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Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. U-M·I University Microfilms International A Bell &Howell Information Company 300 North Zeeb Road. Ann Arbor. M148106-1346 USA 313i761-4700 800/521-0600 Order Number 9118042 Benthic marine algae of the South China Sea: Floristics, community ecology and biogeography Lewis, Jane Elaine, Ph.D. University of Hawaii, 1900 Copyright @1990 by Lewis, Jane Elaine. All rights reserved. U·M·I 300 N. Zeeb Rd. Ann Arbor, MI 48106 ~I rI>!~ ,"'AT··"',' ,,.,,"S'" "·'t,...';"l""C"'''''''''··'''"''j'',\'' ""'f' 'S'" .... .... e '''''''''"S'''jrI''lr:,'g' ... "'w ,,"'''',e' ' ..::I'e'S',,;- I.'M' =V"" ....... BENTHIC MARINE ALGAE OF THE SOUTH CHINA SEA: FLORISTICS, COMMUNITY ECOLOGY AND BIOGEOGRAPHY A DISSERTATION SUBMIITED TO THE GRADUATE DMSION OF THE UNIVERSITY OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN BOTANICAL SCIENCES (BOTANY) DECEMBER 1990 by Jane E. Lewis Dissertation Committee: Isabella A. Abbott, Chair Kent W. Bridges Douglas J. C. Friend E. Alison Kay Sanford M. Siegel © Copyright by Jane E. Lewis 1990 All Rights Reserved Acknowledgements Thanks first go to Dr. Isabella Abbott and C. K. Tseng for assistance throughout this project, from inception through the specimen identifications which they both oversaw, to the completion. Thanks go to the administration, faculty and students of Zhanjiang Fishery College, Zhanjiang, Guangdong Province, China. Liu Sijian, President of the College, provided logistical support. Prof. Li Weixin, Ding Zenfen, Li Liquan were enthusiastic supporters of the project and were all instrumental in the success of the Liezhou Peninsula field work. Students of Prof. Li assisted great­ ly with tasks ofspecimen pressing and processing. Identifications were conducted with the assistance of Chinese specialists, including Dong Meiling (Chlorophyta), Lu Baoren (Phaeophyta) and Xia Bangmei and Zhang Junfu (Rhodophyta), to whom I am extremely grateful. Prof. Zhu at Nanjing University took on the ex­ amination of those green taxa unidentifiable in Qingdao. Field research was par­ tially funded through the East West Center, Honolulu, Hawaii and the Waimea Botanical Garden, Oahu, Hawaii. Dr. Abbott made her encyclopedic knowledge and extensive library on issues ofAsia and seaweeds openly available to me, and I am truly grateful. Dr. Paul Lamed put the species tables into final desktop publish­ ed form. Dr. Kent Bridges assisted with many aspects of this work, not the least of which was assistance with desktop publishing and computer graphics capabilities. Special thanks are due for his strong support, and especially his willingness to spend parts ofhis sabbatical year assisting with this project. Prof. Dieter Mueller­ Dombois provided many useful comments as applications ofcommunity analysis techniques were being worked out. Kanehiro Kitayama provided valuable discus­ sions on theoretical aspects of the analyses and comparison with terrestrial sys­ tems techniques. I acknowledge the permission to reproduce Appendix B, which originally appeared in the Bulletin ofManne Science, from the Bulletin's editor, W. J. Richards. Dr. Sanford Siegel, Chairman ofthe Botany Department, and a mem­ ber on this dissertation committee, passed away the day before the final copy was completed. His rare wisdom and strong support will be missed. iv Abstract Among the many areas of the Pacific Ocean area where the marine algae are poor­ ly known, the continental coast of the Northern South China Sea has stood out be­ cause no floristic investigations had been conducted in an expanse of over 2,000 kIn. A large collection of algal specimens was made from the Leizhou Peninsula in southern Guangdong Province and in the south of Hainan Island. There were 219 species in 99 genera of Chlorophyta (55 species), Phaeophyta (49 species) and Rhodophyta (115 species). The taxa represent a mix of tropical and subtropical species, and contain a number that previously have not been reported for China. Presence/absence data from this collection was subject to community ecology analysis