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) ................................ 3
J. Mauricio Bonifacino, Harold Robinson, Vicki A. Funk,
Hans Walter Lack, Gerhard Wagenitz, Christian Feuillet
and D.J. Nicholas Hind
Introduction ................................................. 3
Pre-Tournefort era ........................................... 3
Joseph Pitton de Tournefort (1656-1708) ...................... 5
Sebastien Vaillant (1669-1722) ............................... 6
Johannes Le Francq van Berkhey (1729-1812) ................... 7
Alexandre-Henri-Gabriel de Cassini (1781-1832) ............... 9
Christian Friedrich Lessing (1809-1862) ..................... 14
Carl (Karl) Heinrich Schultz Bipontinus (1805-1867) ......... 17
George Bentham (1800-1884) .................................. 19
Karl August Hoffmann (1853-1909) ............................ 22
Benjamin Lincoln Robinson (1864-1935) ....................... 22
James Small (1889-1955) ..................................... 25
Sidney Fay Blake (1892-1959) ................................ 26
Hermann Merxmüller (1920-1988) .............................. 27
José Cuatrecasas Arumí (1903-1996) .......................... 29
Ángel Lulio Cabrera (1908-1999) ............................. 31
Arthur Cronquist (1919-1992) ................................ 33
The late 20th and early 21st century ......................... 34
2 The recent history of Compositae systematics: from daisies
to deep achenes, sister groups and metatrees ................ 39
Vernon H. Heywood
Introduction ................................................ 39
The situation in 1975 ....................................... 39
Recent milestones in Compositae classification and
phylogeny ................................................... 40
Revolutions in taxonomy and systematics ..................... 41
So where are we now? ........................................ 42
Looking to the future ....................................... 43
3 Economic importance of Compositae ........................... 45
Beryl B. Simpson
Introduction ................................................ 45
Edible crops ................................................ 45
Non-food crops .............................................. 53
Conclusions ................................................. 57
Section II: Character evolution at the family level
4 A review of chromosome numbers in Asteraceae with
hypotheses on chromosomal base number evolution ............. 61
John C. Semple and Kuniaki Watanabe
Introduction ................................................ 61
Materials and methods ....................................... 62
Results and discussion ...................................... 62
5 Secondary chemistry of Compositae ........................... 73
Lalita M. Calabria, Vicente P. Emerenciano, Marcus
T. Scotti and Tom J. Mabry
Introduction ................................................ 73
Materials and methods ....................................... 75
Results and discussion ...................................... 76
Conclusions ................................................. 86
6 An introduction to micro-characters of Compositae ........... 89
Harold Robinson
Introduction ................................................ 89
A survey of some characters ................................. 90
Examples of micro-character application ..................... 99
Some axioms regarding the use of micro-characters ........... 99
7 Evolution of pollen in Compositae .......................... 101
Stephen Blackmore, Alexandra H. Wortley, John J. Skvarla
and Harold Robinson
Introduction ............................................... 101
Materials and methods ...................................... 103
Results .................................................... 106
Evolution of pollen morphology based on the Compositae
supertree ............................................... 106
Evolution of pollen morphology in exemplar tribes ....... 112
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 flowers ............................ 131
Charles Jeffrey
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 polyploidy ........................ 139
Daniel J. Crawford, Timothy K. Lowrey, Gregory
J. Anderson, Gabriel Bemardello, Arnoldo Santos-Guerra
and Tod F. Stuessy
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 Asterales .............. 157
Johannes Lundberg
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 АРА ....... 164
Stylidiaceae ............................................... 164
The АРА clade: Alseuosmiaceae, Phellinaceae and
Argophyllaceae ............................................. 165
Pentaphragmataceae ......................................... 165
The basal division: Campanulaceae and Rousseaceae .......... 166
11 Classification of Compositae ............................... 171
Vicki A. Funk, Alfonso Susanna, Tod F. Stuessy and Harold
Robinson
Introduction ............................................... 171
Compositae Giseke (1792) [Asteraceae Martynov (1820)J ...... 173
Family description ...................................... 173
Current classification .................................. 176
Part 2: Basal clades
12 The basal grade of Compositae: Mutisieae (sensu Cabrera)
and Carduoideae ............................................ 193
Santiago Ortiz, J. Mauricio Bonifacino, Jorge V. Crisci,
Vicki A. Funk, Hans V. Hansen, D.J. Nicholas Hind,
Liliana Katinas, N&ádia Roque, Gisela Sancho, Alfonso
Susanna and Maria Cristina Tellcria
Introduction ............................................... 193
Mutisieae (sensu Cabrera 1977) ............................. 194
Orphan Clades .............................................. 195
Catamixis incertae sedis ................................... 208
Subfamily Carduoideae Cass, ex Sweet (1826) ................ 210
Conclusion ................................................. 212
13 Barnadesieae (Barnadesioideae) ............................. 215
Tod F. Stuessy, Estrella Urtubey and Michael
Gruenstaeudl
Introduction ............................................... 215
Historical overview ........................................ 216
Description of subfamily ................................... 216
Morphology and anatomy .................................. 217
Pollen .................................................. 221
Chromosome numbers ...................................... 222
Chemistry ............................................... 222
Phylogenetic relationships ................................. 222
