180 CHAPTER 6 EVOLUTION OF FLOWERING PLANTS F REFERENCES FOR FURTHER STUDY Andrews, H. N. 1961. Studies in Paleobotany. Wiley, New York. APG ifi. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: 105—121. Crane, P. R., E. M. Friis, and K. Pedersen. 1995. The origin and early diversification of angiosperms. Nature 374: 27. 7 Crepet, W. L. 1998. The abominable mystery. Science 282: 1653—1654. Cronquist, A. 1981. An integrated system of classification of flowering plants. Columbia University Press, New York. Davies, T. J., T. G. Barraclough, M. W. Chase, P. 5. Soltis, D. E. Soltis, and V. Savolainen. 2004. Darwin’s abominable mystery: Insights from AND CLASSIFICATION a supertree of DIVERSITY the angiosperms. Proceedings of the National Academy of Sciences of the United States of America 101: 1904—1909. Doyle, J. A. 2006. Seed ferns and the origin of angiosperms. Journal of the Torrey Botanical Society 133: 169—209. Doyle, J. A. 2008. Integrating molecular phylogenetic and paleobotanical evidence on origin of the flower. International Journal of Plant FLOWERING PLANTS: Sciences 169: 816—843. OF Eames, A. J. 1961. Morphology of the angiosperms. McGraw Hill, New York. Friedman, W. and E., J. H. Williams. 2004. Developmental evolution of the sexual process in ancient flowering plant lineages. The Plant Cell NYMPHAEALE S, 16, S119—S 132, Supplement. AMBORELLALE S, Friedman, W. E., and K. C. Ryerson. 2009. Reconstructing the ancestral female gametophyte of angiosperms: insights from Amborella and other ancient lineages of flowering plants. American Journal of Botany 96: 129—143. MAGNOLIIDS, Friis, E. M., J. A. Doyle, AUSTROBAILEYALES, P. K. Endress, and Q. Leng. 2003. Archaefructus: Angiosperm precursor or specialized early angiosperm? Trends in Plant Science 8: 369—373. Friis, E. M., K. R. Pedersen, and P. R. Crane. 2000. 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Angiosperm phylogeny and evolution. Sinauer, Sunderland, MA. CDES 182 Stebbins, G. L. 1974. Flowering ]ORANG1OSPE Monoco1ed0flS 202 Plants: Evolution Above the Species Level. Belknap Press of Harvard University Press, Cambridge, MA. Classification of the Sun, G., D. L. Dilcher, FAMILY DESCRIPTIONS 184 S. Zheng, and Z. Zhou. 1998. In search of the first flower: A Jurassic angiosperm, Archaefructus, from Northeast 203 China. Science 282: 1692—1695, 185 ACOLES 203 Sun, G., Ji, D. L. Dilcher, S. Zheng, Nixon, Acoraceae Q. K. C. and X. Wang. 2002. Archaefructaceae, a new basal angiosperm family. Science 296: 899—904. 186 Stewart, W. N., and G. W. Rothwell. 1993. Paleobotany and the evolution of plants. Cambridge Univesity Press, New York. ALISMATALES 205 Takhtajan, A. L. 1991. Evolutionary 187 Trends in Flowering Plants. Columbia University Press, New York. 205 Thomas, s.aceae H. H. 1925. The Caytoniales, a new group of angiospermous plants from the Jurassic rocks of Yorkshire. Philosophical 187 Transactions of Alismataceae 208 the Royal Society of London 213: 299—363. 187 Vejt, B., P.. J. Schmidt, S. Hake, and M. F Yanofsky. 1993. Maize 210 floral development: new genes and old mutants. The Plant Cell 5: 1205—1215. 187 PETROSALVIALES Zaiiis, M. J., P. S. Soltis, Y. L. Qiu, E. Zimmer, and D. E. Soltis. 2003. Phylogenetic analyses and perianth evolution in basal angiosperms. 187 210 Annals of the Missouri Botanical Garden 90: 129—150. DIOSCOREALES 210 MAGNOLIIDS 189 Dioscoreaceae 190 LAUPALES PANDANALES 210 190 Lauraceae Pandanaceae 210 213 MAGNOLIALES 192 LILIALES 213 Annonaceae 192 Liliaceae Magnoliaceae 192 ASPARAGALES 213 192 218 PIPERALES Agavaceae 220 Aristolochiaceae 192 Alliaceae 220 Piperaceae 197 Amaryllidaceae 221 Saururaceae 197 Asphodelaceae 224 Iridaceae 224 CERATOPHYLLALES 197 Orchidaceae 229 Ceratophyllaceae 197 Themidaceae (Continued) 181 © 2010 Elsevier Inc. All right.s reserved. doi: 10, 1016/B970.012.37438O0.OOOOS 1 AND DIVERSITY OF PLANTS 183 UNIT 11 EVOLUTiON 182 CHAPTER 7 DIVERSITY AND CLASSIFICATION OF FLOWERING PLANTS — Angiospermae MagnoliophYta I . 230 POALES 249 COMMELINIDS Eudicots DASYPOGONACEAE 231 Bromeliaceae 250 T Cyperaceae 250 MagnoliidS rj - ARECALES 231 r .— — Eriocaulaceae 254 — I — 231 — Arecaceae (Palmae) Juncaceae 254 — rj — ‘ -.. Poaceae (Gramineae) 255 c. — 232 c’ COMMELINALES, ZINGIBERALES, AND POALES C’, S Restionaceae 258 — 0 232 COMMELINALES Sparganiaceae 262 235 0 :° — .