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A Monograph of the Fern Genus Including an Attempt Towards A

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A Monograph of the Fern Genus Including an Attempt Towards A A monograph of the fern Genus Microsorum (Polypodiaceae) including an attempt towards a reconstruction of the phylogenetic history of the microsoroids M.T.M. Bosman Rijksherbarium /Hortus Botanicus, Leiden, The Netherlands Contents page 1. Summary 3 2. Introduction and Acknowledgements 3 3. Material and Methods 5 3.1. Material 5 3.2. Methods 5 4. Taxonomic history 7 5. Characters 11 5.1. Rhizome 11 5.2. Scales 14 5.3. Fronds 18 5.4. Stipe 19 5.5. Lamina 21 5.6. Venation 25 5.7. Sori 30 5.8. Paraphyses 32 5.9. Sporangia 33 5.10. Spores 35 5.11. Gametophyte 36 5.12. Chromosomes 37 5.13. Chemistry 37 6. Taxa 40 6.1. Theoreticalconcepts and practical consequences 40 6.2. Selectionof taxa for revision and analysis 41 7. Phylogeny 42 7.1. Methods and algorithms 42 7.2. Data 44 7.3. Cladistic analysis 52 7.4. Conclusions 57 1 (contents continued) 8. Classification 59 8.1. Introduction 59 8.2. Delimitationof Microsorum 59 8.3. Subdivision ofMicrosorum 61 8.4. Classification ofthe microsoroids 62 8.5. Provisional key to the genera 63 9. Distribution and habitat 64 10. Notes on the descriptions and the key 68 11. Genus description 69 12. Key to the species 73 13. Species descriptions 76 14. Doubtful species 129 15. Excluded names 129 16. Identification list 136 17. Glossary of botanical terms 145 18. Index to names of taxa 147 19. Literature 152 2 1. Summary This revision is of the fern Microsorum To a monographic treatment genus (Polypodiaceae). the delimitation and of this elucidate problematic generic systematic position genus an additional selection of 26 possibly related species was also studied. Most of these related species are tradition- ally placed in Colysis, Leptochilus, Neocheiropteris,Paraleptochilus, Phymatosorus (‘Phymatodes’), and Podosorus. In order to produce a natural classification an attempt is made to recognize mono- cladistic The methods and theories which phyletic groups of species through analysis. underlying have been employed are made explicit as much as possible. A suitable selection of intrinsic charac- with the ters is used for analysis computer programs CAFCA, PAUP and HENNIG86. The result- inconsistent with the datamatrix. need ad hoc such ing cladograms are very They many hypotheses, to the of distribution of character states the as homoplasies, explain pattern among species. Even when the number of species to be analysed is artificially reduced by using representative species of inferred monophyletic subgroups ofMicrosorum, the resulting cladograms are to be rejected because of their low consistency indices. The conclusion is reached that the evolution of the microsoroids has resulted in a complex pat- tern, which cannot be understood sufficiently with the present set of data and available methods. Ad- ditional analyses of other characters and species, microsoroids as well as other Polypodiaceae, field observations, cytological and isozyme studies are needed. The results presented here form a good basis for the selection of problematic or promising groups of species. the of the the In absence an acceptable hypothesis concerning genealogicalrelationships among formal classification is constructed. Because ofthis of microsoroids, a temporary temporary nature the classification, stability of nomenclature, and recognizability of the taxa, have in this treatment maximum of information. Thus the classification contain and priority over a phylogenetic may para- polyphyletic groups. This results in the recognition of Microsorum in a restricted sense, comprising 20 species (in- Dendroconche and and of five other microsoroid Para- cluding Diblemma) genera: Colysis (including leptochilus), Leptochilus (emended),Neocheiropteris,Phymatosorus, and Podosorus. Two species of Microsorum are newly described, namely M. cinctum and M. sopuense, and two newly transferred. Nine species, initially thought to belong to Microsorum, are fully described and included in the key, but classified under of the other microsoroid in combina- are finally one genera, resulting seven new tions and onenew species (Phymatosorus biseriatus). Many other species of Microsorum (84 names) are because of of this excluded but, the temporary character classification, the formal new combina- tions, necessary for most of these 84 names, are not formally made. 2. Introduction and Acknowledgements This monograph deals with the revision of Microsorum. Especially the generic boundaries and the systematic position of Microsorum appealed to be problematic. Therefore a number of other microsoroid species were also included in this study. These have been usually placed in: Colysis (C.), Leptochilus (Lc.), Neocheiropteris (N.) Paraleptochilus (Pa.), Phymatosorus (Ph.), and Podosorus (Po.). Additionally one species of Lepisorus (Lp.) has been studied as a possible 'outgroup' (see chapter 6). The abbreviationsemployed above will be used in most instances in this revision. This is the first monographic treatment of Microsorum. A few regional revisions have been carried out before, the most extensive of which is that of Ching (1933b) for the Chinese species. 