Biogeography ............................................... 223
Evolution .................................................. 225
Economic uses .............................................. 226
14 Mutisieae sensu stricto (Mutisioideae sensu stricto) ....... 229
Liliana Katinas, Gisela Sancho, María Cristina Tellería
and Jorge V. Crisci
Introduction ............................................... 229
Historical overview ........................................ 230
Systematics ................................................ 236
Morphology ................................................. 238
Floral biology and pollination ............................. 242
Biogeography ............................................... 243
Adaptation to dry environments ............................. 243
Fossil pollen and early evolution .......................... 244
15 Gochnatieae (Gochnatioideae) and Hyalideae
(Wunderlichioideae p.p.) ................................... 249
Gisela Sancho and Susana E. Freire
Historical overview ........................................ 249
Phylogeny .................................................. 249
Tribe Gochnatieae .......................................... 250
Taxonomy and biogeography ............................... 250
Morphology .............................................. 252
Pollen .................................................. 256
Chromosome number ....................................... 256
Chemistry ............................................... 256
Floral biology .......................................... 256
Economic uses ........................................... 256
Tribe Hyalideae ............................................ 256
Taxonomy and biogeography ............................... 256
Morphology .............................................. 257
Pollen .................................................. 258
Chromosome number ....................................... 258
Chemistry ............................................... 258
Discussion and conclusion .................................. 258
16 Hecastocleideae (Hecastocleidoideae) ....................... 261
Vicki A. Funk and D.J. Nicholas Hind
Historical overview and morphology ......................... 261
Phylogeny .................................................. 261
Taxonomy ................................................... 261
Pollen ..................................................... 263
Chromosome number .......................................... 264
Chemistry .................................................. 264
Biogeography ............................................... 264
Biology, ecology, ethnobotany .............................. 264
17 Dicomeae (Carduoideae) ..................................... 267
Santiago Ortiz, Rodrigo Carbajal, Miguel Serrano and
Antonio X.P. Coutinho
Historical overview ........................................ 267
Phylogeny .................................................. 268
Taxonomy ................................................... 269
Morphology ................................................. 274
Pollen ..................................................... 274
Chromosome numbers ......................................... 276
Chemistry .................................................. 276
Ecology .................................................... 276
Biogeography ............................................... 276
Evolution .................................................. 276
Ethnobotany ................................................ 277
18 Tarchonantheae (Carduoideae) ............................... 279
Santiago Ortiz
Historical overview ........................................ 279
Phylogeny .................................................. 279
Taxonomy ................................................... 280
Morphology ................................................. 283
Pollen ..................................................... 283
Chromosome numbers ......................................... 283
Chemistry .................................................. 283
Ecology .................................................... 283
Biogeography ............................................... 283
Ethnobotany ................................................ 284
19 Oldenburgieae (Carduoideae) ................................ 287
Santiago Ortiz
Historical overview ........................................ 287
Phylogeny .................................................. 287
Taxonomy ................................................... 288
Morphology ................................................. 288
Pollen ..................................................... 290
Chromosome numbers ......................................... 290
Chemistry .................................................. 290
Ecology .................................................... 290
Biogeography ............................................... 290
20 Cardueae (Carduoideae) ..................................... 293
Alfonso Susanna and Núria Garcia-Jacas
Historical overview ........................................ 293
Phylogeny .................................................. 294
Taxonomy ................................................... 294
Subtribal classification ................................... 296
Chemistry .................................................. 309
Biogeography ............................................... 309
Origin and age ............................................. 311
Economic uses .............................................. 311
21 Pertyeae (Pertyoideae) ..................................... 315
Susana E. Freire
Historical overview ........................................ 315
Phylogeny .................................................. 315
Taxonomy ................................................... 315
Morphology and anatomy ..................................... 316
Pollen ..................................................... 321
Chromosome numbers ......................................... 321
Hybridization .............................................. 321
Ecology and floral biology ................................. 321
Biogeography and evolution ................................. 321
Ethnobotany ................................................ 324
Appendix 21.1. List of the taxa of Pertyeae and their
distribution ............................................... 325
22 Gymnarrheneae (Gymnarrhenoideae) ........................... 327
Vicki A. Funk and Ori Fragman-Sapir
Introduction ............................................... 327
