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MOSS FLORA of TAIWAN ( Exclusive of Pleurocarpi )

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MOSS FLORA of TAIWAN ( Exclusive of Pleurocarpi ) MOSS FLORA OF TAIWAN ( Exclusive of Pleurocarpi ) by CHING CHANG CHUANG B. Sc., Taiwan Normal University, 1958 M. Sc., National Taiwan University, 1963 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the department of Botany We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA September, 1971 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of /ffivt Jc?***} The University of British Columbia Vancouver 8, Canada Date ytZpxy i ABSTRACT A floristic study is presented of the moss flora of Taiwan; such taxonomic research is the foundation for botanical science. The knowledge of the bryophytes of Taiwan has represented a serious gap in the knowledge of eastern Asia. In the present treatment are included 276 species including varieties and subspecies belonging to eighty-nine genera and twenty-three families. The included mosses belong to the subclasses Sphagnidae, Andreaeidae, Tetraphidae, Polytrichidae, Buxbaumiidae and the majority of the acrocarpous Bryidae. The pleurocarpous Bryidae have been excluded. Extensive field work throughout the island of Formosa was made during the two summers of 19&7 an^ 1968. Approximately 5»000 specimens were collected and studied. All species considered in this flora that were derived from records from the literature have been checked and reviewed. Type specimens and voucher specimens from the various herbaria plus my own extensive collections have been carefully examined microscopically both morphologically and anatomically. During the course of this study, one new species: Andreaea hohuanensis has been discovered and described. Two new combinations: Oligotrichum suzukii and Mastopoma undulata have been made. Four genera: Wilsoniella, Epipterygium, Leptobryum and Mielichhoferia and twenty-eight species are noted as taxa new to the moss flora of Taiwan. Twenty species and four varieties plus two forms are reduced to synonymy, based on superfluous names or misidentificat• ions. Keys are presented essentially for determination of mosses and are provided for all families in each subclass, and to the genera under each family. Keys are also given to the species and varieties in each genus. Thirty-three species and varieties have been excluded from the moss flora of Taiwan, because they represent nomina nuda, lack precise citation, or are doubtful. Detailed discussion has been made concerning each taxon where particular taxonomic problems arose. Ecological and distributional data are given "for each taxon. The vegetation as well as composition of the flora have been discussed and it is concluded that the bulk of the moss flora of Taiwan considered here is derived from that of the Asian mainland. iii TABLE OF CONTENTS page ABSTRACT . i TABLE OF CONTENTS iii LIST OF MAPS iv LIST OF FIGURES v ACKNOWLEDGEMENTS vi INTRODUCTION 1 GEOGRAPHIC SETTING 5 TOPOGRAPHY 8 GEOLOGY 11 SOIL 14 CLIMATE 19 VEGETATION 20 HISTORY OF BRYOLOGICAL COLLECTION AND STUDY 29 PHYTOGEOGRAPHY 35 CONTRIBUTIONS OF THE PRESENT STUDY TO THE BRYOLOGY OF SOUTHEAST ASIA 42 Outline of the classification 42 Taxa new to the country 43 Revision of nomenclature 46 Keys to taxa 47 Excluded species 48 TAXONOMIC TREATMENT 53 BIBLIOGRAPHY 235 iv LIST OF MAPS Map Page I. Regional location of Taiwan 4 II. Administrative districts of Taiwan 6 III. Physiographic regions of Taiwan 7 IV. Geology of Taiwan 10 V. Soils of Taiwan 13 VI. Average annual rainfall and temperature 18 VII. Map showing distribution of tropical species 38 VIII. Map showing distribution of Asian endemic species. 38 IX. Map showing distribution of temperate Eurasian species 39 X. Map showing distribution of pan boreal species ... 39 XI. Map showing disjunctive distribution 40 XII. Map showing distribution of Himalayan species .... 40 XIII. Map showing localities where the author made collections 41 v LIST OF FIGURES AND PLATE Figure Page 1. Subtropical rain forest 23 2. Subtropical rain forest 23 3. Evergreen broad leafed forest 24 4. Evergreen broad leafed forest 24 5. Mixed forest of broad leafed trees and conifers ... 