Systematics, Evolution, and Biogeography of

Compositae

Edited by

Vicki A. Funk

Alfonso Susanna

Tod F. Stuessy

Randall J. Bayer

Contents

Foreword

Peter H. Raven xxi

Preface .' xxiii

Acknowledgements xxvii

Authors and addresses xxix

Section I: Introduction

1 A history of research in Compositae: early beginnings to the Reading Meeting (1975)J. Mauricio Bonifaciiw, Harold Robinson, Vicki A. Funk, Hans Walter Lack, Gerhard Wagenitz,Christian Feuillet and D.J. Nicholas Hind 3

Introduction 3Pre-Tournefort era 3Joseph Pitton de Tournefort (1656-1708) 5Sebastien Vaillant (1669-1722) 6Johannes Le Francq van Berkhey (1729-1812) 7Alexandre-Henri-Gabriel de Cassini (1781-1832) 9Christian Friedrich Lessing (1809-1862) 14Carl (Karl) Heinnch Schultz Bipontinus (1805-1867) 17George Bentham (1800-1884) 19Karl August Hoffmann (1853-1909) 22Benjamin Lincoln Robinson (1864-1935) 22James Small (1889-1955) 25Sidney Fay Blake (1892-1959) 26Hermann Merxmiiller (1920-1988) 27Jose Cuatrecasas Arumi (1903-1996) 29Angel Lulio Cabrera (1908-1999) 31Arthur Cronquist (1919-1992) 33The late 20th and early 21st century 34

VIII Contents

2 The recent history of Compositae systematics: from daisies to deep achenes, sister groupsand metatrees

Vernon H. Heywood 39

Introduction 39The situation in 1975 39Recent milestones in Compositae classification and phylogeny 40Revolutions in taxonomy and systematics 41So where are we now? 42Looking to the future 43

3 Economic importance of CompositaeBeryl B. Simpson 45

Introduction 45Edible crops 45Non-food crops 53Conclusions 57

Section II: Character evolution at the family level

4 A review of chromosome numbers in Asteraceae with hypotheses on chromosomal basenumber evolution

John C. Semple and Kuniaki Watanabe 61

Introduction 61Materials and methods 62Results and discussion 62

5 Secondary chemistry of CompositaeLalita M. Calabria, Vicente P. Emerenciano, Marcus T. Scotti and Tom]. Mabry 73

Introduction 73Materials and methods 75Results and discussion 76Conclusions 86

6 An introduction to micro-characters of CompositaeHarold Robinson 89

Introduction 89A survey of some characters 90Examples of micro-character application 99Some axioms regarding the use of micro-characters 99

7 Evolution of pollen in CompositaeStephen Blackmore, Alexandra H. Worthy, John]. Skvarla and Harold Robinson 101

Introduction 101Materials and methods 103Results 106

Evolution of pollen morphology based on the Compositae supertree 106Evolution of pollen morphology in exemplar tribes 112

Contents IX

Discussion 115

Conclusions 118

Appendix 7.1. Characters examined 121

Appendix 7.2. Matrix of pollen morphological characters for supertree taxa, using tribes as terminals 123

Appendix 7.3. Pollen descriptions for tribes of Compositae 126

8 Evolution of Compositae flowersCharles Jeffrey 131

Introduction 131

Capitulum 131

Receptacle 132

Achenes 132

Anthers 133

Breeding system 133

Florets 133

Pollination 134

Neoteny and evolution of the capitulum 134

Secondary pollen presentation 135

Dispersal, germination and establishment 135

Summary and conclusion 135

9 Genetic diversity in Asteraceae endemic to oceanic islands: Baker's Law and polyploidyDaniel J. Crawford, Timothy K, Lowrey, Gregory J. Andersdn, Gabriel Bernardello,Anioldo Santos-Guerra and Tod F. Stuessy 139

Introduction 139

Breeding system of colonizers 139

Polyploid colonizers 141

Materials and methods 141

Selection of archipelagos and plants 141

Chromosome numbers and polyploidy 141

Breeding systems 142

Results and discussion 142

Chromosome numbers and polyploidy in endemic island lineages 142

The significance of polyploid colonizers in the evolution of insular lineages 142

Sporophytic self-incompatibility and pseudo-self-fertility: general considerations 145

Breeding systems in colonizers of oceanic archipelagos 146

Summary and conclusions 147

Section III: Phylogeny, classification, and biogeography of the tribes

Part 1: Asterales

10 Asteraceae and relationships within AsteralesJohannes Lundberg 157

The search for the sister of Asteraceae 157

Plesiomorphic Asteraceae 157

Calyceraceae and Asteraceae 161

Goodeniaceae, Calyceraceae and Asteraceae 161

The MGCA clade: Menyanthaceae, Goodeniaceae, Calyceraceae and Asteraceae 161

The Core Asterales clade: MGCA, Stylidiaceae and APA 164

X Contents

Stylidiaceae 164The APA clade: Alseuosmiaceae, Phellinaceae and Argophyllaceae 165Pentaphragmataceae 165The basal division: Campanulaceae and Rousseaceae 166