3 This monograph was prepared as a Ph.D. thesis at the Rijksherbarium/Hortus forms of the Botanicus of the Rijksuniversiteit Leiden and part Polypodiaceae Project supervised by Prof. Dr. E. Hennipman (Rijksuniversiteit Utrecht). A preliminary character ofthe venation and related study of this group, a analysis pattern structures, carried in the E. was out same project by W.L.A. Hetterscheid and Hennipman (1984). The aim of this study is twofold: 1. To produce a natural classification, which is as informative, practical and stable as possible. As will be explained, this entails a classification including only mono- phyletic groups which are assigned formal ranks on the basis ofpractical consid- erations. At present the phylogenetic method seems the best way to achieve this aim. 2. To produce a revision in which the methods and underlying theories are made ex- plicit directly, orindirectly by reference to other authors. This may seem obvious but, unfortunately, especially in 'classical' systematics it is still a rare phenomenon. A vast amount of herbarium material was used for detailed species descriptions and character analyses, which together form the necessary basis for a phylogenetic analysis. Additional information fromcultivated living material was used inciden- tally. first natural has been The aim, to produce a classification, not fully attained, as the complex genealogical relationships of the microsoroid fems cannot be unravelled with the data and methods used in this study. However, the results of the present investigation form a firmbasis for a well-founded choice of species and methods for further research (as indicated in chapter 8). This is partly reflected by the accepted classification, which is probably not completely natural and which is therefore tem- and formalized. porary only partly Whether I succeeded in fulfilling the second aim is to be judged by unbiased readers, especially those who are not familiar with the subjects discussed. This revision is first with my encounter pteridological research. It took some time accustomed the idea fems to get to that do not bear flowers, nor produce fruits, but that they show many other characters which are biologically informative, although observed. In fems be beautiful they are not always easily some respects may even as the this needs as most elegant flowering plants, although some persuasion when working for several years on the very plain-looking Microsorums. From with fems well with my systematic experience as as higher plants I have gained the impression that the evolutionary biology of ferns is generally more com- plex than that of flowering plants and consequently their systematic study is often more problematic, interesting and time-consuming. I have learnt at least one thing: never think too lightly of innocent and plain-look- hide and from which ing phenomena, they may swamps, jungles paradises it is diffi- cult to escape without sweat, scratches and sins. 4 Acknowledgements The assistance of keepers and directors of the herbaria listed in chapter 3 who have been so kind considerable time, is as to put their collections at my disposal, often for a gratefully acknowledged. For their I also wish to to the staff members of the help during my stay, express my gratitude visit to the herbaria I visited (B, BM, K, P). They made me feel welcome and comfortable. My from the Netherlands for herbarium in Paris was made possible by financial support Organization the Advancement ofPure Research (Z. W.O.). I wish to mention Wilbert Hetterscheid for his kind permission to use his photographs of vena- Gerda Uffelen the ofthe of M. and tion patterns. van provided photograph spore linguiforme gave and of surface Gerda Bosman-van helpful suggestions on interpretation terminology spore sculpture. den Heuvel kindly corrected the English text of the general chapters in the first version of the manu- in the script and Monique Smits, apprentice technician, helped scanning spores. The diagnoses of newly described species were translated into Latin by Peter Hovenkamp. The habitus drawings were made by Joop Wessendorp; the other drawings were made from my own Peter Welzen sketches by Jan van Os. The lay-out and setting were done by Emma van Nieuwkoop. van assisted me whenever problems with computers and computerprograms occurred. 3. MATERIAL AND METHODS 3.1. Material A fairly large selection of the dried materialofMicrosorum and some alliedgenera in the herbaria
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