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Polypodiaceae BLUMEA 34 (1990) 277-423 A monograph of the fern genus Goniophlebium (Polypodiaceae) Gisela Rödl-Linder Contents 279 1. Summary and Introduction GENERAL PART 2. Material and Methods 281 281 2.1. Material 2.2. Methods 281 283 3. Taxonomic history of Goniophlebium 285 4. Morphology and anatomy of mature sporophytes 4.1. Introduction 285 4.2. Rhizome 290 4.2.1. Morphology 291 4.2.2. Anatomy 293 4.3. Rhizome scales 293 4.4. Fronds 296 4.4.1. Gross morphology 297 4.4.2. Venation pattern 299 4.4.3. Indument 301 4.4.4. Sori, paraphyses and sporangia 302 4.5. Spores 306 307 • 5. Gametophytes • • • 5.1. Introduction 307 5.2. Observations and Discussion; Conclusion 308 6. Juvenile sporophytes 310 6.1. Introduction 310 6.2. Indument 311 6.2.1. Acicularhairs 311 6.2.2 Glandular hairs 311 6.2.3. Scales 312 7. Karyology 315 (continuedon page 278) * Correspondence address: G. ROdl-Linder, 54 Plaza v. Humboldt,La Florida, La Pas, Bolivia. 278 BLUMEA VOL. 34, No. 2, 1990 (Contents continued) 8. Geography and habitat 317 8.1. Distribution 317 8.2. Habitat 318 8.3. Transplantexperiments 328 9. Phylogeny 330 9.1. Methodology; Procedure 330 9.2. Datamatrices 331 9.3. Cladistic analysis of the goniophlebioidPolypodiaceae 334 9.3.1. Cladistic analysis of Goniophlebium 339 Cladistic the 9.3.2. analysis of Goniophlebiumpercussum- group 341 Cladistic of the 9.3.3. analysis Goniophlebium, excluding G. percussum- group 343 9.4. Choice of the cladogram of the genus Goniophlebium 345 9.5. Transformation series 349 10. The monophyly and systematic position of Goniophlebium 351 10.1. of the 351 Delimitation genus of 10.2. Delimitation the groups 353 10.3. Systematic position of Goniophlebium 354 10.4. Delimitation of the species 354 11. Biogeographical aspects 355 12. A scenario for Goniophlebium 358 12.1. View of previous authors 358 12.2. Introduction 358 A scenario for the 359 12.3. Goniophlebiumpercussum-group 12.4. A scenario for the Goniophlebium subauriculatum-group 359 13. References 360 PLATES 366 TAXONOMIC PART 14. Presentation of data 370 15. Goniophlebium: 371 16. Key to the species of Goniophlebium 373 17. Descriptions 377 17.1. Goniophlebiumpercussum- group 377 17.2. subauriculatum- 387 Goniophlebium group and 17.3. Goniophlebium argutum-, hendersonii- niponicum-group 403 17.4. Species dubiaeatqueSpecies exclusae 412 17.5. Polypodium species studied in comparison with Goniophlebium species 414 18. Index of specimens 417 19. Acknowledgements 421 20. Index totaxonomic names 421 ROdl-Linder: the 279 G. Monograph of fern genus Goniophlebium 1. Summary and introduction The present monographic study deals with the phylogenetic systematics of the goniophlebioid Polypodiaceae, with special reference to the Asian representatives. It includes the recognition and full and reconstruction their description of species of Goniophlebium, a synonymy, a of phylogeny. This study concentrates on the analysis of 34 species earlier referred to Polypo- dium Linnaeus (sensu lato) with a (partly) goniophlebioid venationpattern and also includes a selection of free veinedPolypodium species. Status and nomenclature of the goniophlebioids are most ambiguous and show much variation. Several earlier attempts by other authors towards a natural classifi- cation have remained unsuccessful as is elucidated in a paragraph on the taxonomic history of the group. In orderto unravel the phylogenetic relationships within this group an analysis of characters, i.e. series of inferred homologous character states, was undertaken. They concern the morphology of the mature sporophyte, with respect to the rhizome, the rhizome scales, the gross morphology, the venation pattern, the indument, the son, and the and the paraphyses, the sporangia spores. Microscopical submicroscopical features were included. Several of these characters were studied in detail for this for the first time. group A selection of 90 character states was used in the datamatrix, which served as a prerequisite for cladistic analyses. Following the idea of evolution as descent with modification, relationships between (groups of) species are recognized by tracing their possession of shared derived character states (synapomoiphies). The cladistic method primarily selected for the present study is CAFCA. It implies the deduc- tion of transformation series from a chosen cladogram by application of outgroup comparison. In addition two other cladistic programs were applied, viz. PAUP and HENNIG86. Observations regarding the development of gametophytes are reported. During her three in the the author able years stay Philippines, present was to study complete life of the of the concerned. cycles greaterpart Philippine species Special emphasis is given to growth from germination until maturity as well as to