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MJ Donoghue and PD Cantino campanulidae M. J. Donoghue and P. D. Cantino in P. D. Cantino et al. (2007): 837 [M. J. Donoghue and P. D. Cantino], converted clade name Registration Number: 248 and outside of this clade (e.g., flower size, style length). Erbar and Leins (1996) showed that Definition: The largest crown clade containing "early sympetaly" is largely restricted to this Campanula latifolia Linnaeus 1753 (Asterales) clade, but its correlation with inferior ovary but not Lamium purpureum Linnaeus 1753 and reduced calyx should be explored further (Lamiidae!Lamiales) and Cornus mas Linnaeus (Endress, 2001), and its placement on the tree 1753 (Cornales) and Erica carnea Linnaeus 1753 remains uncertain. For example, it may be an (Ericales/Ericaceae). This is a maximum-crown- apomorphy of the less inclusive clade Apiidae clade definition. Abbreviated definition: max (defined in this volume), as suggested by Stevens crown V (Campanula latifolia Linnaeus 1753 ~ (2011). Lamium purpureum Linnaeus 1753 & Cornus mas Linnaeus 1753 & Erica carnea Linnaeus 1753). Synonyms: The informal names "asterid II", "euasterid(s) II", and "campanulids" are approx- Etymology: Derived from Campanula (name imate synonyms (see Comments). of an included taxon), which is Latin for "little bell" (Gledhill, 1989). Comments: Until we published the name Campanulidae (Camino et al., 2007), there Reference Phylogeny: The primary reference was no preexisting scientific name for this phylogeny is Soltis et al. (2011: Figs. 1, 2e-g). clade, which is strongly supported in molecu- See also Soltis et al. (2000: Figs. 1, 12), Karehed lar analyses (Soltis et al., 2000; Bremer et al., (2001: Figs. 1, 2), Bremer et al. (2002: Fig. 1), 2002; Tank and Donoghue, 2010; Soltis et al., Winkworth et al. (2008: Fig. 1), and Tank and 2011) and in an analysis that combined molecu- Donoghue (2010: Figs. 1, 3). lar and morphological data (Karehed, 2001). It has been referred to informally as "asterid II" Composition: Apiidae (this volume) and prob- (Chase et al., 1993), "euasterid(s) II" (APG, ably Aquifoliales sensu APG II (2003). There is 1998; Olmstead et al., 2000; Savolainen et al., a slight possibility that some or all of the taxa 2000; Soltis et al., 2000; Albach et al., 200la,b; chat are currently included in Aquifoliales are Lundberg, 2001; Judd et al., 2002; APG II, nor pan of Campanulidae (see Comments). 2003), and "campanulids" (Bremer et al., 2002; Judd and Olmstead, 2004; APG III, 2009). D· . •agnostic Apomorphies: We know of no The definition used here differs slightly from una b· m iguous non-molecular synapomorphies. our earlier one (Camino et al., 2007) in that Srevens (2011) cited several characters for this we no longer use Garrya elliptica as an external clade, including vessel elements with scalari- specifier. With 100% boostrap support for the form p r • l erroranons, sma 1 flowers, short styles, grouping of Garryales with the rest of Lamiidae copious d en osperm, and short embryos. Several (Soltis et al., 2011), there is no longer any need of thes h a . e c aracters are poorly sampled; others to include two external specifiers representing te ill-defined or highly variable both within Lamiidae. Campanulidae There is a slight possibility that flex APG III (Angiosperm Phylogeny Group III). 2009. (Aquifoliaceae) is a member of Lamiidae (as An update of the Angiosperm Phylogeny defined in this volume), rather than being Group classification for the orders and families plants: APG III. Bot. ]. Linn. Soc. closely related to Apiidae (this volume) as in of flowering 161:105-121. the reference phylogeny. flex was linked with Bremer, B., K. Bremer, N. Heidari, P. Erixon duplications Lamiidae in an analysis of RPB2 R. G. Olmstead, A. A. Anderberg, M'. (Oxelman et al., 2004) and in an analysis of Kallersjo, and E. Barkhordarian. 2002. matK sequences (Hilu et al., 2003). Because Phylogenetics of asterids based on 3 coding these studies did not include any members of and 3 non-coding chloroplast DNA mark- Helwingja, Phyllonoma, Cardiopteridaceae or ers and the utility of non-coding DNA at Mol. Phylogenet. Stemonuraceae, which have been linked strongly higher taxonomic levels. Evol. 24:274-301. with flex (Karehed, 2001; Bremer et al., 2002; Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Tank and Donoghue, 2010; Soltis et al. , 2011) Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis, could in Aquifoliales, these taxa presumably and M. J. Donoghue. 