25 6. Forest of predominant species: Chamaecyparis obtusa var. formosana 25 7. Tsuga forest at Mt. Ta-wu-shan 26 8. Central range of Taiwan high mountains 26 9. Abies kawakamii forest and alpine grassland 27 10. Mt. Sylvia range 27 11. A slope of Mt. Morrison range 28 12. Pure stand of Juniperus squamata var. morrisonicola 28 Plate 1. Andreaea hohuanensis sp. nov 64 vi ACKNOWLEDGEMENTS I am deeply indebted to Dr. W. B. Schofield under whose supervision this study was conducted, his guidance, financial aid, encouragement, and reading of the manuscript throughout the course of this study. I express my sincere appreciation to Dr. Sinske Hattori of the Hattori Botanical Laboratory, Nichinan, Japan, who offered me space to study the herbarium specimens and literature, and to Dr. and Mrs. Zennoske Iwatsuki who gave me considerable assistance during my visit to the Hattori Laboratory in April 1968 and from May to June 1969. I wish to express my grateful thanks to the curators of herbaria for sending me specimens to study, and also to Drs. A. J. Sharp, A. Noguchi, N. Takaki, H. Ando, H. Suzuki, H. Ochi, S. Nakanishi, U. Mizushima, T. Shin, H. Inoue, N. Kitagawa, C. K. Wang, B. Y. Yang, C. G. G. J. van Steenis all of whom kindly send me literatures and specimens. Great thanks to Dr. T. Koponen who send me his recent unpublished paper concerning the Mniaceae of Taiwan. Acknowledgement is made of the financial support from the University of British Columbia in the form of Graduate Fellow• ships during 1967-70 and summer grant for 1971. Support was also given through grants to Dr. W. B. Schofield from the National Research Council, Canada. -1- INTRODUCTION For many years, during my studies of vascular plants in Taiwan, I have noted the astonishing diversity and mysterious beauty of the mosses of the island. The knowledge of the bryophytes remains in a chaotic state in this part of the world. Although some modern branches of botany have been growing rapidly, it is very clear that the study of these modern fields is fundamen• tally based on precise identity of the plant involved. The name of the plant is of basic import to botanical science. Thus, taxonomic research of this flora is essential. The present study makes an effort to solve the difficult critical problems of the moss flora of Taiwan. It is based on my extensive collections as well as the material deposited in various herbaria. In this treatment are included 276 species including varieties and subspecies belonging to eighty-nine genera and twenty-three families. This constitutes all of the known mosses belonging to the subclasses Sphagnidae, Andreaeidae, Tetraphidae, Polytrichidae, Buxbaumiidae and the majority of the acrocarpous Bryidae. The pleurocarpous Bryidae are excluded, although the author has studied these in a preliminary way. Extensive field work throughout the island was made during the summer of 1967 and 1968 (Map XIII. P. 41). Dr. W. B. Schofield also visited Taiwan to assist my field work in May 7 to May 29, 1968. Approximately 5,000 specimens, with complete ecological data, have been collected for this study and are -2- deposited in the herbarium of the University of British Columbia. All species based on records from the literature have been checked and reviewed. Each species has been carefully examined microscopically both morphologically and anatomically verified, evaluated and cited. When this was impossible, this has been indicated. Of special importance was the study and comparison of many type specimens as well as voucher specimens, which are preserved in the following herbaria: H Botanical Museum, University of Helsinki, Finland. HIRO Hiroshima University, Hiroshima, Japan. JE Herbarium Haussknecht, Friedrich-Schiller Universitat, Germany. KU Kobe University, Kobe, Japan. KYO Kyoto University, Kyoto, Japan. MAK Makino Herbarium, Tokyo Metropolitan University, Tokyo, Japan. NICH Hattori Botanical Laboratory, Nichinan, Japan. NOG Kumamoto University, Kumamoto, Japan. NY New York Botanical Garden, New York, USA. TAI National Taiwan University, Taipei, Taiwan. TI Tokyo University, Tokyo, Japan. THU Tunghai University, Taichung, Taiwan. TNS The National Science Museum, Tokyo, Japan. UBC University of British Columbia, Vancouver, Canada. Abbreviations for herbaria are essentially those used by Lanjouw & Stafleu (1964); herbaria not treated by these authors are given the following abbreviations: KU, MAX, NOG, THU and UBC. Keys are provided for all families in each subclass, and to the genera under each family; keys are also given to the species and varieties under each genus to assist in identification of mosses of Taiwan. Critical discussion is given when the material has permitted careful study. The families are arranged according to Brotherus* system in the second edition of Engler and Prantl's Die Naturlichen Pflanzenfamilien (1924) with some modifications based on more recent treatments. The genera and species are dealt with in alphabetical order. Ecological data and distribution are given and phytogeographic notes are included with each taxon. For each taxon the basis for its inclusion is indicated under the designation "Cited specimen". This notes an actual specimen examined, the literature record of those not examined and when possible, the location of the specimen. _4- Map, I. Regional location of Taiwan -5- GEOGRAPHIC SETTING Taiwan is composed of the main island of Formosa plus several small satellite islands or island groups: Pescadores Island, involving over 60 small islands with a total area of only 79 sq.
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