Biology and Evolution of Ferns and Lycophytes

Cover......Page 1
Half-title......Page 3
Title......Page 5
Copyright......Page 6
Dedication......Page 7
Contents......Page 9
Contributors......Page 14
Preface......Page 17
Acknowledgments......Page 20
PART I DEVELOPMENT AND\rMORPHOGENESIS......Page 23
1.1 Introduction......Page 25
1.2 Spore germination......Page 29
1.3 Cell growth......Page 31
1.3.1 Growth cessation......Page 32
1.3.2 Resumption of cell growth......Page 33
1.4 Phototropism and polarotropism......Page 34
1.5 Cell division......Page 38
1.6 Apical cell bulging......Page 41
1.7 Chloroplast movement......Page 42
1.7.1 Cell sensitivity to light......Page 44
1.7.2 Induction of chloroplast movement......Page 45
1.7.3 How chloroplasts sense different fluence rates......Page 47
1.7.4 Signals transferred from a photoreceptor to chloroplasts......Page 48
1.7.5 Life of activated photoreceptors......Page 50
1.7.6 Speed of chloroplast movement......Page 51
1.8 Nuclear movement......Page 52
1.10 Photoreceptors for photomorphogenesis......Page 53
1.10.1 Phytochromes......Page 55
1.10.2 Phototropins......Page 56
1.10.3 Neochrome......Page 57
1.10.4 Cryptochromes......Page 60
1.11 Concluding remarks......Page 61
References......Page 62
2.1 Introduction......Page 71
2.2 ‘ ‘The” fern life cycle......Page 73
2.3 Historical summary......Page 74
2.4 Variations on a theme......Page 77
2.4.2 Facultative apogamy......Page 78
2.4.3 Obligate apogamy......Page 79
2.4.4 Apospory......Page 81
2.4.5 “Conventional” alternation of generations......Page 82
2.5 Sporogenesis......Page 83
2.5.1 Spore release and dispersal......Page 87
References......Page 90
3.1 Introduction......Page 97
3.2.1 Organization of shoot apical meristems of ferns and lycophytes......Page 98
3.2.2 Apical cell versus plural initial cells......Page 102
3.2.3 Evolution of organization of the shoot apical meristem......Page 103
3.3.1 Megaphyll development in ferns (euphyllous)......Page 104
3.3.2 Microphyll development in lycophytes......Page 105
3.3.3 Origin and evolution of megaphylls and microphylls......Page 106
3.4.1 Ferns......Page 108
3.4.2 Lycophytes......Page 111
3.5.1 Initiation and branching of fern roots......Page 112
3.5.2 Lycophyte root apical meristems and their branching......Page 114
3.5.3 Evolution of roots......Page 115
3.6 Psilotalean rhizomes (subterranean axes)......Page 116
3.7 Root-producing organs, rhizophores and rhizomorphs......Page 117
3.8 Summary and future goals......Page 119
References......Page 120
PART II GENETICS AND REPRODUCTION......Page 127
4.1 Introduction......Page 129
4.2 Population genetics and reproductive biology......Page 130
4.3 Genetic structure of populations......Page 143
4.4 Gene flow and divergence......Page 144
4.5 Population genetics of dispersal and colonization......Page 145
4.6 Summary and future prospects......Page 146
References......Page 147
5.2 History of discovery......Page 156
5.3 General effect of antheridiogen......Page 157
5.4 Occurrence of different antheridiogens and their chemical structure......Page 160
5.5 Experiments under laboratory conditions......Page 162
5.6 Dark germination: a further influence of antheridiogen......Page 165
5.7 Antheridiogen in nature......Page 167
5.8 Biological and evolutionary implications of the antheridiogen system......Page 171
5.9 Future goals......Page 172
References......Page 173
6.1 Introduction......Page 181
6.2 The golden age of fern chloroplast genomics......Page 183
6.3 The age of complete plastid genome sequences......Page 184
6.4.1 Background......Page 186
6.4.2 Methods......Page 189
6.4.3 Results......Page 190
References......Page 193
7.1 Introduction......Page 197
7.2 Historical summary......Page 199
7.3 Review of critical recent advances......Page 202
7.4 Synthesis of current perspectives......Page 210
7.5 Future goals and directions......Page 211
References......Page 215
PART III ECOLOGY......Page 221
8.2 Historical summary......Page 223
8.3 Review of critical recent advances......Page 225
8.3.1 Seasonality in tropical ferns......Page 226
Terrestrial ferns, especially tree ferns......Page 228
Climbing ferns and epiphytes......Page 230
Terrestrial species......Page 231
Mangrove species......Page 232
Lithophytes......Page 233
Epiphytes......Page 234
8.4 Synthesis of current perspectives......Page 237
8.6 Importance of long-term studies......Page 238
References......Page 239
9.1 Introduction......Page 244
9.2 Ecomorphology......Page 247
9.2.1 Ecological adaptations of gametophyte types......Page 250
9.2.2 Support for hypotheses of ecomorphological differentiation......Page 251