2007. Towards a phylo- also be related to Lamiidae. Our definition of genetic nomenclature of Tracheophyta. Taxon Campanulidae is designed to include flex and 56:822-846 and El-E44. these relatives (Aquifoliales) if they are more Chase, M. W., M. W. Chase, D. E. Soltis, R. G. closely related to Apiidae than to Lamiidae and Olmstead, D. Morgan, D. H. Les, B. D. to exclude them if that is not the case. If flex Mishler, M. R. Duvall, R. A. Price, H . G. Hills, Qiu, K. A. Kron, J. H . Rettig, E. Conti, and its relatives were to be found to be more Y.-L. J. D. Palmer, J. R. Manhart, K. J. Sytsma, H. closely related to Lamiidae than to Apiidae, then J. Michaels, W. J. Kress, K. G. Karol, W. D. become syn- Campanulidae and Apiidae would Clark, M. Hedren, B. S. Gaut, R. K. Jansen, onyms. As we stated previously (Camino et al., K.-J. Kim, C. F. Wimpee, J. F. Smith, G. R. 2007), it is our intent that Campanulidae have Fumier, S. H . Strauss, Q-Y. Xiang, G. M. precedence over Apiidae in the unlikely event Plunkett, P. S. Soltis, S. M. Swensen, S. E. that both names refer to the same clade. Williams, P. A. Gadek, C. J. Quinn, L. E. Eguiarte, E. Golenberg, G. H. Learn, Jr., S. W. Graham, S. C.H. Barrett, S. Dayanandan, Literature Cited and V. A. Albert. 1993. Phylogenetics of seed plants: an analysis of nucleotide sequences Albach, D. C., P. S. Soltis, and D. E. Soltis. 2001a. from the plastid gene rbcL. Ann. Mo. Bot. Patterns ofembryological and biochemical evo- Gard. 80:528-580. " lution in the asterids. Syst. Bot. 26:242-262. Endress, P. K. 2001. Origins of flower morphology. Albach, D. C., P. S. Soltis, D. E. Soltis, and R. G. J Exp. Zool. 291:105-115. Olmstead. 20016. Phylogenetic analysis of Erbar, C., and P. Leins. 1996. Distribution of asterids based on sequences of four genes. Ann. the character states "early sympetaly" and Mo. Bot. Gard. 88:163-212. "late sympetaly" within the "Sympetalae APG (Angiosperm Phylogeny Group). 1998. An Tetracyclicae" and presumably allied groups. ordinal classification for the families of flower- Bot. Acta 109:427-440. ing plants. Ann. Mo. Bot. Gard. 85:531-553. Gledhill, D. 1989. 1he Names ofPlants. 2nd edition. APG II (Angiosperm Phylogeny Group II). 2003. Cambridge University Press, Cambridge, UK. E. Soltis, An update of the Angiosperm Phylogeny Hilu, K. W., T. Borsch, K. Muller, D. , M. Group classification for the orders and families P. S. Soltis, V. Savolainen, M. W. Chase H . Sauquec, of flowering plants: APG II. Bot. j. Linn. Soc. P. Powell, L. A. Alice, R. Evans, , T. A. B. Slotta, J. G. Rohwer, 141 :399-436. C. Neinhuis 374 Campanulidae c, S. Campbell, and L. W. Chatrou. 2003. M.A. Gitzendanner, K. J. Sytsma, Y.-L. Qiu, Angiosperm phylogeny based on matKsequence K. H. Hilu, C. C. Davis, M. J. Sanderson, R. in form ation. Am. J Bot. 90:1758-1776. S. Beaman, R. G. Olmstead, W. S. Judd, M. J. Judd. W. S., C. S. Campbell, E. A. Kellogg, P. F. Donoghue, and P. S. Soltis. 2011. Angiosperm · Srevens, a nd M. J. Donoghue. 2002. Plant phylogeny: 17 genes, 640 taxa. Am. J Bot. 98: Systematics-A Phylogenetic Approach. 2nd edi- 704-730. tion. Sinauer Associates, Sunderland, MA. Soltis, D. E., P. S. Soltis, M. W. Chase, M. E. Mort, Judd, W. S., and R. G. Olmstead. 2004. A survey of D. C. Albach, M. Zanis, V. Savolainen, W. tricolpate (eudicot) phylogenetic relationships. H. Hahn, S. B. Hoot, M. F. Fay, M. Axtell, Am. J Bot. 91:1627-1644. S. M. Swensen, L. M. Prince, W. J. Kress, K. Kirehed, J. 2001. Multiple origin of the tropi- C. Nixon, and J. S. Farris. 2000. Angiosperm cal forest tree family lcacinaceae. Am. J Bot. phylogeny inferred from 18S rDNA, rbcL, and 88:2259-2274. atpBsequences. Bot.] Linn. Soc. 133:381-461. Lundberg, J. 2001. Phylogenetic studies in the Stevens, P. F. 2011. Angiosperm Phylogeny Website, Euasterids II, with particular reference to Version 9. Available at http://www.mobot.org/ Asterales and Escalloniaceae. Ph.D. dissertation, mobot/research/apweb/, last updated on 25 Uppsala University. Available at http://publi- June 2011. cations.uu.se/theses/abstract.xsql?dbid=1597. Tank, D. C., and M. J. Donoghue. 2010. Phylogeny Olmstead, R. G., K. Kim, R. K. Jansen, and S. and phylogenetic nomenclature of the J. Wagstaff. 2000. The phylogeny of the Campanulidae based on an expanded sample Asteridae sensu lato based on chloroplast ndhF of genes and taxa. Syst. Bot. 35:425-441. gene sequences. Mol. Phylogenet. Evol. 16: Winkworth, R. C., J. Lundberg, and M. J. 96-112. Donoghue. 2008. Toward a resolution of cam- Oxelman, B., N. Yoshikawa, B. L. McConaughy, panulid phylogeny, with special reference to J. Luo, A. L. Denton, and B. D. Hall. 2004. the placement of Dipsacales. Taxon 57:1-13. RPB2 gene phylogeny in flowering plants, with particular emphasis on asterids. Mol. Authors Phylogenet. Evol. 32:462-479. Savolainen, V. , M. W. Chase, S. B. Hoot, C. M. Michael J. Donoghue; Department of Ecology and Morton, D. E. Soltis, D. Bayer, M. F. Fay, Evolutionary Biology; Yale University; P.O.
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