11 Classification of CompositaeVicki A. Funk, Alfonso Susanna, Tod F. Stuessy and Harold Robinson 171

Introduction 171Compositae Giseke (1792) [Asteraceae Martynov (1820)| 173

Family description 173Current classification 176

Part 2: Basal clades

12 The basal grade of Compositae: Mutisieae (sensu Cabrera) and CarduoideaeSantiago Ortiz, J. Mauricio Bonifacino, Jorge V. Crisci, Vicki A. Funk, Hans V. Hanseti,D.J. Nicholas Hind, Liliana Katinas, Nddia Roque, Gisela Sancho, Alfonso Susannaand Maria Cristina Telleria 193

Introduction 193Mutisieae (sensu Cabrera 1977) 194Orphan Clades 195Catamixis incertae sedis 208Subfamily Carduoideae Cass. ex Sweet (1826) 210Conclusion 212

13 Barnadesieae (Barnadesioideae)Tod F. Stuessy, Estrella Urtubey and Michael Gruenstaeudl 215

Introduction 215Historical overview 216Description of subfamily 216

Morphology and anatomy 217Pollen 221Chromosome numbers 222Chemistry 222

Phylogenetic relationships 222Biogeography 223Evolution 225Economic uses 226

14 Mutisieae sensu stricto (Mutisioideae sensu stricto)Liliana Katinas, Gisela Sancho, Maria Cristina Telleria and Jorge V. Crisci 229

Introduction 229Historical overview 230Systematics 236Morphology 238Floral biology and pollination 242Biogeography 243Adaptation to dry environments 243Fossil pollen and early evolution 244

Contents XI

15 Gochnatieae (Gochnatioideae) and Hyalideae (Wunderlichioideae p.p.)Gisela Sancho and Susana E. Freire 249

Historical overview 249Phylogeny 249Tribe Gochnatieae 250

Taxonomy and biogeography 250Morphology 252Pollen 256Chromosome number 256Chemistry 256Floral biology 256Economic uses 256

Tribe Hyalideae 256Taxonomy and biogeography 256Morphology 257Pollen 258Chromosome number 258Chemistry 258

Discussion and conclusion 258

16 Hecastocleideae (Hecastocleidoideae)Vicki A. Funk and D. J. Nicholas Hind 261

Historical overview and morphology 261Phylogeny 261Taxonomy 261Pollen 263Chromosome number 264Chemistry 264Biogeography 264Biology, ecology, ethnobotany 264

17 Dicomeae (Carduoideae)Santiago Ortiz, Rodrigo Carbajal, Miguel Serrano and Antonio X.P. Coutinho 267

Historical overview 267Phylogeny 268Taxonomy 269Morphology 274Pollen 274Chromosome numbers 276Chemistry 276Ecology 276Biogeography 276Evolution 276Ethnobotany 277

18 Tarchonantheae (Carduoideae)Santiago Ortiz 279

Historical overview 279Phylogeny 279Taxonomy 280

XII Contents

Morphology 283Pollen 283Chromosome numbers 283Chemistry 283Ecology 283Biogeography 283Ethnobotany 284

19 Oldenburgieae (Carduoideae)Santiago Ortiz 287

Historical overview 287Phylogeny 287Taxonomy ' 288Morphology 288Pollen 290Chromosome numbers 290Chemistry 290Ecology 290Biogeography 290

20 Cardueae (Carduoideae)Alfonso Susanna and Nuria Garcia-Jacas 293

Historical overview 293Phylogeny 294Taxonomy 294Subtribal classification 296Chemistry 309Biogeography 309Origin and age 311Economic uses 311

21 Pertyeae (Pertyoideae)Susana E. Freire 315

Historical overview 315Phylogeny 315Taxonomy 315Morphology and anatomy 316Pollen 321Chromosome numbers 321Hybridization 321Ecology and floral biology 321Biogeography and evolution 321Ethnobotany 324Appendix 21.1. List of the taxa of Pertyeae and their distribution 325

22 Gymnarrheneae (Gymnarrhenoideae)Vicki A. Funk and Ori Fragman-Sapir 327

Introduction 327Historical overview 327Phylogeny 327

Contents XIII

Taxonomy 329Morphology 329Anatomy 329Pollen 329Chromosome numbers 330Chemistry 330Ecology and reproductive biology 330Applied aspects 331

