angiosperms and gnetophytes (Nixon et al., 2000; Magallón and Sanderson, 2002 al., 1994; Rothwell and Serbet, 1994; Doyle, Kydin et al., 2002; Burleigh and Mathews, 2006: Fig. 6), and it is presumably homologous 2004; Rai et al., The 2008). chloroplast-DNA with in if the latrer are Structural mutation siphonogamy Gnetophyta that characterizes coniters nested in Coniferae. Coniferae Synonyms: Pinopsida, Coniferophyta, our preference for names that are descriptive Coniferopsida, and Coniferales are approximate or end in phyta (meaning plants) rather than synonyms (see Comments). having an ending that is meaningless except in indicating rank, Coniferae and Coniferopyta are Comments: Many molecular analyses (e.g. the best candidate names for this clade. Here Stefanovic et al., 1998; Bowe et al., 2000; Chaw and previously (Cantino et al., 2007), we have et al., 2000; Gugerli et al., 2001; Magallón and chosen Coniferae over Coniferophyta because the Sanderson, 2002; Rydin et al., 2002; Soltis et informal names "coniferophytes" and "conifer- used tradi- al., 2002; Rai et al., 2008; Zhong et al., 2010, opsids" (generally synonymously) 2011; Leslie et al., 2012; Lu et al., 2014; Ruhfel tionally referred to a hypothesized larger group et al., 2014) and a morphologica. an ysis that includes Ginkgo and Cordaitales as well as Coulter and Chamberlain, 1910; (Hart, 1987) of extant conifers agreed that the conifers (e.g., clade Pinaceae or a clade comprising Pinaceae Chamberlain, 1935; Foster and Gifford, 1974). and Gnetophyta (see below) is sister to the rest The name Coniferophyta, which is not defined of the conifers. However, phylogenetic analyses in this volume, is best reserved for this larger based on morphology have variously suggested group in the context of phylogenies in which that the clade Taxaceae is the extant sister to the it is a clade (e.g, Crane, 1985; Doyle and rest (Miller, 1988, 1999; Rothwell and Serbet, Donoghue, 1986; Doyle, 1996, 2008). 1994), that a clade comprising Podocarpaceae Cited and Pinaceae occupies this position (Doyle, Literature 1996; Hilton and Bateman, 2006), or that the Bowe, L. M., G. Coat, and C. W. dePamphilis. of is unresolved relative position Podocarpaceae 2000.