techniques using COENOS, a computer software program. This provided a useful way ofshowing which taxa can be expected to be found together most of the time, some ofthe time, or almost never. These results allow the selection of certain taxa that can be used as indicator species in order to test relationships with other floras. The floristic data in tum are compared to a database which was compiled ofnearly 20,000 records of over 2,000 taxa from other parts of the warm water western Pacific. Comprehensive tables of species citations of green, brown and red algal species are presented, listed by area in the warm water western Pacific, including the northern South China Sea islands, Taiwan, the Ryukyus islands of Japan, Vietnam, Malaysia, the Philippines and Northern Australia. This study represents the first time that algal distribution information for this area has been gathered together. The use ofselective analyses of a taxonomic databases can predict by biogeographical means, the locations most in need of collections and study, can help establish baseline floristic information for applcation in monitoring, and can assist in the selection of appropriate maricul­ ture sites. v Table of Contents Page Acknowledgements iv Abstract .v List of Tables .viii List of Figures .x Literature Review .1 Introduction .1 Section 1. History of Phycology in the warm water western Pacific .3 Section 2. Biogeography . 12 Section 3. Physical parameters affecting distribution and marine biogeographic provinces ..... 16 Section 4. Vegetation classification . 17 Hypotheses ............ 18 Shallow-water marine algae of the northern South China Sea: Taxonomic Results from a Field Study in South Hainan Island and Leizhou Peninsula, China . 19 Abstract . 19 Introduction . 20 Materials and Methods . 21 Results .. 24 Discussion . 27 Summary. .39 Acknowledgements . 40 vi """'0'-°'.. " ".e'·p"i't"''Y·'?7'W'i'('·P·, e- , ...... "1'0:;,.#..." r""3'·"' i""' ....... "mJCM:t ,?er"riOWhYt'., Community Ecology Analysis ofShallow Water Benthic Marine Algae of the Northern South China Sea . 70 Abstract . 70 Introduction . 70 Materials and Methods . 75 Results .. 77 Discussion . 77 An Overview of the Algal Flora ofthe Northern South China Sea and its Relationship to a Biogeographic Interpretation of the Warm Water Western Pacific 82 Appendix A. A History and Annotated Account of the Benthic Marine Algae ofTaiwan ............................. .. 84 Appendix B. Evaluating Taxonomic Databases for Biogeographic Use 126 Appendix C. Database Summary 136 Bibliography 171 vii List of Tables Page 1.1 Published reports of benthic marine algal taxa in the warm water western Pacific. ............................ .6 2.1 Field collection sites. .42 2.2 Field records (Chlorophyta) for taxa collected from more than one site. ...................... .. 43 2.3 Field records (Phaeophyta) from taxa collected from more than one site. ................................. 45 2.4 Field records (Rhodophyta) from taxa collected from more than one site. ......................... .47 2.5 Field records (Chlorophyta) from only one site. .50 2.6 Field records (Phaeophyta) from only one site. .51 2.7 Field records (Rhodophyta) from only one site. .52 2.8 Summary of Genera by Sites and Area. .... .54 2.9 Taxonomic representation by order and family. .55 2.10 Taxonomic representation compared with taxonomic diversity at the level oforder. ..... 56 2.11 Site ranks for diversity, number of unique species and percentage of unique species. ........... .57 2.12 Unique species compared with total per site. .58 viii 2.13 Summary of Unique Species by Area. .... · . .59 2.14 Genera with the highest taxonomic diversity. · . .60 2.15 Genera most frequently collected. ..... · . .61 2.16 Species with 10 or more accession numbers. · . .62 ix List of Figures Page 1.1 The warm-water western Pacific area (bounded by dashed lines). .2 2.1 The warm-water western Pacific area (bounded by dashed lines). 63 2.2 Field collection sites in the northern South China Sea. 64 2.3 Field site habitats, Leizhou Peninsula. 66 2.4 Field site habitats, South Hainan Island. 67 2.5