Part 3: Cichorioideae and Corymbioideae

23 Introduction to CichorioideaeVicki A. Funk and Raymund Chan 335

Introduction 335Molecular phylogenetic analysis 336Phylogeny 337Taxonomy 339Chromosome numbers 340Chemistry 340Biogeography 340Placement of problematic genera 340

24 CichorieaeNorbert Kilian, Birgit Gemeinholzer and Hans Walter Lack 343

Introduction 343Historical overview 343Phylogeny 346Taxonomy 354Morphology and anatomy 355Chromosome numbers 367Chemistry 368Biogeography 368Evolution 372Economic uses 373Appendix 24.1. Subtribal classification of Cichorieae 380

25 ArctotideaePer Ola Karis, Vicki A. Funk, Robert J. McKenzie, Nigel P. Barker and Raymund Chan 385

Historical overview 385Phylogeny 386Subtribal treatments 388Arctotidinae 388

Taxonomy 388Morphology 388Pollen 393Chromosome numbers 393Chemistry 394Ecology 394Biogeography 394Economic uses 395Invasives 395

XIV Contents

Gorteriinae 395Taxonomy 395Morphology 397Pollen 401Chromosome numbers 403Chemistry 405Ecology 405Biogeography 406Economic uses 406Invasives 406

Character evolution 407Conclusion 408

26 EremothamneaeHarold Robinson and Vicki A. Funk 411

Historical overview and morphology 411Phylogeny 415Taxonomy 415Pollen 416Biogeography 416Chromosome numbers, chemistry, biology, ecology, ethnobotany 416

27 LiabeaeMichael O. Dillon, Vicki A. Funk, Harold Robinson and Raymund Chan 417

Historical overview 417Distribution and diversity 418Phylogeny and systematics 422Morphology and anatomy 425Pollen 428Chromosome numbers 431Biogeography 433Evolution 434Conclusions 434Appendix 27.1. Description of subtribe Sinclairiinae 437

28 VernonieaeSterling C. Keeley and Harold Robinson 439

Historical overview 439Phylogeny 440Taxonomy 447Morphology and anatomy 452Pollen 454Chromosome numbers 454Chemistry 456Ecology 456Biogeography 457Evolution 459Weeds 459Ethnobotanical and medicinal uses 460Economic uses 461

Contents XV

29 PlatycarpheaeVicki A. Funk, Marinda Koekemoer, Harold Robinson and John J. Skvarla 471

Historical overview 471Phylogeny 471Taxonomy 471Morphology 473Pollen 473Chromosome numbers 475Chemistry 475Biogeography 475Biology and ecology 475Ethnobotany 475

30 MoquinieaeHarold Robinson 477

Historical overview 477Phylogeny 477Taxonomy 477Pollen 479Chemistry 480Biogeography 481

31 Heterolepis: an unplaced genusVicki A. Funk and Per Ola Karis 483

Historical overview and morphology 483Phylogeny 484Taxonomy 484Pollen 484Chromosome numbers 484Chemistry 484Biogeography 484Ecology, conservation, horticulture 485

32 CorymbieaeBertil Nordenstam and Vicki A. Funk 487

Historical overview and tribal relationships 487Taxonomy 487Pollen 488Chromosome numbers 488Chemistry 490Biogeography 490Biology 490

Part 4: Asteroideae

33 Introduction to AsteroideaePieter B. Pelser and Linda E. Watson 495

Introduction 495The delimitation of Asteroideae and its phylogenetic position 495

XVI Contents

Tribal delimitation 496Tribal phylogeny 497Biogeography and age 498Chromosome numbers 500Chemistry 500

34 SenecioneaeBertil Nordenstam, Pieter B. Pelser, Joachim W. Kadereit and Linda E. Watson 503

Historical overview 503Phylogeny 508Taxonomy 513Morphology 515Pollen 517Chromosome numbers 517Chemistry 517Biogeography 518Evolution 521Economic uses 521Conclusion 521

35 CalenduleaeBertil Nordenstam and Marl Kdllersjo 527

Historical overview 527Phylogeny 528Taxonomy 529Morphology 530Pollen 535Chromosome numbers 535Chemistry 535Biogeography 535Evolution 536Economic uses 536Conclusions 537

36 GnaphalieaeJosephine Ward, Randall J. Bayer, Use Breitwieser, Rob Smissen, Merce Galbany-Casalsand Matthew Unwin 539

Introduction 539Historical overview 539Phylogeny 540Taxonomy 548Infratribal classification 551Morphology and anatomy 576Pollen 577Chromosome numbers 578Chemistry 579Ectomycorrhizal associations 579Biogeography 580Hybridization 580Horticulture 580Invasiveness 581