Historical overview ........................................ 327
Phylogeny .................................................. 327
Taxonomy ................................................... 329
Morphology ................................................. 329
Anatomy .................................................... 329
Pollen ..................................................... 329
Chromosome numbers ......................................... 330
Chemistry .................................................. 330
Ecology and reproductive biology ........................... 330
Applied aspects ............................................ 331
Part 3: Cichorioideae and Corymbioideae
23 Introduction to Cichorioideae .............................. 335
Vicki A. Funk and Raymund Chan
Introduction ............................................... 335
Molecular phylogenetic analysis ............................ 336
Phylogeny .................................................. 337
Taxonomy ................................................... 339
Chromosome numbers ......................................... 340
Chemistry .................................................. 340
Biogeography ............................................... 340
Placement of problematic genera ............................ 340
24 Cichorieae ................................................. 343
Norbert Kilian, Birgit Gemeinholzer and Hans Walter Lack
Introduction ............................................... 343
Historical overview ........................................ 343
Phylogeny .................................................. 346
Taxonomy ................................................... 354
Morphology and anatomy ..................................... 355
Chromosome numbers ......................................... 367
Chemistry .................................................. 368
Biogeography ............................................... 368
Evolution .................................................. 372
Economic uses .............................................. 373
Appendix 24.1. Subtribal classification of Cichorieae ...... 380
25 Arctotideae ................................................ 385
Per Ola Karis, Vicki A. Funk, Robert J. McKenzie, Nigel
P. Barker and Raymund Chan
Historical overview ........................................ 385
Phylogeny .................................................. 386
Subtribal treatments ....................................... 388
Arctotidinae ............................................... 388
Taxonomy ................................................ 388
Morphology .............................................. 388
Pollen .................................................. 393
Chromosome numbers ...................................... 393
Chemistry ............................................... 394
Ecology ................................................. 394
Biogeography ............................................ 394
Economic uses ........................................... 395
Invasives ............................................... 395
Gorteriinae ............................................. 395
Taxonomy ................................................ 395
Morphology .............................................. 397
Pollen .................................................. 401
Chromosome numbers ...................................... 403
Chemistry ............................................... 405
Ecology ................................................. 405
Biogeography ............................................ 406
Economic uses ........................................... 406
Invasives ............................................... 406
Character evolution ........................................ 407
Conclusion ................................................. 408
26 Eremothamneae .............................................. 411
Harold Robinson and Vicki A. Funk
Historical overview and morphology ......................... 411
Phylogeny .................................................. 415
Taxonomy ................................................... 415
Pollen ..................................................... 416
Biogeography ............................................... 416
Chromosome numbers, chemistry, biology, ecology,
ethnobotany ................................................ 416
27 Liabeae .................................................... 417
Michael O. Dillon, Vicki A. Funk, Harold Robinson and
Raymund Chan
Historical overview ........................................ 417
Distribution and diversity ................................. 418
Phylogeny and systetnatics ................................. 422
Morphology and anatomy ..................................... 425
Pollen ..................................................... 428
Chromosome numbers ......................................... 431
Biogeography ............................................... 433
Evolution .................................................. 434
Conclusions ................................................ 434
Appendix 27.1. Description of subtribe Sinclairiinae ....... 437
28 Vernonieae ................................................. 439
Sterling C. Keeley and Harold Robinson
Historical overview ........................................ 439
Phylogeny .................................................. 440
Taxonomy ................................................... 447
Morphology and anatomy ..................................... 452
Pollen ..................................................... 454
Chromosome numbers ......................................... 454
Chemistry .................................................. 456
Ecology .................................................... 456
Biogeography ............................................... 457
Evolution .................................................. 459
Weeds ...................................................... 459
Ethnobotanical and medicinal uses .......................... 460
Economic uses .............................................. 461
29 Platycarpheae .............................................. 471
Vicki A. Funk, Marinda Koekemoer, Harold Robinson and
John J. Skvarla
Historical overview ........................................ 471
Phylogeny .................................................. 471
Taxonomy ................................................... 471
Morphology ................................................. 473