9.2.3 Evolution of growth form in fern gametophytes......Page 254
9.3 Ecophysiology......Page 255
9.3.1 Gametophyte autecology......Page 256
9.3.2 Water relations and desiccation tolerance......Page 258
9.3.3 Gametophyte carbon and light stress relations......Page 262
9.4.1 Morphological comparison of laboratory and field grown gametophytes......Page 264
9.4.2 Culture conditions and sexual development......Page 267
9.4.3 Effects of light, gravity and humidity on archegonial development......Page 269
9.5 Summary......Page 272
References......Page 273
10.1 Introduction......Page 279
10.2 Conservation importance of ferns and lycophytes......Page 280
10.3.1 Habitat fragmentation, degradation, and destruction......Page 282
10.3.2 Commercial collection......Page 284
10.3.4 Climate change......Page 285
10.4 Life cycle challenges for conservation......Page 286
10.6 Regional and ecosystem-level conservation......Page 287
10.7 Conservation of fern and lycophyte taxa......Page 290
10.8 Genetics in fern and lycophyte conservation strategies......Page 291
10.9.1 Planning protected areas......Page 292
10.9.2 Restoration......Page 293
10.10 Future directions in fern and lycophyte conservation......Page 294
References......Page 295
11.1 Introduction......Page 306
11.2.2 Ex situ banking of spores......Page 307
11.3 In vitro cultures and collections......Page 312
11.4 Ex situ cryostorage of gametophytes......Page 313
11.5 Ex situ cryostorage of sporophytes......Page 314
11.6 In vitro collecting for ex situ conservation......Page 315
11.7 Current status and future perspectives......Page 316
References......Page 318
PART IV SYSTEMATICS AND EVOLUTIONARY\rBIOLOGY......Page 323
12.1.1 Are species real or imagined?......Page 325
12.2 Species concepts and definitions......Page 326
12.2.1 Asexual species and cryptic species......Page 329
12.2.2 An example of progress in discovering species boundaries......Page 330
12.3 Speciation......Page 333
12.3.2 Primary speciation through allopatry......Page 334
12.3.3 Primary speciation through adaptation and/or changing\recological preferences......Page 337
12.3.4 Introduction to secondary speciation......Page 338
12.3.5 Secondary speciation through autopolyploidy......Page 339
12.3.7 Secondary speciation through allohomoploidy......Page 340
12.3.8 Secondary speciation through allopolyploidy......Page 342
12.3.9 Tertiary speciation......Page 343
12.4 Summary and future prospects......Page 344
References......Page 346
13.1 Introduction......Page 354
13.3 Systematic relationships among ferns, fern-like plants, and other euphyllophytes......Page 357
13.4 Moniliforms – the most ancient fern-like plants......Page 359
13.4.1 Cladoxylopsids and iridopterids......Page 360
13.4.2 Stauropterids......Page 363
13.4.3 Zygopterids......Page 364
13.5 Ophioglossid ferns......Page 365
13.6 Marattioid ferns......Page 367
13.7.1 Three evolutionary radiations of homosporous leptosporangiate ferns......Page 368
The first radiation......Page 369
The second radiation......Page 372
The third radiation......Page 375
13.7.2 Hydropteridales, the heterosporous leptosporangiate clade......Page 379
13.8 Historical context, popular practices, and the upward outlook......Page 380
References......Page 382
14.2 Historical review......Page 389
14.3 Diversity......Page 390
14.4 Long-distance dispersal......Page 394
14.5 Vicariance......Page 400
14.6 Floristics......Page 401
14.7 Important questions......Page 403
14.8 Future directions......Page 404
References......Page 405
15.1 Introduction......Page 417
15.2 Early vascular plant divergences......Page 418
15.3 Early fern divergences......Page 421
15.4 Early leptosporangiate divergences......Page 425
15.5 Early polypod divergences......Page 427
15.6 Divergences within eupolypods I......Page 428
15.7 Divergences within eupolypods II......Page 429
15.8 Future prospects......Page 430
References......Page 431
16.1 Introduction and historical summary......Page 439
16.2 Review of critical recent advances......Page 442
16.3 Synthesis of current perspectives: the classification of ferns......Page 443
A. Order Ophioglossales......Page 444
D. Order Marattiales (incl. Christenseniales)......Page 445
E. Order Osmundales......Page 446
G. Order Gleicheniales......Page 447
H. Order Schizaeales......Page 449
I. Order Salviniales......Page 450
J. Order Cyatheales......Page 451
K. Order Polypodiales......Page 454
16.4 Synthesis: lessons learned from morphology and molecular\rsystematics, and unexpected surprises......Page 467
References......Page 470
Appendix A: Familial names applied to extant ferns......Page 482
Appendix B: Index to genera......Page 484
Index......Page 490