Contents XVII

Conservation/endangered species 581Ethnobotany 581Conclusions 581Appendix 36.1. Nomenclatural changes made since Bayer et al. (2007) 588

37 AstereaeLuc Brouillet, Timothy K. Lowrey, Lowell Urbatsch, Vesna Karaman-Castro, Gisela Sancho,Steve Wagstaff and John C. Semple 589

Introduction 589Materials and methods 590Results 590Phylogenetic lineages 591Classification 611Morphology, anatomy, palynology 613Chromosome numbers 614Chemistry 614Biology and evolution 615

Biogeography 617Economic uses ' 619Conclusion 620

38 AnthemideaeChristoph Oberprieler, Sven Hinunelreich, Marl Kdllersjo, Joan Valles, Linda E. Watsonand Robert Vogt 631

Historical overview 631Phylogeny 633Subtribal taxonomy 637Morphology 648Anatomy 653Pollen 654Embryology 655Chromosome numbers 655Chemistry 658Biogeography 659Evolution 661Economic uses 662

39 InuleaeAnte A. Anderberg 667

Historical overview 667Phylogeny 669Taxonomy 670Morphology and anatomy 671Pollen 678Chromosome numbers 678Chemistry 678Dispersal 678Biogeography 678Economic uses 679Conclusion 679

XVIII Contents

40 AthroismeaeArne A. Anderberg 681

Historical overview 681Phylogeny 684Taxonomy 685Morphology 685Pollen 686Chromosome numbers 686Chemistry 686Dispersal 686Biogeography 687Economic uses 687

41 Heliantheae allianceBruce G. Baldwin 689

Historical overview 689Reconsideration of Helenieae 691Homoplasy and taxonomic rethinking of epaleate clades 695The closest relatives of Eupatorieae 702Phylogenetic and tribal reassessment of paleate lineages 703Evolution 705Biogeography 707Conclusions 707

42 CoreopsideaeDaniel J. Crawford, Mesfin Tadesse, Mark E. Mort, Rebecca T Kimball and Christopher P. Randle 713

Historical overview and phylogeny 713Taxonomy 720Anatomy 725Pollen 725Chromosome numbers 725Chemistry 727Economic uses 727

43 EupatorieaeHarold Robinson, Edward Schilling and Jose L. Panero 731

Introduction 731Phylogeny 731Subtribal classification 735Evolution 743Conclusion 744

Section IV: Conclusion

44 Compositae metatrees: the next generationVicki A. Funk, Arne A. Anderberg, Bruce G. Baldwin, Randall J. Bayer, J. Mauricio Bonifacino,Use Breitwieser, Luc Brouilht, Rodrigo Carbajal, Raymund Chan, Antonio X. P. Coutinho,Daniel J. Crawford, Jorge V Crisci, Michael O. Dillon, Susana E. Freirc, Merce Galbany-Casals,Nuria Garcia-Jacas, Birgit Gemeinholzer, Michael Gruenstaeudl, Hans V. Hausen, Sven Himmelreich,

Contents XIX

Joachim W. Kadereit, Mari Kdllersjo, Vesna Karamau-Castro, Per Ola Karis, Liliana Katinas,Sterluig C. Keeley, Norbert Kilian, Rebecca T. Kimball, Timothy K. Lowrey, Johannes Lundberg,Robert J. McKenzie, Mesfin Tadesse, Mark E. Mort, Bertil Nordenstam, Christoph Oberprieler,Santiago Ortiz, Pieter B. Pelser, Christopher P. Randle, Harold Robinson, Nadia Roque, Gisela Sancho,John C. Seinple, Miguel Serrano, Tod F. Stuessy, Alfonso Susanna, Matthew Unwin, Lowell Urbatsch,

Estrella Urtubey, Joan Valles, Robert Vogt, Steve Wagstaff, Josephine Ward and Linda E. Watson 747

Introduction 747Materials and methods 748

Construction of the metatree 748Sources of the trees 749Outgroups 749Compositae 749Area optimization analysis using parsimony 754

Results and discussion 755The metatree and its sections 755Odd genera 767Age of origin 768

Conclusions 770

Section V: Appendices

A Illustrated glossary of CompositaeNadia Roque, David J. Keil and Alfonso Susanna 781

B Bibliography of pollen literature in CompositaeAlexandra H. Worthy, Stephen Blackmore and John J. Skvarla 807

Introduction 807Taxonomic listing of supertree genera with references 809Alphabetical bibliography 821List of generic ''synonyms" 863

C Original figure legends for plates in Chapter 1 869

D New names and combinations 873

E Complete list of literature cited 875

Taxon index 941