Pollen ..................................................... 473
Chromosome numbers ......................................... 475
Chemistry .................................................. 475
Biogeography ............................................... 475
Biology and ecology ........................................ 475
Ethnobotany ................................................ 475
30 Moquinieae ................................................. 477
Harold Robinson
Historical overview ........................................ 477
Phylogeny .................................................. 477
Taxonomy ................................................... 477
Pollen ..................................................... 479
Chemistry .................................................. 480
Biogeography ............................................... 481
31 Heterolepis: an unplaced genus ............................. 483
Vicki A. Funk and Per Ola Karis
Historical overview and morphology ......................... 483
Phylogeny .................................................. 484
Taxonomy ................................................... 484
Pollen ..................................................... 484
Chromosome numbers ......................................... 484
Chemistry .................................................. 484
Biogeography ............................................... 484
Ecology, conservation, horticulture ........................ 485
32 Corymbieae ................................................. 487
Bertil Nordenstam and Vicki A. Funk
Historical overview and tribal relationships ............... 487
Taxonomy ................................................... 487
Pollen ..................................................... 488
Chromosome numbers ......................................... 488
Chemistry .................................................. 490
Biogeography ............................................... 490
Biology .................................................... 490
Part 4: Asteroideae
33 Introduction to Asteroideae ................................ 495
Pieter B. Pelser and Linda E. Watson
Introduction ............................................... 495
The delimitation of Asteroideae and its phylogenetic
position ................................................... 495
Tribal delimitation ........................................ 496
Tribal phylogeny ........................................... 497
Biogeography and age ....................................... 498
Chromosome numbers ......................................... 500
Chemistry .................................................. 500
34 Senecioneae ................................................ 503
Bertil Nordenstam, Pieter B. Pelser, Joachim W. Kadereit
and Linda E. Watson
Historical overview ........................................ 503
Phylogeny .................................................. 508
Taxonomy ................................................... 513
Morphology ................................................. 515
Pollen ..................................................... 517
Chromosome numbers ......................................... 517
Chemistry .................................................. 517
Biogeography ............................................... 518
Evolution .................................................. 521
Economic uses .............................................. 521
Conclusion ................................................. 521
35 Calenduleae ................................................ 527
Bertil Nordenstam and Mari Källersjö
Historical overview ........................................ 527
Phylogeny .................................................. 528
Taxonomy ................................................... 529
Morphology ................................................. 530
Pollen ..................................................... 535
Chromosome numbers ......................................... 535
Chemistry .................................................. 535
Biogeography ............................................... 535
Evolution .................................................. 536
Economic uses .............................................. 536
Conclusions ................................................ 537
36 Gnaphalieae ................................................ 539
Josephine Ward, Randall J. Bayer, Use Breitwieser, Rob
Smissen, Mercè Galbany-Casals and Matthew Unwin
Introduction ............................................... 539
Historical overview ........................................ 539
Phylogeny .................................................. 540
Taxonomy ................................................... 548
Infratribal classification ................................. 551
Morphology and anatomy ..................................... 576
Pollen ..................................................... 577
Chromosome numbers ......................................... 578
Chemistry .................................................. 579
Ectomycorrhizal associations ............................... 579
Biogeography ............................................... 580
Hybridization .............................................. 580
Horticulture ............................................... 580
Invasiveness ............................................... 581
Conservation/endangered species ............................ 581
Ethnobotany ................................................ 581
Conclusions ................................................ 581
Appendix 36.1. Nomenclatural changes made since Bayer et
al. (2007) .................................. 588
37 Astereae ................................................... 589
Luc Brouillet, Timothy K. Lowrey, Lowell Urbatsch, Vesna
Karaman-Castro, Gisela Sancho, Steve Wagstaff and John
C. Semple
Introduction ............................................... 589
Materials and methods ...................................... 590
Results .................................................... 590
Phylogenetic lineages ...................................... 591
Classification ............................................. 611
Morphology, anatomy, palynology ............................ 613
Chromosome numbers ......................................... 614
Chemistry .................................................. 614
Biology and evolution ...................................... 615
Biogeography ............................................... 617
