A COMBINED CLADISTIC Owi I. Nandi2, Mark W. Chase3 andANALYSIS OF ANGIOSPERMS Peter K. Endress2USING rbcL ANDNON-MOLECULAR DATASETS'ABSTRACT A combined analysis of 162 extant angiosperm taxa for which rbcL sequence-data and/or an appreciable amount ofnon-molecular information is available was calculated. A non-molecular tree, an rbcL tree, and a combined tree arepresented. Only the rbcL and the combined data set show large numbers of groupings with bootstrap percentages greaterthan 50%, whereas the non-molecular trees show only eleven clades of this kind; this seems due to the number ofmissing cells in the non-molecular matrix. We tried to identify non-molecular characters (including biochemical) thatsupport groups present in these analyses, especially in cases where clades turned out to be new when compared to oneor more "classical" taxonomic systems. New groupings found in the non-molecular analysis that parallel the rbcLtopologies include a grade containing Illiciales, Austrobaileyaceae, and Amborellaceae (magnoliid II); a clade containingMagnoliales, Laurales, Aristolochianae, and monocots (magnoliid 1); a hamamelid group; subgroups of asterids (e.g., asimilar asterid III clade containing Scytopetalaceae, Lecythidaceae, Sapotaceae, Ebenaceae, Theaceae, Primulales,Styracaceae, Marcgraviaceae, Actinidiaceae, Clethraceae, and Ericales); an expanded caryophyllid group; a Malvaless.I. clade; a partial Malpighiales grade containing Quiinaceae, Linales s. str., Passiflorales, Violaceae, Kiggelariaceae,Flacourtiaceae s. str., and Ochnaceae; and some smaller clades, similar to the corresponding groups found in rbcLcladograms (Illiciales-Austrobaileyaceae; Aristolochianae-monocots; Hydrangeaceae-Cornales; Lecythidaceae-Scyto-petalaceae; Pittosporaceae-Araliales; Geissolomataceae-Stachyuraceae; Connaraceae-Oxalidaceae). Capparales s.l. andthe nitrogen-fixing clade, two novel molecular clades, are only found in the rbcL and the combined trees. Cistaceaehave been shown to share important characters with Malvales s.l. and are consistently found within this clade. Thesefindings argue against their previous inclusion in Violales. The rbcL tree contains 38 terminal taxa that are includedfor the first time in a published phylogeny. Considerable progress has been made in assembling a morphological/chemical data set that parallels the broad coverage of angiosperms seen in DNA studies. New opportunities for the study of seed-plantphylogeny have opened due to the continued de-velopment of computer hardware and software. Inaddition, gene sequencing has become reasonablyfast, and large nucleotide data matrices have beenproduced (e.g., Chase et al., 1993; Savolainen etal., 1996; Soltis et al., 1997b). These studies havestimulated even more molecular work on macrosys-tematics, including the addition of more "critical"taxa to the data matrices, comparison with resultsfrom other gene sequences, and the combination ofnucleotide with non-molecular data matrices, ashas been undertaken in this study. Other examplesof broadly sampled combined nucleotide and non-molecular studies are those of Doyle et al. (1994)and Chase et al. (1995). The non-molecular investigations of this studyoriginated from the question of the position of Cis-taceae within eudicots. Cistaceae have been in-cluded in Violales (Takhtajan, 1966; Cronquist,1981) and Malvales (Dahlgren, 1980), yet the mostnatural (i.e., phylogenetic) position has remained amatter of debate (Thorne, 1983, 1992). Thus, it wasa major objective of our non-molecular study toidentify the accurate position of Cistaceae and theirallies (Bixaceae and Cochlospermaceae) within theeudicots. Most families that have commonly beenallied with Malvales or Violales are included in ' We thank Thomas Baumann, Roland Eberwein, Helena Eklund, Mary Endress, Else Marie Friis, Robert Hegnauer,Anton Igersheim, Alexander Kocyan, Reto Nyffeler, Markus Reut, Rolf Rutishauser, Harald Schneider, Jtlrg Scht~nen-berger, Edwin Urmi, and Christian Wagner for their criticism and help, and Phil Ackery, Victor Albert, AbelardoAparicio, Pieter Baas, Harvey Ballard, Gilles Dutartre, James Farris, Jeffrey Harborne, Mark Hershkovitz, Larry Hufford,Donald Les, Richard Olmstead, and Valentina Ukraintseva for useful communications. We gratefully acknowledge UrsJauch, Shuqing He, Rosemarie Siegrist, Annette Nandi-Koch, and the many colleagues involved in rbcL sequencing:Bil Alverson, Diane Bowman, Anette de Bruijn, Kenneth Cameron, Michael Fay, James Hartwell, Harold Hills, SaraHoot, Lola Lled�, Cynthia Morton, Yin-Long Qiu, Vincent Savolainen, and Susan Swensen. Thanks are also due toWalter Judd, Douglas Soltis, and an anonymous reviewer for critically examining the manuscript. 2 Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, 8008 Zurich, Switzerland. : Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3DS, United Kingdom. ANN. MISSOURI BOT. GARD. 85: 137-212. 1998.
BioStor
