Austral Hepaticae, Part I

The proliferation of “species” — often placed in the “wrong” genus, has impacted all taxonamic work in 20th Century hepaticology. The same species may have been described under 3-4, or more, genera; under a single genus, such as Lepidolejeunea, species have been described in some 10-12 genera.
The greatest outstanding need is for a synthesis that attempts to critically define families and, in particular, genera and subgenera. For more than a century nobody has attempted such a synthesis.
This work attempts to answer this need by providing a modern synthesis of what we have learned, mostly in the second half of the twentieth century, about the families and genera of liverworts. It deals primarily with the conceptual bases that underlie a modern classification of the Hepaticae: the boundaries of suborders, families and genera are drawn by evaluating modern criteria. In such an evaluation emphasis is on features largely unused prior to 1950: ramification patterns and branch origins; capsule wall anatomy; seta and stem anatomy; oil-bodies and cell wall criteria.

211 figures, drawn by the author, illustrate these features and also show many taxa in drawings illustrating basic aspects of each genus. Almost all larger groups were studied from living plants and emphasis is on liverworts as living organisms.
The larger genera of Hepaticae are each illustrated with several plates and emphasis is on “key” genera which occupy critical phylogenetic positions, such as Lepidozia, Pseudocephalozia and Paracromastigum. No attempt is made at species diagnoses but generic boundaries, in larger genera, are defined by illustrating several species per genus. Except for genera with numerous species, keys to species are provided. In numbered annotations various problems, such as generic relationships and boundaries, are discussed; key species are often treated at length. As noted in the Introduction, since all taxa cannot be illustrated and/or described, for all larger genera the “type” method is used: boundaries are established by treating (and illustrating) key taxa so as to draw logical boundaries for each genus – – and family.
The author has conducted field work in the Antipodes for over 40 years, starting in Fuegia in 1960. He has not only become familiar with all the chief groups as living plants, but has studied them microscopically in areas as diverse as Fiji, the Prince Edward Islands., Campbell I., Chile and New Zealand, residing for months or sometimes years in these regions. The opportunity to study hepatics in the field has given him a unique opportunity to learn these organisms as living entities; to draw cytological details from living plants, and to place critical material into FAA for subsequent illustration, especially of the ephemeral sporophytes. Unlike most extant taxonamic works, which are based on herbarium study, this work tries to give the reader a “feeling” for the many taxa as living organisms.