the location of indu- antheridiaand Results ment, archegonia. are presented and compared with informa- tions given in the literature. As to juvenile sporophytes, special attention is paid to the venation and to the indument.The presence, the period of development, the shape, the location and the persistency of acicular and glandular hairs as well as of laminar scales are the main observations reported on. Indumentof juvenile and adult sporophytes is compared and differencesare stated. A comparison between laminar and rhizome scales shows a numberof morphological similarities. Known data from literatureabout chromosome numbers are presented in a sum- marized form. Geographical and ecological particularities were investigated. Areas of endemism are listedwith theirrespective species. Collecting localities are indicatedin detailed All in relation with defined climatic rainfall maps. species are set and patterns result- 280 BLUMEA VOL. 34, No. 2, 1990 considerable cladistic Altitudinal ing in support for groupings suggested by analyses. in for each Results of ranges are presented separate diagrams species. transplant experiments which were undertaken in the Philippines point out that morphological characters may show a great plasticity. These experiments also added important de- tails to the author's species delineation. The phylogenetic relationships between the species are correlated with the distri- The close between bution areas. biogeographical analysis suggests a relationship New Guinea, Celebes and the Philippine Islands. The Malay Peninsula and Sumatra related each other well both these MainlandAsia. appear closely to as as areas to As a result of the morphological, cladistic, biological and geographical interpreta- tion the Asian genus Goniophlebium, comprising 23 species, is recognized. Phylo- genetic relationships within the inferredmonophyletic genus are discussed. Cladistic resulted in the of of nine This analyses recognition a monophyletic group species. includes the of the J. group (G. percussum- group) type genusSchellolepis Smith, S. percussa (Cavanilles) Pichi Sermolli. Eight other species are accommodatedin the G. subauriculatum-group, which includes the type of the genus Goniophlebium, G. subauriculatum (Blume) Presl. The remaining taxa buildpairs of species of which the relationship to the G. percussum-group and G. subauriculatum-group could not be further resolved. Therefore, the present author has refrained from assigning a for- mal taxonomic rank to the G. percussum - group and the G. subauriculatum-group, although they are here proposed for the first time. It is expected that an even better phylogenetic resolution including the six remaining species will be obtained from further studies with modem experimental approaches. It is noteworthy that species which have always been regarded as closely related, e.g. G. percussum and G. sub- auriculatum, are now assigned to different groups. Uppermost frond segments and characters for the delimitationof the well defined G. spores provide important per- the new all cussum-group.. According to generic description pinnatifid species, ex- cluded by Copeland (1947) and Holttum (1968), are now accommodated in the genus Goniophlebium. Therefore, four species are newly combined. The other eleven species studied are retained in Polypodium. A scenario for Goniophlebium is given. The incorporation of the biogeographical of characters in and leads the history (evolution morphological space time) to postu- lation of a scenario on the two major monophyletic groups withinthe genus Gonio- phlebium. Copeland's (1915) perception of Polypodium, which was taken up by De la Sota (1973) and Ching (1978), is largely confirmed and the goniophlebioids from the Neotropics, North America and Eurasia are regarded to have evolved sepa- rately from those in the Asian (sub)tropics. of In the taxonomic part an emended description the genus is given. The typifi- cation of the and had receive much attention. The genus appeared most complex to argumentation for the lectotypification of Goniophlebium is published separately. The species are described in detail and identificationkeys to all taxa are provided. taxonomic contains also the The part further full synonymy for specific and infra- specific taxa, information on the distribution and the habitat and last but not least notes on field and other relevant observations. Indices to taxonomic names as well as of the specimens studied are included. 281 G. Rodl-Linder: Monograph of the fern genus Goniophlebium GENERAL PART 2. MATERIAL AND METHODS 2.1. MATERIAL Dried specimens from Asia of approximately 1500 collection numbers, partly from the herbaria abbreviations with many duplicates, were studied following (the follow the Index Herbariorum): B
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