Economic uses .............................................. 619
Conclusion ................................................. 620
38 Anthemideae ................................................ 631
Christoph Oberprieler, Sven Himmelreich, Mari Kälkrsjö,
Joan Vallès, Linda E. Watson and Robert Vogt
Historical overview ........................................ 631
Phytogeny .................................................. 633
Subtribal taxonomy ......................................... 637
Morphology ................................................. 648
Anatomy .................................................... 653
Pollen ..................................................... 654
Embryology ................................................. 655
Chromosome numbers ......................................... 655
Chemistry .................................................. 658
Biogeography ............................................... 659
Evolution .................................................. 661
Economic uses .............................................. 662
39 Inuleae .................................................... 667
Arne A. Anderberg
Historical overview ........................................ 667
Phylogeny .................................................. 669
Taxonomy ................................................... 670
Morphology and anatomy ..................................... 671
Pollen ..................................................... 678
Chromosome numbers ......................................... 678
Chemistry .................................................. 678
Dispersal .................................................. 678
Biogeography ............................................... 678
Economic uses .............................................. 679
Conclusion ................................................. 679
40 Athroismeae ................................................ 681
Arne A. Anderberg
Historical overview ........................................ 681
Phylogeny .................................................. 684
Taxonomy ................................................... 685
Morphology ................................................. 685
Pollen ..................................................... 686
Chromosome numbers ......................................... 686
Chemistry .................................................. 686
Dispersal .................................................. 686
Biogeography ............................................... 687
Economic uses .............................................. 687
41 Heliantheae alliance ....................................... 689
Bruce G. Baldwin
Historical overview ........................................ 689
Reconsideration of Helenieae ............................... 691
Homoplasy and taxonomic rethinking of epaleate clades ...... 695
The closest relatives of Eupatorieae ....................... 702
Phylogenetic and tribal reassessment of paleate lineages ... 703
Evolution .................................................. 705
Biogeography ............................................... 707
Conclusions ................................................ 707
42 Coreopsideae ............................................... 713
Daniel J. Crawford, Mesfin Tadesse, Mark E. Mort,
Rebecca T. Kimball and Christopher P. Randle
Historical overview and phylogeny .......................... 713
Taxonomy ................................................... 720
Anatomy .................................................... 725
Pollen ..................................................... 725
Chromosome numbers ......................................... 725
Chemistry .................................................. 727
Economic uses .............................................. 727
43 Eupatorieae ................................................ 731
Harold Robinson, Edward Schilling and José L. Panero
Introduction ............................................... 731
Phylogeny .................................................. 731
Subtribal classification ................................... 735
Evolution .................................................. 743
Conclusion ................................................. 744
Section IV: Conclusion
44 Compositae metatrees: the next generation .................. 747
Vicki A. Funk, Arne A. Anderberg, Bruce G. Baldwin,
Randall J. Bayer, J. Mauricio Bonifacino, Ilse
Breitwieser, Luc Brouillet, Rodrigo Carbajal,
Raymund Chan, Antonio X.P. Coutinho, Daniel J.
Crawford, Jorge V. Crisci, Michael O. Dillon, Susana
E. Freire, Mercè Galbany-Casals, Núria Garcia-Jacas,
Birgit Gemeinholzer, Michael Gruenstaeudl, Hans V.
Hansen, Sven Himmelreich, Joachim W. Kadereit, Mari
Källersjö, Vesna Karaman-Castro, Per Ola Karis,
Liliana Katinas, Sterling 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, Nádia Roque, Gisela
Sancho, John C. Semple, Miguel Serrano, Tod F.
Stuessy, Alfonso Susanna, Matthew Unwin, Lowell
Urbatsch, Estrella Urtubey, Joan Vallès, Robert
Vogt, Steve Wagstaff, Josephine Ward and Linda
E. Watson
Introduction ............................................... 747
Materials and methods ...................................... 748
Construction of the metatree ............................ 748
Sources of the trees .................................... 749
Outgroups ............................................... 749
Compositae .............................................. 749
Area optimization analysis using parsimony .............. 754
Results and discussion ..................................... 755
The metatree and its sections ........................... 755
Odd genera .............................................. 767
Age of origin ........................................... 768
Conclusions ................................................ 770
Section V: Appendices
A Illustrated glossary of Compositae
Nádia Roque, David J. Keil and Alfonso Susanna ............. 781
В Bibliography of pollen literature in Compositae
Alexandra H. Wortley, Stephen Blackmore and John
J. Skvarla ................................................. 807
Introduction ............................................... 807
Taxonomic listing of supertree genera with references ...... 809
Alphabetical bibliography .................................. 821
List of generic "synonyms" ................................. 863
С 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
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