— Commelinaceae Typhaceae 262 — — Haemodoraceae 235 Xyridaceae 262 Pontederiaceae 238 265 ZINGIBERALES 241 REVIEW O.LJESTIONS 266 Musaceae 241 EXERCISES Strelitziaceae 242 266 REFERENCES FOR FURTHER STUDY cotyledon one Zingiberaceae 245 245 WEB SITES 274 Cannaceae leaf venation parallel vasculature atactostelic, absent s.l.). The APG III system classifies one to several families into vascular cambium pollen tncolpate INTRODUCTION or tricolpate-derived orders (thus, each group having the ending “-ales”), where sieve tube plastids order is monophyletic. It proteiflaceoUS/Cuneate The phylogenetic relationships within the angiosperms has been strong evidence suggests that the that the designated orders are and continues to be a field of active research in plant systematics. must be understood, however, are not indicative of a Much progress has been made with the use of cladistic method not comparable evolutionary units and Chapter 2). For exam ology and the incorporation of morphological, anatomical, hierarchical classification system (see monophyletic group embryological, palynological, karyological, chemical, and ple, a single “order” may be sister to a orders. The orders can be viewed simply as molecular data (see Chapters 9—14). The more recent use of mul containing several families that appear tiple gene sequence data has been particularly useful in assessing convenient placeholders for one or more to comprise a monophyletic group with relatively high cer higher-level angiosperm relationships. However, the phyloge taxa) containing several orders vessels (secondarily lost in some netic relationships and classification presented in this chapter tainty. Some monophyletic groups Magnoliids, Monocotyledons can be viewed as somewhat preliminary, to be further refined are given names, such as (eudicots), Rosids, with continued research. For a more precise understanding of (monocots), Commelinids, Eudicotyledons group, there is no substitute for Fabids, Malvids, Asterids, Lamiids, and Campanulids. (2007), with selected apomor relationships within a particular APG ifi (2009) and Soltis et al. precise interrelationships of the major groups of angio of major angiosperm clades, after consulting the most recent, primary scientific literature. The FIGURE 7.1 Phylogenetic relationships uncertainty, but recent results have sperms still show some phies (see Chapter 6). begun to converge. Figure 7.1 illustrates higher-level phylo are paraphyletic and relationships from various analyses that are summa angiospermS other than monocots), MAJORANGIOSPERM CL4DES genetic assemblage, not to be recog (all angiospermS are a paraphyletic as a formal taxonomic unit. rized in APG III and modified from Soltis et al. (2007). Note orders are must be abandoned taxon. The families within the chapter and in Chapter 8, occur; further research may, in time, nized as a formal the descriptions in this Portrayal of the relationships of major angiosperm groups is that some polytomies monocots), Tables 7.2 In group. listed in Table 7.1 (all except the for each order or other major modeled (with very few exceptions) after the system of the resolve many of these. In particular, the elucidation of the eudicot exemplars are used and 7.3 (commeliflid monocots); limited in the context of of the flowering plants may yield insight (“basal” monocots) choice of these exemplars is very Angiosperm Phylogeny Group, 2009 (referred to as “APG III most basal branches Chapter 8. The not are listed in Tables 8.1—8.3 of angiospermS. These treatments are and APG II 2003. The into early angiosperm evolution and radiation. families the huge diversity of the 2009”), which supersedes APG 1998 angiospermS in terms of species diver on flower The great bulk of the substitute for the many fine references APG III system is based on published cladistic analyses pri As seen in Figure 7.1, the angiosperms can be broadly The mono- designed as a within the monocots and eudicots. the references at the end of delimited into several groups: the Amborellales, Nymphaeales, sity are contained plant family characteristics (see marily utilizing molecular data (e.g., Chase et al. 1993, 2000; approximatelY