Table of Contents
Preface 1
Introduction 3
Geographical Area Covered 6
Methodology and Approaches 7
Acknowledgements 8
Abbreviations 9
Classification of Hepaticae 10
Subclasses and Synopsis of Orders of Hepaticae 15
Subclass Jungermanniidae Schust.
Order I. Calobryales Campb. ex Schuster 17
Family l.Haplomitriaceae Dedecek (1886) 19
1. Haplomitrium Nees 19
Key to Species 21
Annotations 22
Subg. Haplomitrium Nees 26
Annotations 26
Subg. Calobryau (Nees) Schust. 27
Annotations 27
Order II. Jungermanniales 32
Synopsis of Suborders of Jungermanniales 38
Artificial Key to Genera of Jungermanniales 44
Suborder I. Lepicoleineae Schust. 75
Key to Families 77
Family 1. Vetaformaceae Fulf. & Taylor. 77
1. Vetaforma Fulf. & Taylor. 79
Annotations 79
Family 2. Lepicoleaceae Schust. 79
1. Lepicolea Dumort. 81
Key to Species 81
Annotations 82
Suborder II. Herbertineae Schust. 84
Key to Families 85
Family 1. Herbertaceae K. Müll. 86
Key to Genera 89
1. Olgantha Schust. 89
2. Triandrophyllum Fulf. & Hatch 90
Key to Subgenera and Species 92
Annotations 94
3. Herbertus SF Gray 94
Annotations 97
Family 2. Grolleaceae (Solari) Schust. 99
1. Grollea Schust. 101
Annotations 101
Family 3. Trichocoleaceae Nakai 101
Key to Subfamilies and Genera 104
Subfamily Temnomoideae Schust. 106
1. Archeophylla Schust. 107
Key to Species 110
Annotations 110
2. Archeochaete Schust. 113
Key to Species 116
Annotations 116
3. Temnoma Mitt. 119
Key to Subgenera and Species of Temnoma 123
Annotations 126
4. Pseudolepicolea Fulf. & Taylor. 133
Annotations 137
5. Herzogiaria Fulf. 137
6. Isophyllaria Hodgs. & Allis 139
Key to [tentative] Species of Isophyllaria 141
Subfamily Chaetocoleoideae Schust. 141
1. Chaetocolea Spr. 141
Annotations 1444
Subfamily Trichocoleoideae Nakai 144
1. Leiomitra Lindb. 146
Provisional Key to Subgenera and Species 147
Annotations 149
2. Trichocolea Dumort. 152
Tentative Key to Species 154
Annotations 156
3. Eotrichocolea Schust. 159
Subfamily Blepharostomatoideae Schust. ex Grolle 162
1. Blepharostoma (Dumort.) Dumort. 162
Annotations 165
Family 4. Trichotemnomaceae Schust. 166
1. Trichemnoma Schust. 166
Suborder III. Antheliineae Schust. 169
Family 1. Antheliaceae Schust. 170
1. Anthelia (Dumort.) Dumort. 170
Suborder IV. Lepidoziineae Schust. 172
Key to Families of Lepidoziineae 174
Family 1. Lepidoziaceae Limpr. 174
Key to Subfamilies and Genera of Lepidoziaceae 176
Subfamily 1. Lepidozioideae 180
1. Lepidozia (Dumort.) Dumort. 181
Provisional Key to Subgenera and [Most] Sections 183
Subg. 1. Dendrolepidozia Schust. 185
Subsections 2. Mastigolepidozia Schust., subg. n 187
Key to Species 187
Annotations 189
Subg. 3. Glaucolepidozia Schust. 189
Key to Species 192
Annotations 192
Subg. 4. Lepidozia 193
1. Sectio Setigerae Schust., sect. n. 193
2. Sectio Pendulinae Schust., sect. n . 197
3. Sectio Kirkii Schust., sect. n. 197
4. Sectio Austrolepidozia, sect. n. 197
Subg. 5. Chaetolepidozia Schust. 202
Subg. 6. Cladolepidozia Schust. 202
Annotations 206
2. Telaranea Spr. ex Schiffn. 207
Key to Subgenera 208
Subg. 1. Neolepidozia (Fulf. & Tayl.) Schust. 209
Synopsis of Taxa 211
Annotations 216
Subg. 2. Telaranea Spr. ex Schiffn. 221
Key to Species 222
Annotations 223
Subg. 3. Tricholepidozia Schust. 236
Annotations 236
3. Sprucella Steph. 240
Annotations 242
4. Kurzia v. Mart 244
Synopsis of Subgenera 246
Subg. 1. Micrisophylla (Fulf.) Engel ex Schust. 247
Key to Sections and Species 249
Annotations 249
Subg. 2. Microlepidozia (Spr.) Schust. 250
Key to Sections and Species 255
Annotations 259
Subg. 3. Nanolepidozia Schust. 263
Key to Species 264
Annotations 264
Subg. 4. Kurzia v. Mart 267
Key to Sections and Species 268
Annotations 268
Subg. 5. Austrolepidozia (Schust.) Schust., subg. n. 273
Key to Species 274
Annotations 274
Subg. 6. Dendrolembidium (Herz. emend. Schust.) Grolle 274
Key to Species 277
Annotations 278
5. Megalembidium Schust. 279
6. Psiloclada Mitt. 283
Key to Subspecies 285
Annotations 286
Subfamily 2. Drucelloideae (Bosen) Schust. 287
1. Drucella Hodgson 289
Annotations 289
Subfamily 3. Bazzanioideae Rodw. 291
Key to Genera 293
1. Acromastigum Evans 293
Key to Subgenera (and some Sections and Species) 295
Annotations 297
2. Bazzania SF Gray 304
Annotations 306
3. Mastigopelma Mitt. 310
Key to Species of Mastigopelma 312
Annotations 312
Subfamily 4. Neogrolleoideae Schust. 313
1. Neogrollea Hodgs. 316
Subfamily 5. Lembidioideae Schust. 316
Key to Genera and Subgenera of Lembidioideae 318
1. Isolembidium Schust. 319
2. Lembidium Mitt. 321
Key to Species of Lembidium 324
Annotations 324
3. Hygrolembidium Schust. 326
Key to Subgenera and Species of Hygrolembidium 329
Annotations 330
4. Chlorantelia Schust. 332
Key to Subgenera and Species of Chloranthelia 334
Annotations 336
Subfamily 6. Zoopsidoideae Schust. 338
Key to Genera of Zoopsidoideae 339
Annotations to Key. 341
1. Pseudocephalozia Schust 341
Key to Subgenera and Species of Pseudocephalozia 343
Annotations 345
2. Paracromastigum Fulf. & Taylor. 356
Key to Species 360
Annotations 364
Synopsis of the Neotropical P. granatensis-bifidum Complex 369
3. Hyalolepidozia S. Arn. ex Grolle 384
Annotations 387
4. Zoopsis Hook. f. & Tayl. ex GL & N. 390
Key to Subgenera and Species 395
Annotations 396
5. Zoopsidella Schust. 414
Key to Subgenera and Species 415
Annotations 418
6. Monodactylopsis (Schust.) Schust. 434
Annotations 435
7. Pteropsiella Spr. 439
Annotations 441
8. Arachniopsis Spr. 446
Key to Subgenera, Sections and Species of Arachniopis 450
Annotations 452
9. Odontoseries Fulf. 469
Subfamily 7. Micropterygioideae Grolle 470
1. Micropterygium Lindenb. in GL & N. 471
Key to Subgenera and Species of Micropterygium 472
Annotations 474
2. Mytilopsis Spruce 479
Annotations 481
Subfamily 8. Protocephalezioideae Schust. 483
1. Protocephalozia (Spr.) Goebel 484
Annotations 484
Family 2. Phycolepidoziaceae Schust. 486
1. Phycolepidozia Schust. 486
Annotations 488
Family 3. Calypogeiaceae HW Arn. 489
Key to Austral Genera 490
1. Calypogeia Raddi 490
Key to Subgenera of Calypogeia 492
Annotations 493
2. Mnioloma Herz 501
Key to Subgenera 503
Subg. 1. Caracoma (Bischl.) Schust. 504
Key to Species of Mnioloma subg. Caracoma 504
Annotations 505
Subg. 2. Mnioloma Heart 513
Annotations 515